CROP ECOLOGY,

CULTIVATION
AND USES OF
CACTUS PEAR
Advance draft prepared for the
IX INTERNATIONAL CONGRESS ON CACTUS PEAR AND COCHINEAL
CAM crops for a hotter and drier world
Coquimbo, Chile, 26-30 March 2017
CROP ECOLOGY,
CULTIVATION
AND USES OF
CACTUS PEAR
Editorial team
Prof. Paolo Inglese, Università degli Studi di Palermo, Italy;
General Coordinator Of the Cactusnet
Dr. Candelario Mondragon, INIFAP, Mexico
Dr. Ali Nefzaoui, ICARDA, Tunisia
Prof. Carmen Sáenz, Universidad de Chile, Chile

Coordination team
Makiko Taguchi, FAO
Harinder Makkar, FAO
Mounir Louhaichi, ICARDA

Editorial support
Ruth Duffy

Book design and layout

Davide Moretti, Art&Design − Rome

Published by
the Food and Agriculture Organization of the United Nations
and
the International Center for Agricultural Research in the Dry Areas
Rome, 2017
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© FAO, 2017

Chapter photographs:
© Fotolia, iStock photo
CONTENTS

Foreword IX
Acknowledgements X
Acronyms XI

A History and economic and agro−ecological importance 1

María Judith Ochoa and Giuseppe Barbera.

Introduction 2
History 2
Current situation 4
Americas 5
Africa 7
West Asia 9
Australia 10
Europe 10
Perspectives 11

2 Origin and taxonomy of Opuntia ficus−indica 13

Roberto Kiesling and Detlev Metzing

Introduction 14
Origin and taxonomy of Opuntia ficus−indica 14
Chromosome numbers 15
Molecular studies 15
Distribution and naturalization 16
Vernacular names 17
The role of cochineal 17
Taxonomical and nomenclatural conspectus 18
Conclusions 19
IV

3 Morphology and anatomy of Platyopuntiae 21

Loreto Prat, Nicolás Franck and Fusa Sudzuki
CONTENTS
Introduction 22
Root system 22
Cladode 23
Bark 26
Wood 26
Flower buds 26
Pollen grain 28
Fruit 28

4 Ecophysiology and reproductive biology of cultivated cacti 29

Paolo Inglese,Giorgia Liguori and Erick de la Barrera

Introduction 30
Cam Cycle 30
Soil suitability in the cultivation of cactus pear 32
CO2 uptake and water availability 33
Response to temperature 34
Response to light 35
Reproductive biology 36
Biomass productivity 38
Potential responses to climate change 39

5 Nopsal (Opuntia spp.) genetic resources 43

Paolo Inglese,Giorgia Liguori and Erick de la Barrera

Introduction 44
Genetic resources exploration 44
Germplasm banks, core collections and cultivar trials 46
Genetic resources documentation 46
Using modern molecular tools to explain cactus pear variability 47
Conservation of cactus pear genetic resources 48
Utilization: germplasm enhancement and breeding 48
Towards sustainable utilization of cactus pear 49
 V

6 Fruit production and post−harvest management 51

Johan Potgieter and Salvatore D’Aquino
CONTENTS
Introduction 52
Orchard planning and establishment 52
Orchard layout and design 55
Establishment 56
Orchard management 57
Harvesting 65
Ready−to−eat cactus pear 68
Conclusions and future perspectives 69

7 Forage production and supply for animal nutrition 73

Jose C.B. Dubeux Jr., Hichem Ben Salem and Ali Nefzaoui

Introduction 74
Cactus−based forage production systems 74
Agronomic practices 79
Forage quality 82
Utilization 86
Animal performance and product quality 89
Future perspectives and research needs 90

8 Nopalitos or vegetable cactus production and utilization 93

Candelario Mondragón Jacobo and Santiago de Jesus Méndez Gallegos

Introduction 94
Early utilization and domestication 94
Vegetable nopalito varieties 95
Agricultural importance of the nopalito crop 96
Nopalito production systems 97
Production practice 100
Basic preparation techniques 102

9 Cochineal breeding 105

Liberato Portillo and Ana Lilia Vigueras

Introduction 106
Biology 107
Cochineal breeding 108
Ecology 111
 J Diseases of cactus pear 115
Giovanni Granata, Roberto Faedda and María Judith Ochoa
CONTENTS
Introduction 116
Fungal diseases 116
Bacterial diseases 121
Phytoplasma and virus diseases 122
Abiotic disorders 122

K Insect pests of cactus pear 125

Jaime Mena Covarrubias

Introduction 126
Insects 126
Integrated pest management for cactus insect pests 132

L Processing and utilization of fruit cladodes and seeds 135

Carmen Sáenz

Introduction 136
Chemical composition and bioactive compounds 136
Technological characteristics 137
Processing technologies 138
Other technologies 144
Other products 146
Conclusion 149

M Nutritional properties and medicinal derivative of fruits and cladodes 151

Mónica Azucena Nazareno

Introduction 152
Nutritional aspects 152
Bioactive phytochemicals in cactus plant 154
Medicinal properties 155
Perspectives 158
 VII

N Cactus ecosystem goods and services 159

Mounir Louhaichi, Ali Nefzaoui and Juan Carlos Guevara
CONTENTS
Introduction 160
Rangeland improvement 161
Alleviation of soil erosion 162
Biological fencing/vegetative barriers 164
Carbon sequestration potential 165
Alley cropping 166
Biodiversity conservation 167
Conclusion and recommendations 169

O Global invasions of cacti (Opuntia sp.): 171

Control, management and conflicts of interest
Helmuth Zimmermann
Introduction 172
Cacti as a problem 172
Utilization and conflicts of interest 183
Conclusions 184

P Biogas production 187

Maria Teresa Varnero and Ian Homer

Introduction 188
Using cactus waste in biogas production 188
Opuntia spp. plantations for biogas production 189
Biodigesters design and operation 191
Economic aspects 192
Other bioenergy uses 193

Q Marketing and communication constraints and strategies 195

Marcos Mora

Introduction 196
Production and marketing background 197
Perceived quality in cactus pears: from intrinsic to extrinsic attributes 200
Value attributes and consumer preferences 200
Proposed commercial development strategy for fresh cactus pear and its products 201
Conclusions 201

Bibliography 203
 IX

Foreword

C
limate change is one of the biggest challenges the world must meet today and in the future.
Prolonged droughts and desertification are among the issues faced by many countries,
especially in Africa and Asia, where the rural poor and smallholders are most heavily affected.
If people are to survive in these ever harsher conditions, their crops need to withstand
drought, high temperatures and poor soils.

Cactus crops are gaining increasing interest across the globe, in particular cactus pear (Opuntia ficus−
indica), because of its unique characteristics which provide resilience to the above mentioned harsh
conditions. Cactus pear is able to grow on land where no other crops are able to grow; it can be used
to restore degraded land and in many countries, such as Ethiopia, it is the only crop that can be relied
on when everything else fails. The crop originates in Mexico – still the largest producing and consuming
country in the world – but other countries, including Morocco, Ethiopia, South Africa, Kenya, India and
Pakistan, are increasing their production and use.

In addition to its resilience as a crop, cactus pear is also increasingly appreciated for its multiple purposes.
The fruit and young cladodes can be consumed by humans, and there is growing interest in its use as
fodder. In Brazil, a plantation of over 400 000 ha in the northeast region serves as a key component in
supporting livestock production in the country’s semi−arid regions. The cultivation of cactus pear for
use as fodder is also being adopted in sub−Saharan Africa and South Asia. Furthermore, its medicinal
properties and industrial uses are being researched and promoted.

The FAO–ICARDA International Technical Cooperation Network on Cactus (CactusNet) was established
in 1993 to support the promotion of this underutilized crop. The first edition of this book, Agro−ecology,
cultivation and uses of cactus pear, was published in 1995. During the last 20 years much knowledge on
cactus pear has been generated, and this is reflected in this extensively revised edition in 2017.

The dedication of all the experts who have contributed to this edition is greatly appreciated and it
is hoped that the book will become a useful resource in those countries interested in developing or
increasing cactus production. Further, we acknowledge the excellent work of CactusNet, and encourage
others to join and contribute to expanding the outputs generated by the Network and its partners.

Hans Dreyer Andrew Noble

Director Deputy Director General – Research
Plant Production and Protection Division International Center for Agricultural Research
Food and Agriculture Organization in the Dry Areas (ICARDA)
of the United Nations (FAO)
X

Acknowledgements

I
t was August 1993 when the International Cactus Pear Network (CactusNet) was established
following initial discussions in Santiago, in September 1992, during the International Prickly Pear
Congress organized by Fusa Sudzuki and Carmen Sáenz at the University of Chile. Eulogio Pi-
mienta, one of the editors of the first edition of this book, was also the first General Coordinator
of the Network. In 1995, the International Society for Horticultural Science (ISHS) Working Group on
Cactus Pear and Cochineal was established and has made a tremendous contribution to the develop-
ment of the knowledge of this crop and its utilization. Papers presented in the Acta Horticulturae of
the Working Group symposia and in refereed journals have had a valuable impact on the advancement
of cactus pear. Now, 25 years later, we are back in Chile, proud to present the second edition of Crop
Ecology, Cultivation and Uses of Cactus Pear. The first edition was a miracle, considered a “milestone
in the literature of cactus pear”. We were at the earliest stage of the Web. No emails available! Only
floppy disks and airmail! Twenty−six authors from 18 universities and research institutions in Chile,
Germany, Israel, Italy, Mexico, Peru, South Africa and the United States of America contributed. Sim-
ilar to the first edition, the second is the result of international cooperation. Some of the authors of
the first edition are still involved: Giuseppe Barbera, Carmen Sáenz, Candelario Mondragón, Giovanni
Granata, Maria T. Varnero, Helmuth Zimmermann. We thank all of them for the long run we have
made together. But we also thank all the authors for their devotion to this crop and their enthusiastic
contribution to this book. In total, we have brought together 30 authors from 10 countries, surpassing
the first edition.

During the last 25 years we have had the dedicated support of the Food and Agriculture Organization
of the United Nations (FAO) and the International Center for Agricultural Research in the Dry Areas
(ICARDA), and we have grown considerably. Enrique Arias, from 1992 until he retired from FAO in
2012, was for CactusNet like a lighthouse in the storm. He guided us and served science and technical
development for the rural poor with devotion and daily commitment. Umberto Menini in the early
years, and later Alison Hodder, both of the Plant Production and Protection Division, FAO, also leant
their full support. Today, we have the good fortune to share our work with Makiko Taguchi who has
led this edition of the book. We are deeply indebted to her for her “firm” kindness and determina-
tion. ICARDA became a partner of our Network when Ali Nefzaoui became the General Coordinator;
the FAO–ICARDA partnership created the conditions for further development of CactusNet activities.
This book was made possible by support from FAO and ICARDA, with timely contributions from the
German Agency for International Cooperation (GIZ).

We thank all the members of CactusNet. We are proud to belong to this community and to share
science and friendship within it. Indeed, this book is dedicated to Enza Chessa, a member of our
community who passionately championed science along with friendship and loyalty. She was able to
smile until the end, and we smile back to her in our soul.

Let us finish with the same words used for the first edition which we firmly believe are still valid:
We sincerely hope that this book will play an important role in increasing the knowledge and usage
of cactus pear, given its potential in the agriculture and economy of the world’s arid and semi−arid
zones.

The Editors
 XI

Acronyms

ADF Acid detergent fibre

ADG Average daily gain
ADL Acid detergent lignin
AFLP Amplified fragment length polymorphism
AGR Absolute growth rate
ANADEC National Association of Cactus Development (Morocco)
ARS Agricultural Research Service (United States of America)
aw water activity
BCE Before the Common Era [BC]
CactusNet International Cactus Pear Network (FAO)
CAI Cladode area index
CAM Crassulacean acid metabolism
CBTA Agricultural Technology School (Mexico)
CECCAM Center for Studies for Change in the Mexican Countryside
CENARGEN National Research Center for Genetic Resources and Biotechnology
CEZA Agricultural Centre for Arid Zones (Chile)
CFU Colony−forming units
CIF Forage Research Center (Bolivia)
CIREN Natural Resources Centre (Chile)
CIRNO Research Center for the Northwest Region (Mexico)
CMC Carboxymethyl cellulose
CNERNA Coordination of Studies and Research on Nutrition and Food (France)
CNR National Research Council (Italy)
CNRS National Center for Scientific Research (France)
CONACYT National Council of Science and Technology (Mexico)
CONICET National Scientific and Technical Research Council (Argentina)
CP Crude protein
cp−SSR Chloroplast simple sequence repeat
CREZAS Regional Center for Arid and Semi−arid Zones (Mexico)
CRUCEN Regional Center North Center University (Mexico)
DB Dry basis
DE Dextrose equivalent
DESA Department of Economic and Social Affairs (United Nations)
DM Dry matter
DNA Desoxyribonucleic acid
DW Dry weight
EC Electrical conductivity
EDW Estimated dry weight
EPDM Ethylene propylene diene terpolymer
EST Expressed sequence tag
FAUANL Faculty of Agronomy of the Autonomous University of Nuevo León (Mexico)
FDP Fruit development period
FI First flush (flowers)
FIA Foundation for Agrarian Innovation (Chile)
XII

FII Second flush (flowers)

FM Fodder machine−sliced (cladodes)
FOB Free on board
FW Fresh weight
GA Gibberellic acid
GAE Gallic acid equivalent
GDH Growing degree hours
GIZ German Agency for International Cooperation
GRIN Germplasm Resources Information Network
HDPE High density polyethylene
HI Harvest index
HTST High−temperature short−time
IADIZA Argentine Dryland Research Institute
IBA Indole−butyric acid
ICARDA International Center for Agricultural Research in the Dry Areas
IFAD International Fund for Agricultural Development
IIZD Institute for Desert Area Research (Mexico)
IMZ Imazalil
INCA National Institute of Agricultural Sciences (Mexico)
INEGI National Institute of Statistics and Geography (Mexico)
INIFAP National Institute for Forestry, Agricultural and Animal Husbandry Research (Mexico)
INRAT National Institute of Agricultural Research of Tunisia
IPCC Intergovernmental Panel on Climate Change
IPM Integrated pest management
IQF Individually quick frozen
ISHS International Society for Horticultural Science
ISPA Institute of Sciences of Food Production (Italy)
ISSR Inter−simple sequence repeat
ITS Internal transcribed spacer
Kc Crop factor
KC Knife−chopped (cladodes)
LAI Leaf area index
LPG Liquefied petroleum gas
ME Metabolizable energy
MF Microfiltration
NAFTA North American Free Trade Agreement
NALPGRU National Arid Land Plant Genetic Resources Unit (United States of America)
NCRE Non−conventional renewable energy
NDF Neutral detergent fibre
NEMA National Environment Management Authority
NF Nanofiltration
NGO Non−governmental organization
nrITS Nuclear ribosomal internal transcribed spacer
NTU Nephelometric Turbidity Unit
OD Osmotic distillation
ODEPA Office of Agricultural Studies and Policies (Ministry of Agriculture, Chile)
OEP Office of Livestock and Pasture (Tunisia)
OM Organic matter
 XIII

OP Oviposition period
PAR Photosynthetically active radiation
PCR Polymerase chain reaction
PDO Protected designation of origin
PEP Phosphoenolpyruvate
Pn Net photosynthesis
PPFD Photosynthetic photon flux density
PPO Polyphenol oxidase
PROAGRO Sustainable Agricultural Development Program (Bolivia)
PUFA Polyunsaturated fatty acids
PVA Polyvinyl alcohol
PVC Polyvinyl chloride
RAPD Random amplified polymorphic DNA
RCPs Representative Concentration Pathways
RH Relative humidity
RNA Ribonucleic acid
RO Reverse osmosis
ROS Reactive oxygen species
RUE Rain−use efficiency
SAGARPA Secretariat of Agriculture, Livestock, Rural Development, Fisheries and Food (Mexico)
SAI Stem area index
SC Separate concentrate (feed)
SFA Saturated fatty acids
SFR Spring flush removal
SIAP Agricultural and Fishery Information System (Mexico)
SINAREFI National System of Plant Genetic Resources for Food and Agriculture (Mexico)
SLP San Luis Potosí
SNICS National Service Seed Inspection and Certification (Mexico)
SOMECH Mexican Society of Horticultural Sciences
SSR Simple sequence repeats
TBZ Thiabendazole
TDM Total dry matter
TMR Total mixed ration (feed)
TN Total nitrogen
TSS Total soluble solids
UF Ultrafiltration
UFRPE Federal Rural University of Pernambuco (Brazil)
UNAM National Autonomous University of Mexico
UNDP United Nations Development Programme
UNSO Office to Combat Desertification and Drought
USDA United States Department of Agriculture
UV Ultraviolet
VFA Volatile fatty acids
WBT Wide−band tracheid
WUE Water−use efficiency
ww Weight of water
 01

History and economic

and agro-ecological
importance
María Judith Ochoaa and Giuseppe Barberab
a
National University of Santiago del Estero, Argentina
b
University of Palermo, Italy
 01 History and economic
and agro-ecological importance

INTRODUCTION HISTORY

“The most important thing since the creation of When the Spaniards arrived in 1492 on Hispaniola
the world ... is the discovery of the Indies.” These Island (now Haiti and the Dominican Republic) in
words, addressed in 1552 to the Emperor Charles the Caribbean Sea, the natives introduced them
V by Francisco Lopez de Gomara, author of the to Opuntia red fruits, then called “tuna”, from
the Caribbean word tun. An amusing anecdote
famous Historia General de las Indias (General
recorded by the chronicler refers to how the new-
History of the Indies, 1552), testify to the Eu-
comers, upon observing the colour of their urine,
ropean conquerors’ awareness of the possible
thought they had been poisoned and were bleed-
outcomes of the encounter between the Old and
ing to death (Kiesling, 1999a). The first Europeans
the New World. According to Crosby (l972), the
to land on the American continent appreciated
consequences of the “Columbian exchange” were the important role – both cultural and economic
particularly significant for – of opuntias in the complex pre−Hispanic world.
the flora and fauna of When the first conquistador, H. Cortés, reached
the various regions of the the Mexican Plateau in 1519, they could not fail
planet and especially for to notice the presence everywhere of nopalli (the
the agriculture. Through- Nahuatl word for the plant), and on entering Tlax-
out the centuries, this ex- cala (Diaz del CastiIlo, 1991), they were welcomed
change has continued and with cactus pear fruits (nochtli). Cactus pear was
does not cease to amaze part of the Aztec culture: the Aztec army flag
us. During the decades shows an eagle over a cactus pear eating a snake;
immediately following Co- and the empire capital was called Tenochtitlan, i.e.
lumbus’ first trip, the rate “Cactus pear over a rock” (now Mexico City). This
of exchange of animals and same scene is depicted on the first page of Codex
plants between the two Mendoza, representing the centre of the universe
Worlds varied. The con- (Berdan and Anwart, 1992) (Figure 1).
quistadors quickly imposed
The conquistadors began to eat the fruits. Ovie-
their traditional crops in do and Valdés – the first author to describe the
America: they had the fruit and the plant – wrote that his fellows “knew
scientific and technological and ate that fruit with pleasure” in Hispaniola in
edge and wished to recre- 1515. He also recorded an accurate description (in
Figure 1
Representation of ate the conditions and habits of their homeland. particular in a later work, written in 1535) of the
Tenochtitlan’s emblem Transfer in the opposite direction took longer, since morphology of the plant. According to this de-
the Codex Mendoza, the Europeans were mainly interested in products scription, the plant grew not only on Hispaniola,
the cactus pear as
capable of bringing immediate profit. When they but also on other islands and in other areas of the
centre of the universe
did eventually become interested in the local flora, Indies. Moreover, “its seeds and skin are like those
it was through scientific curiosity or a taste for the of the fig and are tasty too”.
exotic; they were not aware of the potential of
Other authors mention the success of the fruit
those plants as food or as goods of economic rele- with both the local populations and the Spaniards
vance. On the contrary, the Spaniards were initially (Donkin, 1977). In particular, Oviedo y Valdés, To-
suspicious of the culture of the peoples they had ribio de Motolinia and Galeotto Cei (1539–1553)
conquered, and they approached their food hab- all described various species, often highlighting
its with much caution and out of mere necessity the various uses in addition to food: as herbs (the
(Doria, 1992). For these reasons, it was centuries “welding tree” variety was used to treat frac-
before opuntias – one of the most relevant eco- tures), as a water source and as barriers or fences.
nomic plants of the Aztec civilization – were fully The presence of different species is underlined
appreciated in the various regions of the world. by Toribio de Motolinia, who mentions “many
History and economic and agro−ecological importance 3

species”, and by Bernardino de Sahagun and Fran- Neither his words nor his paintings, however,
cisco Hermindez (around 1570), who cite 13 and are adequate to provide an accurate description:
9 species, respectively. Despite this interest, there “Maybe Berreguete or other great painters such
was no real economic assessment of the fruits. as Leonardo da Vinci who I met in Milan, or An-
drea Mantegna would better draw it...”
On the other hand, the highly prized red dye, no-
cheztli, was greatly appreciated by the Spaniards, Their introduction to Europe is not documented;
who called it grana cochinilla. The dye is rich in however it did take place before 1552, when Lopez
carminic acid and is extracted from the body of de Gomara wrote about the “nopal”, taking for
the Dactylopius coccus cochineal that lives in the granted that it was already well known in Spain
cladodes (Reyes Agüero et al., 2005). The red (Donkin, 1977). The plant was O. ficus−indica or
dye was highly rated for its commercial qualities, − as maintained by Berger (1912a) − O. amyclaea.
and was probably shipped in small quantities to The first opuntias probably grew in the vicinity of
Europe early on. It is certain that Spain was re- Seville or Cadiz, the terminus for traffic with the
questing it by 1521 and that in 1544, according to Indies (Donkin, 1977); they then spread to the lush
Cervantes de Salazar, “a great quantity at a high estates of aristocratic mansions and to botanic
cost is imported by Spain” (Donkin, 1977). gardens. It is reported to have been present in Italy
around 1560, in Germany and the Netherlands in
The new product was successful because of its
1583, and in England in 1596 (Donkin, 1977).
intense red colour. It was the most desired import
commodity from America at the end of the six- It was immediately well received as a curiosity and
teenth century, surpassed only by precious metals. decoration. Pier Andrea Mattioli − who in 1558
The precise nature of the powder (grana) was for was the first to
a long time the subject of debate, until Hartsoeker provide a draw-
(1694), Van Leenwenhoek (1704) and De Ruuscher ing based on
(1729) specified its animal origin (Donkin, 1977). plants grown in
Europe − wrote
However, despite the economic importance of
that “this plant
grana, the host plant was not introduced into Eu-
rightly deserves
rope. There was a ban on exporting the infested
to be included
cladodes and the Spanish maintained a monopoly
among the mir-
for two centuries, until a French doctor, Thiery
acles of nature”.
de Menonville, brought the insect to the French
The 1568 edition
dominions (Haiti) in 1777. The insect was reported
of his Discorsi
in India in 1795 and in Europe at the beginning of
included two ta-
the nineteenth century, where attempts to breed
bles testifying to
it failed in France, Spain and Italy. In 1853, howev-
the general curi-
er, there were 14 breeding plants in Algeria, and
osity surround-
in 1850–1860 the Canary Islands were already
ing the species.
exporting twice the quantity coming from Ameri-
The fruits were
ca (Donkin, 1977). Hence, since the plant was not
probably not
introduced for economic reasons, it may be con- Figure 2
appreciated − they could barely ripen in northern
cluded that the Europeans were mainly attracted Drawings of columnar
Italy − and Mattioli reported the words of Oviedo cacti and two species of
by its shape, which was completely new to them.
y Valdés citing one of its supposedly terrible ef- cactus pear by Oviedo
Indeed, European travellers went in search of fects: on eating them, the urine turns red. Valdés, 1535
unusual plants (Europe was in the period of the
The Spaniards took the cactus pear − and most
“Cabinet of Curiosities” and Wunderkammem)
likely also Dactylopius coccus − to Peru, although
and cactus pears fitted the bill. The first depic-
other cochineal were used for dying before the
tions of cactus pear and other columnar cacti,
Hispanic period (De Acosta, 1590). In Argentina,
are found in the Historia Natural y General de las
red fabric dye, clothes and “ponchos” (De Acos-
Indias (Natural and General History of the Indies),
ta, 1590) were made using native cochineals.
written by the Spanish officer Gonzalo Fernando
The medical use of cacti and its derivates was
Oviedo y Valdés in 1536 (Figure 2) (written in-
documented in Medicina en el Paraguay natural
formation had been recorded earlier on Colon´s
(Medicine in Natural Paraguay), 1771−1776, writ-
second Caribbean trip). Oviedo y Valdés wrote: “I
ten by José Sánchez Labrador; he described the
cannot make up my mind whether it is a tree or
use of cochinilla gusano de la grana (cochineal red
one of the most frightful monsters among trees.”
worm) for fevers and insanity, how the fruit was
 4 Crop ecology, cultivation and uses of cactus pear

used as refreshment and seeds as an astringent placed in greenhouses. In Mediterranean regions,

(Ruiz Moreno, 1948). on the other hand, the climate was ideal and
opuntias soon naturalized, becoming one of the
A John Arrowsmith publication (London, 1842)
most common and representative elements of the
about the topography of Plata and its provinces
landscape. The plant spread along the Mediterra-
indicates the Santiago del Estero province in
nean coast: birds ate its fruits, the Moors returned
northern Argentina as a great grana producer
to North Africa from Spain, and it was also shipped
(Figure 3).
to combat scurvy (caused by lack of vitamin C),
In 1580, Gian Vettori o Soderini recalled the fruit’s as cladodes easily tolerate long journeys without
Mexican origin and how it roused the same won- losing their ability to take root (Kiesling, 1999a).
der as peacocks. Towards the end of the century, During the eighteenth century, opuntias spread
Agostin del Riccio named cactus pear as a plant to South Africa (1772), India (1780), China (1700)
ideal for a king’s garden (Tangiorgi Tomasi e Tosi, and Indochina, although it is also possible that the
1990). Numerous works of art incorporate the plant was introduced in these countries before
cactus pear. A wonderful watercolour painting of that time (Donkin, 1977). The iconic use of the
the Iconographia Plantarum by Ulisse Aldrovrandi plant became very popular during the nineteenth
also dates from the end of the sixteenth century. century in arts, cartoons, marketing and even in
In 1600, the Bishop of Eichstatt commissioned politics (Barbera and Inglese, 2001). Numerous
the Hortus Eystehensis to comprise all the plants artists, including Diego Rivera, Frida Kahlo, Roy
of the palace garden, and it includes a drawing of Lichtenstein, Felice Casorati and Renato Guttuso,
the species. Bruegel the Elder’s “Land of Plenty” used it in their works.
(1567, now in the Alte Pinakhotek of Munich) also
includes an Opuntia. In Rome’s magnificent Piazza
Navona, in 1648–1651, Gian Lorenzo Bernini sculp- CURRENT SITUATION
tured the Fours Rivers Fountain, designed by Bor-
romini, depicting cactus pear cladodes beside the
Opuntias are now part of the natural environment
Rio de la Plata. By the seventeenth century, there
and agricultural systems, and Opuntia ficus−indica
is evidence of the presence of Opuntia in Europe: a
(L.) Mill. is the most important economic cactus
xylograph by A. Switzer (1650, now in the National
species worldwide (Kiesling, 1999a). It is cultivat-
Central Library in Florence) shows the cactus pear
ed in America, Africa, Asia, Europe and Oceania
among other plants in the Garden of Eden; and
(Casas and Barbera, 2002). It can be found from
in 1650/1651, J. Bahllin wrote: “it is grown in the
Canada to Patagonia, Argentina, from sea level to
pensile gardens of noblemen’s mansions”.
5 100 m asl in Peru (Bravo Hollis and Scheinvar,
In northern regions, cultivation was indeed only 1995). When first discovered by the European
possible in privileged spaces, since to survive the explorers, opuntias were distributed from Mes-
winter the plants had to be well sheltered or oamerica to Cuba and other Caribbean islands

Figure 3
Geographical atlas of
Santiago del Estero

In the greatest
deserts you can
find an abundance
of cactus
species where
insectcochinillas
live, around
10 000 pounds
of this colour are
exported.
History and economic and agro−ecological importance 5

(Griffith, 2004). It became invasive in areas with a wet highest level of production and technology in intensive
season characterized by high temperatures, for exam- forage production. In this area, O. ficus−indica reaches
ple, in South Africa and Australia (Wessels, 1988a; the highest level of fresh and dry mass production in
Zimmermann et al., 2009). In Mediterranean climates, the world.
natural invasion is limited by the humidity and cold win-
ter temperatures that contrast the warm, dry conditions Chile
of summer (Barbera, 1995).
Cactus pear production zones in Chile cover 934.4 ha,
The traditional and popular uses of cactus pear in a
with no significant changes in the last decade. The
wide range of countries and its multiple functions have
main areas of fruit production are the Metropolitana
caught the imagination of farmers, cattlemen and the
area and the regions of Coquimbo and Valparaíso
scientific community.
(ODEPA, 2016). Opuntia ficus−indica gives two crops
per year without technical intervention: the first in Feb-
ruary–April, the second in July–September. The second
AMERICAS crop produces just one−third the volume of the first,
but it reaches higher prices (Sudzuki Hills et al., 1993;
Argentina Sáenz et al., eds, 2006). Production is destined mainly
for fresh fruit consumption on the national market. As
Argentina has a cactus pear production surface area a result of projects in recent years, there is growing
of 1 650 ha; 80% is in the northeastern region, the re- interest in the use of cladodes as food − marmalades or
mainder scattered in the centre and Cuyo (Targa et al., nopalitos − in the Chilean diet.
2013). The main cultivar is a local selection called ‘Am-
arilla sin espinas’ (yellow without spines) (Ochoa, 2003). Peru
Production goes to the national markets in Córdoba and
Buenos Aires. In the last 5 years, during the dry season, At present, 60% of cactus pear farming is destined for
producers have introduced Opuntia ficus−indica as cochineal production and 40% for the fresh fruit mar-
forage. Research institutions have begun incorporating ket. In recent years, the processing industry has grown
cactus pear in their programmes and research projects. with fruits transformed into nectar juice.

Bolivia The Peruvian cactus pear crop is grown mainly for do-
mestic consumption rather than export. Cochineal red
Opuntia ficus−indica is well known in Bolivia, where is dye is produced on 10 000 ha under intensive (80% of
has multiple uses. Cactus pear production was devel- the area) and semi−wild plantations in Huarochirí and
oped in Cochabamba, Chuquisaca, Sucre, Tarija and La Ayacucho districts. In 2012, the association for cactus
Paz, in areas with 350−640 mm year−1 rainfall and at producers and cochineal in Huarochirí produced more
altitudes of 1 500−3 000 m asl. than 5 000 tonnes of cactus pear fruits for local markets;
‘Blanca’ and ‘Morada’ are the most important cultivars.
In recent years, cactus pear has been promoted in the
districts of Pasorap and Cochabamba, supported by the Mexico
Forage Research Center (CIF) and San Simón Univer-
sity, Pasorapa City, under the Sustainable Agricultural Opuntias in Mexico have the highest degree of genetic
Development Program (PROAGRO). In the valley region diversity and the greatest level of consumption (Reyes
and Chaco, cactus pear production is currently being Agüero et al., 2005). The main species utilized in Mex-
increased to meet the demand for forage for livestock ico are: O. ficus−indica, O. joconoxtle, O. megacantha
in arid and semi−arid zones of Bolivia (M. Ochoa, per- and O. streptocantha. In addition, O. robusta, O. strep-
sonal communication). tacantha, O. leucotricha, O. hyptiacantha and O. chave-
na are harvested in the wild on 3 million ha in arid and
Brazil semi−arid zones with poor soil in Sonora, Baja Califor-
nia, Baja California Sur, Sinaloa, Chihuahua, Coahuila,
Brazil has a production area of 500 000 ha (J. Dubeaux, Nuevo León, Tamaulipas, Durango, Zacatecas, San Luis
personal communication) in the northeast region in the Potosí, Jalisco, Guanajuato, Querétaro and Hidalgo.
states of Paraiba, Pernambuco, Sergipe and Alagoas,
and recently extended to other states, such as Bahia, The first modern farming techniques with the best
Río Grande del Norte and Ceara. There are a large cactus pear varieties were established in the 1940s and
number of small growers and the majority of the pro- 1950s in the states of Mexico, Zacatecas, San Luis Po-
duction is destined for forage. Brazil boasts the largest tosí, Aguascalientes, Jalisco and Guanajuato. This sys-
cactus pear system in South America, and it has the tem peaked in 1985, as a result of social programmes
6 Crop ecology, cultivation and uses of cactus pear

launched towards the end of the 1970s (Pimienta Barri- mercial presentations – given the widespread pres-
os, 1990). Today, cactus pear production is distributed ence of cactus fruits in supermarkets in large and
among various states mainly in Federal District, State of medium−sized cities and the renewed interest in the
Mexico, San Luis Potosí, Zacatecas, Tamaulipas, Aguas- export market.
calientes and Guanajuato (SIAP, 2014). Cactus pear is
A keen interest in the nutritional properties of cactus
the sixth fruit crop in Mexico, after orange, avocado,
pear and the positive effects on human health attrib-
banana, mango and apple; it is more important than
uted to the regular consumption of fruits and tender
peach, guava or table grapes (SIAP, 2014).
cladodes has resulted in a myriad of scientific studies
The recent history of cactus pear as a crop in Mexico since the beginning of the century. Forage production,
presents vicissitudes. During the 1970s and early 1980s, on the other hand, has proven beneficial at experi-
the crop was heavily subsidized by government pro- mental and semi−commercial level, but has not taken
grammes and promoted extensively as an alternative to off – perhaps because of the abundance of standard
dry beans and corn. Extension was intense and exper- feedstuffs, the low revenue of traditional livestock pro-
imental, based on the limited technical developments duction systems and the loss of rangelands.
available. Many farmers began planting and producing
fruits without proper market development. In this peri- Cactus pears in family gardens or
od, the predominantly national market was subject to nopaleras de solar
extreme vagaries in prices. In 1985, the Government These gardens are found in rural environments, since
promoted farmers’ organizations based on the assess- cactus pear fruits are an important component of the
ment of crop needs and the evaluation of problems, diet in the Central Plateau and the north. Nopaleras de
involving all the actors in the production chain: grow- solar are grown in Zacatecas, San Luis Potosí, Guana-
ers, wholesale and retail vendors, and agro−industry juato and Aguascalientes, mostly for self−consumption
leaders. Research increased, but was held back by the or sale on local markets.
recurrent lack of proper funding. These developments
were in line with a general trend: the Mexican grow- Vegetable nopalitos production
er entered global markets and was exposed to open The farming of nopalitos − young tender cladodes for
competition from foreign growers while in a position to human consumption used in Mexican cooking − de-
exploit export markets. pends on a selection of the best varieties. The biggest
cultivation area is in Mexico City in the Milpa Alta bor-
In the semi−arid regions of central Mexico, cactus pear
ough, where research programmes have supported and
is now the most reliable and even profitable option
developed production since 1950. This area is the most
for utilizing rainfed semi−arid lands. In areas exposed
important in the country in terms of high production
to drought, it is the crop of choice over corn or dry
levels and quality. Opuntia ficus−indica (L.) Mill. is the
beans, while in slightly more benign sites, cactus pear
most important species for nopalitos production.
supplements the income obtained from these staples.
The available figures illustrate its importance: the plant-
United States of America
ed area covers 53 876 ha; it is the fifth fruit crop in
the country; about 20 000 families obtain income from
In the United States of America at time of the coloniza-
cactus pear cultivation; and gross annual production
tion of California, the Franciscan friars established the
has reached 428 763 tonnes year−1 (Gallegos Vázquez
first plantation. At the beginning of the twentieth cen-
et al., 2013). Vegetable production − characterized by
tury, selections created by Luther Burbank were widely
the utilization of small tracts of land − accounts for an
used in the diets of both men and animals. Today cac-
additional 12 000 ha of planted area.
tus pear is cultivated for fruit production in California,
The last two decades have seen significant changes in where there are 120 ha under intensive production
cactus pear production in Mexico: (“D’Arrigo Bros” trademarked cactus pears in 1928)
• Slow but steady adoption of drip irrigation in semi− (P. Felker, personal communication). In Texas, Arizona
arid areas – a practice unheard of in the last century. and parts of California, O. lindheimeri are often used
as emergency forage (Russel and Felker, 1987a). There
• Introduction of cactus pear to more benign areas,
are also some small industries based on fruit processing
with less limiting rainfall patterns and better quality
for jelly.
soils – located in the central highlands and some
subtropical regions in the central south and western Stephen D’Arrigo emigrated to the East Coast in 1911
part of the country. at the age of 17. He began growing vegetables such as
• Increased utilization of mechanical fruit−cleaning broccoli and fennel (common in his homeland but not
technologies. easily available in America) in San Jose, and became a
distributor shipping cactus pears east in the late 1920s.
• Use of improved packing materials and novel com-
History and economic and agro−ecological importance 7

In 1930, he registered a trademark, “Andy Boy”, with ha, 60% of which in the municipality of Sidi−Fredj (45
a logo depicting his 6−year−old son, Andrew. When km north of Souk Ahras) and the remainder in Ouled
Stephen died in 1951, Andrew took over the family Mimoune, Taoura, Dréa and Ouilène (Huffpost Algeria,
business; he still heads the company today, aged 87, 2015). The area was originally extended by the High
and takes a particular interest in the firm’s cactus pears. Commission for Steppe Development, the Directorate
Cactus pear cultivation in the Santa Clara Valley went of Agricultural Services and Forest Conservation in order
through several cycles of contraction and expansion to control desert progress (Belgacem, 2012).
during the Depression, the Second World War and the In the north, O. ficus−indica is used as a fence around
post−war years. As the area developed and land be- houses and small towns; fence plants are also used for
came more valuable, Marco and Sal LoBue, the grand- fruit production and, in the dry season, as a source of
sons of Rancadore, moved most of their production forage. Fruits are harvested from wild plantations and
south to Gilroy. In 1968, the LoBue family was ready are used for human consumption or sold on local mar-
to give up growing cactus pears, but Andrew D’Arrigo kets. The Opuntia cladodes in the south are used as
stepped in and bought the business, including the pro- emergent feed for small ruminants and camels.
duction orchards.
As in other African countries, the crop is gaining in
A few years later, D’Arrigo changed the name of interest in Algeria, which now has its first cactus pear
the product from “prickly pear” – which he thought processing unit. The infrastructure − based in Sidi−Fredj
sounded off−putting – to “cactus pear”. As labour be- and covering 5 000 m2 − can transform about 2 tonnes
came scarce, he discontinued the traditional practice of hour1. Its principal functions are the packaging of cactus
wrapping each fruit in tissue paper. In the 1980s, he pear and the production of essential oils, pharmaceuti-
moved his plantations to the Gonzales area southeast cals, juice, jam and livestock feed. The processing plant
of Salinas. represents an important means of income improvement
This fertile valley, with its equable maritime climate, is for the inhabitants of the wilaya of Souk Ahras (Agence
best known as the summer “salad bowl” of America, Ecofin, 2015).
and it might seem curious that cactus, most associated
in the public mind with broiling deserts, would fare
Ethiopia
well here. Moreover, the main season for cactus pear On the Eritrean border, the Ethiopian region of Tigray
in Mexico is summer, when the country exports large cultivates cactus pear, known as beles. The crop has
quantities of inexpensive fruit to the United States of an important economic and cultural role, reflected in
America. However, in the cooler northern climate, and traditional songs and sayings: “Oh my beles you spare
with the aid of the old Sicilian practice of scozzolatura me this summer till barley has cheerfully come to rescue
(meaning “take the berries away”), involving the re- me.” More than 85% of the population of Tigray deri-
moval of flowers from the spring flush, D’Arrigo har- ve their livelihood directly from agriculture. Given the
vests from October to March, obtaining higher prices ever−increasing pressures of humans and livestock on
when there is less competition. The D’Arrigo Brothers the land, combined with a decline in soil productivity
recently selected some new cultivars, thanks also to the and recurrent episodes of drought and famine, there is
work of R. Bunch and P. Felker increasing reliance on cactus pear to minimize risk and
ensure crop and food security. Cactus pear has a crucial
economic role: it is a source of food, animal feed and
AFRICA fuelwood; in some cases, it is a means of additional in-
come, increasing the efficiency and economic viability
The introduction of cactus to North Africa was furthe- of small and low−income farmers. Tigray covers about
red by the Spanish expansion during the sixteenth and 80 000 km2 and is estimated to have about 360 000
seventeenth centuries and also by the return of the Mo- ha of cactus pear, two−thirds of which are spiny plants.
Approximately half the existing area of opuntias was
ors to their homeland when they were finally depor-
planted; the remainder has been invaded by naturali-
ted from Spain in 1610 (Diguet, 1928). They took with
zed cacti. Various projects have been developed to raise
them the “Indian fig tree” with its succulent fruits and
awareness of the uses of cactus by local communities;
planted them around their villages (Diguet, 1928).
however, the introduction of Dactylopius coccus created
serious problems when it became aggressive to the wild
Algeria
stands. Cactus pear is mainly used for fruit production,
Historically, the introduction of the cactus in Algeria was although not in any intensive or rational way. Food and
similar to that in Morocco and Tunisia. Today, the area de- Agriculture Organization of the United Nation (FAO) co-
voted to the cultivation of Opuntia extends over 30 000 operation projects in the area, carried out by International
8 Crop ecology, cultivation and uses of cactus pear

Cactus Pear Network (CactusNet) members, have led to South Africa alone, with serious consequences for agri-
increased use of nopalitos. culture, despite its utilization as fruit and forage. In the
cooler parts of the country, the cactus was less aggressive
and farmers adopted it extensively (Walters et al., 2011).
Morocco
To solve the problem of uncontrollable invasions of
In Morocco, cactus was introduced in 1770 and it is now
cactus pear, South Africa relied on biological control.
well represented throughout the national landscape. As
About 80% of the infestations were brought under
a result of drought, the cactus area has evolved signifi-
control as a result of the serious damage caused by two
cantly in the past two decades: from 50 000 ha in 1998
cactus−feeding insects of American origin: the cactus
to > 120 000 ha at present. The region of Guelmin−Sidi
moth, Cactoblastis cactorum and the cochineal, Dacty-
Ifni accounts for over 50% (> 50 000 ha) of the na-
lopius opuntiae (Zimmermann et al., 2009). An estimat-
tional surface area, followed by Haouz−El Kelaa of the
ed 1 500 ha are cultivated for fruit and another 3 000
Sraghnas with 30% (about 33 000 ha). Khourigba is in
ha exclusively for forage. An estimated 150 000 ha of
third place, Doukkala in fourth. This increase in cactus
the original cactus pear infestations are now also exten-
area over the last two decades is the result of abundant
sively utilized, mainly for their fruit, which are sold on
planting operations carried out by farmers in collabora-
informal markets.
tion with development agencies and extension services
as part of national drought control programmes (Arba, Germplasm banks have been established in the Lim-
2009a). Currently, the modernization of Moroccan agri- popo, Free State, Eastern and Western Cape provinces,
culture (Green Morocco Plan) encourages cactus plan- with 42 varieties or types of O. ficus−indica established
tation as an alternative crop in less favourable regions. for research purposes. Research originally focused ex-
Each year, over 4 000 ha are planted in the centre and tensively on the use of cactus as drought fodder (Men-
south of the country (Ait Hamou, 2007). In the past 10 ezes et al., 2010). However, the focus has recently shift-
years, several firms and cooperatives of rural women ed to the intensive production of cactus for animal feed
have been created for the transformation of cactus. The and human consumption.
main products manufactured are: jam, vinegar, silage,
flour and seed oil. However, the main interest is in its Tunisia
role as a source of forage for livestock during the fre-
quent dry years (M. Ochoa, personal communication). Beginning in 1920–1930, cultivation for fodder produc-
tion has been gradually established in Tunisia, based
A recent threat to cactus in Morocco and throughout the
mainly on Opuntia ficus−indica f. inermis. For example,
Mediterranean Basin is the advance of cochineal (Dacty-
prior to the expansion of irrigated farming in the 1950s
lopius opuntiae). The origin of the infestation is hard to
and 1960s, the dairy operations around Tunis and other
establish, but it probably came from Spain where it was
cities in North Africa used fresh cactus as green stuff
reported in Murcia in 2006 and in Almeria in 2013. The
(particularly in summer and autumn) in addition to hay,
cochineal is today present in Sidi Bennour, Rhamna and
straw and cereal grain. Fodder plantations were sys-
Youssoufia; if no action is taken, it will proliferate, destroy
tematically developed following the expertise provided
Moroccan plantations and migrate to Algeria and Tunisia.
by Griffith to the Government of Tunisia in 1932 (Le
The authorities carried out an emergency intervention,
Houérou, 2002). In the early 1930s, under the coloni-
uprooting and incinerating all cactus plantations on more
al land allotment in Sidi Bouzid (central Tunisia), where
than 400 ha at Sidi Bennour in Doukkala (Abdelouahed
average rainfall is 250 mm, land ownership was only
Kidiss, 2016). A control strategy has been put in place in
granted by the Government on condition that contract-
collaboration with the International Center for Agricul-
ing beneficiaries, inter alia, planted 10% of the land
tural Research in the Dry Areas (ICARDA).
allocated to spineless cactus as an emergency fodder
crop reserve. This was a very wise decision, considering
South Africa
the frequent droughts faced by farmers (Le Houérou,
Cactus pear, Opuntia ficus−indica, was introduced in the 1965). With the support of international organizations
Cape well over 300 years ago. The plant was carried by (World Food Programme [WFP] and FAO), large areas
settlers to all the arid and semi−arid parts of the sub- (70 000 ha) in arid Tunisia were planted with cactus dur-
continent, where it was cultivated as a living fence and ing the 1960s and 1970s to serve as fodder reserves
for its delicious fruit. The plants gradually reverted to the (Le Houérou, 2002). Meanwhile the Tunisian Govern-
spiny forms over a period of about 150 years, contribut- ment provided support, based on state incentives and
ing significantly to its invasive properties and resulting in soft bank loans, to boost cactus plantations. In addi-
dense impenetrable thickets in certain regions, mainly in tion, in 1990 the country launched a national strategy
the Eastern Cape (Beinart and Wotshela, 2011). The cacti for rangeland improvement; 142 000 ha of cactus were
invaded > 2 million ha in the early twentieth century in planted in this period (OEP, 2016).
History and economic and agro−ecological importance 9

The current area covered by cactus is around 600 000 gation. Cactus is also planted under mixed plantation
ha under different forms: defensive hedges, groves sur- systems, intercropped with other fruit trees, such as fig
rounding rural houses, consolidation of erosion control and olive; however, due to the negative effect of cactus
structures, extensive fruit production and, more recent- spines on the fruits, this system is gradually disappear-
ly, intensive fruit production in favourable agroclimates. ing. In the high rocky areas of Jordan, spiny cactus are
Most of these plantations are located in arid areas with still planted, while spineless varieties cannot survive in
< 300 mm rainfall and production is destined for fodder these environments (Nasr, 2015). Given the severe live-
and fruit. In the last decade, in intensive cactus orchards stock feed shortage in Jordan, more effort is required
where all required inputs are applied, the practice of to raise awareness of cactus as fodder.
scozzalatura has been adopted to produce fruits for
both the local market and export. In winter, the price Lebanon
of cactus fruits reaches double that of apple, pear or
In Lebanon, cactus pear was introduced long ago in
banana. Tunisia uses the varieties used in Sicily (‘Rossa’,
coastal and internal areas at altitudes of 0–900 m asl.
‘Gialla’ and ‘Bianca’); indeed, it was Italian farmers who
Cactus pear is destined mainly for fruit production, but
introduced scozzolatura in the early 1990s. In recent
it is also grown as a natural fence for the production of
years, the agro−industry of cactus pear transformation
“arak” (unsweetened anise−flavoured distilled alcoholic
has expanded. Several enterprises, such as “Nopal Tuni-
drink). Cactus plantations are common under mixed
sia”, specialize in the production and export of organic
plantation systems growing alongside other fruit trees,
cactus pear fruit, cosmetics (seed oil, soap and sham-
as well as in family gardens for self−consumption.
poo), foods (jam, frozen pulp and juice) and dried flour
Extensive production systems for commercial fruit pro-
made from cactus pads.
duction exist in a very limited area in south and north
Lebanon (Chalak et al., 2012). Cactus pear cultivars
grown in Lebanon are ‘Baldi’ (spiny cactus) and im-
WEST ASIA ported cultivars from Europe, Brazil etc. The imported
cultivars include both spiny and spineless ones (Chalak
In most West Asian countries, cactus is long established, et al., 2012).
mainly for fruit production and as hedges around hous-
es. In recent years, there has been a growing interest in Syria
the crop, but as a secondary crop planted in marginal In Syria, as in Jordan and Lebanon, cactus pear has long
lands and only for fruit production. However, given the been cultivated, planted extensively in the rural areas of
low input requirements and the high level of adaptation Damascus mostly for fruit production. At a later stage,
to harsh environments, governments and farmers are and as a consequence of research projects, many culti-
becoming increasingly interested in growing cactus as vars were introduced to Syria from North African coun-
a high water−use efficiency crop to enhance local fruit tries. However, these accessions were kept in research
consumption. Nevertheless, despite the severe feed nurseries and not disseminated on farms for two main
shortages and the negative effects of climate change reasons. First, there was a lack of awareness of the im-
in these countries, cactus is still not recognized as a portance of this crop; second, the Ministry of Agricul-
good source of fodder, capable of making a significant ture was focused on the improvement and breeding of
contribution to alleviating the feed gap. On the other barley as the main livestock feed resource. In the last 2
hand, cactus was recently promoted in Syria for feed years, with the Syrian crisis and the shortage of fodder,
production, and a high level of adoption was observed. interest has increased in cactus cultivation in relative-
ly stable parts of the country, such as Swidaa, Homs
Jordan and the coastal areas. Cultivars sent by ICARDA were
In Jordan, spiny cactus plantations were started 60 planted as mother plants in two research stations in the
years ago as hedges around homes and orchards. The coastal area and the Damascus rural area; the new pads
spineless cactus was later introduced and planted ex- are now being distributed to farmers. A recent Interna-
tensively under a sole cropping system in the semi−arid tional Fund for Agricultural Development (IFAD)−fund-
areas in the middle of Jordan. In general, in addition ed project for “livestock development” planted 100
to the local spiny and spineless Khadri cactus, most ha for feed and fruit production. The results are very
available cultivars are imported to Jordan. The current promising, with a huge demand for pads and a high
area under cactus is approximately 300 ha, comprising level of adoption.
extensive production systems, hedges and mixed plan-
tations (Nasr, 2015). Extensive production systems are Gulf countries and Yemen
located mainly in Madaba, where farmers grow cactus Within the framework of its regional programme in the
for fruit production in rows using supplementary irri- Arabian Peninsula, ICARDA introduced 38 cactus ac-
10 Crop ecology, cultivation and uses of cactus pear

cessions from Tunisia in 2005 and established a mother pear has been exploited since the eighteenth century,
nursery in Oman. The pads of these accessions were when its multiple uses included fences in farming sys-
then introduced in research stations in the Gulf coun- tems and emergency fodder. It was known as “bread
tries and Yemen. Studies on production performances for the poor” (Barbera et al., 1991).
and adaptation to agro−ecological conditions in the
On the coast and close to the main cities, fruit pro-
Arabian Peninsula were conducted in Oman, Qatar,
duction was exported to the continent. This economic
Saudi Arabia, the United Arab Emirates and Yemen.
success was reinforced by the scozzolatura technique,
Since 2012, dissemination has been underway. In 2016,
developed by the Italians to be able to harvest in au-
a total of 47 farmers adopted spineless cactus: 15 in
tumn, producing better quality fruit than in the regular
Qatar, 10 in Oman, 9 in Yemen, 7 in Saudi Arabia and 6
August season. In 1975, modern plantations where es-
in the United Arab Emirates.
tablished for the fruit industry. Cactus pear production
thrives in environments that are environmentally too
Israel
poor for other crops.
In Israel, O. ficus−indica has been the subject of several
In Italy, cactus pear cultivation is concentrated on the
projects, including investigations to induce blooming
island of Sicily, which accounts for 90% of total produc-
and ripening in summer, autumn and winter. Today,
tion (3 500 ha of specialized area, 15 000 ha in total).
fruit is sold almost year−round, and at high prices
There are four important regions: San Cono, Mount
when off−season (Y. Mizrahi, personal communica-
Etna, Roccapalumba and Santa Margherita Belice; rain-
tion). ‘Ofer’ is cultivated on 350 ha in the Negev and
fall is 600 mm year−1. The cultivars, ‘Gialla’, ‘Rossa’ and
Arava valley areas, close to the Egyptian border in the
‘Bianca’, are grown; summer fruits account for 10% of
south and near Jordan in the west. In these regions,
production, while the winter harvest (August−Novem-
fruit ripens from late June to late August. Adoption
ber) accounts for the remaining 90%. Under irrigation,
of appropriate cultural techniques produces a second
yield can reach 25 tonnes ha−1 (Basile, 1990).
bloom, with fruits ripening between December and
April. There is also a limited production of dried flowers Portugal
harvested after pollination and sold as natural medicine
for non−malignant prostate growth (Jonas et al., 1998). In Portugal, the private sector has recently begun to
plant commercial plantations of cactus pear for fruit
production in the semi−arid areas of Alentejo and Al-
garve. More than 200 ha have been planted and 500
AUSTRALIA ha will be planted in the coming years, thanks also to
Australia has a long and interesting history relating to a programme to help unemployed young people who
the commercial use of Opuntia, which was first intro- want to work in agriculture (Gonçalves, personal com-
duced in the early 1840s for cochineal production and munication).
for fencing. Cochineal insects were imported, but they
did not thrive because of the abundance of bird and Spain
insect pests; the fledgling cochineal industry failed. In Spain − Andalusia, Murcia, Almeria and the Balearic
Nevertheless, Opuntia thrived thanks to the suitable Islands − cactus pear is only found on family planta-
climate and the lack of natural Opuntia pests and tions. Lanzarote (Canary Islands) has a small production
grazing animals. By 1925, the introduced species had of red dye.
naturalized and covered 24 million ha. Biological pests
were introduced and they successfully eradicated the PERSPECTIVES
vast feral populations.
“Green gold”, “fruit for the poor”, “treasure under its
The modern cactus pear industry in Australia is very spines”, “world vegetable dromedary”, “future plant”,
small with no more than 200 ha currently grown in the “sacred plant” and “monster tree” are just some of the
whole continent. Most production and varietal diversity epithets used for the plant and the fruit (Arias Jiménez,
is found in home gardens among Australians of Med- 2013a). These names convey the importance of opun-
iterranean heritage (S. Prowse, personal communica- tias in the work and lives of people, because of their
tion). resistance to drought and high temperatures and their
adaptability to poor soils. However, there is still much
EUROPE scope for development and improvement:
• Greenhouse effect. The increasing concentrations
Italy
of CO2 and other gases in the atmosphere will lead
Italy, in particular Sicily, represents an atypical example to a greater distribution of the species and increased
of the appreciation of Opuntia ficus−indica. Cactus productivity surpassing all previous levels (Nobel and
History and economic and agro−ecological importance 11

García de Cortázar, 1991). Moreover, the increase in

cactus pear plantations could be part of a strategy to
• Glochids and seeds: marketing obstacles to
lessen the accumulation of CO2 in the atmosphere.
overcome. The glochids on the rind can be removed
Cactus pear plantations can function, not only as a
effectively after harvesting and consumers need
water reserve, but as a carbon reservoir in arid and
to be informed accordingly. Reduced seed content,
semi−arid regions, where climate is more erratic.
on the other hand, is a major challenge: there have
More research is required.
been no important developments in recent decades,
• Plant diversity. Based on the current understanding although some selections of the ‘Bianca’ and ‘Ofer’
of the evolutive patterns of opuntias, further investi- cultivars (El Behi et al., 2015; Weiss et al., 1993a) are
gation is needed to address issues of generic delim- available; further research is required.
itation and species identity. The integration of mo- • Cladodes as food. Nopalitos offer good prospects
lecular, morphological and biogeographical data will for the future, but they are currently used only in
generate a clearer picture of the relations between Mexico and further efforts are required for their
the genus and produce a more stable and reliable dissemination. CactusNet has introduced nopalitos in
source of biological information about the diversity regional diets and gastronomy, and in Ethiopia the
of this exceptional plant family (Chessa, 2010). “Mums for Mums” project promotes the use of no-
• Marketing. Since the sixteenth century, opuntias palitos.
– mainly Opuntia ficus−indica − have been used • Multiple uses. There is clearly vast potential for
in subsistence gardening in many communities of non−food uses of cactus pear, exploiting its biofunc-
Africa, Asia, Europe and America. However, fruit tional, medicinal, nutraceutic and cosmetic proper-
consumption remains limited to local ethnic markets ties. Many products have been patented in recent
and there is little export. Cactus pear benefits from years (http://www.bionap.com/it/).
good marketing strategies only in Italy, Mexico, the
United States of America and South Africa (Inglese et • Forage. The exploitation of Opuntia for forage must
al., 2002a). Marketing and promotion campaigns are respond to the economic needs of each country and
therefore needed, as well as consumer−education technologies must adapt to livestock requirements;
strategies at local and international level (Caplan, Brazil is a clear example of progress in this regard.
1990).
The legacy of Park S. Nobel on the environmental biol-
• Fruit quality. How can people be encouraged to ogy of opuntias, combined with the cooperation and
consume cactus pear? Despite the efforts of the last technical assistance of the FAO–ICARDA International
20 years, current knowledge remains scarce and only Cactus Pear Network (CactusNet – created in 1993),
limited scientific information is available to growers paved the way for the current understanding of the
(Inglese et al., 2002a). It is essential to do away with potential of the species and its prospective role in mar-
the common belief that cactus pear needs few inputs ginal semi−arid areas. The last 25 years have seen: the
for good performance. This misconception has led publication of technical and scientific books, leaflets,
to bad management of cactus pear plantations and acta and proceedings; the organization of International
poor fruit quality. The Italian experience has shown Society for Horticultural Science (ISHS) symposia, re-
that rational orchard management gives better re- gional congresses and seminars; and the implementa-
turns and higher quality fruit with low management tion of projects, thanks to the efforts of the Network,
costs. To improve productivity and fruit quality, there its coordinators and FAO and ICARDA officers. It is vital
must be greater awareness of the influence of the to continue moving towards these clear objectives for
environment and orchard management on fertility, the benefit of the poorest regions in the world. Cactus
fruit growth and ripening (Ochoa, 2003). It is also pear is the “world vegetable dromedary” and one of
important to establish fruit quality standards and the most valuable legacies given to the world by Ameri-
implement proper orchard design and management. ca’s native population over five centuries ago.
 02

Origin and taxonomy

of Opuntia ficus−indica
Roberto Kiesling a and Detlev Metzing b
a
Argentine Dryland Research Institute, National Scientific and Technical Research Council,
Mendoza, Argentina
b
Vegetation Science and Nature Conservation Group, Institute for Biology and Environmental Sciences,
University of Oldenburg, Germany
 02 Origin and taxonomy
of Opuntia ficus−indica

INTRODUCTION and several other taxa and names are often con-
fused due to ambiguous descriptions and a lack of
types (Leuenberger, 1988).
Opuntia ficus−indica is the cactus species of
greatest agronomic importance, due to its deli- Griffiths (1914) considered O. megacantha Salm−
cious fruits, but also to its stems, which are used Dyck to be the wild thorny form of cultivated O.
as fodder for livestock or as a vegetable for human ficus−indica (in the narrow sense, or O. ficus−ind-
consumption (Alkämper, 1984; Kiesling, 1999a; ica f. ficus−indica); this was later corroborated by
Casas and Barbera, 2002). The use and cultivation molecular studies (Griffith, 2004). However, the
of opuntias dates back to prehistoric times, long most likely explanation is that they have a com-
before the Spaniards arrived in the Americas. The mon ancestor. Both arose from natural hybridiza-
Indian chroniclers were the first to record these tion and multiple sporadic interbreeding. Benson
plants and its fruits, which were carried to Spain (1982) considers O. megacantha as a cultivat-
and initially used as ornamental plants (Casas and ed taxon and a synonym of O. ficus−indica in the
Barbera, 2002). It is probable that opuntias were “spiny form”, and discards the category of varie-
brought back after the first or second visit of ty or form. Other authors (e.g. Gibson and Nobel,
Columbus to the Caribbean, although the first de- 1986; Brutsch and Zimmermann, 1993) follow this
finitive record is from Mexico in 1515 (in the chron- reasoning. According to Kiesling (1999a), O. meg-
icle of Fernández de Oviedo, reproduced by López acantha is a reversion to spined plants from es-
Piñeiro et al., 1992). The long history of the use, caped, spineless O. ficus−indica (see below).
cultivation and domestication of O. ficus−indica
One of the few authors mentioning O. ficus−indica
and related species resulted in taxonomical and no-
solely in the spineless form is Bravo Hollis (1978),
menclatural problems, summarized in this chapter.
who uses only morphological characters for the
delimitation; however, she contradicts herself more
than once (1978: 288, 322). In the same book she
ORIGIN AND TAXONOMY OF lists six variedades hortícolas (which correspond to
OPUNTIA FICUS−INDICA the concept of cultivar) based solely on the fruit
characters. Scheinvar (1995) accepts three separate
The spineless form of O. ficus−indica − common species: O. ficus−indica, O. streptacantha and O.
in agriculture today − is the result of a long selec- amyclaea, but also considers O. ficus−indica as a
tion process in cultivation and it is absent in wild
stocks. According to Bravo Hollis and Sánchez
Mejorada (1991), its domestication began about
Figure 1 8 000 years ago. Reyes Agüero et al. (2005) main-
Opuntia ficus-indica
tain that domestication took place in the south of
the meridional Mexican highlands. Archaeological
references indicate that the opuntias used 8 000
years ago cannot be associated directly with O.
ficus−indica. Callen (1965) studied the food habits
of pre−Columbian Mexican Indians and found ep-
idermis remnants of Opuntia in coprolites on the
floor of caves, indicating that the consumption of
Opuntia dated back thousands of years to at least
5200 BCE. According to Casas and Barbera (2002),
archaeological remains of opuntias were found in
caves of the Ajuereado phase (14000–8000 BCE).
Several taxa are mentioned as putative ancestors
of O. ficus−indica, in particular O. megacantha
Salm−Dyck and O. streptacantha Lemaire. These
Origin and taxonomy of Opuntia ficus−indica 15

cultivated form originating from O. streptacantha and tetra−, hexa− or octoploids in the putative relationship
keeps it separate at a specific rank for practical reasons of O. ficus−indica (spiny form as O. megacantha) n = 44
(L. Scheinvar, personal communication). (Pinkava et al., 1973); O. streptacantha n = 44 (Pinkava
and Parfitt, 1982); O. streptacantha 2n = 88 (Palomino
Several names of the series Streptacanthae and Ficus−
and Heras, 2001); O. amyclaea and O. megacantha 2n
indicae (Britton and Rose, 1919) correspond to minor
= 88 (Sosa and Acosta, 1966); O. polyacantha 2n =
morphological variations of O. megacantha. A descrip-
44, 66 (Stockwell, 1935). Octoploids (2n = 88) are also
tion by Britton and Rose (1919) of the series Ficus−indi-
reported for other taxa of the series Streptacanthae (in-
cae, which includes the more or less spineless forms of
cluding ser. Ficus−indicae) (Segura et al., 2007; Majure
the group discussed here, stated: “None of the species
et al., 2012a). Polyploidy is favoured by hybridization.
is definitively known in the wild state, but all doubtless
Natural, interspecific hybridization in the genus Opun-
originated from tropical ancestors, and they may all
tia has been proved by several studies (e.g. Benson and
represent spineless races of plants here included in our
Walkington, 1965; Grant and Grant, 1982; Griffith,
series Streptacanthae.”
2003; McLeod, 1975) and hybridization in cultivation
On the other hand, others maintain that there are is common. The occurrence of higher ploidy levels of
enough differences to keep the series Streptacanthae cultivars in comparison with wild relatives is obviously
and Ficus−indicae separate. Colunga Garcia et al. (1986) true for cultivated O. ficus−indica (Mondragón Jacobo
wrote: “Thus, these two groups can be differentiated and Bordelon, 1996). For O. ficus−indica alone, many
based on cladode size and areole length, fruit and seed chromosome counts show that both the spiny and
length, as well as the length and weight of pulp of the the spineless forms are octoploid (Pinkava et al., 1973,
fruit (the edible portion of the fruit). In our opinion, the 1992). Cultivated plants of O. ficus−indica in Italy were
cladode size, the weight of fruit pulp, as well the areole found to be octoploid (Barbera and Inglese, 1993).
form may differ caused by the selection process that However, this species is also reported as hepta−, pen-
aims to improve the quality for use as animal fodder or ta−, hexa− and diploid, so there exists a variation of
for human consumption.” chromosome numbers, depending on the provenance
(Spencer, 1955; Weedin and Powell, 1978; Pinkava,
The spines − presence and size − represent another very
2002; Majure et al., 2012a) − unless the different num-
variable character. Although the development of spine-
bers are the result of misidentified study material or
less forms was encouraged during the domestication
anomalies in the meiosis. McLeod (1975) indicates the
process, the opposite − from spineless to spination − is
presence of hybrid specimens with 2n = 77, in between
also possible. Such reversions, where some branches of
O. ficus−indica megacantha (octoploid: 2n = 88) and
spineless forms produce spines after drought stress or
O. phaeacantha var. major Engelm. (hexaploid: 2n =
other sorts of stress, are mentioned by several authors
66). Carpio (1952) mentions also n = 44 for O. ficus−
(Griffiths, 1912, 1914; Le Houérou, 1996a; Kiesling,
indica. He suggests that the anomalous meiosis and
1999a). Moreover, when seeds of the spineless form
the existence of tetravalentes show that O. ficus−indica
are sowed, a small percentage of the seedlings devel-
is either an allopolyploid originating from two species
op spines; and vice versa, sowing of seeds from spiny
with 2n = 44 or an autopolyploid. Allopolyploidy of O.
plants results in a small proportion of spineless plants
ficus−indica is confirmed by Griffith (2004).
(Berger, 1905; M. Ochoa, personal communication; I.
Chessa, personal communication; authors’ experience).
The reversion of spineless cultivars to spiny plants was
also observed in South Africa and Sicily (Zimmermann, MOLECULAR STUDIES
2011; Leuenberger and Arroyo Leuenberger, 2014). The
presence of spines is not a valuable character in Opuntia Given that morphological studies resulted in different
taxonomy, because the formation of spines is not inde- taxonomic hypotheses, greater insight was expected
pendent of environmental factors (Labra et al., 2003). from molecular studies (mainly of DNA) concerning var-
iability, relationship and origin of the Opuntia species
and O. ficus−indica in particular. Although several stud-
CHROMOSOME NUMBERS ies focus mainly on the genetic diversity of cultivars (e.g.
Bendhifi et al., 2013; El Finti et al., 2013; Ganopoulos et
al., 2015), some give insight into the differentiation of
Determination of chromosome number and ploidy level
O. ficus−indica. Wang et al. (1999) studied five cactus
is a useful tool in plant taxonomy. The basic chromo-
fruit cultivars from Mexico and Chile, two ornamental
some number in the cactus family is n = 11, and the
Texas accessions, and one vegetable accession from
number in somatic cells is mostly 2n = 22. In the sub-
Mexico. The DNA analysis revealed significant differ-
family Opuntioideae, 64.3% of the taxa are polyploid
ences between the market accessions, but only slight
(Pinkava et al., 1985). Several karyotypic studies show
differences between the fruit cultivars (including spined
16 Crop ecology, cultivation and uses of cactus pear

and spineless forms). The genetic diversity of cultivated clustered within different groups. While the study
cacti seems to be low in general, probably because failed to identify the ancestor, O. hyptiacantha could be
they originate from a narrow germplasm base (Boyle related with the majority of the O. ficus−indica samples
and Anderson, 2002). studied. Srikanth and Whang (2015) compared three
taxa of Opuntia cultivated in Korea and found that the
In the study by Labra et al. (2003), molecular data re-
Korean O. ficus−indica is closely related to O. engel-
vealed a high genetic similarity between O. ficus−indica
and O. megacantha. The only (morphological) differ- mannii and O. ellisiana, but not to the O. ficus−indica
ence between the units is the presence of spines. The samples taken from the GenBank database. Molecular
authors conclude that O. ficus−indica should be consid- studies reveal the faultiness of the current taxonomy
ered a domesticated form of O. megacantha. for the species and cultivar complex of O. ficus−indica,
and question whether these problems are caused by
Griffith (2004), when studying the origin of O. ficus− hybridization, adaptive genetic responses, phenotypic
indica using molecular data, found a well−supported plasticity, epigenetic bases or other factors (Valadez
clade including O. ficus−indica, O. streptacantha, O. Moctezuma et al., 2014).
tomentosa, O. leucotricha and O. hyptiacantha, all from
southern and central Mexico (diversity centres of Opun-
tia − Barthlott et al., 2015). The analysis supports the
DISTRIBUTION AND NATURALIZATION
hypothesis that the centre of domestication was central
Mexico and O. ficus−indica may be polyphyletic, i.e.
While its ancestors originate in central Mexico (Griffith,
descended from different lineages. This could be due
2004), O. ficus−indica has been taken by humans to oth-
to hybridization (in nature or during cultivation), deri-
er areas of the world with warm climates. Following the
vation of multiple unique clones from various parental
introduction of O. ficus−indica in Spain around 1500,
stock, or lineage sorting of multiple internal transcribed
the species (and others of the same genus) spread and
spacer (ITS) copies in an ancestral population from
naturalized throughout the Mediterranean area, soon
which O. ficus−indica and closely related species may
becoming a characteristic element of the landscape. It
have descended (Griffith, 2004). Caruso et al. (2010)
was already widespread in Europe in 1550 (Mottram,
studied the genetic diversity of O. ficus−indica cultivat-
2013). Therefore, it is not surprising that one species, O.
ed genotypes. Their analysis supports the hypothesis
that O. ficus−indica consists of a group of multiple un- amyclaea Ten., was described in 1826 as coming from
related clones, derived from different parental species Italy, where it had been found near the town of Amyclæ
and selected for different agronomical features. (today Monticelli). It corresponds to the spiny form of
O. ficus−indica. O. amyclaea was considered a form of
Majure et al. (2012b) concluded that O. ficus−indica is O. ficus−indica by Schelle (1907); for this reason, in the
one of several species originating from allopolyploidiza- taxonomic rank of form its previous name was O. ficus−
tion events caused by the hybridization of species be- indica f. amyclaea (Ten.) Schelle. Berger (1905, 1912b)
longing to different clades. The O. ficus−indica samples also assumed that this Opuntia established in Italy must
studied by Caruso et al. (2010) did not cluster separate- be the original form of O. ficus−indica, i.e. an ancestral
ly from other species (O. amyclaea, O. megacantha, O. form. A form of O. ficus−indica from Argentina was also
streptacantha, O. fusicaulis and O. albicarpa), indicating described as a new species (O. cordobensis Speg.), and
that the current taxonomical position and the genetic similarly a form from Bolivia (O. arcei Cárdenas) (Kiesling,
patterns do not fit very well. Lyra et al. (2013a) studied 2013). At the beginning of the twentieth century, the
characteristics of cultivars of four species (O. ficus−in- American botanist and agronomist David Griffiths stud-
dica, O. albicarpa, O. streptacantha and O. robusta), ied and cultivated opuntias in Texas for taxonomic and
but with the used marker (ITS) it was not possible to agronomic evaluation (Benson and Walkington, 1965;
assign the samples of these species to separate clades. Walkington, 1968). He described several species from
This difficulty may arise from the fact that the sam- cultivated specimens, and some of these species − better
ples are of hybrid origin or have a common ancestry.
treated as cultivars − are considered synonyms or hybrids
Valadez Moctezuma et al. (2015) advanced this latter
of O. ficus−indica today (e.g. O. fusicaulis Griffiths, 1908;
assumption when O. ficus−indica, O. albicarpa and O.
Kiesling et al., 2008). In the eighteenth century, O. ficus−
megacantha proved impossible to separate in different
indica was introduced to other continents by navigators
clades. Similarly, Samah et al. (2015) could not detect
who − given its vitamin C content and low perishability
clear boundaries between O. ficus−indica, O. albicarpa,
− used it as a vegetable to prevent scurvy (Diguet, 1928).
O. megacantha, O. streptacantha, O. lasiacantha and
They also transported it to:
O. hyptiacantha. Astello Garcia et al. (2015), in a study
of the molecular composition of five Opuntia species, • meet the demand for carmine dye from cochineal,
could not verify a proposed domestication gradient for which feeds on Opuntia;
O. ficus−indica, when different cultivars of this species • use as fodder;
Origin and taxonomy of Opuntia ficus−indica 17

• incorporate in the human diet; and before Linné, to designate several more or less similar
• make living fences. species. In other languages, similar vernacular names
Adapted to harsh and dry conditions, opuntias could are used: figo da India (Portuguese); Indian fig (English);
easily escape and naturalize in arid areas of Africa, Asia figuier d’Inde (French); Indianische Feige (German); fico
and Australia. O. ficus−indica had already been intro- d’India (Italian) (Reynolds and Arias, 2001).
duced to South America by the Spanish conquerors, for Another widely known name is tuna de Castilla (or
example, in Bolivia (Hoffmann, 1955). Opuntias spread nopal de Castilla), obviously derived from the name of
rapidly in many regions, sometimes becoming invasive: the former Spanish kingdom of Castilla, from where
a threat to native biodiversity and to agricultural land use Opuntia was distributed to other countries. Le Houérou
(Brutsch and Zimmermann, 1993; Barbera and Inglese, (1996a) mentions Andalusia as the first propagation
1993: 11). Today O. ficus−indica is naturalized in 26 coun- centre in that continent − the region Christopher Co-
tries outside its native range (Novoa et al., 2014). lumbus returned to after his voyages. Following the
dissemination of Opuntia in Spain, it was introduced
In all tropical arid countries where it is cultivated or
to North Africa, where it was called higo de los cris-
naturalized, O. ficus−indica has undergone genetic
tianos. The spiny form is currently widespread in Mo-
alterations and phenetic modifications, resulting in
rocco, where it is called tapia (from the Spanish word
new forms that have been distinguished and formally
for fence), a reference to its use as a hedge (A. Prina,
named. Sometimes they are classified as species or vari-
personal communication). The name “sabra” − a word
eties, even when a classification and naming as cultivars
used to refer both to the native people and to the
of the two forms (O. ficus−indica f. amyclaea and O.
prickly pear plant − illustrates the extent of its distribu-
ficus−indica f. ficus−indica) would be more appropriate
tion in the Mediterranean area. Indeed, the species is
(Brickell et al., eds, 2009).
frequently used to illustrate postage stamps in several
While there are numerous recordings of the introduc- countries around the Mediterranean Sea.
tion of the spineless O. ficus−indica f. ficus−indica to
In 1769, the Franciscan missionaries took the cultivated
different countries, it should be noted that there are
form from Mexico to California, where it is called “mis-
almost no references concerning the introduction of sion cactus” (Benson and Walkington, 1965; Walk-
the spiny form. ington, 1968). However, it is not certain whether the
natives already cultivated this species before the arrival
of the Franciscans. Walkington (1968) used the name
VERNACULAR NAMES O. ficus−indica in a wide sense.
The species is very important for the economy of
Given the importance of O. ficus−indica and its numerous
northeast Brazil, where it is mainly used as forage
benefits, it has been given many names in its native range
throughout the year and is called palma forrageira. It
and in the regions where it has been introduced (Reyes
is not known when it was introduced to that country
Aguero et al., 2005). Some of these names are a good
(Domingues, 1963). Palma−de−gado is another com-
illustration of the origin of introduction and distribution.
mon name for O. ficus−indica in northeast Brazil. Sev-
The name “tuna” is of Caribbean origin (Bravo Hollis eral other vernacular names beginning with “palma”
and Sánchez Mejorada, 1991: 505) and was used by are applied to the species in regions of the Brazilian
the first Spaniards to arrive in the Americas. To be pre- state Bahia, where the pads are used as forage and the
cise, it is a Tain name (Moringo, 1966). It usually refers fruits, mucilage and roots have several uses in human
to the fruits, but is also used for the vegetative parts of nutrition and medicine (Andrade, 2008).
the Opuntia species. The name is currently in use in a
very extensive area, which suggests that it was the first
name known by the Spaniards, even before the Mex- THE ROLE OF COCHINEAL
ican names (since they reached the Caribbean islands
before the mainland). The cochineals of the genus Dactylopius parasite on
many cactus species, including those of the genus
“Nopal” is a Mexican name derived from the Nahuatl
Opuntia. Cochineals have great species−specificity.
Nopalli (Bravo Hollis and Sánchez Mejorada, 1991: 558;
Dactylopius coccus, also known as grana, has an ab-
Moringo, 1966: 424), and is used for several species. Ten-
solute preference for O. ficus−indica and for taxa that
ochtli is the original name used in large parts of Mexico.
are considered its synonyms or are closely related (e.g.
The first Spanish name is higo de las Indias, a reference O. megacantha, O. streptacantha, O. cordobensis).
to its origin, the New Indies; it gave rise to the first José de Acosta (1590, cited after Di Lullo, 1944) wrote
scientific name: Cactus ficus−indica Linné. The epithet, about tunales domésticos in High and Low Peru (today,
ficus−indica, was used as a “diagnostic phrase” long Bolivia and Peru), and his observations are in line with
18 Crop ecology, cultivation and uses of cactus pear

current knowledge of O. ficus−indica; it can therefore

be deduced that this cactus was probably present in
those areas in that early period. In Peru, the use of
grana dates to the pre−Hispanic period, probably as
far back as the time of Christ (Marín, 1991; Sáenz et
al., 2002a); however, it is not clear whether it was D.
coccus or another cochineal species. Fester (1941) and
Fester and Lexow (1943) mentioned a spectrometric
analysis of the colours of pre−Hispanic tissues (Paracas,
from Peru) and of tissues from northern Argentina,
demonstrating that the red colorant probably origi-
nates not from D. coccus in Mexico and Central Ameri-
ca, but from other species of Dactylopius.
The economic importance of cochineal production in
the nineteenth and twentieth centuries led to the in-
troduction of both host plant and parasite into several
countries outside their native distribution area. Accord-
ing to Piña (1981), D. coccus was introduced to Peru in
the nineteenth century, before the country began to
export grana in 1830.
For Argentina, Lafone Quevedo (1927) described how
grana was harvested from quiscaloro (vernacular name
for the wild species Opuntia sulphurea and O. ana-
cantha, among others). According to the Argentinean
entomologists, Claps and de Haro (2001), five wild and
red−dye−producing species of Dacytylopius parasite on
several cactus species native to Argentina. A sixth spe-
cies is D. coccus, found on cultivated and naturalized
O. ficus−indica. During the 1980s, the former President
Menem introduced D. coccus to encourage the produc-
tion of grana in Argentina. While this attempt at grana
production was not successful, naturalized D. coccus
was recorded for the first time in 1999 in La Rioja, Ar-
gentina (De Haro and Claps, 1999). Later, there were
also recordings in Salta (Van Dam et al. 2015). Hence,
it is plausible that the red colorant extracted in the past
was obtained from other Dactylopius species. The same
indica is not a natural species (Kiesling, 2013), rather a
researchers state that while the native cochineal live
complex of cultivars and naturalized clones.
on different Cactaceae species, they live neither on O.
ficus−indica nor on O. cordobensis − a strong indication O. ficus−indica can be distinguished from other species
in favour of its species−specificity. According to Van by several characters. The receptacle of the flower and
Dam et al. (2015), D. coccus is a domesticated form, as later the fruits have many areoles (≥ 38), with a very
is also assumed for O. ficus−indica. small number of cultivars having fewer (Pinkava et al.,
From the known records it can be deduced that D. coc- 1992; Kiesling, 1999a); the areoles are situated mostly
cus was not present in Andean South America before on very notable tubercles. Other Opuntia species have
the arrival of the Spanish conquerors. fewer areoles at the flowers and fruits, situated on less
prominent tubercles.
The name Opuntia ficus−indica has priority over other
TAXONOMICAL AND names given to this species (Kiesling, 1999b). Nomen-
NOMENCLATURAL CONSPECTUS clatural types based on herbarium specimens, which
determine the application of names, were not desig-
The different entities considered and denominated un- nated for O. ficus−indica until 1991, and for O. strep-
der several scientific names as species correspond to a tacantha and O. megacantha until 2010 (Leuenberger,
single biological entity. In the strictest sense, O. ficus− 1991; Scheinvar et al., 2010), although all three names
Origin and taxonomy of Opuntia ficus−indica 19

were published in the eighteenth or nineteenth centu- CONCLUSIONS

ry and widely used afterwards. The specimen chosen
as nomenclatural type (lectotype) of O. ficus−indica Based on the facts presented and biological knowledge,
(Leuenberger, 1991) corresponds to a plant without (or the following conclusions can be drawn:
with very small) spines. However, the presence/absence
• O. ficus−indica f. ficus−indica is the result of a long−
of spines is not a useful character for distinguishing
lasting selection process during cultivation. A result
O. ficus−indica from other species, and the spiny and
of this process is the development of polyploid (up to
spineless forms must be considered different pheno-
octoploid) and more vigorous forms or cultivars, more
types of one species. Nevertheless, this character is
convenient for human use than their wild relatives.
used here to separate the spiny and spineless plants of
The selection of less spiny plants has led to spine−
O. ficus−indica formally in the rank of form (the lowest
free forms. Fruit size and quality also influenced the
level of the taxonomic categories), even if both forms
selection process, which had begun before the arrival
can arise from each other.
of the Spaniards in Mexico. Intraspecific as well as
Opuntia ficus−indica (L.) Mill., Gard. Dict. ed. 8, interspecific hybridization suggests a polyphyletic
Nr. 2; 1768 origin.

Basionym: Cactus ficus−indica L., Sp. pl.: 468. 1753. • The cultivated, spineless form was introduced to
1753. Spain, probably a few years after the discovery of
Plants are shrubby or tree−like, up to 6 m high, usually America (c. 1500), and initially used as an ornamental
with well−developed trunks. Stem segments are varia- plant and a curiosity in the gardens of the nobility.
ble, broadly obovate or oblong to spatulate, flattened, From there it was taken to other countries in the
20–50 cm long, 20–30 cm wide, about 2 cm thick, matt Mediterranean as well as to South America, South
green, covered by a very thin waxy layer, areoles 2–5 Africa, India and Australia. Navigators increased its
cm apart. Glochids falling away early, spines absent or distribution by taking it as a fresh vegetable to guard
2 (–7) per areole, 0.5–1.0 cm long, weak whitish. Flow- against scurvy. The main reasons for the further dis-
ers yellow, rarely orange, 6–8 cm long and 5–10 cm in semination of O. ficus−indica by humans were the
diameter during anthesis. Fruit with numerous (approx. production of fodder in arid areas, the use of the
30–40) areoles, with glochids, rarely with spines, tuber- fruits or pads for human consumption, and cochineal
culate, ovoid to oblong, 6 (–8) cm long, 3 (–5) cm in production.
diameter, yellow, orange, pink–green or reddish. • In several countries with a suitable warm and arid
climate, the species was introduced and cultivated,
Opuntia ficus−indica f. ficus−indica spreading by vegetative and generative reproduction
Synonyms: Opuntia ficus−indica var. gymnocarpa until it became naturalized. This process happened in-
(F.A.C. Weber) Speg., Anales Mus. Nac. Buenos Aires dependently and more than once in several places, in
ser. 3, 4: 512. 1905. Opuntia ficus−indica var. decuma- different countries, and on all continents, in the native
na (Haw.) Speg., Anales Mus. Nac. Buenos Aires ser. 3, as well as the new distribution areas. This resulted in
4: 512. 1905. Opuntia ficus−barbarica A. Berger. Mon- new centres of infraspecific differentiation, with the
atsschr. Kakteenk. 22: 181. 1912. Opuntia tuna−blanca emergence of cultivars and subsequently naturalized
Speg. An. Soc. Cient. Arg. 99: 107. 1925. clones and hybrids. The naturalized forms developed
slightly different morphological and physiological
Areoles without spines or with only small and weak spines. characters compared with the cultivated clones. Spiny
forms emerged repeatedly from the spineless plants.
Opuntia ficus−indica f. amyclaea (Ten.)
• O. ficus−indica is considered a species, or a group
Schelle, Handb. Kakteenkultur: 51. 1907
of multiple unrelated clones derived from different
Basionym: Opuntia amyclaea Ten., Fl. Neap. Prod. App.: parental species. The native distribution area of the
15. 1826. ancestral taxa is central Mexico.
Synonyms: Opuntia ficus−indica var. amyclaea (Ten.) A.
Berger, Hort. Mortol: 411. 1912. Opuntia megacantha
Salm−Dyck, Hort. Dyck.: 363. 1834. Opuntia strept-
acantha Lem., Cact. Gen. Sp. Nov. 62. 1839. Opuntia
cordobensis Speg., Anales Mus. Nac. Buenos Aires ser.
3, 4: 513. 1905. Opuntia arcei Cárdenas, Cact. Succ. J.
(Los Angeles) 28: 113. 1956.
Areoles with notable spines.
 03

Morphology and
anatomy of Platyopuntiae
Loreto Prat, Nicolás Franck and Fusa Sudzuki - University of Chile, Santiago de Chile, Chile
03 Morphology and anatomy
of Platyopuntiae

INTRODUCTION 4−8 m. It has also been observed that plants period-

ically fertilized with manure develop succulent and
The species of Opuntia, particularly those in the sub- unbranched roots, while they usually develop more
genus Platyopuntia, have developed anatomical, mor- lateral roots that soon develop a surface layer of loose
phological and physiological adaptations to grow and exfoliating cork (North and Nobel, 1992).
survive in arid environments where severe water stress
hinders the survival of other plant species (Beccaro et According to Snyman (2004, 2005), the root system
al., 2015). This chapter describes some of the most of O. ficus−indica is very complex and can exhibit four
important anatomical and morphological adaptations kinds of roots:
present in the root, shoot and reproductive structures • Skeletal roots: formed from a primary skeleton of
of Opuntia ficus−indica (prickly pear) − the Opuntia scarcely fibrous roots, 20–30 cm long, which very
species with the largest economic value. soon increase in thickness by secondary growth to
form a periderm. When the root skeleton is kept
dry for some time and then remoistened, absorbing
ROOT SYSTEM roots are formed from latent buds within a few hours
to quickly respond to the advent of moisture (North
Cactaceae roots receive little attention. They differ et al., 1993; Dubrovsky et al., 1998). The initiation of
from those of other plants, as they develop xeromor- lateral roots is always endogenous from parenchyma
phic characteristics that enable the plant to survive cells belonging to the secondary phloem. Research
prolonged periods of drought. Roots can contribute into the regularities of adventitious root formation in
to drought tolerance in various ways: O. ficus−indica has shown that the fine lateral roots
on the tap−root die off with age. This process stimu-
• Restriction of the root surface and decrease in their per-
lates cell division in the parenchyma root tissues and
meability to water − fine roots are covered with a layer
the formation of meristem spots with adventitious
relatively impermeable to water, or roots abscise at a
roots (Gibson and Nobel, 1986). This fine and fragile
cicatrization layer in order to avoid water loss to dry soil.
mass of roots is formed from short and branched
• Rapid absorption of the small quantity of water sup- rootlets, completely covered with root hairs. Also,
plied by light rains − “rain roots” develop within a different kinds of ectomycorrhizae, mostly the vesic-
few hours after a shower and disappear as soon as ular−arbuscular type, grow together with the short
the soil dries up, or there is a reduction in the root and branched rootlets.
surface from which water flows off.
• Absorbing roots: formed within a few hours as the
• Decrease in shoot transpiration due to root high neg- lateral buds rapidly respond to the advent of mois-
ative potential. ture. Gibson and Nobel (1986) named these absorb-
The above characteristics mean that these drought−re- ing roots “rain roots”. They develop from the hidden
sistant water savers can have high hydraulic resistance latent bud in the cortex of the older roots. These rain
(Passioura, 1988); this, in turn, decreases water flow to roots die off as soon as the soil dries.
the shoot. • Root spurs: developed from the bulkiest mass of
Opuntia ficus−indica has a shallow and fleshy root sys- roots as clusters (Boke, 1980). The spur base of O.
tem, which spreads horizontally. Root distribution may ficus−indica exhibits a crown of appending−like
depend on the type of soil and cultural management bracts and, contrary to Boke’s description, the roots
(Snyman, 2005). Under favourable soil conditions, a developed from the spur in O. ficus−indica are of two
tap−root develops, penetrating nearly 30 cm into the classes: short, gross and fleshy, with plenty of root
soil. Under drought conditions, such as those found in hairs; and two or three slender and long ones, similar
arid and semi−arid regions, fleshy side roots develop to absorbing roots. It is not known whether the short
from the tap−root to take up soil moisture at a greater roots die off or mature with time.
depth. Nevertheless, in all kinds of soil, bulk masses of • Roots developing from areoles: developed when
absorbing roots are found in the topmost centimetres, the areoles are in contact with the soil. At the onset
at a maximum depth of 30 cm, but spreading outwards of their development, they are gross and without
Morphology and anatomy of Platyopuntiae 23

root hairs; they have a prominent calyptra, with the • Regulate the movement of carbon dioxide into − and
epidermal cells forming bract−like appendages. The oxygen out of − the plant.
growth of the young roots is very rapid; they become • Retain water within the plant body.
slender with a cortex 3−4 cells thick, and are covered
• Protect the plant from fungi, insects and intense sun-
with root hairs. In some of these cells, water deficit
light (Mauseth, 1984).
induces the formation of a higher number of endo-
dermal cells with Casparian bands closer to the root The outermost wall of the epidermis is impregnated
tip (De Micco and Aronne, 2012). and covered with wax of a fatty substance called cutin.
The cuticle of cactus, in comparison with the cuticle
of most plants, is relatively thick and in O. ficus−indica
can reach 8–20 μm. In chemical terms, cutin is not a
CLADODE pure product like cellulose; it is a mixture of fatty acids,
which polymerize spontaneously in the presence of
O. ficus−indica is a CAM (crassulacean acid metabolism)
oxygen. Not only is cutin water−resistant, it is also one
plant, commonly considered drought resistant, because
of the most natural chemicals known (Mauseth, 1984).
it stores considerable amounts of water in its shoots
(Nobel, 1994, 1995). The morphology and anatomy of Examination of the surface of O. ficus−indica cladodes,
the shoots have evolved to serve this function. using scanning and transmission electron microscopy,
reveals a slender aggregate coating of a basic wax
According to Buxbaum (1955), Cactaceae are broadly
structure of semi−vertical plates. This coating gives the
characterized by short shoots (“areoles”, bearing hairs
cladodes their glaucous appearance. According to Met-
and spines), a succulent stem with a green cortex, and
calf and Chalk (1979), the epicuticular wax present in
the absence of foliage leaves. According to Hunt and
O. ficus−indica corresponds to the basic group III type
Taylor (1986), the areoles of Opuntioideae, in addition classified as plates and scales.
to or instead of spines, develop short hair−like prickles
called glochids. In O. ficus−indica, the stem−like organs, The waxy, rough surface around the epidermal cell
known as cladodes, are succulent and their shape is serves a number of functions. As far as water equilib-
typically oblong to spatulate−oblong, usually 30−40 cm rium is concerned, the cuticle prevents the escape of
long, sometimes longer (70−80 cm), and 18−25 cm wide. water vapour from the plants and repels surface water.
Anatomically, the cladode in a cross−section is a eustele, The white cuticle reflects solar radiation, resulting in a
formed by: skin; cortex; vascular tissue, in a ring and made reduced stem temperature (Gibson and Nobel, 1986).
from collateral bundles separated by parenchyma tissue; According to Mauseth (2006a), the epidermis in most
and pith, which is the major succulent tissue. Cactaceae persists for decades as a living, transparent
tissue capable of facilitating gas exchange. This is in
Skin contrast with the ephemeral stem epidermis in most
seed plants, which lasts only as long as a plant lives or
The skin consists of one layer of epidermal cells and dies when bark is formed on perennial plants. The ep-
6−7 layers of hypodermal cells, with thick primary walls idermis in cactus is present and alive, as long as green
resembling a laminar collenchyma tissue. The epidermal colour is visible in the underlying cortex.
cells are flat, thin and shaped much like paving stones.
The epidermis and hypodermis provide an effective bar- Mauseth (2006a) also mentions the fact that the epi-
rier to prevent physical damage and maintain mechan- dermis in Cactaceae presents a folded surface, rather
ical integrity. The thick cells of the hypodermis are very than a smooth one, affecting the shoot’s strength,
flexibility and ability to swell without tearing. A plicate
strong and act as the first line of defence against fungi,
surface means that, as water is absorbed and shoot
bacteria and foraging by small organisms. The skin re-
volume increases, the rib bases spread and the volume
mains intact for a long time and is eventually replaced
of the rib is increased without requiring any increase in
by bark (periderm).
surface area. As the rib bases widen, the inner cortex
The periderm is formed from epidermal cells, either as can also expand. As a result, there is a change in the
part of the natural ageing process, or from the deeper volume − but not the surface area − of the shoots, and
tissues of cortex when injury causes the skin to break. there is no damage to the epidermis or hypodermis
When bark forms from the thick cell walls of the hypo- (Mauseth, 2000).
dermis, it breaks off.
Stomata are abundant in the stem epidermis of most
The epidermis constitutes the outermost layer of cells other Cactaceae and are evenly distributed over both
in the cactus body. It is a continuous protective cell sides of the entire stem surface (Sajeva and Mauseth,
layer, which also bears the stomata (Figure 1). The epi- 1991). In O. ficus−indica, they are randomly spread, not
dermis has three major functions: great in number and are usually 15–35 mm2 (Mauseth,
 24 Crop ecology, cultivation and uses of cactus pear

1984). The guard cells do not differ from other ephemeral structure corresponds to a leaf (Figure
flowering plants; they are 40 μ and sunken, so the 3). The podarium is prominent in the first stage
pore is hidden from the stem surface. A prominent of cladode development, and becomes lost as
substomatal canal, through the sclerenchymatous the stem ages. The areole begins to form at the
hypodermis tissue, forms a passageway for gas very base of the short apical meristem, and very
exchange between the atmosphere and the pho- soon the spines develop from the basal meristem
tosynthetic tissue below the hypodermis; the en- (Mauseth, 1984); the number of spines varies, but
dostomatic chamber is in the chlorenchyma layer there are often one or two long central spines
(Figure 2). The pair of guard cells are surrounded and another shorter lateral spine. The long central
by 3–4 rows of subsidiary cells. The epicuticular ones grow for a longer period than the others;
surface brightens these cells, and the stomata can they are thicker because they are produced by
therefore be observed with low magnification. more robust primordia (Gibson and Nobel, 1986)
Within the epidermal cells and hypodermis lies a and have elongate cells with lignified cell walls. As
very conspicuous crystal aggregate of calcium ox- they grow, glochids also appear.
alate, named drusas (Figure 1).
Spines and glochids

The presence of spines is the particular character-

istic of the areoles and their morphology has po-
tential taxonomic significance (Robinson, 1974). It is
possible to distinguish two types: spines and spine−
hairs (glochids). According to Boke (1944), glochids
and spines are best regarded as morphological
equivalents of leaves and the differences between
them are quantitative; both types are derived from
tunica and corpus, as are leaf primordia.
The number and duration of the spines and
glochids in O. ficus−indica depend on the type.
Usually, spines are present in the first stage of
cladode growth, and most of them fall off as the
Figure 1 According to Mauseth (2006a), in O. ficus−indica,
temperature rises, only occasionally remaining at
Stomata in the epidermis cell walls tend to be thin, not thickened
epidermis of a fully
the base of the cladodes for a long period.
and not lignified. The epidermis is often thinner
developed cladode.
Also druses present than expected for a xerophyte and long, multicel- According to Robinson (1974), it is very easy to dis-
in the hypodermic are lular uniseriate trichomes are abundant at every tinguish the subfamily Opuntioideae by the rough
shown, surrounding the node, but the internode epidermis has almost surface of the glochids and spines. In O. ficus−in-
stomata.
no outgrowth. A hypodermis of several layers dica, the spines have a rough, and the glochids
of cells, with extremely thick and hard walls, is a smooth surface; the glochids are arranged in
present (Gibson and Horak, 1978; Mauseth and clusters of 7−12 in the cavities of the areoles.
Ross, 1988; Mauseth, 1996, 1999b; Mauseth and Spines are white; one or two are long (1.0−1.5
Kiesling, 1997; Terrazas Salgado and Mauseth, cm); two smaller ones accompany each of them.
2002; Loza Cornejo and Terrazas, 2003; Terrazas As the tops of the spines harden, the basal meris-
Salgado and Arias, 2003). tem continues to lengthen. According to Gamong,
cited by Buxbaum (1950), the spines are formed
Areoles in series. Glochids are tiny, short and multicel-
lular; it is suggested that the first two spines of
Areoles are oval, 2 mm below the skin surface. the areole could be stipules, where the elongated
Under appropriate environmental conditions, new smooth cells (14−16 in number) form a chain and
cladodes, flowers or roots will arise from the mer- do not become sclerified. They are brown, and
istem tissue of the areoles. In O. ficus−indica, the give this colour to the areoles. The surface of the
areoles are distributed in a helical formation, and spine−hair is covered with calcium carbonate and
they develop spines (rather than leaves, as most pectic substances (Buxbaum, 1950). Glochids are
plants do). When the cladode is young, the areole grouped in 4−6 tight clusters, from which it is
forms at the base of the podarium (cladode). The possible to distinguish 2−3 somewhat longer ones
podarium holds a small subulate short−lived green with a spine−like structure. They are hard, scler-
structure, which soon shrivels and falls off. This ified and pointed; their surface is covered with
Morphology and anatomy of Platyopuntiae 25

barbed scales, that permit them to lodge in the ma) parts both contain idioblasts, filled with mu-
skin and which cannot usually be removed easily. cilage or with crystals smaller than those of the
While growing, the glochids are fastened to the epidermal cells. While the green part containing
areole, but a phellogen then develops in the outer the chloroplasts has the main function of photo-
cortical region around the base of each cluster. As synthesis, the white part is a simple water−storage
a result, in O. ficus−indica, they are deciduous and bag. The parenchyma is the “chemical factory”
easily removed when either touched or blown by that produces the hormones, alkaloids and other
the wind. Their precise function remains unclear, products formed during plant metabolism (Mau-
but they may condense water from the air (Bux- seth, 1984; Sajeva and Mauseth, 1991). It does not
baum, 1950). Moreover, spines help to reduce have chloroplasts, but does have other organelles
daytime stem temperature, and their presence − amyloplasts − for starch storage. It is made from
also diminishes light interception by the cladode. homogeneous parenchymatous tissue, with small
intercellular spaces.
Corte
The precise function of the mucilage is not known;
however, it is generally believed that it helps to
Most Opuntioideae have a thick, chlorophyllous
retain water inside the plant. Mauseth (1980) gives
palisade cortex overlying a less or non−chloro-
a detailed explanation of the development of mu-
phyllous inner cortex (Mauseth, 2005). Those with
cilage cells. Very active dictyosomes produce small
laterally flattened cladodes lack surface plications.
sacs or “vesicles” of mucilage along their sides,
In Opuntia, below the hypodermis lies the chlor-
and this mucilage substance is deposited outside
enchyma, which constitutes the tissue between
the cytoplasm as the amount of mucilage increas-
the hypoderm and the ring of vascular bundles.
es. Thereafter, the cytoplasm dies and the orga-
Opuntia chlorenchyma is thick and succulent and
nelles break down, leaving only mucilage, where
consists of a primary cortex, formed from a rather
once there was a living cell. The mucilage cells are
homogeneous mass of parenchymatous cells. It
present in all types of tissue: hypodermic, cortical,
is chlorophyllous in the outer part and arranged
vascular and pith. Mucilage is very slippery, com-
in long radial rows to form a prominent palisade
plex and formed by indigestible carbohydrates
chlorenchyma, similar to a leaf palisade. The inner
(Gibson and Nobel, 1986).
part is white and formed from spheroidal parenchy-
matous cells, similar to those of spongy mesophyll.
Pith
All species of Opuntioideae with flattened cla-
dodes have wide (660–3 600 mm) canals filled The pith of O. ficus−indica is made from large
with mucilage cells floating freely in extracellular spheroidal parenchymatous cells with thin cell
mucilage, unattached by middle lamellas. The ca- walls, similar to those of the inner cortex. Most
nals are lined with a multilayered epithelium. Mu- pith cells in close proximity to the vascular bun-
cilage−cell canals occur just outside the primary dles contain numerous spheroidal starch grains
phloem, widening as inner epithelium cells detach or mucilage and some small druses. Surprisingly,
and float into the lumen (Mauseth, 1980, 2005). the perimedullary zone (outermost pith adjacent
The green (chlorenchyma) and white (parenchy- to protoxylem and medullary rays) of some Opun-

Figure 2
The epidermis with 4−5
layers of the
hypodermis cells

Figure 3
Areole in the young
cladode showing two
spines, ephemeral leaf
and several brownish
glochids
26 Crop ecology, cultivation and uses of cactus pear

tioideae consists of wide−band tracheid (Mauseth, WOOD

2006b), a cell type associated with xylem rather than
ground tissue, such as pith or cortex. In Opuntioideae, wood has tremendous diversity: some
species have ordinary fibrous wood, while others have
Vascular tissue wide−band tracheid (WBT) wood (Gibson, 1977, 1978a,
b; Mauseth, 2006b). There are also two variations in
Beneath the cortex, O. ficus−indica has a ring of col- WBT in Opuntioideae, since in some cases WBTs are
lateral vascular bundles with soft tissue between them located in the rays themselves. The distinction between
(eustele), corresponding to fascicular cambium or med- those WBTs located in the ray system and those in the
ullary rays. The vascular bundles are connected to the axial system is not clear because WBTs occur in both.
meristematic tissue of the areoles, and form an elab- Axial regions are defined by the presence of vessels
orate, very fine fenestrate vascular tissue network of (and by sieve tube members in corresponding regions
accessory bundles. As Freeman (1970) mentions for O. of secondary phloem), and rays are areas with no ves-
basilaris, anastomosing of stem bundles is common and sels (and corresponding regions of secondary phloem
results in the formation of a complex cylinder around lack sieve tube members). Ray WBTs can be distinctly
the pith. A vascular network is formed at each node larger than axial WBTs in the same shoot. In the other
by the fusion of two sympodia and a leaf trace with an variation in Opuntioideae, WBTs occur only in rays, not
areole trace and numerous accessory bundles; a closed in the axial system. Rays are very wide, consisting of
system is thus created. Between these bundles, large WBTs and parenchyma. The fusiform initial−derived
rhomboidal parenchymatous gaps are formed above axial systems consist of just one or two rows of vessels
the point at which each leaf trace diverges from the and a few axial parenchyma cells.
vascular network towards the areole.
The xylem is simple, and its vessel elements (75 μ wide) are
greater in number than the scattered tracheary elements FLOWER BUDS
(40 μ wide). Primary vessels, as well as tracheary elements
of vascular tissue, are characterized by helical and annular The axillary buds in cactus are areoles, which have
thickenings, with scalleriform or reticulate perforations in developed very early in the axil of the leaf primordia.
the secondary walls (Hamilton, 1970). Mucilage cells and The areole begins to evolve as a mass of cells, which
rapidly become a regular shoot apical meristem with
spheroidal starch grains are abundant around the vascular
all the proper zones: tunical, central mother cells, pe-
bundles. Freeman (1969) puts forward the theory that
ripheral zones and pith rib meristem. The spines and
mucilaginous canals or ducts are not actually formed;
glochids develop at a very early stage. After a period
rather, there is a frequent lysogenic breakdown of the
(variably long or short), the meristem at the centre of
walls separating the mucilaginous cells.
the depression of the areoles goes dormant. When it
All cactus stems, even the most flattened cladodes in becomes active again and grows as a long shoot in-
Opuntioideae, have a single ring of collateral vascular stead of a short shoot, it forms a branch with leaves
bundles (a eustele, typical of all non−monocot seed and areoles or produces a kind of super−modified long
plants). These primary vascular tissues have few or no shoot, which is the flower (Pimienta Barrios, 1990).
unusual features in Cactaceae (Bailey, 1961; Gibson,
A review by Reyes Agüero and Valiente Banuet (2006)
1976; Mauseth, 2004).
on the reproductive biology of Opuntia indicates that,
in contrast with all other Cactaceae, the meristem that
can produce either a new cladode or a flower will not
BARK produce again. Furthermore, 74% of floral buds in O.
ficus−indica grow on 1−year cladodes, with most veg-
With few exceptions, cork cambium of cacti arises from etative buds sprouting from 2−year cladodes. O. ficus−
epidermis cells (Mauseth and Ross, 1988; Mauseth, indica develops 20 or more floral buds per cladode,
1989, 1996, 1999a, b; Mauseth and Kiesling, 1997; and during flower development, the pericarp maintains
Terrazas Salgado and Mauseth, 2002). Because the its photosynthetic activity and areolar development
one and only cork cambium is superficial, cactus bark of glochids, spines and leaves. When the floral bud
consists simply of alternating bands of cork cells and of O. ficus−indica is 0.5 cm long, the gynoecium and
sclereids (both produced by the cork cambium); it never the androecium can be distinguished microscopically.
contains the hypodermis, cortex, primary or secondary Stamens rapidly differentiate into pairs of anthers, each
phloem typical of bark in many seed plants. The wa- having two pollen sacs. The same authors mention that
ter−storing cortex is never shed. Lenticels appear to be in O. ficus−indica the ovary has many carpels, varying
absent (Terrazas et al., 2005). in number from 6 to 12 (usually 8), that grow together
Morphology and anatomy of Platyopuntiae 27

at the base and become separated at the stigmatic other fruit trees (apple, pear etc.) where floral dif-
region. The unilocular ovary contains up to 270 ferentiation begins the year before (Pimienta Bar-
ovules or seed primordia weighing 0.44–3.01 mg. rios and Engleman, 1985). The sterile part of the
In O. ficus−indica, flower anthesis occurs after flower is represented by the perianth and there
noon, and flowers close at sunset and open again are few differences between sepals and petals:
the following day. At the onset of anthesis, sta- the sepals are smaller, but both are oblong and
mens are grouped together around the style but fused at the base, with brilliant yellow or pink pet-
become separated later. Anther dehiscence can als. The yellow flowers change colour to orange
occur 12 hours before or at the time of blooming. or pink after fecundation. The numerous stamens
are base−fixed and inserted in the cavity of the re-
Many characteristics make the cactus flower
ceptacle in a fasciculate arrangement, developing
unique (Figure 4):
centrifugally in such a way that the inner whorl
• Perianth segments, weakly differentiated as petals. is under the pistil and the upper one is at the al-
• Spirally arranged stamens. titude of the stigma. The filaments are free and
• Pistil of four or more fused carpels, enclosed in a the anthers produce a large amount of dry pollen
floral cup. grains, which shed before the ovules (protandry).
• Unilocular ovary with parietal placentation, According to Pimienta Barrios (1990), the anthers
apparently embedded in the end of a modified have a thigmotropic movement. At the beginning
branch (Boke, 1980). of anthesis, the stamens are close to the style and
Furthermore, cacti are among the few plants in the anthers are in contact with the base of the pis-
which the exterior of an inferior ovary, called re- til. This process is known as cleistogamy, and the
ceptacle, displays leaves and perfect areoles. This protandry of the flower explains the autogamous
structure later becomes the peel of the fruit. As nature of O. ficus−indica (Rosas and Pimienta
the bud emerges, it is possible to see whether it is Barrios, 1986). Pollination is entomophilous when
vegetative or reproductive from its spatial volume. the flower is open; the stigma is placed above the
The reproductive bud is more spherical (Figure 5), stamens (Pimienta Barrios, 1990).
while the vegetative is flatter. The pistil has a wide style at the base, and a
The proportion of flowering to vegetative buds prominent sectioned, granular and viscous stigma
is 3 : 1, and 10% of the cladodes may have both formed from ten septs and shaped like an anther.
kinds of bud in equal proportions (Sudzuki Hills et The ovary is syncarpous, unilocular, formed from
al., 1993). The flowers are hermaphrodite and ac- five carpels and has a parietal placentation.
tinomorphic. They develop on the upper part of a
In Opuntia, the funicle is slender and relatively long;
1− or 2−year cladode, and occasionally on a 3−year
the ovules have true integuments (Archibald, 1935),
cladode. Both kinds of bud usually develop over the
with the inner one beyond the outer. The third in-
cladode surface with greatest exposure to light.
tegument is a modified part of one of the two nor-
Floral differentiation occurs over a short period, mal integuments and it surrounds the ovule com-
from 50−60 days after the meristem starts to be pletely (Eames, 1961). Ovules are circinotropous,
active, through to anthesis, in contrast with many since the funicle surrounds the ovule and covers the

Figure 4
Longitudinal section of
a fully developed flow-
er, showing an inferior
ovary with numerous
ovules and the cortex,
petals, stamen and pistil

Figure 5
Floral bud at the edge
of a fully developed
cladode.
28 Crop ecology, cultivation and uses of cactus pear

micropyle like a thick third integument. It is possible to of the receptacle. The peel originates in the receptacle
observe this structure in seeds of O. ficus−indica, where and has the same morphology as the cladode:
it is present with the micropyle pore quite free. • Epidermis with ephemeral leaves and perfects are-
The inner surface of the funicle is covered with short oles; the glochids are more permanent than in the
hairs − “papillae” − especially at the point where the cladodes.
funicle touches the micropyle. The ovules are arranged in • Slender hypodermis and bulky cortex.
two rows in each ventral prominent suture. This situation • Abundance of mucilage cells, but no crystals.
may suggest that the carpel union is margin to margin,
and dorsal bundles extend to the ovules (Boke, 1980). The pulp is formed from the outgrowth of the
trichomes that originate in the epidermal cells of the fu-
niculi and the funicular envelope (Pimienta Barrios and
POLLEN GRAIN Engleman, 1985). Fruit size depends on the number of
fecundated ovules and the number of seeds that abort
The pollen grain has a thick outer layer, the exine, (Archibald, 1935; Pimienta Barrios, 1990; Barbera et al.,
which has a protective function. Its volume changes 1994). It has not yet been established why seeds abort.
depending on the level of humidity (Eames, 1961). The The fruits often have two kinds of sterile seeds, one
inner layer is the intine; it is thin and readily adapts itself of which sometimes predominates. The lack of viability
to changes in size. The patterns of exine have consider- of the seeds is related to an abortion of the ovule or
able taxonomic and phylogenetic value, as the wall may of the young adventitious embryo (Archibald, 1935).
become very thick and complex in structure and the In the latter case, the nucellus continues to grow, even
outer layer is sculptured with projecting ridges, spines after the embryo is formed, and at the time the funicle
and granules. In Opuntia, the pollen grains are spheroi- is formed, it changes into a hard lignified coat. In Chile,
dal, reticulate and policolpate; they have 18 colpos – i.e. many fruits of O. ficus−indica have revealed a third kind
similar to the pollen grain of O. fusicaulis described by of non−viable seed. It is an anomalous form, possibly
Scheinvar (1995), which has 20 colpos. There are a high also resulting from the transformation of the funicle
number of pollen grains and it is possible to observe into a hard cover, but with the active development of
germinated pollen tubes over the glandular epidermal the nucellus, which eventually emerges through the mi-
surface of the broad stylar canal 24 hours after anthe- cropylar pore that was not blocked by the ovule. In O.
sis. The fecundation of the ovules gradually begins 48 ficus−indica cultivars, the presence of a high number of
hours after anthesis and continues for 10 days. This pat- normal seeds in the fruit is considered an obstacle for
tern of fertilization was named by Rosas and Pimienta its commercialization. Italian cultivars have an average
Barrios (1986) as “progamyc”. The average number of of 273 seeds per fruit, of which 146 are normal and 127
fecundated ovules per flower depends on the cultivar sterile. Israeli cultivars have an average of 268 normal
(Rosas and Pimienta Barrios, 1986). seeds per fruit and Mexican cultivars 203 normal seeds
In O. ficus−indica, 18.4−30.1% of pollen grains deposit- per fruit (Reyes Agüero and Valiente Banuet, 2006).
ed on the stigma germinate and form pollen tubes. Up The equatorial diameter best represents the fruit’s fresh
to 397 tubes may be formed in this species, and they de- and dry weight. Comparisons between the develop-
velop over the glandular epidermis of the inner channel ment of the peel and the edible portion reveal that the
of the style. Most tubes are initiated on the stigma and growth of the peel is greater during the first weeks
grow towards the base of the style. Pollen tubes reach after flowering, whereas the edible portion begins to
the style base about 48 hours after pollination and ovule expand more in the last 5–6 weeks before ripening.
fecundation begins 72 hours later. After 4 days, 48% of According to the review of Reyes Agüero and Valiente
ovules are fertilized (Rosas and Pimienta Barrios, 1986). Banuet (2006) on the reproductive biology of Opuntia,
Polyembryony of nucellar origin is common in Opuntia. O. ficus−indica fruit grows vigorously in terms of length,
This explains the development of two or three plants width, weight and volume in the first 20–30 days after
from only one seed. Archibald (1935) estimated that anthesis; it stops growing approximately 59–90 days
nucellar polyembryony, with the development of en- after anthesis. The peel grows fastest during phase 1 of
dosperm and without fertilization, occurs in O. ficus−in- fruit growth, the seed has maximum growth in phase
dica and other cacti. 2, and maximum pulp growth occurs during phases 2
and 3. Fruit weight is affected by the order of produc-
FRUIT tion of the flower bud and the number of fruits on the
cladode. Thus, floral buds that sprout earlier usually
The fruit of O. ficus−indica is a simple fleshy berry become heavier fruits. Furthermore, the heaviest fruits
formed from an inferior ovary sunk into the stem tissues are obtained from cladodes with only six fruits.
 04

Ecophysiology and
reproductive biology
of cultivated cacti
Paolo Inglesea,Giorgia Liguoria and Erick de la Barrerab
a
Department of Agricultural and Forestry Sciences, University of Palermo, Italy
b
Institute of Research in Ecosystems, National Autonomous University of Mexico,
Mexico City, Mexico
04 Ecophysiology and reproductive biology
of cultivated cacti

INTRODUCTION Mojave Desert in California, or the Rajasthan Desert in

India (Felker and Inglese, 2003). Le Houérou (2002) de-
Cactus pear − Opuntia ficus−indica (L.) Mill. − is a CAM scribes plantations in Aziza (Lybia) where the maximum
(crassulacean acid metabolism) plant cultivated in a wide temperature may exceed 50° C. O. ficus−indica cladode
range of environments, resulting in major differences in cannot survive at 70° C (Nobel, 2002).
plant survival and development, and in crop potential. The fruits of O. ficus−indica can be harvested from July
The ecological success of opuntias, specifically O. fi- to November in the Northern Hemisphere − Mediterra-
cus−indica, is due largely to their peculiar daily pattern nean Basin, California, and Mexico − and from January
of carbon uptake and water loss, both of which occur to April in the Southern Hemisphere, depending on
mostly at night. Like other CAM plants, cactus pear the genotype and genotype × environment interac-
opens its stomata at night to fix CO2 and accumulate tion. Natural or induced reflowering may extend the
and store malate in the vacuoles of the chlorenchyma ripening period to January−February in the Northern
cells. Since night−time temperatures are lower than di- Hemisphere and to September−October in the South-
urnal ones, and relative humidity is generally higher, the ern Hemisphere. Almost continuous flowering has been
transpiration of CAM plants is three to five times lower reported in Salinas, California (Bunch, 1996), resulting
than that of C3 and C4 plants (Nobel, 1988). The result in an extended fruit ripening period.
is a tremendous increase in water−use efficiency and in
the plant’s ability to thrive in semi−arid environments
characterized by a restricted water supply (200–300 mm
CAM CYCLE
of annual rainfall) or where long periods of drought and
relatively high temperatures occur. The mechanisms of
Originally attributed to the Crassulaceae family, CAM
adaptation to aridity are not necessarily valid in relation
represents a mechanism of concentration of CO2 which
to high temperatures. Although they occur at night, CO2
has evolved in response to dryness in terrestrial envi-
uptake and acid accumulation are strongly influenced by
ronments and deficiency of inorganic carbon in water
environmental variables such as air temperature, light,
environments (Keeley, 1998).
plant water status, nutrients and soil salinity (Nobel,
1988). Indeed, in the native area of the Central Mexican CAM occurs in 16 000 species (6–7% of plant species),
Plateau (1 800–2 200 m asl), rainfall is < 500 mm, the belonging to over 300 genera from around 40 families,
mean annual temperatures range from 16 to 18° C, and ranging from the deciduous tropical forest to desert
the daily maximum temperature for the hottest month platyopuntias and columnar cacti. The vast majority
does not exceed 35° C (Pimienta Barrios, 1990). of plants using CAM are angiosperms. Most of them
are either epiphytes (e.g. orchids, bromeliads) or suc-
In the Mexican mesas (tablelands), the dry season coin-
culent xerophytes (e.g. cacti, cactoid Euphorbias); but
cides with a cold winter, while the hot season, during
CAM is also found in lithophytes, terrestrial bromeliads,
which fruit and vegetative growth take place, is very
the halophyte Mesembryanthenum crystallynum, one
humid and rainy; exactly the opposite occurs in the
non−succulent terrestrial plant (Dodonaea viscosa) and
Mediterranean Basin, where the dry season coincides
one mangrove associate (Sesuvium portulacastrum).
with the hottest days, when the fruit develops and veg-
etative growth occurs (Inglese et al., 2002b). In Sicily,
Characteristics of CAM and succulence
where O. ficus−indica is grown for fruit production,
annual rainfall is around 500 mm and the mean annual
• Separation of CO2 uptake and decarboxylation −
temperature ranges from 15 to 18° C, with peaks of
temporally or spatially.
37° C in August, during the fruit development period
(Inglese, 1995). O. ficus−indica occurs extensively in • Reduction of transpiration due to night−time stomatal
northern Africa (Monjauze and Le Houérou, 1965a), in opening − and water storage organs contain 90−95%
the highlands (2 000–2 500 m asl) of Tigray, Ethiopia water compared with 40−70% of non−succulent wood.
and in South Africa. In contrast, it is absent in regions • Reduced water loss − CAM plants lose 20–30% of
with rainfall < 350 mm and daily summer maximum the water lost by C3 o C4 plants for the same degree
temperatures > 42° C, such as the Sahelian belt, the of stomatal opening during daytime CO2 uptake.
Ecophysiology and reproductive biology of cultivated cacti 31

• Low root: shoot ratios and fast root growth dur- • Phase 2. Early in the morning, the transition
ing wet conditions. takes place from PEPc to Rubisco activity.
• Suberization of cortical cells, with the formation • Phase 3. During the day, plants close their guard
of root–soil gaps during soil drying. cells, stomata are tightly closed and the pH in-
• Internal recycle of water from parenchyma vs creases. Malate diffuses out of the vacuole and
chlorenchyma − maintains turgor of the photosyn- is decarboxylated. CO2 is released (rising from 0.2
thetic tissue over a wide range of water content. to 2.5%) in the cytosol and fixed into the Calvin
cycle in the chloroplasts by the ribulose−1,5−bis-
• Supply of water and solutes to fruit through phosphate carboxylase/oxygenase (Rubisco) lead-
phloem (phloem osmotic pressure is relatively ing to the synthesis of starch or other glucans.
low: 0.94 MPa (2–3 times less than for most
• Phase 4. In the late afternoon PEPc becomes
other vascular plants).
active. Under very dry environmental conditions,
The CAM pathway can be summarized as follows this may be the only phase of the CAM cycle
(Figure 1): that takes place.
• Phase 1. At night, CO2 fixation occurs when sto- Measurements of gas exchange in O. ficus−indica
mata open and CO2 diffuses into the mesophyll’s began in the early 1980s, when Nobel and Hart-
intracellular spaces and then into the cytosol, sock (1983) measured CO2 uptake on single clado-
where it is bound to phosphoenolpyruvate (PEP), des. At optimal temperature and intercepted radi-
a 3−C compound, through PEP carboxylase. The ation, instantaneous values of net CO2 uptake of
enzyme catalyzes the formation of oxaloacetate, 1−year cladodes may reach 18 µmol m –2 s–1, with
which can be transformed into malate by NAD+ a total daily CO2 uptake of 680 mmol m –2 (Nobel
malate dehydrogenase. To avoid inhibition, and Bobich, 2002). However, although individu-
malate is actively transported from the cytosol to al−cladode net photosynthesis (Pn) determinations
the vacuole, where it is converted into malic acid, are useful for estimating gas exchange rate per unit
leading to a noticeable increase in acidity. The area, they have limitations when used to scale up
vacuoles of the cells of the chlorenchyma occupy to whole canopy gas exchange, because there may
more than 90% of the cell volume because of be wide variability in carbon assimilation within the
nocturnal accumulation of organic acids. canopy, due to differences in cladode age (Samish

Figure 1
Light Dark Light Dark Light Dark Crassulacean acid
metabolism (CAM)
Carboydrate

Triose phosphate
Carboydrate
breakdown
Starch

PCR
cycle Phosphoglycerate

Phosphoenolpyruvate pH
Malate CO2 Pyruvate
Total organic acids
Oxaloacetate
Malic
Krebs
cycle
Malate

Citric

Stomata Stomata CO2

closed open
32 Crop ecology, cultivation and uses of cactus pear

and Ellern, 1975), intercepted radiation (Nobel, 1988), • Root: shoot ratio ranges from 0.12 (Nobel, 1988) to
crop load (Inglese et al., 1994b), source–sink relation- 0.09 (Inglese et al., 2012) and 0.20 (Inglese et al.,
ships (Pimienta Barrios et al., 2005) and response to 2012) on mature irrigated trees.
abiotic stress (Nobel and Bobich, 2002). • Wax layer in the epidermis is 10–50 μm vs 0.2–2 μm
Few data are available on cladode net CO2 uptake ac- in leaves of C3 or C4.
cording to cladode age. Samish and Ellern (1975) noted • The chlorenchyma of O. ficus−indica can reversibly lose
that titratable acidity decreases linearly with increasing 70% of its water content at full turgor and the parenchy-
age, and 1−year cladodes have a level of acidity three ma can lose 82%, both still being able to fully recover.
times higher than 2−year cladodes, located in a basi- • After a 15−week drought period, the parenchyma
petal position. Carbon translocation from 2−year cla- loses 60% of its water content and the chlorenchyma
dodes to meet the photo−assimilate demand of 1−year 25% (Goldstein et al., 1991).
fruiting cladodes has been hypothesized (Inglese et al.,
1994b; De la Barrera and Nobel, 2004), and carbon Sunny better than cloudy
partitioning between mother and daughter cladodes
changes depending on the cladode developmental • Light and CO2 uptake are shifted in time and noctur-
stage and environmental conditions (shading, water nal stomata opening depends on daily rather than
availability) (Luo and Nobel, 1993; Pimienta Barrios et instantaneous PPF values.
al., 2005). CO2 uptake of 2−year cladodes throughout • Compensation point at PPF = 3 moles m−2 day−1.
the season can be 40% lower than for 1−year cladodes • Saturation point at PPF = 30 moles m−2 day−1.
(Liguori et al., 2013a).
• At PPF = 22 moles m−2 day−1, CO2 uptake is at 90% of
its maximal.
CAM−related statistics

• O. ficus−indica may assimilate 344–680 mmol m−2

day−1 of CO2 (C = 35%). SOIL SUITABILITY IN THE CULTIVATION
• Maximum instantaneous values of CO2 uptake by OF CACTUS PEAR
cladodes = 18 μmol m−2 s−1.
The species grows in all types of soil. It frequently adapts
• Fruit 2 weeks after bloom may reach a net CO2 up- even to soils limited by continuous hard rock in the first
take of 42 mmol m−2 day−1, i.e. 20% of the cladodes 25 cm, which rest on materials with a calcium carbonate
at the same place and time of the year. content > 40% or which contain < 10% by weight of
• 10–30 stomata mm−2 in CAM plants vs 100–300 fine soil (Nobel, 2002). Generally, O. ficus−indica is very
mm−2 in C3 plants. versatile, but it is adverse to high salinity and to water-

TABLE 1 Functional characteristics and classes of soil capacity for O. ficus−indica cultivation
Functional characteristics Classes
S1 Most suitable S2 Suitable S3 Less suitable N Not suitable
Minimum temperature °C >3 >3 > −3; < 3 < −5
Medium temperature °C 18–23 15–18 10–15 < 10
Annual rainfall mm > 400 > 400 200–400 < 100
Texture sandy to mixed clay–silt clay–silt
Skeleton indifferent indifferent indifferent abundant
Depth indifferent indifferent indifferent indifferent
Carbonates indifferent indifferent indifferent indifferent
Reaction pH KCl 5–8 5–8 5–8 < 5; > 8
Organic matter % < 0.5 < 0.5 > 0.5 > 0.5
Ca available elevated elevated mean insufficient
K available elevated elevated mean insufficient
ECe a
dS m −1
<2 2–4 4–7 >7
Subsuperficial groundwater absent absent absent present
a
ECe = electrical conductivity of the saturate extract of the soil
Ecophysiology and reproductive biology of cultivated cacti 33

logging, since the root system is very sensitive to anoxia under conditions of water stress, an efficient “buffer”
(Table 1). For commercial production of the species, effect in response to different physiological needs
the salt concentration threshold of the soil solution is (Nobel, 2002). Parenchyma and chlorenchyma osmotic
50 mM NaCl (Nobel, 2002). A concentration of 30 mol pressure is little affected after 3 months of drought,
m−3 (1.76 ppt NaCl) reduces growth by 40%, compared whereas turgor pressure is reduced by 86% compared
with the non−saline control, while a concentration of with well−watered conditions (Goldstein et al., 1991).
100 mol m−3 (5.85 ppt NaCl) reduces growth by 93%
Diel fluctuations in cladode thickness may also represent
(Gersani et al., 1993). It tolerates well high air capacities
an early indicator of dehydration stress. Young cladodes
(high textural or structural macroporosity) and a modest
show more pronounced diel thickness fluctuations com-
or meagre organic content. With regard to the edaphic
pared with older cladodes, and therefore serve as a suita-
requirements, ample availability of calcium and potassi-
ble model for assessing plant responses to environmental
um is advantageous for a good crop.
factors. Under well−watered conditions, diel fluctuations
of cladode thickness are directly related to temperature
CO2 UPTAKE AND WATER AVAILABILITY variations, but not under severe drought stress (Scalisi
et al., 2015). In addition, cladode growth is extremely
O. ficus−indica water−use efficiency (WUE) reaches responsive to rehydration after long periods of drought,
values of approximately 3.3–4.0 g dry matter (DM) kg−1 suggesting that a regulated reduction of irrigation would
water (De Kock, 1980; Le Houérou, 2002). In young cla- not significantly affect plant biomass.
dodes, water initially moves through the phloem, while
xylematic water uptake starts about 1 month after their Surprisingly, measurements integrating whole−tree
appearance (Wang et al., 1997). Sink−to−source tran- gas exchange prove that O. ficus−indica trees continue
sition is also accompanied by an inversion of water po- their photosynthetic activity 60 days after irrigation is
tential difference between young and mother cladodes. stopped and when soil water content is < 5% (Figure
Specifically, water potential is initially higher in the very 2). At this stage, current−year and 1−year cladodes
young than in the mother cladodes, and it becomes low- of non−irrigated trees have become flaccid, but daily
er in the young cladodes after 4 weeks of growth (Luo net CO2 uptake of the whole tree still maintains the
and Nobel, 1993; Wang et al., 1997). Cladode succulence same rate as at the beginning of the experiment while,
acts as a buffer to maintain turgor in the photosynthetic during the season, well−watered trees double their
tissue (chlorenchyma), making it possible for the cladode net CO2 uptake. Eventually, the whole−tree daily CO2
uptake, which integrates cladodes of different age and
to continue photosynthesizing during dry periods.
photosynthetic capacity, is well below values measured
Long drought periods have several physiological effects for individual cladodes under similar conditions (wet
on cactus pear. Under optimal conditions (25/15° C or dry) (Nobel and Bobich, 2002; Pimienta Barrios et
day/night) and light saturation, O. ficus−indica may al., 2005) − indication of the different contribution
assimilate 3.44 g m−2 day−1 CO2; Nobel and Hartsock of single cladodes to whole−tree photosynthesis.
(1984) showed that 3 weeks of drought are required The capacity of O. ficus−indica trees to substantially
to halve single cladode net CO2 uptake over 24−hour maintain their photosynthetic activity, even when they
periods; after 50 days of drought, net CO2 uptake of have lost most of the water from the parenchyma, is
cladodes is around zero (Acevedo et al., 1983). Moreo- probably related to water movement (from mother to
ver, 3 months of drought decrease nocturnal acid accu- daughter cladodes) and intertissue water transfer (from
mulation by 73%, essentially abolish transpiration, and parenchyma to chlorenchyma) resulting in reduced wa-
lead to a decrease in the fresh weight (FW) of the pa- ter loss of the photosynthetic tissue (Barcikowski and
renchyma equal to 61%, while the chlorenchyma lose Nobel, 1984; Nobel, 2006). According to Goldstein
only 27% of water (Acevedo et al., 1983; Goldstein et et al. (1991) and Nerd and Nobel (1991), following a
al., 1991). Optimal temperatures for cladode nocturnal 15−week drought period, cladode water content de-
net CO2 uptake are reduced if plants are under drought creases by around 60% and the water−storage paren-
conditions (Nobel and Hartsock, 1984), especially chyma loses a greater fraction of water (65%) than the
when water deficit is prolonged (Nobel, 2001). After chlorenchyma (27%), which most likely helps maintain
prolonged drought, relative water content, cladode nocturnal acid accumulation in the tissue of the chlor-
(parenchyma) thickness and chlorophyll content of enchyma. The chlorenchyma of O. ficus−indica can
single cladodes decline sharply (Pimienta Barrios et al., reversibly lose 70% of its water content at full turgor,
2007). In conditions of water stress, the parenchyma while the water storage parenchyma can lose 82%
is barely distinguishable from the surrounding chlor- (Andrade et al., 2009); indeed, when rainfall occurs fol-
enchyma (Barcikowski and Nobel, 1984; Liguori et al., lowing a long dry period, re−watered cladodes show
2013a). The ability of the parenchyma tissue to store complete recovery in terms of fresh and dry weight
water and to transfer it to chlorenchyma provides, and gas exchange rate pattern (Pimienta Barrios et
 34 Crop ecology, cultivation and uses of cactus pear

Net CO2 uptake

(µ mol m-2 s-1) dry wet
12

10

10 15 20 01 06 11 10 15 20 01 06 11 10 15 20 01 06 11 16
A B C time of day
Daily PPF: 27.3 mol m-2 d-1 Daily PPF: 35.1 mol m-2 d-1 Daily PPF: 39.2 mol m-2 d-1 (hour)
Day/Night temp: 26°/19°C Day/Night temp: 28°/20°C Day/Night temp: 28°/23°C
Soil water content: Soil water content: Soil water content:
· irrigated: 18%; · irrigated: 18%; · irrigated: 19%;
· non irrigated: 6% · non irrigated: 5% · non irrigated: 2%

Measurements made with open chambers covering three single trees for each treatment (dry and wet) (a: June, b: July, c: August). Dotted bar
coincides with the night−time period.

Figure 2 al., 2005). The accumulated C, in the absence of et al., 2002a). Indeed, the plant is able to recy-
Daily net CO2 uptake in fruit, is partitioned in well−watered cladodes to cle internal water and supply water to the fruit
irrigated and non−irri-
gated O. ficus−indica
the parenchyma of 1− and 2−year cladodes and through the phloem (Nobel, 2002). Nerd et al.
potted trees the current−year growth. On the other hand, dry (1989) report that in the Negev Desert, where
matter accumulated during the drought period is the annual rainfall is < 300 mm, winter irrigation
likely to be destined to older stems and the root regulates plant fertility and spring burst timing in
system, which are responsible for plant recovery spring. However, irrigation alone cannot counter-
following a long period of drought (Nobel, 1988). act the effect of very high temperatures, which
The roots of 18−month non−irrigated O. ficus− affect carbon exchange rate more than transpi-
indica trees incorporate 23% more carbon from ration and always result in reduced fruit growth
the basal cladodes than do well−watered ones and advanced maturity, even under irrigation.
(Luo and Nobel, 1993). Shifting the carbon alloca- Consoli et al. (2013) − in areas with no summer
tion pattern in favour of the roots contributes to rains (June−September) and with annual rainfall
drought resistance in perennials and woody trees of 500 mm − measured Kc values of 0.45−0.50.
and facilitates their recovery after long periods of
water stress (Da Costa and Huang, 2009; Faust,
1989). Moreover, the soluble sugar content usu- RESPONSE TO TEMPERATURE
ally decreases with drought, associated with an
equivalent increase in polysaccharides, presuma-
Dry but not too hot: this could be the motto for O.
bly starch, in the water−storage parenchyma, but
ficus−indica. CAM metabolism enables the plant
not in the chlorenchyma (Nerd and Nobel, 1991).
to reach maximum photosynthetic productivity
In areas where no rains (< 30−50 mm) occur dur- with daylight temperatures of 25° C and noctur-
ing the fruit development period or where annual nal temperatures of 15° C. Higher day or lower
rainfall is < 300 mm (Gugliuzza et al., 2002a), O. night temperatures produce a sharp decrease in
ficus−indica requires supplementary irrigation carbon assimilation, leading to poor plant growth,
to obtain adequate yields and good fruit quality reduced production (Nobel, 2002) and eventually
(> 120 g FW) (Barbera, 1984; Gugliuzza et al., low crop value.
2002a; Van der Merwe et al., 1997). Fruit size Temperatures > 30° C cause reductions of up to
is reduced by water deficit long before the tree 70% in photosynthetic activity; they affect fruit
shows any symptoms of water stress (Gugliuzza shape when occurring during the initial stages
Ecophysiology and reproductive biology of cultivated cacti 35

of fruit growth; and they shorten the third stage of to the spring flush removal (SFR) is greatly affected by
fruit growth, when most of the growth of the edible the prevailing temperatures at removal time (Barbera et
flesh occurs, leading to advanced and early ripening, al., 1991; Brutsch and Scott, 1991; Nieddu and Spano,
and reduced fruit size, firmness and sugar content. 1992). High temperatures (> 30/20° C day/night) pro-
High temperatures during fruit development enhance duce more new cladodes than fruits; lower temperature
fruit sensitivity to low temperatures (< 8° C) during (< 20/15° C day/night), on the other hand, may not
post−harvest storage (Inglese et al., 2002b). On the result in rebudding.
other hand, daily temperatures < 15° C slow down fruit
growth, delay fruit ripening time and result in thicker
fruit peel and lower soluble solid content and poor RESPONSE TO LIGHT
peel colour (Inglese, 1995; Liguori et al., 2006; Nerd
et al., 1991a). Temperatures < 0° C, even for 4 hours, Fruits of O. ficus−indica do not generally grow on
produce irreversible damage to the cladode tissue and shaded cladodes (García de Cortázar and Nobel, 1992),
the fruit. High temperatures are a major constraint for probably because in such conditions cladodes do not
the production of high quality fruits in areas with a hot accumulate sufficient dry weight to support reproduc-
and dry summer. As a matter of fact, the optimal daily tive development. Net assimilation rate of cladodes of
temperature for CO2 uptake decreases from 17° C un- O. ficus−indica becomes negative for a total daily pho-
der wet conditions to 14° C after 7 weeks of drought tosynthetically active radiation (PAR) of 5 mol m−2 day−1
(Nobel, 2002). The number of days required to reach and reaches 90% of the maximum at 20 mol m−2 day−1
commercial harvest maturity changes with the time of (Nobel, 1988). Shading affects flower evocation, as was
bloom and with prevailing temperatures during the fruit clearly demonstrated by Cicala et al. (1997) and Deidda
development period, but the thermal time measured in et al. (1992). At least 80% of flowering seems to be
terms of growing degree hours (GDH) from bloom to inhibited if light is withheld the last 2 months before
harvest does not change (40−43 × 103) (Inglese et al., bloom − indication that flower evocation occurs close
1999; Liguori et al., 2006). Nerd and Mizrahi (1995a) to bud sprouting. Sufficient light intensity during the
found that detached cladodes that experienced low winter rest period is therefore essential for a regular
winter temperatures produce the most fruit buds the bloom. Shading applied within 5 days after the removal
following spring. Similar results were reported by Gut- of the spring flush also inhibits the return bloom (Bar-
terman (1995), who examined 18 light/temperature bera et al., 1993a).
combinations of detached cladodes and found that de-
tached cladodes produce significantly more fruit with 8 Shading also affects fruit quality, depending on the ex-
hours of light grown outside in cool temperatures than tent of shading and sucrose exchange within the tree.
with 8 hours of light in a heated greenhouse. Nobel and Shading completely the main source of photo−assimi-
Castañeda (1998) indicate an increase in fruit produc- late for the fruit for 45–75 days after bloom has no in-
tion on detached cladodes held at 15/5° C compared fluence on fruit weight, quality or ripening time. On the
with cladodes left at 25/15° C day/night temperature. other hand, a short period (15 days) of imposed shade
during earlier stages of fruit growth significantly affects
Potgieter and Smith (2006) report a strong environmen-
fruit weight, but not the total soluble solids content or
tal influence on fruit yield, with the highest fruit yield
fruit firmness and ripening time. The fruits do not fully
obtained in areas of South Africa with warm summers
recover to maximum growth, even when the fruiting
and cool winters. They also report a strong genotype
cladode is shaded for a short period before bloom. As
× environment interaction, indicating variation in the
with other fruit trees, such as peach and apple, fruit
plasticity of cultivars in terms of temperature require-
size at harvest seems to be related to the fruit growth
ment for optimal fruit production. However, cactus
rate attained in the early stages of fruit development.
pear produce fruits in the Valley of Catamarca, north-
At this stage, photo−assimilate supply from the fruiting
west Argentina, and in the Canary Islands, where no
cladode becomes crucial to support fruit growth, prob-
more than 100 chilling units accumulate in winter; it is
ably because of the competitive demand of different
also able to reflower several times in the same season
and actively growing vegetative and reproductive sinks
− naturally, in the case of Chile or California, artificially
(Inglese et al., 1999).
induced, in the case of Italy and Israel (Inglese, 1995;
Liguori et al., 2006). These out−of−season blooms ap- This is in line with the fact that thinning is most effec-
pear to have no relation with endodormancy; the rest tive in increasing fruit size when applied no later than
period could result from ecodormancy rather than true 3 weeks after bloom (Inglese et al., 1995a). Since net
rest or endodormancy. In O. ficus−indica, another flush assimilation of heavy shaded cladodes is negligible, fruit
of fruits may take away the first spring flush of flowers growth in cladodes shaded for long periods during
and cladodes (Inglese et al., 2002b). Temperature af- the fruit development period must depend on exten-
fects plant reflowering aptitude. In fact, plant response sive photosynthate translocation from non−shaded
36 Crop ecology, cultivation and uses of cactus pear

branches (Inglese et al., 1994b; Luo and Nobel, 1993). weight (EDW) exceeds the minimum dry weight for a
This mobilization of storage carbohydrates makes fruit particular surface area by at least 33 g. Values of EDW be-
growth only partially dependent on reduced photo- come positive after 60−70 days of cladode development
synthetic photon flux density (PPFD) − wavelengths of and in November they reach 13.1 ± 2.4 g and 10.6 ± 1.2
400−700 nm − at fruit canopy location. Fruit ripening g, respectively, for spring cladodes and cladodes from the
in shaded cladodes occurs later than in sunlit ones. The second flush of the previous season.
pattern of PPFD distribution within the canopy enhanc-
Flower buds emerge during the spring (Pimienta Barri-
es the variability in fruit quality in terms of size, and
os, 1990; Gutterman, 1995; Nerd and Mizrahi, 1995a).
fruits located in the lower part of the canopy are small-
Photoperiod and/or low winter temperature may be the
er than those at the top. Fruit ripening varies within
environmental signal for such bud initiation. In addition,
the plant and the fruiting cladode, mainly because of
nutrients can also affect bud initiation, since exposing
a different time course of flower bud formation, de-
plants to nitrogen fertilization (using ammonium nitrate)
velopment and flowering (Barbera and Inglese, 1993).
can increase the number of buds per plant (Nerd et al.,
The pattern of PPFD distribution within the canopy en-
1993a). Most of the flowers occur on 1−year terminal
hances this variability, since shade delays fruit ripening
cladodes, and new cladodes usually develop on 2−year
according to the length of the shading period.
or even older cladodes (Inglese et al., 1994a). Vegetative
In conclusion, the wide within−tree variability of fruit and reproductive buds appear contemporarily in spring
quality and ripening time that occurs in cactus pear or early summer when the spring flush is removed to
(Barbera and Inglese, 1993) can be partially explained induce reflowering (Barbera et al., 1991). The flowers
in terms of PPFD distribution within the canopy. generally develop from the areolae disposed along the
crown of the cladode, but production of flowers from
REPRODUCTIVE BIOLOGY the areolae on the planar surface exposed to the sun is
not infrequent. A cladode can produce 35−40 flowers;
Fruit production by O. ficus−indica is extremely variable. during abundant flowering in conditions of high poten-
tial evapotranspiration, each can lose ≤ 3 g of water
Yields of 20−30 tonnes ha−1 are reported in Israel and
day−1, equivalent to 15% of its weight at the moment of
Italy (Barbera and Inglese, 1993; Nerd and Mizrahi, 1993)
the anthesis (De la Barrera and Nobel, 2004).
and 10−30 tonnes ha−1 in South Africa (Wessels, 1988a;
Brutsch and Zimmerman, 1993). The wide variability in
yield depends on orchard design (plant spacing), cultural
Floral biology
practices, environmental conditions and cultivar fertility.
Flower buds develop quickly, starting with a meristem
Plants begin to yield 2−3 years after planting; they reach
covered by polycellular trichomes above the petiolar
their maximum potential 6−8 years after planting and
scar (Nieddu and Spano, 1992). When the flower bud
remain productive for 25−30 years and even longer,
reaches a length of 0.5 cm (Wessels and Swart, 1990),
depending on pruning and overall orchard management.
the gyneaceum and the androecium become visible
inside. The ovary, which is inferior and unilocular, is
Cladode fertility
multicarpellate (6–12 carpels, usually 8 growing togeth-
er at the base and separating at the stigmatic region)
For a mature plant, most (80–90%) 1−year cladodes bear
(Nieddu and Spano, 1992). The stamens differentiate
fruits accounting for 90% of the annual yield. However,
two techae with two pollen sacs each; another dehis-
they show a wide fertility range, depending on plant age,
cence occurs before anthesis (Nieddu and Spano, 1992).
environmental conditions and their state of growth. The
Pollen is presented in a spherical, apolar radio−symmet-
average number of flowers produced by a single, 1−year
rical form with a reticulated and poly−panto−porated
cladode is 6−9, with peaks of 30−35. Two−year clado-
surface, with a circular polygonal perimeter (6–8 sides)
des are generally responsible for vegetative renewal and
(El Behi et al., 2015).
a small proportion (10−30%) produce fruits, but with
reduced fertility compared with 1−year cladodes. The Aguilar and Chavez (1995) reported that the hormone
number of fruiting cladodes occurring on a plant every gibberellic acid (GA) promotes floral bud development
year depends on plant spacing, and ranges from 100–120 and indole−butyric acid (IBA) stimulates vegetative
for 350−400 plants ha−1 to 20−30 for 1 000−1 200 sprouting. In subtropical regions, Opuntia floral buds
plants ha−1. A mature cactus pear tree O. ficus−indica (L.) start when the mean monthly temperature exceeds
Mill. produces new fruits and cladodes at a ratio of 4 : 1 16°C − in March or April in the Northern Hemisphere
every year (Barbera and Inglese, 1993). Cladode fertility and in September or October in the Southern Hem-
is related to the amount of dry weight accumulated per isphere (Nerd and Mizrahi, 1995). The floral bud
unit surface area (García de Cortázar and Nobel, 1992). production period fluctuates between 3 and 5 weeks
Cladodes become productive when their estimated dry (Reyes Agüero et al., 2006). Floral development from
Ecophysiology and reproductive biology of cultivated cacti 37

bud to anthesis requires 21−47 days, but can last as among cladodes of different ages. Young developing cla-
long as 75 days (Wessels and Swart, 1990). Generally, dodes apparently compete with fruits − as indicated by
in the Northern Hemisphere, anthesis takes place be- their higher absolute growth rate (AGR). However, they
tween May and July, and in the Southern Hemisphere become a source of carbohydrates at an early stage of
between February and August (Reyes Agüero et al., their development (Luo and Nobel, 1993), a time which
2006). Flowers are mostly hermaphrodite. Autogamic coincides with the development of flowers or the earliest
and xenogamic pollination both occur in O. ficus−indica stage of FDP. Relative sink strength changes in line with
(Nerd and Mizrahi, 1995a). Different bee species visit the developmental stages of the seasonal growth of fruit
Opuntia flowers during pollination. and cladodes (Inglese et al., 1999). Fruits become the
major sink during Stage III − as indicated by the sharp
Apomixis occurs frequently in Opuntia (Pimienta Barrios,
reduction in cladode AGR at that stage.
1990; Mondragón and Pimienta Barrios, 1995). Gil and
Espinoza (1980) obtained normal−sized parthenocarpic Competition between fruit and cladode growth, as
fruits containing empty seeds from fertilized flowers well as the reduction in the number of new cladodes
treated by gibberellin and auxin in pre−anthesis, but for following SFR can be sources of plant alternate bearing
some clones of Opuntia, the occurrence of pollination behaviour that also changes with genotype (Inglese et
is not necessary to obtain seedless cactus pear fruits. In al., 2002b). Alternate bearing of cactus pear depends
fact, Weiss et al. (1993b) reported vegetative partheno- on the reduction in the fertile cladodes following scarce
carpy in some Opuntia ficus−indica (L.) clones. vegetative activity the previous year. The number of
flowers per fertile cladode becomes increasingly stable
El Behi et al. (2015), in a study of a seedless cultivar
each year; it depends on the cladode age and is highest
‘Bianca’ from Sicilian germplasm, hypothesized that
in 1−year cladodes. Fruit size varies between trees and
its seedlessness could be based on a partial stenosper-
depends on factors such as plant architecture and crop
mocarpy, as reported for grape and Citrus (Vardi et al.,
yield per plant and per cladode.
2008). Opuntia fruits are unilocular and polyspermic. In
O. ficus−indica, the unripened green peel has stomatal
Out−of−season crop
and photosynthetic activity, contributing 8−10% to
fruit growth (Inglese et al., 1994b). Fruit areoles always
The most powerful tool for achieving a longer fruit
have glochids and may have spines and bristles or hairs,
marketing season is the cladodes’ ability to reflower
which can persist from the flower stage (Bravo, 1978).
several times within the same season, either naturally
Seed growth and ripening occur 30−70 days after an- or after applying inductive practices (Nerd and Mizrahi,
thesis; seeds are small and ovoid or lens−shaped (Bravo, 1997). Double flowering occurs naturally in the Salinas
1978). Italian O. ficus−indica cultivars have an average of area in California, where fruits are picked from Septem-
273 seeds per fruit, of which 146 are normal and 127 ster- ber to March (Inglese, 1995), and in the central region
ile (Barbera et al., 1991); Israeli cultivars have an average of in Chile, where the fruit harvest lasts from February to
268 normal seeds per fruit (140−430) (Nerd and Mizrahi, April and from July to September (Sudzuki Hills et al.,
1995a) and Mexican cultivars an average of 203 normal 1993). In Italy, a second flowering is obtained as a result
seeds per fruit (10−448) (Reyes Agüero et al., 2006). of the complete removal of the spring flush of flowers
and cladodes. SFR takes place when the main bloom
Fruit growth occurs, between the end of May and the last week in
June in the Northern Hemisphere, and in October in the
Fruit growth follows a double sigmoid pattern in terms Southern Hemisphere (Inglese, 1995). New flower buds
of fresh weight; there is a pronounced gain in dry develop on the fertile cladodes of the natural flush, and
weight for the peel during Stage I, for the seeds during the reflowering index − defined by the ratio of second
Stage II and for the core during Stage III of the fruit versus first flush flowers − is highest for cladodes with a
development period (FDP) (Barbera et al., 1992b; Nerd natural fertility of 5−10 flowers, sharply decreasing with
and Mizrahi, 1997). The growth of fruits and daughter each additional flower that develops during the first
cladodes implies a substantial translocation of stored flower flush. Removal time affects cladode reflowering
carbohydrates from the basal cladodes (Luo and Nobel, rate (Barbera et al., 1991; Brutsch and Scott, 1991). For
1993; Inglese et al., 1994b). In fact, when more than instance, removing flowers at pre−bloom stage results
five fruits develop on a 1−year fruiting cladode, an ex- in the highest reflowering rate, while removing the
tensive import of assimilates occurs, particularly during spring flush after petal shedding reduces reflowering
Stage III of fruit growth (Inglese et al., 1994b). by up to 50−70% (Barbera et al., 1991; Inglese et al.,
1998). This decrease in reflowering that occurs when
Inglese et al. (1994b) and Luo and Nobel (1993) inves-
flowers are removed at full bloom or after petal shed-
tigated the source−sink relationship on mature fruiting
ding is related to an inhibitory effect on flower bud
plants and found a massive carbon flux of assimilates
initiation (Barbera et al., 1993a) of GA3 diffused from
 38 Crop ecology, cultivation and uses of cactus pear

the flowers to the mother cladode (Inglese et al., cember to March depending on covering time and
1998). Indeed, Barbera et al. (1993a) demonstrat- environmental conditions. The length of the fruit
ed that GA3 applied within 6 days after SFR totally development period increases from 100−120 days
inhibits reflowering − indication that the flower to 160−190 days for the out−of−season winter
bud, induced the preceding spring, is still revers- crop, depending on a longer third stage of fruit
ible at that stage. Current−season developing cla- growth which occurs when temperatures are un-
dodes also inhibit reflowering if they are not fully der the optimal values for fruit growth (Figure 3).
removed (Inglese et al., 1994a). Pre−blooming However, winter temperatures inhibit fruit growth
fruits have the shortest fruit development period, and ripening; it is therefore necessary to cover the
and ripen 15−20 days earlier than full−bloom and plants with PVC tunnels to create appropriate con-
30−40 days earlier than post−bloom fruits. ditions for regular fruit development. Winter fruit
production occurs in the field in Israel (Nerd et al.,
The reflowering rate depends also on the pre-
1993a). In this case, the second crop follows the
vailing environmental conditions at removal time
main one and occurs on current−year cladodes
(mainly soil water content and air temperature).
developed since May. After the harvest of the first
Indeed, the extent of reflowering greatly differs
from year to year and depends on orchard loca- crop, a second bloom is induced through extensive
tion (Barbera et al., 1991; Brutsch and Scott, 1991; N fertilization and irrigation. The second bloom
Nieddu and Spano, 1992). High temperatures occurs in September−October and fruit develop
during bud initiation induce vegetative rather than from January to March. Fruit have a regular size
reproductive activity, resulting in a lower reflower- but a reduced percentage of flesh (50−55%).
ing rate. Liguori et al. (2006) developed a strategy
based on the potential of the plant to reflower BIOMASS PRODUCTIVITY
even after a double spring flush removal: the first
in early June with the removal of the spring flush High planting densities lead to an extremely high
and the second in late June with the removal of accumulation of dry matter in the vegetative
the second, induced flush. The rate of this second growth, but deeply affect the allocation of re-
reflowering (third flush) is very uneven, ranging sources to the fruit (García de Cortázar and Nobel,
from 25 to 40%. The plant blooms during the first 1992). O. ficus−indica can have an annual dry
10 days of August and the fruit ripens from De- matter productivity that exceeds that of nearly all

Diameter
(mm)
Figure 3 55
Cumulative growth
curve for fruits of O.
ficus−indica cv. ‘Gialla’
coming from the 2nd 50
and 3rd flush of bloom

45

40

35

30
second flush fruits third flush fruits
25
0 20 40 60 80 100 120 140 160 180
Days after bloom
Ecophysiology and reproductive biology of cultivated cacti 39

cultivated C3 and C4 species. For instance, a productivity interception by mother cladodes (Inglese et al., 1999).
of 47 tonnes dry weight (DW) ha –1 year–1 of cladodes For instance, the photosynthetically active canopy com-
(99%) and fruit (1%) has been predicted for high ponents of a commercial orchard for fruit production
density plantings (24 plants m –2), while a productivity with 420 trees ha−1 (Liguori et al., 2013a) account for
of nearly 15 tonnes ha –1 was measured for lower spac- 60% of canopy surface, with SAI−orchard = 2.6 and
ings (0.24 plants m –2), resulting in a lower vegetative SAI−tree = 4.0 − i.e. values well below those reported
vs reproductive growth, with early fruit cropping and by Nobel (1988). In these conditions, the HI was about
an alternate bearing behaviour (García de Cortázar and 29%, considering both first and second flush, or 26%
Nobel, 1992). Acevedo et al. (1983) measured a clado- if only the reproductive and vegetative growth of the
de dry mass production of 1 kg m –2 ground area and second flush are considered. Solar radiation (40%) is
0.3 kg fruit m –2 year–1. Recently Pinos Rodríguez et al. intercepted by the less photosynthetically efficient cla-
(2010) reported a dry mass production of 1.39 kg m –2 dodes and by lignified scaffold. Carbon stock among
for O. ficus−indica grown to produce fresh biomass for canopy and root component shows that a mature plant
cattle use. Inglese et al. (2012) − in commercial plan- can have 9−10 fruits m−2 of cladode surface area (cur-
tations for fruit production with a much lower plant rent−year to 2−year cladodes), which means 1.1−1.4 kg
density (333 trees ha –1) − found very similar values to fruits m−2. A commercial orchard for fruit production can
those found by Nobel (1988), but for a much lower accumulate 7.5 tonnes DM ha−1 year−1, which means 3.4
total biomass and a commercial fruit weight (1.2 kg m –2 tonnes C ha−1 stored in the canopy components (includ-
ground area, considering the first and second flush of ing fruits). These values are similar to those reported for
fruits and cladodes). By the end of the first season of deciduous orchards (e.g. peach) and for evergreens (e.g.
growth, current−year cladodes almost cover their final orange) (Tagliavini et al., 2008).
surface area, but they only reach 65% of the dry weight
of 1−year cladodes. During the second year of growth,
they show a marginal increase in terms of surface area, POTENTIAL RESPONSES TO CLIMATE
but a significant accumulation of dry weight (+30%). CHANGE
Thereafter, cladodes develop only marginally in terms
of surface area, but continue to accumulate dry weight. Agricultural, vehicular and industrial activities that rely
The high specific dry weight of scaffold cladodes is on fossil fuels release greenhouse gases, mainly CO2.
twice that of 1−year cladodes. The absolute growth Indeed, in just 150 years, the atmospheric concentra-
rate for cladodes and fruits can reach, respectively, 0.12 tion of carbon dioxide has rapidly increased from the
day–1 and 0.16 day–1. pre−industrial level of 280 ppm to the current level of
400 ppm (IPCC, 2014). In turn, increased CO2 leads to
The fruit component (harvest index [HI]) has not been
higher photosynthetic rates as a result of a steeper con-
systematically studied in O. ficus−indica. García de
centration gradient between the atmosphere and the
Cortázar and Nobel (1992) showed that yearly variations
sites of carboxylation in plant tissues. Also, given that
in cladode vs fruit dry matter allocation also depend on
higher partial pressures lead to a decreased carbon “de-
planting densities. An increase in dry matter allocated
mand” by leaves, a reduction in stomatal conductance
to the fruit was accompanied by a reduction in cladode
is observed for plants exposed to elevated CO2 (Smith
count and dry weight accumulation.
et al., 2009). Despite having an inherent mechanism
Light interception, CO2 uptake and plant productivity for concentrating CO2, which could make CAM plants
depend on canopy architecture and stem area index insensitive to higher atmospheric concentrations of this
(SAI) per ground area (taking account of both sides of gas, an actual fertilization effect has been observed.
the cladode for SAI) − SAI is the equivalent of the leaf This is the case of O. ficus−indica exposed to doubled
area index (LAI) (Nobel, 1988). Despite their importance, CO2, either in controlled environmental chambers or in
SAI values for O. ficus−indica orchards have been poorly rooted plants kept in open−top chambers. In particu-
investigated. SAI values of 6.5−8.5 have been measured, lar, plants accumulate 20−40% more dry mass under
considering all the cladodes in the canopy and the sur- doubled CO2 than under 350 ppm (Nobel and García
face area covered by the tree canopy. Maximum produc- de Cortázar, 1991; Nobel and Israel, 1994). In addition,
tivity (2 kg DM m–2 ground area year–1) has been predict- plant development is enhanced by carbon fertilization.
ed for SAI = 4−6; while for SAI < 3, with 20 000 and 6 For instance, a doubling of the production of second
000 plants ha –1, the total net CO2 uptake is about linear order daughter cladodes is observed after 3 months of
with SAI (Nobel, 1988). These calculations include all exposing mature cladodes to elevated CO2 (Nobel and
cladodes; they do not take into account that the single Israel, 1994). In addition, root elongation can increase
cladode contribution to CO2 uptake changes with cla- up to 25% under elevated CO2 given optimal air tem-
dode age (Nobel, 1988), nor that optimal SAI definition peratures averaging 30ºC (Drennan and Nobel, 1998).
must include fruit quality, which also depends on PAR These fertilization effects may be driven by anatomical
40 Crop ecology, cultivation and uses of cactus pear

TABLE 2 RCPs considered in global climate change models

Radiative forcing
Equivalent previous Greenhouse gas Year 2100 Temperature
RCP relative to 1750
IPCC scenario emissions (CO2−eq.) (ppm) increase (ºC)
(W m−2)
RCP4.5 B1 Stringent mitigation 430−480 2.6 < 1.5
RCP6.0 B2 Intermediate 580−720 4.6 2−3
RCP2.6 n/a Intermediate 720−1 000 6.0 3−4
RCP8.5 A2/A1FI Very large > 1 000 8.5 >4
Source: IPCC (2014).

responses of O. ficus−indica, such as a chlorenchyma • RCP8.5 is the most extreme scenario, which assumes
that is 31% thicker under doubled CO2 than under that despite reduction targets and international agree-
ambient concentrations, despite a decrease in stomatal ments, the world will still heavily rely on fossil fuels.
frequency of 20% and a 30% thickening of the cuticle
that has more epicuticular wax (North et al., 1995). This The extent of environmental change occurring during
fertilization effect resulting from exposure to elevated this century depends on compliance with the targets
CO2 has also been observed for other cacti (e.g. the of reduction of greenhouse gas emissions agreed to by
hemiepiphytic Hylocereus undatus). For this tropical most countries in 2015, including the two main contribu-
crop, cultivated in over 20 countries, the photosynthetic tors (the United States of America and China).
rates increase by 34% under doubled CO2 (Raveh et al.,
1995; Nobel and De la Barrera, 2004). However, high This imminent climate change is likely to be detrimental
CO2 concentrations can also lead to lower respiration to global agricultural productivity (Monterroso Rivas et
rates, decreased maximum activity of cytochrome c al., 2011; Lobell and Gourdji, 2012; IPCC, 2014; Ovalle
oxidase, and lower mitochondrial numbers for O. ficus− Rivera et al., 2015). However, for certain cultivated
indica individuals (Gomez Casanovas et al., 2007). Thus, CAM plants, the new environmental conditions may
while the increasing concentrations of CO2 can enhance result in an increase in the area suitable for their cul-
plant productivity, this might be true up to certain levels tivation (García Moya et al., 2010; Lobell and Goudji,
when plant metabolism will be reduced. 2012). Naturally, a major adaptation of CAM plants to a
warmer and often drier environment is the fact that gas
In addition to the direct effects on plant physiology
exchange occurs at night when air temperature is low-
and anatomy, elevated greenhouse gas concentrations
er. In addition, CAM plant productivity is predominantly
are likely to impact plant productivity by substantially
driven by night−time air temperature (when gas ex-
changing the planet’s climate. In this respect, four
change occurs), rather than by the diurnal temperature
greenhouse gas emission scenarios are being consid-
(when the stomata are closed) (Andrade et al., 2007). In
ered by the Intergovernmental Panel on Climate Change
(ICPP) that could lead to different climate outcomes most cases, the average nocturnal temperature leading
during this century (Table 2). These so−called Repre- to maximum CO2 uptake by CAM plants ranges from 10
sentative Concentration Pathways (RCPs) consider to 20ºC (Nobel, 1988). Thus, as long as the nocturnal
scenarios leading to a given increase in radiative forcing air temperature is appropriate, substantial CO2 uptake
relative to that from 1750 and differ from the previous can occur even under increasing diurnal temperatures.
emission scenarios (A1, B1, A2, B2) in that they explicitly Indeed, given the expected increase in potential pro-
consider international environmental and development ductivity of various species of Agave in the tropical and
policies (IPCC, 2014). subtropical regions of the Western Hemisphere, their
cultivation has been proposed for carbon sequestra-
• RCP2.6 is the most optimistic scenario. With no pre-
tion, mitigation of soil erosion, and the production of
vious equivalent considered by the ICPP, RCP2.6 relies
biomass for use in biofuels (García Moya et al., 2010).
on stringent mitigation policies that would result in
a net removal of greenhouse gases from the atmos- Higher air temperatures could also be similarly advanta-
phere and could lead to a global temperature of ≤ 2 geous for cultivated cacti. For instance, while the mean
ºC higher than the pre−industrial climate. nocturnal air temperature resulting in maximum gas
• RCP4.5 and RCP6.0 are intermediate and plausible exchange by O. ficus−indica is about 12ºC, this plant
emissions scenarios. Their previous equivalent IPCC is able to adequately perform under a wide range of
scenarios are those of the B family, which consider conditions; indeed, at least 80% of the maximum CO2
the adoption of green technology with a consequent uptake can be achieved between 6 and 20ºC (Figure 4;
reduction in reliance on fossil fuels, while differing in Nobel, 1988; Nobel et al., 2002). The case of Hylocere-
the degree of international cooperation. us undatus is slightly different. For this tropical crop, a
Ecophysiology and reproductive biology of cultivated cacti 41

Daily net CO2 uptake per unit stem area

(fraction of maximum)
Opuntia ficus-indica Hylocereus undatus Figure 4
Responses of total
daily net CO2 uptake
by the cactus crops
1.0 Opuntia ficus−indica
and Hylocereus undatus
to mean night−time air
temperatures
0.8 Used with permission of
the California Botanical
Society from Nobel et
al. (2002).
0.6

0.4

0.2

0.0

-0.2
0 5 10 15 20 25 30 35
Mean night time air temperature (°C)

higher air temperature will lead to more potential 2009). O. ficus−indica can usually survive exposure
area for cultivation, as the optimum temperature to temperatures up to 66º C, while H. undatus can
for CO2 uptake is approximately 20ºC (Figure 4; only resist up to 55º C − the lowest high−temper-
Nobel et al., 2002). However, for this plant, the ature tolerance measured for a cactus (Drennan,
range of nocturnal air temperatures leading to 2009; Nobel et al., 2002). For many species, the
high gas exchange rates is rather narrow, especial- actual temperature tolerance is determined by the
ly compared with O. ficus−indica, as mean night− prevailing mean air temperature. Indeed, a 10º C
time air temperatures between 14 and 23ºC are increase in the air temperature to which plants are
required to achieve at least 80% of maximum CO2 exposed leads to an average increase of 5.3º C
uptake (Figure 4; Nobel et al., 2002). for the lethal temperature of 18 species of cacti
(Nobel, 1988; Drennan, 2009). For example, for O.
Mean changes in air temperature will determine ficus−indica individuals incubated under a mean
the average productivity of crops, including cactus temperature of 35º C, the lethal temperature is
crops such as O. ficus−indica and H. undatus. How- 62º C, but for plants incubated under 45º C, the
ever, the rate of warming and the severity of ex- lethal temperature is 66 ºC; thus acclimation for
treme weather events − i.e. infrequent heat spells, this species amounts to 4.2 ºC per 10º C increase
frosts, droughts or unusually heavy rains, which in air temperature (Nobel, 1988; Nobel et al.,
are likely to become more frequent and severe 2002). In contrast, the acclimation ability of H. un-
owing to climate change − will determine actual datus is the lowest measured for a cactus. For this
plant survival and limit crop cultivation (IPCC, 2014; neotropical vine, increasing the average air tem-
Nobel, 1988). Whether a plant species will be able perature from 20 to 30º C merely improves the
to fare increasingly frequent and severe episodes high−temperature tolerance from 54.0 to 55.4º
of high temperature depends on its temperature C; in addition, higher average temperatures lead
tolerance. For 18 species of cacti, the lethal high to an inhibition of gas exchange and even plant
temperature is about 57º C (Nobel, 1988; Drennan, death (Nobel et al., 2002).
 05

Nopal (Opuntia spp.)

genetic resources
Candelario Mondragón Jacoboa and Innocenza Chessab
a
Autonomous University of Querétaro, Juriquilla Querétaro, Mexico
b
University of Sassari, Italy
05 Nopal (Opuntia spp.)
genetic resources

INTRODUCTION costs of running a germplasm collection. It is then imper-

ative to research bibliographical sources to understand
World population predictions for 2050 are grim, the origins and dispersal routes, in order to verify the
estimated to reach 9 billion at the current birth rate. need for closer study before embarking on collection.
Countries in arid and semi−arid areas have some of the It is well documented that Opuntia originated and was
fastest−growing populations; they are also the driest domesticated in central Mexico, a country that still has
regions of the world, and water will be twice as scarce the widest range of wild and domesticated species and
by 2050 (UN−DESA, 2015). Climate change will also varieties. It is also known that during the colonial period,
affect the acreage of land under semi−arid conditions. domesticated cactus pear was introduced to Europe and
Populations are expanding fastest in regions where it then to North Africa, mainly to countries occupied by
is most difficult to grow food. For this reason, govern- Spain, France and Italy. Other historical and economic
ments must give immediate and sustained attention to events explain the presence of cactus pear in the Near
the effects of climate change, whether their countries East and central Asia and other parts of the world during
are already affected or potentially affected. the sixteenth to nineteenth centuries. Since the 1980s,
Effective water and land use, wise selection of cultivated there have been significant efforts by governments and
species matching agroclimatic requirements to specific non−governmental organizations (NGOs) to curtail the
sites, sound strategies to increase yields, and improved effects of desertification in North Africa and other areas;
plant and animal genotypes are among the tools for cactus pear cultivation is among the measures promoted
coping with this catastrophe. Multipurpose indigenous to control erosion and boost vegetation recovery.
plants such as cactus pear (Opuntia spp.) may be includ-
ed in the list of choice crops, given their drought toler- Germplasm variability
ance and wide range of potential uses: as fruit, forage
or vegetable crop, in addition to their utilization as living There is a direct connection between the above−men-
fences to protect family households in dry regions, or for tioned historical events and germplasm variability: the
erosion control (Mondragón Jacobo and Chessa, 2013). earliest introductions were based on a restricted ger-
Germplasm resources are destined to be the corner- mplasm base, including mostly domesticated accessions,
stone of all future applications of cactus pear; intensive while recent introductions used an even more limited
exploration, effective in situ and ex situ conservation, dy- number of entries, considering only a few spineless va-
namic evaluation on new sites and vigorous projects of rieties. Despite evidence of excellent adaptation in areas
germplasm exchange and improvement are needed to such as eastern Africa, cross−breeding of close geno-
realize the full potential of this valuable resource. Scien- types does not yield new alleles immediately.
tists must identify the traits for use in intensive breeding It is difficult to generate new variability in Opuntia ger-
programmes: old and new traits useful for coping with mplasm. This is due to the high rate of apomictic seedlings
drought, frost, and indigenous and foreign pests and and the length of the juvenile period (reported to be 4−7
diseases, and for improving nutritional content. years − Mondragón Jacobo, 2001); these biological fea-
This chapter summarizes the progress made in activities tures of cactus pear reduce the chances of cross−breeding
related to germplasm in those countries with the widest to generate new genotypes through recombination. On
variability of cactus pear, whether native or introduced. the other hand, all domesticated genotypes and most
It also describes the basics of Opuntia genetic resources wild genotypes are polyploid, a genetic feature that di-
exploration, documentation, in situ and ex situ conser- minishes the likelihood of expression of natural mutations.
vation, and utilization. The combination of these traits significantly reduces the
potential for generation of new variability on sites other
than the centre of origin and domestication.
GENETIC RESOURCES EXPLORATION
Exploration of germplasm
Before investing time and resources in the collection of
cactus germplasm, it is necessary to put into perspective For successful exploration, it is important to consider
the prospect of finding real, useful genetic variability, the the following:
potential uses and − most important − the long−term • Sources of local knowledge should be sought regard-
Nopal (Opuntia spp.) genetic resources 45

ing the plant, local names, forms of utilization, inte- There have been extensive efforts in Mexico to collect
gration of the plant into the local culture and history. domesticated and semi−domesticated cactus pears
It is always advisable to invite local guides to assist on with the support of a federal government initiative fo-
field trips, not only extension agents, but also herders, cused on native crops: National System of Plant Genetic
wood collectors, and fruit and herb gatherers, who Resources for Food and Agriculture (SINAREFI). A more
are usually better acquainted with the local flora. modest project was the collection of wild ancestors
• Knowledge of the phenology of cactus in the area of undertaken by the National Autonomous University of
interest is crucial, since all descriptors are based on Mexico (UNAM); this collection can be visited at UNAM
in Mexico City. Germplasm prospection reached a pla-
visual traits concerning all parts of the plant; howev-
teau and research teams then turned their attention to
er, roots, flowers and fruits are essential for accurate
conservation and utilization.
botanical identification. A common trait used to
identify accessions and varieties is fruit colour, both What is left to collect? The Tigray region of eastern
internal and external. However, fruit colour is a dy- Africa certainly deserves a closer look; it covers part
namic trait − changing with the ripening phase, while of Ethiopia and Eritrea and boasts the largest area of
the colour of the pulp changes at ripening − and for “naturalized” cactus pear in Africa, originated from
this reason the exploration should be guided by an introductions made during the nineteenth century;
experienced leader who is less likely to be deceived. efforts to characterize the local variability are reported
• An accurate geographical description of the area is by Mondragón Jacobo and Tegegne (2006). Cactus
pear spread to distant regions during its early dispersal,
indispensable: altitude, sunlight exposure and soil
reaching such unlikely places as China, India and Mad-
moisture all affect climate, and climate in turn can
agascar, and it is necessary to explore and characterize
affect the ripening season. Lack of geographical
the variability present in all locations, in order to com-
knowledge is probably one of the main reasons
pare with the genotypes available in germplasm banks
for the presence of redundant accessions in ex situ
and to evaluate the conservation needs and assess the
collections. Accessions of early or late ripening, but
presence of valuable traits for breeding.
collected in different locations, will flower and bear
fruit at the same time when planted side by side. Another case in point could be the recovery of the
• The presence or absence of spines is another interest- Burbank collection, acquired by South Africa at the
ing and important trait. Modern production systems beginning of the twentieth century. The collection
contains most of the spineless varieties developed by
demand mostly spineless cultivars; however, there is
Luther Burbank in California, as well as some acces-
less variability in spineless cultivars compared with
sions of Mexican origin, and is currently under the
spiny, which probably arose as a mutation of spiny
responsibility of the University of Free State, which has
cultivars. Furthermore, the identification of spineless
efficiently linked genetic resources studies to animal
accessions is more difficult, since the process cannot
use and human nutrition.
be completed unless the plants have fruits.
The prospection of cactus genetic resources has been a Maintenance of germplasm
topic of interest since the beginning of the twenty−first
century. Germplasm studies have been conducted in Maintenance of a cactus pear collection is expensive,
Morocco (Arba et al., 2002), Turkey (Bekir, 2006), Tunisia considering that cactus pear is a perennial crop requiring
(Zoghlami et al., 2007), Argentina and Ethiopia (Haile a long time to stabilize productivity: 5−7 years in the
and Belay, 2002; Mondragón Jacobo and Tegegne, case of fruit production. Several years are also required
2006), allowing the identification of accessions for local to obtain meaningful results with regard to forage
consumption mostly as fresh fruit. These studies have fo- and vegetable utilization; nevertheless, useful data on
cused on domesticated and semi−domesticated stocks. productivity and quality are already available after the
Historical sources indicate a common origin for the cac- second year.
tus pears present in these areas; domesticated cactus Another important factor is the agroclimatic variabil-
pears brought from Mexico after the discovery of Amer- ity of production areas: for a thorough investigation,
ica were dispersed by the colonial activity of Italy, Spain more than one experimental site is required, increasing
and other European countries, and by Arab countries in substantially the cost of research or extension−research
North Africa and the Near East. Field observations sup- projects, and funds may not be readily available, de-
port the hypothesis that the original pool was restricted pending on the country providing financial support.
mostly to domesticated accessions, which segregated There is a global trend of long−term conservation
and adapted to these new environments. For this rea- projects per se becoming difficult to sell. Maintenance
son, although reported to be ample, variability is actually may be facilitated through a combination of goals and
relatively narrow compared with the known pool. collaboration between interdisciplinary teams, using
46 Crop ecology, cultivation and uses of cactus pear

the same trials or collections for simultaneous purpos- documentation for existing accessions and new entries
es; nevertheless, uncertainty remains, as each donor is necessary.
agency sets its own terms and operations calendar. The information associated with collected accessions is
as important as the actual plant material held in gene
banks (Given, 1994). For every accession, there should
GERMPLASM BANKS, CORE be descriptive information to optimize utilization, meet
COLLECTIONS AND CULTIVAR TRIALS specific needs of the users, and support decisions con-
cerning the choice of genotype.
It is advisable for all cactus−pear−producing countries As with major crops, characterization and evaluation
to acquire, collect, conserve and evaluate genotypes of the available gene pool is essential for cactus pear
that are different from those in commercial use. Such breeding programmes. Phenotypic and genotypic
collections serve as exploratory sites and as reservoirs of characterization of germplasm provides opportunities
new variability if needed. They are also a valuable source to identify genetically diverse, agronomically superior
of information about the performance of individual ac- accessions for the improvement of Opuntia species as a
cessions on specific sites, since adaptation and produc- multiple−uses crop (Chessa, 2010).
tivity are difficult traits to extrapolate.
Simplicity, speed and relatively low cost make mor-
The extent and nature of these collections vary: from phological markers/traits the most widely used genetic
cultivar trials to formal germplasm banks. Germplasm markers for germplasm characterization. To facilitate
banks are not only recommended, they are mandatory and standardize characterization of collected accessions,
in countries which are the centre of origin and secondary a descriptors list for cactus pear is available, compiled
centres. Core collections are smaller in size and contain (Chessa and Nieddu, 1997) according to the internation-
specific genotypes useful for breeding and preliminary al format currently endorsed by Bioversity International.
agronomical evaluation. Cultivar trials should include Characterization based on morphology has also been
promising new genotypes − whether strictly new or achieved, following descriptor lists and catalogues pro-
simply new to the area − and usually the design imposes posed to improve utilization of accessions of Mexican
the need for replicates. The extent to which germplasm and South African collections (Gallegos Vázquez et al.,
evaluation is useful depends on several factors: 2005; Potgieter and Mashope, 2009). A classification
• Crop − how important is it? scheme for the identification of cultivated and wild vari-
ants of Opuntia species of Mexican origin was reported
• Site − how well does it represent the agroclimate?
by Reyes Agüero et al. (2009). A statistical approach to
• Extension services − how effective are the services identify the characters of differential diagnostic value
linked to cactus pear production? has been considered to provide greater standardization
between collections and to adapt the descriptor list to
specific purposes (Erre and Chessa, 2013).
GENETIC RESOURCES DOCUMENTATION
The identification and description of domesticated
individuals present in low numbers in family orchards
The genetic resources maintained in a collection
for domestic use has been undertaken by Mexican re-
increase in value if complete and accurate data are
searchers. This particular subset is endangered by unu-
assembled. In addition to adequate passport data, re-
sually harsh frosts and hail − probably linked to climate
cords of accession performance through the years may
change − land−use changes, migration, and other so-
represent a valuable source of information for research-
cial and economic factors. These landraces express the
ers and extensionists. Management and maintenance
effects of long−term selection; they are also the carriers
records must be kept in order to:
of valuable traits useful for immediate application by
• monitor the status of conserved germplasm across all breeders. Assembling them in breeding collections may
phases of conservation; save valuable time in the process of genetic enhance-
• optimize the efficiency of conservation programmes; ment. A description of outstanding local and regional
and “varieties” as well as some restricted populations ob-
• facilitate the exchange of materials and information tained from central Mexico is provided by Gallegos and
among germplasm banks and collections. Mondragón Jacobo (2011). Descriptions include data,
pictures, distribution maps and brief information on the
The main concern about genetic resources manage- commercial traits of these “varieties”. The data are val-
ment is the increase in the number of accessions usu- uable and of interest not only to researchers but also to
ally associated with redundancy. To discard duplicates, farmers, merchants and consumers, on the assumption
and to prevent or reduce the addition of duplicative that the wider the audience, the greater the effect on
accessions to germplasm collections, comprehensive conservation interest.
Nopal (Opuntia spp.) genetic resources 47

A common platform, based on the same traits, scales RAPDs combined with morphological and physiological
and units of measurement is needed on a global basis data. The germplasm bank collection of the Faculty
to facilitate exchange of information and plant material. of Agronomy of the Autonomous University of Nuevo
The Germplasm Resources Information Network (GRIN) León (FAUANL) was characterized and duplicates were
− a Web server (http://www.ars−grin.gov/) providing revealed by means of RAPD markers (Garcia Zambra-
germplasm information about plants, animals, microbes no et al., 2006); however, they were not confirmed
and invertebrates − could be useful for cactus pear. The using amplified fragment length polymorphism (AFLP)
FAO database Hortivar (http://www.fao.org/hortivar/), markers (Garcia Zambrano et al., 2009). O. ficus−indica
involving the direct collaboration of users, focuses on ecotypes from Tunisia were characterized by means of
commercial cultivars of horticultural crops; data on cac- RAPD markers, and 13 main groups were identified
tus pear cultivars have yet to be uploaded. with no relationship to different geographical regions
(Zoghlami et al., 2007).
Using AFLP, Labra et al. (2003) suggested that O. ficus−
USING MODERN MOLECULAR TOOLS TO indica should be considered a domesticated form of the
EXPLAIN CACTUS PEAR VARIABILITY spiny O. megacantha. AFLP markers were also applied
AND REDUCE REDUNDANCY to investigate genetic relationships between species in
three Opuntia collections in Tunisia (Snoussi Trifa et al.,
Phenotypic identification based only on morphological 2009). The level of genetic diversity in cactus clones in
markers can be misleading because of the complex gen- Brazil was evaluated by ISSR (inter−simple sequence
otype and environment interaction that governs most repeat) markers, revealing low genetic differentiation
of the traits of interest. The first step towards the ap- among them (Souto Alves et al., 2009). In order to assess
plication of molecular tools to study cactus pear genetic diversity, different molecular markers were applied, for
resources − redundancy, phylogeny and inheritance of example, RAPD−PCR and chloroplast simple sequence
valuable traits − was the optimization of DNA extrac- repeat (cp−SSR) (Chessa, 2010) and RAPD−PCR together
tion. Since the 1990s, protocols have been developed with ISSR using DNA from seeds (Luna Paez et al., 2007).
to extract DNA from cladodes and roots (De la Cruz et
The evolutionary history of O. ficus−indica has been
al., 1997; Griffith and Porter, 2003; Mondragón Jacobo
investigated with the use of Bayesian phylogenetic
and Doudareva, 2000), circumventing the problem of
analyses of nrITS (nuclear ribosomal ITS) DNA se-
mucilage. The adaptation of commercial kits has greatly
quences (Griffith, 2004).
facilitated the processing of large numbers of samples,
Molecular evidence for the hybrid origin of various Opun-
as reported by Fehlberg et al. (2013), who evaluated the
tia species and the ease of introgression has been docu-
effectiveness of the commercial DNeasy Plant Mini Kit
mented, and the polyphyletic origin of O. ficus−indica
(QIAGEN, Valencia, California, United States of America)
has been assumed (Griffith, 2003, 2009). The Griffith hy-
to extract DNA from the spines of eight species of cactus.
pothesis considering O. ficus−indica as a group of multiple
The best tool for the assessment of genetic diversity are unrelated clones derived from different parental species
markers based on DNA polymorphism. A wide range and selected for common agronomical features was also
of molecular marker techniques have been developed supported by Caruso et al. (2010) by means of microsat-
and are commonly used for genotyping individuals and ellites. Six highly polymorphic simple sequence repeats
inferring information on the genetic structure of ger- (SSR) developed by Helsen et al. (2009) and two expressed
mplasm collections, discovery of synonymy, and kinship. sequence tag (EST)−SSR loci were used to fingerprint 62
However, in the case of minor crop species like cactus Opuntia genotypes of Mexican, Israeli and Sicilian origin.
pear, the development and application of molecular A clear separation between cultivated accessions and cac-
markers are actually still limited. tus−pear−related species was documented.
The grouping of different Opuntia varieties and biotypes Helsen et al. (2009) cite the usefulness of both molec-
was achieved by means of isozymes banding pattern on ular and morphological data in conservation planning
pollen (Chessa et al., 1997). Due to the low reliability of − based on the relatively high morphological divergence
these markers, affected by environmental conditions, tis- found on the Galapagos endemic Opuntia species asso-
sue type and plant stage, more informative polymerase ciated with low genetic variability − as evidence for di-
chain reaction (PCR)−derived markers were applied. vergent selection and adaptation to local environments.
The same authors gave evidence that the current mor-
Random amplified polymorphic DNA (RAPD) markers
phology−based taxonomic differentiation between the
were successfully applied to verify the somatic origin
Opuntia taxa was not supported by molecular data.
within some Mexican accessions (Mondragón Jacobo,
2002). In order to identify fruit cultivars, vegetable The high variability and the codominant nature of SSRs
and ornamental accessions, Wang et al. (1998) applied have made them the marker of choice for DNA finger-
48 Crop ecology, cultivation and uses of cactus pear

printing in the characterization of Opuntia accessions in It- culture. In the Western Hemisphere, live collections
aly (Sardinia). A novel set of microsatellite loci were isolat- are present in Argentina, Chile, Brazil and Mexico; the
ed in different species and varieties of Opuntia (Erre et al., collection in Kingsville, Texas is no longer active, but
2013). Five out of the ten SSR loci developed were used to the United States Department of Agriculture (USDA)
characterize two field collections from Italy and Argentina. has assembled a new collection under the umbrella of
The level of polymorphism and the relatively high number the National Clonal Germplasm Repository in Parlier,
of alleles detected suggested that these markers can be California.
used for both inter− and intraspecific studies, as well as to
While Brazil has oriented its conservation efforts to-
provide a more reliable tool in the classification of Opuntia
wards forage accessions, other collections contain fruit,
species, based on their allelic profiles.
forage and double−purpose entries. Mexico, as the
largest centre of diversity, maintains several sites vary-
ing in extension, age and number of accessions (Table
CONSERVATION OF CACTUS 1). In 2010, it started a project on in vitro conservation
PEAR GENETIC RESOURCES of rare genotypes of immediate horticultural value.
Brazil reports 1 417 accessions, including genotypes
In situ conservation
from several countries, as well as segregants and hy-
brids of controlled crosses. Italy reports around 2 200
In situ conservation should be encouraged at the cen-
entries obtained from Mexico, Chile, the United States
tres of origin and diversity of cactus pear, with attention
of America, Morocco, Argentina and South Africa; they
focused on wild ancestors and semi−domesticated
include wild types and ecotypes, local varieties, selec-
types. Central Mexico is undergoing significant land−use
tions and hybrid populations of controlled crosses and
changes, which directly endanger diversity hot spots; the
plants derived from embryo culture.
main threats are incorporation of land to grazing, con-
struction of new roads and growth of urban areas. Land In early 2011, the Mexican Government launched the
tenure allows private individuals and communities to de- National Research Center for Genetic Resources and Bi-
cide the actual or future fate of wild lands, complicating otechnology (CENARGEN) in Tepatitlan, Jalisco, to serve
long−term conservation agreements. as a national repository; its facilities are designed for the
long−term storage of all crops and related organisms
Natural threats in the form of unusual frosts, snow and
relevant to national agriculture, including cactus pear.
hail – believed to be associated with climate change
– are also jeopardizing the survival of wild and semi−
domesticated cactus pear plants, in particular frost−
sensitive varieties. A late, harsh winter was recorded in
UTILIZATION: GERMPLASM
spring 2016, and some of the most important states for
ENHANCEMENT AND BREEDING
wild cactus pear in north central Mexico were covered
Attempts at cactus pear breeding go back to the late
in snow and ice; the final effects are being assessed.
nineteenth century. Results have been mixed for two
Different approaches to encourage conservation have main reasons:
been promoted in Mexico with varying levels of success: • Biological complexity − all opuntias of horticultural
natural protected areas (federal, state and municipal value are polyploid and present apomixes.
level), nature tourism corridors, and other schemes of
• Long−term juvenility − associated with the limited
sustainable utilization. Such policies must balance the
output of breeding programmes and costly projects
expected effects with the economic and social interests
(features directly related to funding).
of the human groups involved. All these initiatives are
under scrutiny due to budget cuts and political chang- As a result, breeding programmes are irregular, short−
es, risking the long−term protection of the resource. lived and poorly funded. Four programmes supported
by the Mexican, Italian, Brazilian and American gov-
Ex situ conservation ernments and one private company conduct breeding.
They are briefly described below.
In the Eastern Hemisphere, ex situ conservation is the
most active approach, with field collections in Tunisia, Italy
Morocco and South Africa. Italy has the largest and
oldest collection outside Mexico. It was started in 1992 The Italian breeding programme has been involved in
and acts as a germplasm bank and breeding collec- germplasm collection, description and documentation
tion; it hosts Opuntia and Nopalea specimens and the since 1992. It has performed crosses and pioneered the
number of accessions is currently 2 200. The collection embryo culture of cactus pear. The products obtained in-
includes local varieties and selections, hybrids derived clude 12 selections of green, yellow and red peel (four of
from open pollination, controlled crosses and embryo each) suitable for cultivation in Mediterranean countries.
Nopal (Opuntia spp.) genetic resources 49

Mexico Arid Land Plant Genetic Resources Unit (NALPGRU) in

Parlier, California maintains and evaluates plant ger-
The programme started in 1995 and has conducted hy- mplasm adapted to arid conditions and high levels of
bridizations and selections using the best Mexican geno- salt, selenium and boron in soil. The result is four new
types for fruit production to obtain multipurpose varieties, cultivar patents, co−owned by Fresno State, Diener’s
and improve fruit quality and adaptation. To date, three Red Rock Ranch and the USDA. Red Rock Ranch in-
improved varieties have been reported (Gallegos and tends to develop and market products in the form of
Mondragón Jacobo, 2011). Since 2013, programme activi- juice and powder blends derived from the new plants
ties have been reduced due to lack of reliable funding. (https://www.fresnostate.edu/jcast/cati/update/2013−
fall/opuntia−study.html).
Brazil

Located in the semi−arid tropical area of Pernambu- TOWARDS SUSTAINABLE UTILIZATION

co, the programme focuses on the development of OF CACTUS PEAR
improved varieties for forage production. Its inventory
includes the varieties ‘Gigante’, ‘Redonda’ and ‘Miuda’, New varieties are the axis of any sustainable system; they
1 061 clones of open−pollinated ‘Palma Gigante’, 171 are under development in Mexico, Brazil and Italy and to
clones of open−pollinated ‘Palma Miuda’, and 159 a limited extent in the United States of America. Since
clones obtained from Chapingo, Mexico, plus 17 clones underdeveloped countries can benefit most from cactus
from CPATSA, 5 from Rio Grande do Norte and 4 from pear cultivation, international collaboration needs to
Petrolina, with a total of 1 417 accessions. be encouraged, with countries sharing expertise, genetic
resources and facilities. In order to achieve this, interna-
United States of America tional centres must steer collaborative projects.
In the meantime, countries with native or naturalized
D´Arrigo Bros., a private American company interested in
stocks should intensify the collection, characterization
cactus pear production and breeding in Salinas, California,
and assessment of best entries, followed by propagation
has reported four improved varieties developed for private
of outstanding selections under strict phytosanitary pro-
use: ‘Sweet Emerald’, ‘Sweet Crimson’, ‘Sweet Purple’
tocols. For other countries new to cactus cultivation, it is
and ‘Sweet Sunset’ (http://www.andyboy.com/products/
recommended to introduce selected genotypes, test them
cactus−pear), reinforcing its line of fresh novel products.
and propagate the best performers under local conditions
The USDA Agricultural Research Service (ARS) National before launching large−scale cultivation programmes.

TABLE 1
Inventory of Mexican germplasm banks of cactus pears, updated to 2014

Use CRUZENa IIZDb CBTAc 38 INIFAP–SLPd

Fresh fruit 357 16 302 136 908
Fruit and forage 5 − − − 17
Forage 7 2 3 3 47
Vegetable 39 5 30 3 86
Triple use 2 − − − 2
Not reported (n.a.) − 23 − − 28
Animal feed − − − − 29
Ornamental − − − − 4
Condiment − − − − 15
Total 410 46 335 142 1 021
a
CRUCEN: Regional Center North Center University, Autonomous University of Chapingo, El Orito, Zacatecas, Mexico.
b
IIZD: Institute for Desert Area Research, Autonomous University of San Luis Potosí, Mexico.
c
CBTA: Agricultural Tech0,5ogy School, Ojocaliente, Zacatecas, Mexico.
d
INIFAP−SLP: National Institute for Forestry, Agricultural, and Animal Husbandry Research − Experimental Field, San
Luis Potosí, Mexico.
 06

Fruit production and

post-harvest management
Johan Potgietera and Salvatore D’Aquinob
a
Horticultural Consultant, Elands Bay, Western Cape, South Africa
b
National Research Council, Institute of Sciences of Food Production, Sassari, Italy
06 Fruit production
and post-harvest management

INTRODUCTION decisions must be made prior to planting. Many aspects

require consideration:
A worldwide expansion of cactus pear cultivation was • meso− and microclimate of the growing site;
predicted by Nobel in 1988. Predictions were partially • physical and chemical soil analysis;
based on the particular environmental biology of the
• cultivar choice;
cactus pear plant, which makes it possible to develop sus-
tainable agricultural production systems in areas with lim- • soil preparation;
ited water and poor soils (Inglese et al., 1995a). Further- • ordering of plant material;
more, there was an increased interest in cactus pear fruit • planting of windbreaks;
production due to a greater demand for exotic fruit on • determination of planting distances and row orienta-
the world’s major markets (Inglese et al., 1995a). In addi- tion; and
tion, limited irrigation water for expansion of mainstream
crop production in arid and semi−arid areas has forced • installation of an irrigation system.
agriculturalists to look at alternative crops with improved
drought tolerance and water−use efficiency (WUE). The Site selection
plant is cultivated in a wide range of environments, re-
The worldwide distribution of cactus pear is indicative of
sulting in differences in orchard practices (depending on
its extensive adaptability to various climatic and soil fac-
the country), fruit quality and crop productivity (Inglese
tors (Brutsch, 1979). Cactus pear plants can tolerate very
et al., 2009; Liguori and Inglese, 2015). Fortunately, since
high temperatures with no negative consequences (Nobel
the 1990s, much more information has become available
et al., 1986); on the other hand, planting sites where the
on orchard management and other cultivation aspects of
minimum temperature drops to below −5° C should be
cactus pear (Felker and Inglese, 2003).
avoided, as most fruit cultivars are killed at these tem-
Currently, cactus pear cultivation for fruit production peratures (Inglese, 1995). Low−lying areas where frost
takes place in at least 18 countries in semi−arid areas occurs in late winter to early spring may cause flower
in both hemispheres on more than 100 000 ha (Inglese bud abortion (Wessels, 1988b; Inglese, 1995; Nerd
et al., 2002a); this figure does not include naturalized and Mizrahi, 1995a) with partial to total crop failure. The
plants, nor plants cultivated for home consumption, optimal temperature range for nocturnal CO2 uptake is
commonly found in many countries (Inglese et al., 25/15° C day/night for cactus pear, because higher day
2002a). The country with the largest area under cultiva- or lower night temperatures result in reduced carbon
tion of cactus fruit is Mexico (51 112 ha) (Reyes Agüero uptake, leading to poor plant growth and production
et al., 2013), while other countries, such as Italy, Chile, (Nobel 1994; Nobel and Bobich, 2002). Temperature
South Africa, Argentina and Israel, also commercially patterns during floral budburst (late winter to spring)
cultivate the fruit (Inglese et al., 2002a). and fruit development period (FDP) have a particularly
significant effect on productivity and fruit quality. Most
The innaccurate perception that cactus pear does not
Opuntia species are poorly adapted to long periods of
need to be cultivated like other crops and that little or no
cool moist weather (Brutsch, 1979).
care still gives high productivity persists today to some
degree. Even the early cactus pear researchers − Griffiths In most countries characterized by predominant sum-
(1909) and Turpin and Gill (1928) − ascribed this theory mer rainfall, cactus pear is grown under rainfed con-
to the ease with which the plant grows in virtually any ditions; however irrigation is necessary where the dry
environment. Today we know that this is untrue: in order time of year coincides with the FDP or where annual
to be profitable, the plant requires appropriate care. rainfall is < 300 mm (Inglese, 1995). Cactus pear is suc-
cessfully grown in winter rainfall areas when provision
is made for irrigation in the hot dry summer months.
ORCHARD PLANNING AND The maximum rainfall limit is 1 000 mm per year (Le
ESTABLISHMENT Houérou, 1992, 1996a), but high rainfall is less ideal
as it results in poor pollination, increased presence of
Thorough planning is crucial when establishing a com- fungal diseases, fruit cracking and reduced fruit quality
mercial cactus pear fruit orchard and many important (Wessels, 1989).
Fruit production and post−harvest management 53

Given the succulence of the plant and fruit, it is easily Adverse soil conditions, such as impenetrable layers, per-
damaged by hail, and commercial fruit production ennial weed infestation and shallow soil depth, need to
should be avoided where hail is a regular occurrence be addressed before planting (Wessels, 1988b; Inglese et
(Wessels, 1988a; Brutsch, 1997a). Physical damage as a al., 2002a). Deep soil cultivation (at least 500 mm) with
result of hail facilitates the entry of pathogenic fungi into a ripper/subsoiler on the plant row is needed to break up
the plant (Granata, 1995; Swart et al., 2003), and causes any hardpan layers, and to improve drainage, aeration
cosmetic damage making the fruit unmarketable. and water−holding capacity. In heavier compacted soils,
Snyman (2004, 2005) showed that at 2 years, O. ficus− deep cultivation across the rows may also be beneficial:
indica roots comprise only 7% of the total plant biomass it helps the plant survive during low rainfall years when
– indication of the large difference between below− and it can utilize soil water from deeper soil depths (Inglese,
above−ground biomass in the early years of plant growth. 1995; Potgieter, 2001). On very shallow soils, ridging is
As a result, strong winds accompanied by rain may cause recommended (Singh, 2003). It is important to remove
plants to lodge, especially when plants are young and the perennial weeds before establishment − either mechan-
root system is not yet fully developed to keep it upright. ically or chemically − as they compete strongly with the
cactus pear plant, particularly during the early stages of
It is important to carefully examine the microclimatic plant growth (Wessels, 1988b).
features when selecting the most appropriate plant-
ing site for orchard establishment. For example, a Pre−plant fertilization
north−facing site is warmer than a south−facing one
(Southern Hemisphere), resulting in earlier harvest and
Soil amendments to correct soil nutrient imbalances and
improved fruit quality.
soil pH must be carried out prior to establishment. A soil
While cactus pear can be grown in a wide range of soil pH (water) of 6.5–7.5 is considered optimum (Wessels,
types (Inglese et al., 2002a), it is important to select 1988a, b; Singh, 2003). Fertilizer applications should be
the best soils available for high productivity (Wessels, based on the results of soil analysis, which indicate the
1988a). Soils with a sandy to sandy loam texture are levels of plant nutrients in the soil (Table 1).
ideal (Vazquez−Alvarado et al., 2006), but cactus pear
can grow equally well on heavier soils. Wessels (1988b) TABLE 1
noted that soil drainage rather than soil type is the con- Suggested optimum soil nutrient levels for cactus pear
ditioning factor. Cactus pear is very sensitive to a lack fruit production for rainfed cultivation in summer rain-
of oxygen in the soil (Brutsch, 1979; Le Houérou, 1992, fall areas of South Africa
1996a), but very sandy soils have poor water−holding
capacity and are more prone to leaching of nutrients Element Optimal soil level (mg kg−1)
(Wessels, 1988a). The plant prefers a neutral to slightly
P 20−30
alkaline soil pH (water) (Wessels, 1988b; Zegbe et al.,
2015). Soil selected for planting should be at least K 80−100
300 mm deep (Wessels, 1988a), although deeper soils Ca > 400
(600–700 mm) are preferred for commercial produc- Mg a
100−150
tion (Inglese et al., 2002a). Most Opuntia species do a
Mg levels should not be higher than Ca levels.
not tolerate even moderate levels of dissolved salts in
Source: Potgieter, 2001, 2007 (adapted).
the rooting zone. Since net CO2 uptake is inhibited, it
negatively affects the vegetative growth of cacti in soils
high in sodium chloride or calcium carbonate (Nobel, Representative soil samples of the topsoil (0−300 mm)
1994, 1995). Gersani et al. (1993) report a 40% reduc- and subsoil (300−600 mm) need to be taken for chemi-
tion in plant growth with a concentration of 30 mol cal and physical analysis. The secondary effects of soil pH
m−3 (1.76 g litre−1 NaCl), and a 93% reduction with a on the availability of other plant nutrients are probably
concentration of 100 mol m−3 (5.85 g litre−1 NaCl). more important than soil pH per se. For example, P be-
comes less available to plants at a low soil pH (Nobel,
Soil preparation 1988). Lime and P are relatively immobile in soils and as
a result these need to be thoroughly mixed with the soil
Pre−plant soil preparation is essential for successful prior to planting. In addition to decreasing vegetative
cactus pear production and cannot be adequately per- growth and dry mass (Berry and Nobel, 1985), high soil
formed after orchard establishment. Land clearing and salinity also decreases water content of the cladode,
levelling may be required (Inglese, 1995); irrigation sup- uptake of K and Ca, and root−shoot ratio (Nerd et al.,
ply lines may be installed and plant rows marked out. In 1991c). The addition of gypsum helps to neutralize excess
areas where game and domestic animals can damage salts in the soil solution. Cactus pear reacts very well to
young plants, fencing is required. organic manures which also improve the soil structure,
54 Crop ecology, cultivation and uses of cactus pear

nutrient content and water−holding capacity (Inglese, depends primarily on the climatic conditions of the
1995; Singh, 2003). As a general guideline, 6–10 tonnes planting site and on market demand. Opuntia species
ha−1 of well−composted animal manure needs to be and cultivars differ greatly with regard to their potential
incorporated into the soil before planting. fruit yield, quality characteristics and adaptability to
environmental conditions. Not all cultivars are equally
Windbreaks adapted to a particular area; indeed, most cultivars
exhibit strong genotype–environment interaction (G ×
In windy areas, it may be beneficial to plant live wind- E) (Potgieter, 2007). Cultivars must be chosen wisely;
breaks to minimize the negative effects of wind in changing cultivar after establishment entails very high
cactus pear orchards. Heavy rainfall accompanied by costs, because plants are self−rooted and grafting or
strong wind can cause young plants to lodge and can top−working is not economically viable.
even cause branches and cladodes to break off (Felker
In addition, cultivar characteristics must be carefully
et al., 2005). Pollination and plant protection sprays
considered (Table 2) before a final decision is made:
are all negatively affected by high winds; it is difficult
there may be specific cultivar preferences for supplying
to perform orchard practices (pruning, fruit thinning,
a particular market. For example, cultivars with a red,
harvesting) under windy conditions as the glochids tend
to become airborne (Wessels, 1989). When developing pink, orange or yellow pulp are preferred on most Euro-
cladodes and fruit are too close together on a cladode, pean and North American markets, whereas white and
fruits can be damaged by chafing during windy spells. green pulp fruit are favoured by South African (Wessels,
Live windbreaks must be adapted to the area of plant- 1988) and Mexican consumers (Mondragón Jacobo and
ing and they should receive water and plant nutrition Perez Gonzalez, 1994, 1996). If growers are contem-
to ensure that they do not compete with the orchard. plating exporting fruit, the cultivar choice becomes even
A popular tree for live windbreaks in orchards is the more important as characteristics such as appearance,
Australian beefwood (Casuarina spp.). post−harvest resistance to handling and shelf−life play a
major role in successful exports. Producers should take
Cultivar choice into account that some cultivar characteristics can be
influenced by the environment or by orchard manage-
The number of cactus pear fruit cultivars available varies ment practices. For example, although fruit mass may
significantly between countries. Mexico and South be genetically controlled, it can be influenced by fruit
Africa have numerous cultivars, while in most other thinning, irrigation, fertilization and pruning. Annual
producing countries the cultivar choice is rather limited variation in fruit yield and quality is also evident in most
(Inglese et al., 2002a). The choice of a fresh fruit cultivar commercial orchards.

TABLE 2 Main cactus pear fruit cultivar characteristics in producing countries of the world

Cultivar Country Species Spininess Peel/pulp Ripening Fruit Pulp TSS Post−harvest
name colour season mass handling
50% peel N. g % o
Brix
colour break Hemisphere
Algerian SA O. ficus−indica none red/dark pink Jul./Aug. 162 59.4 13.9 moderate
Gymno
SA O. ficus−indica none Yellow/Orange Aug./Sept. 170 61.7 11.2 moderate
Carpo
Meyers SA O. ficus−indica none red/dark pink Jul./Aug. 176 60.7 13.6 good
Morado SA O. ficus−indica none Green/White Jul./Aug. 146 60.0 14.4 poor
Nudosa SA Unknown slight Red/red Aug./Sept. 236 60.8 11.2 good
Roedtan SA O. ficus−indica none Yellow/Orange Aug./Sept. 171 60.7 14.2 moderate
Turpin SA O. ficus−indica none Yellow/Orange Jul./Aug. 181 55.0 13.6 moderate
Light green/
Zastron SA O. ficus−indica none Jun./Jul. 137 57.2 13.5 good
White
Light green/
Reyna Mexico O. albicarpa spiny Jul./Aug. 120 63.7 16.4 poor
light green
Light green/
Cristalina Mexico O. albicarpa spiny Aug./Sept. 207 60.2 12.7 good
light green
Fruit production and post−harvest management 55

(Continued)

Cultivar Country Species Spininess Peel/pulp Ripening Fruit Pulp TSS Post−harvest
name colour season mass handling
50% peel N. g % o
Brix
colour break Hemisphere
Green/light
Villanueva Mexico O. albicarpa spiny Jun./Jul. 129 60.9 14.4 poor
green
Light green/
Burrona Mexico O. amyclaea spiny Sept./Oct. 217 59.8 12.7 very good
light green
Roja San Red purple/
Mexico O. ficus−indica slight Jul./Aug. 116 44.5 13.7 poor
Martin purple
Naranjona Mexico O. megacantha slight Yellow/Orange Aug./Sept. 170 51.7 13.1 very good
Roja Vigor Mexico O. ficus−indica Bright red/red Jul./Aug. 174 moderate
Light green/
Dellahia Morocco O. robusta none 100 53.1 14.0
Light green
Moussa Morocco O. ficus−indica none Yellow/Orange Dec. 101 51.0 14.4
Aissa Morocco O. ficus−indica none Yellow/Orange 96 50.0
Gialla Italy O. ficus−indica none 103 51.0 13.0 good
Rossa Italy O. ficus−indica none
Bianca Italy O. ficus−indica none 112 53.6
Amarilla sin
Argentina O. ficus−indica none Yellow/Green 13.9
Espinas
Ofer Israel O. ficus−indica none Yellow/Orange
Andy Boy USA
Verde
Chile O. ficus−indica spiny Green/Green Feb−Apr. 132 49.6 13.3 good
(Criolla)

SA = South Africa; USA = United States of America.

ORCHARD LAYOUT AND DESIGN

Planting systems
Row orientation
Choice of planting design and planting distance depends
Row orientation, unlike tree density, cannot be changed on farm size, farm management, implements, climate,
and is fixed for the lifespan of the orchard. According to soil fertility, cultivar growth habit, plant training system
Nobel (1982), cactus pear cladodes are inclined to face and occurrence of pests such as cochineal (Wessels,
east−west, except at lower latitudes (> 27°) and in areas 1988b; Inglese, 1995).
where vegetative growth occurs during winter. In these
situations, row orientation is less important (Nobel, 1982). Hedgerow
Once row orientation is decided, it is important to achieve Various planting systems can be employed for cactus
optimal light utilization over the whole tree canopy during pear fruit production, but in recent years, high−density
the day (García de Cortázar and Nobel, 1991; Stassen et hedgerows planted in a rectangular system have become
al., 1995). While the generally preferred row orientation is the norm in large commercial orchards in South Africa
north−south to capture equal daily solar irradiance under (Unterpertinger, 2006), Israel (Nerd and Mizrahi, 1993),
sunlit and cloudy weather conditions, it can be adapted the United States of America (California) (Bunch, 1996)
to suit the latitude, site and incidence of sunburn. There and Argentina (Felker and Guevara, 2001). Hedgerows
are other practical considerations, such as sloping fields, have the advantage of lower input costs. They also en-
where it is recommended to plant on the contour to pre- able more thorough spray coverage, and solar radiation
vent soil erosion (Inglese, 1995; Stassen et al., 1995). capture is maximized, resulting in higher productivity.
56 Crop ecology, cultivation and uses of cactus pear

Rectangular or square planting (free−standing trees) years. Therefore, further pruning and tree removal
In Italy, trees are usually widely spaced in a square or every few years is necessary to avoid a reduction in
rectangular layout; however, similar fruit yields have yield and fruit quality (Inglese et al., 1995a; Inglese et
been reported in Israel and Italy under different plant al., 2002a). Barbera and Inglese (1993) recommend
spacings (Inglese et al., 1995a). more cuttings per planting station, spaced 4 m apart,
resulting in a rapid SAI (stem area index) increase. This
Spacing results in trees with most of the production on the
outer cladodes. However, trees rapidly become 3−3.5
Plant spacing for cactus pear fruit production varies m high and 4−4.5 m wide, entailing higher pruning
greatly, depending on the hemisphere, country and en- and harvest costs (Inglese et al., 1995a). In addition
vironment. In Italy, planting distances are generally wide to the problem of overshading, dense canopies also
and vary from 6 × 6 m (278 plants ha−1 ) to 4 × 6 m (416 aid cochineal infestations and reduce pest control
plants ha−1) (Inglese, 1995; Inglese et al., 2002b; Tudisca efficiency (Inglese, 1995b). High plant density (2 500
et al., 2015). In other parts of the Mediterranean, plants plants ha−1) gives high fruit yield (> 30 tonnes ha−1)
are spaced much closer. For instance, in Israel, 1.5 m in the from 3−year plants; however, more vegetative than
row and 4 m between rows (1 666 plants ha−1) is used to reproductive growth occurs as the plant ages (García
increase fruit yield in the early stages of orchard life (Nerd de Cortázar and Nobel, 1992).
and Mizrahi, 1993). A plant density range of 500−2000 • Plant trees according to a realistic high−density
plants ha−1 has been reported in Jordan where plants in spacing and utilize tree management systems to
the rows are 1−2 m apart, with rows 5−10 m apart and shape and contain the tree within the allocated
usually positioned on the contour line (Nasr, 2015). In space (Stassen et al., 1995). Close spacing within a
California, plant spacing is 1.5−4 m in rows 4−6 m apart row maximizes fruit yield in young plants as the num-
(830−1 666 plants ha−1) (Bunch, 1996). On large farms ber of fertile cladodes per unit area increase (Inglese
in Mexico, plants are spaced 4 × 5 m (500 plants ha−1) et al., 2002a). This approach is dependent on the use
(Pimienta Barrios, 1990). In the Southern Hemisphere, of manipulation techniques to maintain a specific
most new orchards in South Africa are established at 2 × plant volume. Tree height should not surpass 80%
5 to 2 × 4 m in and between rows (1 000−1 250 plants of the width between rows to avoid shading of the
ha−1) in a hedgerow system. Targa et al. (2013) reports lower parts of the hedgerow (Stassen et al., 1995),
plant spacings of 1.5 m in row and 6 m between rows (1 which makes these areas less fruitful. Provided tree
111 plants ha−1) as continuous hedgerows in Argentina. management is done on an annual basis, this strategy
is more efficient than lower plant densities. However,
Whatever spacing is used, it is important to keep open a if tree management cannot for some reason be per-
working row of approximately 1.8 m for farm machinery formed, it is advisable to maintain a low tree density.
movement (Wessels, 1988b; Gittens, 1993).

High−density planting
ESTABLISHMENT
A general problem in cactus pear cultivation occurs
Cladode rooting and subsequent vegetative growth
when plants are allowed to become too large, resulting
in the field depend on many factors. Observations in
in “forests” rather than productive orchards. Large
many commercial orchards have revealed a high degree
portions of these big trees, especially in the lower parts
of variability despite the use of vegetative plant material
of the canopy, become completely shaded out (Liguori
(Brutsch, 1979; Wessels, 1988a), indicating a need for
and Inglese, 2015). As a result, fruit−set is limited to the
careful selection of planting material.
outside of the plant canopy and labour costs are high
at harvest. Given the ever−increasing costs of land and Plant material selection and preparation
production inputs, it is necessary to seriously re−evalu-
ate planting systems, density and tree management in Cactus pear for fruit production purposes is primarily
relation to fruit yield and quality (Stassen et al., 1995; propagated by clonal means to ensure true−to−type-
Liguori and Inglese, 2015). Low planting densities are ness (Wessels, 1988b). Where farmers do not have their
not economically justified; nevertheless, there are two own cactus pear plants, it is advisable to order plant
schools of thought on plant spacing: material well in advance from producers that have good
• Plant trees on an intensive scale and remove quality plant material. Both single and multiple cladode
some as encroachment occurs. The aim is to have cuttings are used worldwide for orchard establishment.
an early heavy crop. Close spacing (1−1.5 m) within
the row maximizes yield in the early years after Single cuttings
planting, but may result in overshading after a few One− or two−year single cuttings (cladodes) can be
Fruit production and post−harvest management 57

used, although 1−year cuttings develop more and lon- The upright position is the most commonly adopted
ger roots than 2−year cuttings (Arba, 2009b). In gener- (Inglese, 1995) and is preferred for fruit production.
al, single, mature, large, terminal cladodes of uniform Cuttings are planted upright (vertically) with the cut end
appearance, visually free of any defects, insects and pointing downwards into the soil. Plants root quickly
diseases, should be selected as plant material (Potgieter, and a sturdy plant develops (Arba, 2009b; Arba and
2007). Barbera et al. (1993b) and Wessels et al. (1997) Benrachid, 2013). Cladodes should be positioned with
report that the surface area and dry mass of a cutting the flat sides towards the working row. To ensure prop-
have a significant influence on successful rooting and er contact between the cutting and soil, the soil needs
budding in the field. A surface area of 500 cm2 or dry to be firmed around the cladode after planting. The only
mass of 70−100 g allows good plant growth. Inglese disadvantage of this planting method is possible rotting
(1995) recommends placing two parallel cuttings spaced at the cut end (Wessels, 1988b).
0.4 m apart at a single planting station for rapid canopy
development, or alternatively 3−4 single cuttings posi- Planting time
tioned in a triangle or square and spaced 0.3 m apart.
Although this method has the advantage of faster can- Roots and cladodes reach their highest growth rate
opy development, it results in wider within−row spacing during late spring to early summer (Wessels, 1988b).
and requires large quantities of planting material (Ing- In summer rainfall areas, newly planted cladodes
lese, 1995; Mondragón Jacobo and Pimienta Barrios, benefit from rains that occur after planting (Pimienta
1995). Single cladodes as propagation units have the Barrios, 1990; Singh, 2006). Planting can be extended
advantages of lower transport costs and easier handling to midsummer in areas with mild winters. The idea is
during the planting process for the plant to become well established and survive
colder winter conditions. However, autumn planting is
Multiple cuttings recommended in Morocco (Nasr and Jamjoum, 2002;
Cactus pear can also be propagated using mature Nasr, 2015) and Jordan (Homrani Bakali, 2013), where
attached (one, two or more) cladodes; this is common it resulted in well−established root systems in winter,
practice in Sicily where 2−3−year multiple cuttings are strong growth in summer and earlier fruiting.
used (Tudisca et al., 2015). Most of the basal cladode
is placed underground to ensure plant stability (Inglese, Care of newly planted cladodes
1995). Multiple cladode cuttings allow more rapid plant
development and earlier fruiting than single cladodes One or two light irrigations (10 litres plant−1) in a small
after planting (Homrani Bakali, 2013; Nasr, 2015). How- earthen dam around the plant promote root devel-
ever, due to their size and mass they are more difficult opment, but care should be taken not to overirrigate
to handle and transport. young plants (Potgieter, 2001). Newly established cla-
Cuttings can be cured for 4−6 weeks in partial shade dodes may develop fruit soon after planting. Due to the
on a dry surface to allow the cutting wound to callus high sink demands these fruit have on the plant, it is
(Potgieter, 2001; Inglese et al., 2002a). Alternatively, best to remove them in the establishment year (Wessels,
various copper−based fungicides can be used to treat 1988b; Inglese, 1995). From the first production year a
the cutting wound before planting (Inglese, 1995). light crop may be left to mature.
To prevent the introduction of insects (e.g. cochineal
and cactus pear moth) to new planting areas, it is
recommended to disinfect cladodes thoroughly with a ORCHARD MANAGEMENT
registered insecticide before planting. Immature stages
of the cochineal insect are barely visible with the naked Weed control
eye and unsuspecting producers may bring cochineal
into an area where it did not previously occur with po- While cactus pear is well adapted to arid and semi−arid
tentially devastating consequences. Cladodes should be regions and can survive severe drought conditions,
washed with the insecticide mixture using a soft brush weed control has been shown to enhance productivity
to destroy any possible insects. (Felker and Russel, 1988), especially in young plantations
(Inglese et al., 1995a). The plant’s sensitivity to weed
Planting depth and methods competition is due to the very shallow root system (Felk-
er and Russel, 1988; Snyman, 2005), where it competes
In sandy soils, cuttings need to be planted deeper at the same soil level as weeds for nutrients and water.
than in heavier soils to prevent lodging; likewise, small Nobel and De la Barrera (2003) showed that 95% of
cladodes should be planted deeper in order to ensure the roots of mature cactus pear plants develop at a soil
adequate rooting. There are three ways to plant cactus depth of 40−470 mm, while Snyman (2006a) reports
pear: upright; on the side at a 30−45° angle; or flat. that roots can spread as far as 2.5 m from the stem of
58 Crop ecology, cultivation and uses of cactus pear

the plant in 2 years. Various methods of weed control exposed to adequate sunlight will produce flower buds
can be used, but soil cultivation should be restricted to (Nerd and Mizrahi, 1995b), while shaded cladodes are
a minimum, in order to avoid damaging the shallow root usually low−yielding or even infertile (Wessels, 1988a;
system (Inglese, 1995). Weed control is best performed Pimienta Barrios, 1990; Inglese et al., 2010). Therefore,
at an early stage of growth, when competition with the to ensure high CO2 uptake and cladode fertility, it is im-
cactus pear crop is minimal (Wessels, 1988a; Inglese, portant to prevent excessive cladode shading (Pimienta
1995). Barrios, 1990; Inglese et al., 1994a) particularly during
the last 8 weeks before spring floral budburst (Barbera
Chemical weed control et al., 1993a; Cicala et al., 1997). Other benefits of prun-
Due to high labour costs, chemical weed control is the ing include: controlled plant size, training of the plant
norm in commercial fruit production. A range of herbi- into a hedgerow, increased fruit yield, improved fruit
cides may be used, but farmers are urged to only use size, easier pest detection and control, easier harvesting
products that are registered in their respective countries, and rejuvenation of old plants (Hester and Cacho, 2003;
especially when the fuit is produced for export. Weeds Inglese et al., 2009, 2010). Between 20 and 50% of
should preferably be controlled to 1−1.5 m on both the terminal cladodes should be removed by pruning
sides of the planting rows (Potgieter, 2001); where the (Oelofse et al., 2006). However, excessive pruning will
danger of soil erosion is limited, complete weed control reduce yield and contribute to strong vegetative growth
between rows can be performed (Brutsch, 1979; Felker the following season (Inglese et al., 2002b). All diseased,
et al., 2005). Because the early growth of cactus pear is small and damaged cladodes should be removed.
extremely sensitive to herbicide sprays, spraying should
be avoided on windy days. Where lower trunk sections Formative pruning
contain chlorophyll, stems should be shielded during Formative pruning begins in the first year of establish-
spraying to prevent herbicide damage. ment and is changed to production pruning when the
plants start bearing (Targa et al., 2013; Nasr, 2015). The
Mechanical weed control aim of formative pruning is to direct vegetative growth
In a hedgerow planting system, in−row weed control into the desired plant shape. In countries where high−
by mechanical means is preferred when cactus pear density hedgerows are used, plants are pruned to a
plants are young due to their sensitivity to herbicides pyramidal shape (Potgieter, 2001). Where square plant-
(Potgieter, 2007). However, in square planting systems ing systems with wider spacing are common, vase− or
with free−standing globe−trained plants, weed control globed−shaped plants are formed (Inglese, 1995). These
remains manual because this training system makes it plants do not have main stems, resulting in large plants
difficult to work with ordinary farm implements be- with a high number of terminal cladodes distributed
tween trees (Inglese and Barbera, 1993). In smaller or around the outer portion of the canopy (Inglese et al.,
more traditional farming systems with limited access 2002a).
to herbicides, the soil between the plant rows may
be ploughed to clear the fields of weeds (Nasr, 2015; Productive pruning
Tudisca et al., 2015). Although mechanical weed control Productive pruning is used to maintain a good balance
is not ideal, it is better than no weed control at all (Felker between vegetative and reproductive growth with an
and Russel, 1988). adequate number of new terminal cladodes for the
subsequent year’s blooming (Mulas and D’Hallewin,
Permanent grass strips 1992). Environmental conditions, cultivar growth habit
Where orchards are planted along slopes, it is recom- and plant spacing all affect canopy density (Inglese et
mended that a regularly mowed grass strip be main- al., 2002a). Reduction of the canopy density through
tained between rows to prevent soil erosion (Potgieter, pruning facilitates orchard practices (e.g. fruit thinning,
2001). Weeds can be mowed and left on the soil surface scozzolatura, harvesting) and contributes to improved
as mulch to retain moisture and reduce weed growth fruit quality (Inglese et al., 2002a, 2010). Plant height
(Inglese, 1995). should preferably not exceed 1.8 m in order to avoid the
use of ladders to perform orchard practices (Potgieter,
Pruning and training systems 2001; Nasr, 2015).

The selection of an appropriate pruning and training Renewal pruning

system for cactus pear is closely related to the planting In the case of old cactus pear plantations, senescence of
system, layout and spacing chosen in the planning phase. the canopy and yield reduction with noticeable alternate
The main reason for pruning in cactus pear is to ensure yielding are common (Mulas and D’Hallewin, 1992).
maximum photosynthetically active radiation (PAR) in- Rejuvenation of old plants can be achieved by cutting
terception by terminal cladodes. Most terminal cladodes the plant back to a height of 0.5 m above soil level.
Fruit production and post−harvest management 59

Only 3−4 well−spaced main scaffold branches should will result in cladodes not being exposed to sufficient
be left for development of the new plant. To prevent PAR to make them fertile. In South Africa, pruning takes
sunburn, the whole plant should be painted with white place from April to July (autumn/winter), when the plant
polyvinyl alcohol (PVA) paint, mixed 1 : 1 with water. is no longer actively growing (Wessels, 1988c; Potgieter,
With a large established root system, the plant resumes 2001). Similarly, in Mexico, Pimienta Barrios (1986, 1990)
fruiting within 2−3 years after rejuvenation pruning suggests pruning from November to March (winter).
(Wessels, 1988b; Mulas and D’Hallewin, 1992). The
newly developing cladodes must be thinned to prevent Fertilization
cladode overcrowding.
Deficiencies in mineral nutrients affect cactus pear
Summer pruning plant metabolism with a resultant negative impact on
Complete removal of newly developing cladodes in fruit yield and quality (Nerd and Mizrahi, 1992; Zegbe
spring is common practice in Sicily to reduce compe- Dominguez et al., 2014). In order to make fertilizer rec-
tition between fruit and vegetative growth. However, ommendations for cactus pear, it is essential to consider
this practice may result in an alternate bearing pattern the plant nutrient status of the terminal cladodes as well
(Inglese and Barbera, 1993; Inglese et al., 2002b). Sum- as the available nutrient reserves in the soil.
mer pruning is not advisable in areas with cold winters,
because cladodes developing late in the season would Plant nutrition results reported for cactus pear fruit yield
not have sufficient time to harden−off before winter and quality are highly inconsistent and contradictory
when they would be subject to frost damage (Wessels, making fertilization recommendations challenging. Cac-
1988b). However, in South Africa, thinning of some tus pear plants differ from most other crop plants, both
of the excess developing cladodes in spring and early physiologically and morphologically; for this reason,
summer is performed. Newly developing cladodes close fertilizer recommendations applied to other crops are
to flower buds may cause chafing of the fruit epidermis,
of little use (Nobel, 1983, 1988; Magallanes Quintanar
making it unmarketable due to cosmetic damage (Wes-
et al., 2006). In the absence of agreement on cactus
sels, 1988b; Potgieter, 2001).
pear fertilization, Table 3 provides growers with broad
The best time to prune is after fruit harvest but not provisional soil and plant tissue analysis norms, while
later than 2 months before floral bud break (Wessels, Table 4 lays down general fertilization guidelines where
1988b). Late pruning, especially in overcrowded trees, no soil analysis results are available.

TABLE 3 Provisional soil and plant tissue norms for cactus pear fruit production

Soil parameter Norm Terminal cladode Norm (% DM)

pH (water) 6.5–7.5 N b

P (mg kg−1) 20–30 P 0.1–0.3

K (mg kg ) −1 a
80–100 K 1.5–4.0
Ca (mg kg−1) > 400 Ca 2.0–4.5
Mg (mg kg−1) 100–150 Mg 1.0–1.5
Na (mg kg ) −1
< 200 Na 0.02–0.03
Mn (mg kg−1) 30–70
Ca/N 4.0
K/N 3.4
N/P 4.5
Ca/Mg 3.0
a
If (Ca + Mg)/K > 8 (sandy soils) or < 5 (clay soils), apply K.
b
Young trees (0–2 years): 0.6–0.8%. Mature trees (≥ 3 years): 0.9–1.3%.
60 Crop ecology, cultivation and uses of cactus pear

TABLE 4 Provisional fertilizer recommendations for fruit production where no soil analysis results are available
(plant nutrient amounts in kg ha−1 year−1)

Nutrient element Year 1 Year 2 Year 3 > Year 4

N 50 50 70 100
P 10 10 20 30
K 20 20 30 50
Mg 10 10 20 40

Mineral nutrition research on cactus pear in various in winter (Barbera et al., 1992a; Inglese, 1995) with fer-
countries shows that fertilizer application, both organic tigation applied throughout the year in Israel (Nerd et
and inorganic, is generally beneficial in fruit production. al., 1991b). Nerd et al. (1989, 1991b) and Ochoa and
Nutrient elements influence vegetative and reproductive Uhart (2006c) report that the application of NPK fer-
phenology, fruit yield and quality in cactus pear, with tilizer in winter increases the production of floral buds
macroelements having the greatest effect on fruit pro- the following spring. However, according to García de
duction (Zegbe Dominguez et al., 2014; Arba et al., Cortázar and Nobel (1991), the best time to apply fer-
2015b). Of all the plant nutrients, N is the most limit- tilizer is during the warmer months due to higher pho-
ing nutrient in cacti (Nobel, 1983), with the highest N tosynthetic photon flux density (PPFD) in summer. In
values found in young fertile cladodes (Nobel, 1988). Mexico, half the N and all the P and K are applied with
However, very high N concentrations (> 2.2%) in 2− and irrigation at the onset of floral budburst, and the other
3−year old cladodes may result in excessive vegetative half of the N is applied after harvest (Zegbe Dominguez
growth with accompanying higher input cost, reduced et al., 2014). Under rainfed conditions (summer rainfall),
cladode fertility, poor fruit colour development and un- half the N and K and all the P can be applied directly
even ripening (Potgieter and Mkhari, 2000; Inglese et after fruit harvest and the remainder towards the end
al., 2002a). Reported P and K plant tissue concentra- of March (Wessels, 1988b), while liming is carried out
tions range from 0.06−0.3 to 0.06−3.5%, respectively at any time of the year, but preferably at least 1 month
(Nobel, 1983, 1988; Arba et al., 2015b). Ca and K are after N fertilization (Claassens and Wessels, 1997).
the most plentiful mineral elements in the cladodes, po-
tentially more abundant than N (Galizzi et al., 2004). Irrigation
Furthermore, Mg in young cladodes can reach levels of
1.47% (Magallanes Quintanar et al., 2006). Therefore, The exceptional drought tolerance and high WUE of cac-
N, P, K, Ca and Mg are all potentially limiting factors in tus pear plants (Han and Felker, 1997; Zegbe−Domin-
cactus pear fruit production if cultivated in nutrient−de- guez et al., 2015) are the primary reasons for its popu-
ficient soils (Magallanes Quintanar et al., 2006). Nutrient larity as a rainfed crop in many areas of the world with
concentration in cladodes is affected by fruit crop load, low rainfall and a shortage of irrigation water. Although
cladode position, plant age, plant tissue analysed and the plant can survive in areas receiving 200 mm year−1
season (Nerd and Nobel, 1995; Gugluizza et al., 2002a). (Acevedo et al, 1983), the optimal rainfall range for cac-
tus pear production is 400−600 mm year−1, but soil type
In cactus pear, an additional flower flush can be induced
also plays a role in the actual plant water requirement
with the application of N just after the removal of the sum-
(Le Houérou, 1992, 1994). Although considerable fruit
mer harvest. According to Nerd et al. (1993b), the number
yields can be achieved under low rainfall, rainfed condi-
of flower buds increases with increasing N levels up to 120
tions (Potgieter, 2007), supplementary irrigation of cac-
kg N ha−1, while the N concentration in the cladode tissue
tus pear is advisable in summer rainfall areas where <
is positively correlated to the number of flowers formed.
300 mm year−1 is received (Mulas and D’Hallewin, 1997;
Nerd and Mizrahi (1995b) further found that the autumn
Van der Merwe et al., 1997). In addition, irrigation
flower flush is higher in younger (< 6 years) than in older
during periods of unfavourable climate − such as dry
plants. However, high production systems with two fruit
spells during the rainy season, or when spring rains are
harvests in one year from the same plant may have addi-
late − is advantageous (Wessels, 1988d). In a Mediter-
tional nutritional requirements (Groenewald, 1996).
ranean climate, where most rainfall is in winter, supple-
Due to the synergistic relationship between fertilization mentary irrigation in summer is indispensable for high
and irrigation, fertilizer application should take place yield and good quality (Mulas and D’Hallewin, 1997;
when adequate rainfall or irrigation is available (Nerd Homrani Bakali, 2013). Irrigation of cactus pear is there-
et al., 1989; Mondragón Jacobo, 1999). In countries fore common practice in Italy, Israel, Jordan, Morocco,
with a Mediterranean climate, fertilization takes place Chile and the winter rainfall areas of South Africa.
Fruit production and post−harvest management 61

There are definite advantages with supplementary ir- surface area with small volumes are very suitable for cactus
rigation of cactus pear, especially during certain critical pear with its shallow and widespread root system (Potgiet-
phases of plant growth and development. García de er, 2001; Snyman 2004, 2005), and they positively influ-
Cortázar and Nobel (1992), Mulas and D’Hallewin (1997) ence fruit size and quality (Inglese et al., 2010). Although
and Liguori et al. (2013b) all report the beneficial effects traditional irrigation methods, such as basin irrigation, may
of irrigation on vegetative plant growth, cladode number be less efficient, such systems may provide an easy and
and canopy size. Fruit yield per plant is generally higher cheap solution for cash−strapped farmers if irrigation is re-
in irrigated than in non−irrigated plants, and researchers quired only 2−3 times per season (Wessels 1988b).
ascribe higher yield to higher average number of fruit per
Cactus pear is sensitive to dissolved salts in its rooting
cladode rather than to increase in fruit size (Mulas and
zone and therefore the quality of irrigation water needs
D’Hallewin, 1997; Mondragón Jacobo et al., 1995).
to be tested to determine its suitability. Nerd et al. (1991c)
According to Nerd et al. (1989), delay in irrigation during recommend that NaCl in irrigation water for cactus pear
winter when annual rainfall is < 300 mm results in a not exceed 25 mol m−3 to avoid salinity problems. Water
substantial reduction of cladode fertility, and off−sea- harvesting and mulching to improve the productivity of
son winter crop water shortages, particularly during FDP, cactus pear are sound agricultural strategies to be used
may adversely affect fruit quality. Application of 2−3 ir- in arid areas with limited scope for irrigation and to con-
rigations of 30−50 mm each during FDP increase fruit serve soil water. Mondragón Jacobo (1999) showed that
size and fruit pulp percentage of cactus pear (Barbera, where small water microcatchments are made in the area
1984, 1994; Zegbe Dominguez et al., 2015). However, between plant rows, a great deal of runoff is prevented,
irrigation alone cannot compensate for a reduced fruit leaving more water available for the plant and conse-
size when there is a high number of a fruits per cladode, quently resulting in higher fruit yields.
making fruit thinning essential to achieve good fruit size
(La Mantia et al., 1998; Gugliuzza et al., 2002a). Fruit thinning
Nobel (1995) points out that as little as 10 mm rainfall is
adequate to wet the soil in the root zone of cactus pear, Fruit size in cactus pear depends on water availability
resulting in the plant being able to efficiently utilize small (Barbera, 1984), cultivar differences (Potgieter, 2007;
quantities of rainfall. Wessels (1988b) notes that due to Zegbe Dominguez and Mena Covarrubias, 2010b),
the shallow root system of the plant, irrigation amounts length of FDP (Barbera et al., 1992a), mineral nutrition
of 20−25 mm at a time should be adequate. There is lim- (Ochoa et al., 2002) and, most importantly, cladode fruit
ited information on field parameters for appropriate irri- load (Brutsch, 1992; Inglese et al., 1995a). In contrast
gation scheduling, and the amount and timing of water with many other fruit crops, very few cactus pear flow-
application vary substantially between countries (Felker ers abscise and 95% of the flowers that set become
and Inglese, 2003). Recently, Consoli et al. (2013) deter- fruit, unless damaged by late winter frost. However, if
mined the crop factor (Kc) for cactus pear within a range crop load is not reduced by fruit thinning, individual
of 0.5−0.6. Drip irrigation of 150−200 mm water per fruit size is low and whole branches and cladodes may
year seems adequate for the main summer crop in Argen- even break off due to the excessive weight. Fruit prices
tina, while 4−5 irrigations per year are given for a scoz- on local and export markets generally depend on fruit
zolatura crop (Ochoa and Uhart, 2006a). Gugluizza et al. size, with larger fruits selling for higher prices. However,
(2002b) report that 2−3 irrigations (60−100 mm) applied heavy thinning to four fruits per cladode may substan-
during FDP increase productivity and improve fruit quality, tially reduce total fruit yield by as much as 58% without
while two 50−80 mm applications of water during FDP any fruit size increase, and it could even cause a second
are essential to achieve export fruit size in the scozzola- reflowering (Zegbe Dominguez and Mena Covarrubias,
tura crop (Inglese et al., 1995a). According to Homrani 2010a). According to Brutsch (1992), thinned cladodes
Bakali (2013), 3 and 6 irrigations per year produce more produce larger fruit than unthinned cladodes, regardless
fruit than just 1 per year in Morocco. In South Africa, of the number of fruit per cladode. Thus, good fruit size
Haulik (1988) suggess ≤ 3 supplementary irrigations per is achieved with a high fruit−set per cladode followed by
year: beginning in August to stimulate the reproductive timely fruit thinning to reduce the crop load.
flush, then at anthesis, and again in the early stages of
fruit development. Where scozzolatura is practised, the In addition to a higher individual fruit mass (Inglese et al.,
first irrigation is given at floral induction (40 days before 1995a; Nasr, 2015), fruit thinning has other advantages:
budburst), the next after flowering, then ≤ 5 weeks after • easier harvesting (Wessels, 1989);
fruit−set and during fruit maturation (Targa et al., 2013). • prevention of branches from breaking due to a heavy
Different irrigation systems are used in cactus pear orchards crop load (Wessels, 1988a);
with drip and microsprinklers common in modern orchards • reduction of alternate bearing (Wessels, 1988b; Hes-
(Inglese, 1995). Microsprinklers covering a fairly large soil ter and Cacho, 2003);
62 Crop ecology, cultivation and uses of cactus pear

• regular and earlier ripening (Inglese et al., 2002b); • adopting crop manipulation techniques (Barbera et
• increase in total soluble solids (TSS); and al., 1992a; Brutsch and Scott, 1991).
• higher percentage of first class fruits (Zegbe Domin-
Scozzolatura
guez and Mena Covarrubias, 2009, 2010a, b).
The scozzolatura technique − discovered by chance in
Fruit thinning can take place as soon as the spherical the early nineteenth century (Coppoler, 1827, cited by
fruit buds are distinguishable from the elongated vege- Barbera et al., 1991, 1992a) – has since become stan-
tative buds (Wessels, 1988), but no later than 3 weeks dard crop practice in the cactus pear fruit industry in
after anthesis, as later thinning does not improve fruit Italy (Barbera, 1995). Complete removal of all newly
size (Inglese et al., 1995b; La Mantia et al., 1998; Gug- developing flowers and cladodes of the spring flush re-
liuzza et al., 2002a). sults in a second reflowering approximately 12−16 days
later with fruits ripening 6−8 weeks after the spring
Investigations showed that export−sized fruits (> 120
flush (Barbera et al., 1988, 1991, 1992b; Brutsch and
g) can only be produced if no more than 6 fruit per
Scott, 1991). Although the second flush normally sets
cladode are retained (Inglese et al., 1994b). Since not all
fewer flowers than the spring flush, fruit are marketed
cladodes are the same size, the norm in the South Afri-
when prices are higher (Brutsch and Scott, 1991; Bar-
can commercial sector is to thin fruit to approximately
bera and Inglese, 1993; Boujghagh and Bouharroud,
50−70 mm between fruitlets, rather than to a specific
2015), which to an extent compensates for the lower
number per cladode (Potgieter, 2001). Leaving adequate
fruit yield. Scozzolatura brings numerous advantages,
space between developing fruits ensures less damage to
including:
adjacent fruits during the harvesting process, especially
where specialized harvesting secateurs are used. Fruits • increased prices;
that develop on the flat sides of the cladode need to be • improved fruit quality − in particular, improved fruit
removed as they tend to have a long “fruit stalk”, mak- size, lower seed−to−pulp ratio and higher pulp per-
ing packing more difficult. Excess fruits can be removed centage (Barbera et al., 1992b; Hammami et al.,
by hand using a polyvinyl chloride (PVC) glove and a 2015; Boujghagh and Bouharroud, 2015);
sharp knife or pruning secateurs. • greater flesh firmness and more intense pulp color-
ation (Mondragón Jacoboet al., 1995);
Out−of−season cropping
• more complex and compact plant architecture in ad-
dition to more fertile terminal cladodes and higher
Floral induction in most perennial fruit trees is largely
fruit yield (when practised from an early plant age).
synchronized, resulting in a single harvest at a specific
time of the year (Liguori and Inglese, 2015). However, The reflowering index, as defined by the ratio of sec-
one of the most remarkable characteristics of cactus ond flush to first flush flowers (FII : FI), may vary greatly
pear is the capability of the plant to reflower at different depending on the timing of spring flush removal (SFR)
times in the same season (Inglese, 1995; Inglese et al., and the environmental conditions at removal (Inglese,
2002a), naturally or after inductive practices have been 1995). The number of cladodes produced with scoz-
applied (Nerd and Mizrahi, 1997). These out−of−season zolatura may be 10−40% less than the spring flush and
fruit are sold at substantially higher prices than those fruit yield can be as much as 50% lower than in the
of the normal summer season (Mondragón Jacobo et summer season (Nerd et al., 1991b; Inglese, 1995). In-
al., 2009). Successful application of crop manipulation deed, scozzolatura can also have disadvantages, in-
techniques, such as scozzolatura and winter production, cluding:
has considerably increased the provision of fruit from • reduced yield;
5 to 9 months of the year on the local fresh produce • higher peel percentage (Mondragón Jacobo et al.,
markets of South Africa, although the volume is limited 1995) − possibly due to reduced temperatures during
from May to September. Nevertheless, cactus pear fruit FDP (Inglese, 1995; Hammami et al., 2015);
marketing is generally highly seasonal, with cultivars
• lower TSS;
available for approximately 4 months per season (Ing-
lese, 1995; Liguori and Inglese, 2015). Increased fruit • increased peel cracking;
availability on markets could be achieved by: • lower titratable acids; and
• growing cactus pear in diverse agroclimatic areas (Mon- • poorly coloured fruit (Inglese, 1995; Mulas, 1997).
dragón Jacobo et al., 2009; Liguori and Inglese, 2015);
A maximum of 25% of the spring season cladodes
• using cultivars with different ripening periods (Brutsch, should be kept on the plant after scozzolatura, as a
1992; Gallegos Vazques et al., 2006); higher percentage reduces the reflowering rate of the
• improving post−harvest technology (Liguori and Ing- following spring and promotes biennial bearing (Inglese
lese, 2015); and et al., 2002b, 1994a).
Fruit production and post−harvest management 63

Climatic conditions, cultivar response and timing of (Inglese, 1995), California (Curtis, 1977) and Chile (Sud-
flush removal are important factors affecting scozzola- zuki Hills et al., 1993), as well as in the hot subtropical
tura. Environmental conditions at removal time influ- areas of Limpopo, South Africa (Groenewald, 1996; Pot-
ence the degree of reflowering and may cause large gieter, 2001). In addition to the natural out−of−season
annual variation in the reflowering response (Barbera budburst in areas with mild winters, a second flowering
et al., 1991; Nieddu and Spano, 1992). For example, flush can be obtained: Nerd et al. (1993b) and Nerd and
a lower reflowering rate may be obtained if high tem- Mizrahi (1994) showed that following the main summer
peratures coincide with bud initiation which will result crop harvest, immediate irrigation and N application at
in more vegetative than reproductive buds (Nerd et al., a rate of 120 kg ha−1 produced an autumn budburst;
1989; Nobel and Castaneda, 1998). In some countries, production of flower buds increased with increasing
scozzolatura is performed with irrigation (Inglese et al., rates of N application and was highly correlated with
2002a) and N fertilization (Flores Valdez, 2003), or farm- the soluble reduced N content in the terminal cladodes
ers may apply a once−off fertigation at SFR (Nerd et al., (Nerd and Mizrahi, 1994).
1993b). It is essential to select the most suitable cultivar
for the technique, as reflowering may be low or even Although the winter crop gives yields 50–80% smaller
absent in some cultivars (Mondragón Jacobo, 2001; Tar- than the main summer crop (Nerd et al., 1993b), higher
ga et al., 2013). The timing of the SFR affects the extent prices are obtained (Mondragón Jacobo and Bordelon,
of reflowering, the ripening time and fruit characteris- 1996). Groenewald (1996) reported that even without
tics (Barbera et al, 1992b). Inglese (1995) reported re- irrigation, this technique can be successfully applied un-
flowering rates of between 0.7 for pre−anthesis flower der rainfed conditions such as in summer rainfall areas
removal and 0.5−0.3 for post−anthesis flower removal. of South Africa.
Pre−bloom removal produces the highest reflowering Furthermore, the flowering response to N is affected
rate (Brutsch and Scott, 1991), but the latest stage of by the age of the plants. Floral bud production is much
removal normally gives the highest economic returns, higher in young plants (≤ 6 years) than in older plants
although fruit yield may be lower than for other SFR (Nerd and Mizrahi, 1994). However, this technique is
times (Mulas, 1992; Boujghagh and Bouharroud, 2015). only feasible where winter temperatures are sufficient-
In addition to Italy (Barbera et al., 1991), scozzolatura ly high for fruit development (Nerd et al., 1993b). The
is also regularly practised in South Africa (Brutsch and peel−to−pulp ratio is higher in winter than in summer
Scott, 1991), Morocco (Boujghagh and Bouharroud, fruit, due to the thicker peel (Nerd et al., 1993b; Groe-
2015) and Tunisia (Aounallah et al., 2005; Hammami newald, 1996). Producers should take note that cultivars
et al., 2015). In other parts of the world, scozzolatura producing high fruit yields in summer (e.g. ‘American
produced poor results. For example, scozzolatura under Giant’) do not respond at all to the applied N (Groe-
Mexican conditions with ‘Cristalina’and ‘Reyna’ gave newald, 1996). In addition, pruning needs to be delayed
negative results (Mondragón Jacobo et al., 1995); simi- until after winter fruit ripening, by which time flower
larly, Ochoa et al. (2009) reported a very low reflowering buds of the main summer crop have already appeared,
index of 0.05 in Argentina with ‘Amarilla sin espinas’. making pruning difficult (Groenewald, 1996).
Following the scozzolatura performed at SFR, the pro-
cess can be repeated with the complete removal of Orchard sanitation
the first scozzolatura cladodes and fruit (Inglese et al.,
2010). Liguori et al. (2006) demonstrated that the dou- Winter cladode prunings, cladodes that break off during
ble removal of new fruits and cladodes induced a third normal orchard practices and fruitlets removed during
flush of flowers and cladodes in late August with fruit thinning need to be removed from the orchard on a reg-
production ripening in winter (January−March) in the ular basis and destroyed. Pruned cladodes should not
Northern Hemisphere. Winter fruits obtained by double be dumped near the orchard, as detached cladodes and
scozzolatura and covered under PVC polymeric film in thinned fruitlets form roots and begin to grow, serving
late autumn were regular in size and flesh percentage, as host plants for cochineal, Cactoblastis and various
but with slightly lower TSS. However, the rate of the other diseases, which results in increased plant protec-
second reflowering was low (20−40%) (Liguori and In- tion costs (Potgieter, 2001).
glese, 2015). Low temperatures in December stop fruit
growth and ripening; for fuit to develop normally, it is Productivity
necessary to cover the plants with PVC tunnels (Liguori
et al., 2006). The fruit yield of cactus pear is extremely erratic and yields
vary greatly, not only between and within countries, but
Winter fruit production within orchards of the same cultivar. Fruit yields vary from
Flowering of the cactus pear plant is not restricted to 1−5 tonnes ha−1 under traditional methods to 15−30
spring. A smaller budburst occurs naturally in Argentina tonnes ha−1 with intensive orchard practices under rain-
64 Crop ecology, cultivation and uses of cactus pear

fed conditions of 400−600 mm per year (Monjauze and Potgieter (2007) showed that soil P and soil N levels had
Le Houérou, 1965a). Fruit yield is relatively low in most the largest influence on fruit yield of 11 cactus pear cul-
of the plantings in Mexico (2−8 tonnes ha−1 − Pimienta tivars. It is also well known that more than one crop
Barrios, 1990, 1994); however, some irrigated orchards can be obtained for the same environment by natural
may yield 25 tonnes ha−1 (Gallegos Vazques et al., 2009). or artificially induced reflowering (Barbera et al., 1991;
In Chile, fruit yields are generally low (6−9 tonnes ha−1 – Brutsch and Scott, 1991; Nerd et al., 1993b; Sudzuki
Saenz, 1985), while in Argentina they range from 8−11 Hills et al., 1993).
ha−1 (rainfed) to 22 tonnes ha−1 (irrigated) (Ochoa, 2003).
Total fruit yields in excess of 50 tonnes ha−1 have been Genotype characteristics and interaction
reported in the Karoo, South Africa (Brutsch, 1979) and Brutsch (1979), Pimienta Barrios (1990, 1994) and Wes-
Texas, the United States of America (Parish and Felker, sels (1988a, 1989) indicated that cultivars differ in their
1997). In other rainfed areas of South Africa, such as the reproductive vigour and cladode fertility. According to
Free State Province, the highest mean fruit yield obtained Wessels (1989) and Pimienta Barrios (1990), this wide
was 17.44 tonnes ha−1 in a trial comparing 42 cultivars variation in productivity between cultivars is due
(Coetzer and Fouche, 2015). In Israel and Italy, yields of to:
20−30 tonnes ha−1 are regularly reported (Barbera and • inherent genetic fertility differences;
Inglese, 1993; Nerd et al., 1993b).
• fertility of the mother plant; and
If well managed, orchards can have a life span of > • cladode fertility of the mother plant.
100 years, as witnessed in North Africa (Le Houérou, On the other hand, variation within a cultivar is due
1994). Fruit yield is expected to increase yearly from to:
planting through to approximately the fifth production
• variation in fertility of the mother plant; and
year when plants have reached full maturity (Potgieter,
2007). Most of the flowers develop on 1−year terminal • fertility differences between cladodes according to
cladodes, while new cladodes usually develop on 2− position on the mother plant.
year or older cladodes (Inglese et al., 1994a; Wessels, According to Barbera (1995), large variances in cactus
1988a). The fertility of the cladodes depends on envi- pear fruit yield are due to inadequate understanding of
ronmental conditions, plant age and dry matter (DM) plant × environment interaction. In an 8−year field
accumulation (García de Cortázar and Nobel, 1990; In- trial with 11 cactus pear cultivars in three diverse agro-
glese et al., 2002b; Valdez Cepeda et al., 2013). Clado- climatic areas in South Africa, Potgieter (2007) showed
des with a higher than average DM content tend to pro- that there were significant differences between cultivars,
duce more fruits (García de Cortázar and Nobel, 1992). diverse environments and production years in terms of
The possible reasons for high variability in fruit yield lie fruit yield and its components. The variance observed in
in four main areas: environmental conditions, genotype fruit yield was due first to soil P levels and second to ap-
characteristics and their interactions, orchard planning plied N. The study results demonstrate clearly not only
and design, and orchard practices (Nerd et al., 1991b; that environmental factors have a definite influence on
Inglese, 1995; Inglese et al., 2002a; Potgieter 2007). fruit yield, but that there is strong interaction between
the 11 cultivars tested and the environmental condi-
Environmental conditions tions. Broad cultivar adaptation was only seen in one
In contrast to vegetative growth, little is known about cultivar and yielding ability in some cultivars is a genetic
the influence of environmental factors on cladode fer- trait rather than a G × E response (Potgieter, 2007).
tility and fruit yield in cactus pear (Inglese et al., 1995a;
Nerd and Mizrahi, 1995b). According to Barbera et al. Orchard planning and design
(1991), García de Cortázar and Nobel (1991), Nerd and Fruit productivity in cactus pear can be improved by in-
Mizrahi (1995b) and Inglese et al. (2002a), cladode fer- creasing the number of fertile cladodes per plant and/or
tility depends on environmental conditions such as plant by increasing the plant population (Inglese et al., 2002a).
water status, temperature, photosynthetic photo flux Extremely high fruit yields at an early plant age can be
density (PPFD) and soil nutrients. Wessels (1989) related obtained at high plant densities. In Israel, spacing plants
seasonal variation in fruit yield to: 1.5 m in the row and 4 m between rows (1 666 plants
ha−1) substantially increased the number of fertile clado-
• differences in agroclimatic conditions (chill require-
des in the early stages of orchard life, with fruit yields of
ment, rainfall, temperature);
18 tonnes ha−1 for 4−year trees reported (Nerd and Miz-
• differences in soil fertility status; and rahi, 1993). According to Inglese et al. (2002a), to obtain
• poor pollination and fertilization due to absence of an annual yield of 20 tonnes ha−1 − with an average fruit
pollinators and unsuitable climatic conditions during weight of 100−120 g and cladode fertility of six fruit per
the pollination period (cold, rain). cladode after thinning − 28 000−30 000 fertile cladodes
Fruit production and post−harvest management 65

per hectare are needed. This implies 80−90 fertile clado- HARVESTING
des per plant on free−standing plants spaced 6 × 5 m
apart (335 plants ha−1) or 28−30 fertile cladodes per plant Quality is a very important factor in fruit production,
for high−density hedgerow plantings with plants spaced since consumers prefer attractive fruit with a good taste
5 × 2 m apart (1 000 plants ha−1) (Inglese et al., 2002a). In and high nutritional and functional value. The overall
order to further increase fruit yield, it would be necessary quality is generally highest at harvest; it then declines at
to increase the number of fertile cladodes rather than in- rates which vary according to genetic background, pre−
crease cladode fertility (Inglese, 1995). harvest treatments, environmental conditions, degree of
maturity at harvest, handling processes, post−harvest
Orchard practices treatments, and storage and distribution conditions.
Large variations in fruit yield are regularly observed, even Overall quality includes a very complex set of features
in well−managed orchards of the same cultivar (Potgiet- not always positively correlated: depending on the
er, 2007). The low fruit yields reported in Mexico are targeted consumers, market destination and planned
partially due to the fact that a large percentage of grow- storage time, the importance of the various qualitative
ers do not use cultural practices such as fertilization and aspects varies. As the fruits mature, their nutritional val-
pruning, resulting in poor vegetative growth and low ue, flavour and taste improve, but the tissue’s natural
fruit yields (Pimienta Barrios, 1994). In comparison, the defence mechanisms against pathogens, susceptibility
relatively high yields obtained in Sicily (14 tonnes ha−1) to some physiological disorders and potential life span
are mainly due to irrigation, fertilization and thinning all decrease. Therefore – as with other species (Crisosto
(Tudisca et al., 2015). and Valero, 2008) – for direct delivery to local markets,
Biennial or alternate bearing has been reported in cactus harvest should take place when the highest eating qual-
pear (Brutsch, 1979; Pimienta Barrios, 1990) and it is ity is reached; for delivery to distant markets, earlier har-
one of the reasons for large differences in annual fruit vesting is more appropriate to prolong the post−harvest
yield. Inglese et al. (1995) and Brutsch (1979) noted that life span.
the possible reasons for alternate bearing are: In order to identify the best harvest time, objective and
• incorrect pruning; subjective maturity indexes have been developed, based
on factors such as cultivar, producing country, fruit des-
• cultivar differences;
tination and utilization. The most popular harvest matu-
• plant age; rity indexes include:
• competition between floral and vegetative growth; • percentage of peel colour−break;
and
• total soluble solids (TSS) ≥ 13%;
• bud induction timing.
• pulp firmness (measured with an 8−mm plunger) ≥ 8
Nevertheless, farmers prefer a cultivar that bears consis- kg cm−2 (Pimienta Barrios, 1990; Barbera et al., 1992);
tently − even if at a lower yield level − rather than a cul- • reducing sugar level around 90% of that of full ripe
tivar that bears well one year and poorly the next, as this fruit – however, in some cultivars the reducing sugars
pattern has a serious economic impact which negatively never exceed 50% of the total sugars (Pimienta Barri-
affects the cash flow position of the enterprise (Potgiet- os and Mauricio, 1989);
er and Smith, 2006; Potgieter, 2007). Indeed, compe-
• abscission of glochids;
tition between reproductive and vegetative growth, as
well as reduction in the number of new cladodes follow- • flattening of the floral cavity of the receptacle;
ing SFR, are potential sources of plant alternate bearing • percentage of pulp;
behaviour (Inglese et al., 2002b). Although Barbera et • peel thickness and ease of removal; and
al. (1991) found that alternating plants in the off−year • peel resistance to physical handling (Cantwell, 1995).
had the same number of 1−year cladodes as in the on−
year, most of these cladodes were unfertile under scoz- Cactus pears are particularly difficult to harvest because
zolatura conditions. Practical approaches for reducing of the presence of glochids and spines, which can pierce
alternate bearing are to: the skin and enter the eyes and respiratory tract. Fruit
• adopt appropriate pruning systems (García de are therefore harvested in the morning when humidity
Cortázar and Nobel, 1992); is sufficiently high to prevent glochids from dislodging
and floating in the air. Pickers should be provided with
• ensure fruit thinning regimes (Wessels, 1989; Hester protective clothing (gloves and safety glasses). Despite
and Cacho, 2003); and the plant’s tough appearance and its ability to withstand
• avoid scozzolatura and winter production on the same harsh environmental conditions, the fruit are very tender
orchards every year. and cannot withstand rough treatment (Wessels, 1992a).
66 Crop ecology, cultivation and uses of cactus pear

For most cultivars, the physiological loosening of the Larger packing houses comprise the following:
articulation connecting the fruit to the mother clado- • dumping area;
de is low at harvest time, and injury at the stem end is
• moving conveyer belt − workers preselect fruit;
inevitable if the fruit is harvested by snapping, pulling
or twisting. Therefore, for commercial purposes, a knife • despining section − designed with the same criteria as
must be used, a sharp cut made at the base of the fruit for a small−scale packing house;
and a small piece of cladode left attached. Pickers usu- • sizing devise − either mechanical or electronic;
ally collect the fruit in plastic baskets or lugs and empty • delivery and packing devise − for final sorting and
them into plastic boxes holding 15−20 kg of fruit for packing.
transport to the packing house.
Fruit are usually packaged on the day of harvest and
Physical damage during harvest and transport can mark- directly delivered to destination markets in refrigerated
edly compromise fruit quality and storage length and conditions. They can be transported alone or in com-
increase fruit susceptibility to physiological disorders bination with other commodities, by truck, ship or air-
and decay. Bruises and wounds can occur due to finger craft. When fruit are handled a few days after harvest,
pressure when cutting the fruit or impact when fruit are they can be stored at ambient conditions for curing
dropped into the baskets, and also during transport and or in storage rooms at 6−10 °C. In a small number of
handling in the packing house. Injuries are also inflict- countries (e.g. South Africa), fruit are waxed to replace
ed by cutting and by glochids. The susceptibility of fruit
the natural waxes lost with despination, in order to re-
to physical injuries increases with maturity. High cellu-
duce transpiration and enhance skin gloss. No synthetic
lar pressure can also cause cracks and microcracks of
fungicide is registered for post−harvest purposes; there-
the peel, especially in second−crop fruit, which ripen in
fore, much care must be taken to avoid injuries and pre-
more humid environmental conditions.
vent microbiological decay.
Post−harvest handling In Italy, fruit grading is based on:
• cultivar (‘Gialla’ or ‘Surfarina’; ‘Red’ or ‘Sanguigna’;
The presence of spines and glochids is one of the main and ‘Bianca’ or ‘Muscaredda’ or ‘Sciannarina’);
constraints limiting cactus pear consumption and mar-
• category (EXTRA and I); and
ketability; worldwide, their removal is the primary post−
harvest operation before commercialization. In many • weight (class B, 105−140 g; class C, 140−190 g; class
countries, especially with fruit destined for the local D, 190−270 g).
market and consumed within a few days of harvest, de- Depending on the cultivar, peel shades can range from
spination is still done manually: the fruit are spread on green to orange–yellow for the yellow cultivar (‘Gial-
the grass or areas covered with straw, and then the fruit la’), from green to ruby−red for the red cultivar (‘Rossa’)
are brushed with brooms (Cantwell, 1995). However, and from green to straw−white for the white cultivar
with fruit destined for distant markets, despination is (‘Bianca’). Depending on producing area, fruit destined
done in packing lines. for the fresh market must have the following charac-
Unlike other kinds of fruit, post−harvest operations of teristics:
cactus pear are quite simple and restricted to despina-
• weight ≥ 120 g
tion, grading and packing. Regardless of the scale of the
packing house, despination is generally accomplished • TSS ≥ 13−14% °Brix
by dry brushing. Given the limited post−harvest treat- • flesh firmness ≥ 6 kg cm2
ments, the production of several associated growers
Larger fruit can be packed in one−layer plastic nest trays
can be handled in a small−scale on−farm packing house
inserted in carton or plastic trays or directly in carton
with a small, simple packing line comprising the fol-
trays. Small fruit, generally destined for local markets,
lowing components:
are packaged in plastic trays or punnets containing 6−8
• dumping devise − fruit are dry−dumped before pass- fruits.
ing onto a series of rollers;
• tunnel − a series of brushes, each one rotating in the Post−harvest physiology
opposite direction to the next, remove the glochids,
which are either vacuum−sucked out of the unit and Cactus pears are classified as non−climacteric, as they
deposited in a disposable bag or left to drop beneath do not exhibit a rise in respiratory activity during the rip-
the rollers; ening period. Respiration rates are generally considered
• second set of rollers − fruit are conveyed onto a large quite low compared with other fruits (Lakshminarayana
round rotating table, where workers sort, grade and and Estrella, 1978; Cantwell, 1995). However, respirato-
package them. ry intensity is strongly affected by:
Fruit production and post−harvest management 67

• genetic background (quite wide in cactus pear); Physiological disorders

• ripening stage at harvest; Similar to most species of tropical origin, cactus pear
is susceptible to chilling injury when exposed for pro-
• crop type; and
longed periods to temperatures below 10−12° C. In-
• environmental conditions. deed, the intrinsic sensitivity of fruit to low tem-
In a study of different varieties from the Mexican germ- perature is markedly affected by:
plasm conducted by López Castañeda et al. (2010), re- • environmental conditions;
spiratory activity ranged from 22 in ‘Rojo Pelón’ to 31 ml • agronomic practices;
CO2 kg−1 hour−1 in ‘Sangre de Toro’ and ‘Alfajayucan’. In
• crop type (summer or late crop); ripening stage at har-
studies conducted in Italy with fruit from the first crop of
vest; cultivar; and
‘Gialla’, respiration intensity showed very pronounced
variability from year to year, ranging from 4−14 mg CO2 • post−harvest treatment.
kg−1 hour−1 (Chessa and Schirra, 1992) to 13 ml CO2 Generally fruit at an advanced stage of maturity are less
kg1 hour−1 (D’Aquino et al., 2014) and 60−92 mg CO2 susceptible to chilling injury than less ripe fruit, while
kg−1 hour−1 (Piga et al., 1996). As in other non−climac- pre−harvest treatment with gibberellic acid (Schirra et
teric species, respiration declines gradually when fruit al., 1999a) or calcium chloride (Schirra et al., 1999b) do
are held at warm temperatures (D’Aquino et al., 2014), not affect or increase fruit susceptibility to chilling injury.
but when moved from cold storage to room tempera- Symptoms of chilling injury can manifest on the peel as
ture, respiration increases greatly (Schirra et al., 1999b) scalded or bronzed areas varying in size and intensity,
in response to chilling temperature. Incipient infec- black−brownish pits and sunken brown spots (D’Aqui-
tions from microorganisms, and physical or mechanical no et al., 2012). However, chilling injury can also cause
stresses caused by bruising, impact or wounds, can also metabolic alterations in addition to visible symptoms;
lead to sudden increases in respiratory activity. Ethylene likewise, the appearance and severity of peel disorders
production is very low, generally 0.1−0.2 µl C2H4 kg−1 are accompanied by non−visible qualitative alterations.
hour−1 and, as for respiration, significant increases may Indeed, peel alterations are usually a combination of
occur after prolonged exposure to chilling temperatures, disorders induced by low temperature and other caus-
or because of pathogen infections or abiotic stresses es (superficial wounds inflicted by glochids, excessive
(D’Aquino et al., 2104; Schirra et al., 1996, 1997). transpiration, dry rots) that can appear at non−chilling
As with most non−climacteric fruit, cactus pear does temperatures (D’Aquino et al., 2012, 2014). Metabolic
not contain starch. Therefore, after harvest, TSS, sug- imbalances may alter the respiratory metabolism, induce
ars and organic acids tend to decrease, depending on the production of undesirable volatiles (acetaldehyde,
storage conditions, ripening stage and cultivars; the pat- ethanol) and ethylene, and lower the overall host de-
tern of decline can be gradual or uneven. For example, fence mechanisms to pathogens (Schirra et al., 1999a;
in a comparative study with six cultivars of cactus pear D’Aquino et al., 2014). As a result, chill−injured fruit
stored at 9 °C for 1, 2 or 3 months plus 4 days at room may show peel disorders without any change in eating
temperature, the loss of TSS was very high in ‘Amaril- quality or chemical composition (D’Aquino et al., 2014);
lo Montesa’, ‘Copena T−5’ and Copena−Torreoja’, but or they may show negligible symptoms of chilling injury
gradual in ‘Cristalina’ and ‘Pinochulo’, while in ‘Burrona’ at the end of cold storage but become highly suscepti-
losses were highest during the first month of storage ble to decay when moved to warm temperatures (Schir-
(Corrales Garcìa et al., 1997). ra et al. 1996, 1997, 1999b; D’Aquino et al., 2014).

Vitamin C content at harvest ranges from about 10 to Post−harvest decay

80 mg 100 g−1 depending on the cultivar (Butera et al., Fruit composition and tissue consistency make cac-
2002; Kuti, 2004). It can vary greatly from year to year tus pear highly susceptible to decay incited by various
(Sumaya Martìnez et al., 2011), but also depends on the pathogenic fungi, yeast and bacteria. The stem end is
ripening stage. In fruit of yellow and orange cultivars of the main site of infection due to wounding caused by
Opuntia megacantha harvested at 2−week intervals, 4 harvesting operations. However, a curing treatment of
weeks before commercial maturity and then 4 weeks 1−2 days at room temperature in ventilated conditions
after commercial maturity, the vitamin C content in- helps heal wounded tissue. Even a slight dehydration
creased constantly even when fruit were overripe (Coria of the little piece of cladode left at harvest can effec-
Cayupàn et al., 2011). In fruit stored at low tempera- tively reduce the risk of decay (Cantwell, 1995; Inglese
ture, the vitamin C content is quite stable despite the et al., 2002a). Other sites of infection are the microw-
relatively high pH of the juice (Schirra et al., 1996), but ounds caused by glochids and microcracks in the peel,
it declines rapidly at room temperature (D’Aquino et especially if second−crop fruit are harvested under wet
al., 2014) or after fruit are moved from cold storage to conditions. The main filamentous fungi that cause decay
warm temperatures (Schirra et al., 1996). are Botrytis cinerea, Fusarium spp., Aspergillus spp. and
68 Crop ecology, cultivation and uses of cactus pear

various Penicillium species, including P. digitatum, P. ital- controlled decay when applied at 20 or 50 °C before
icum, P. expansum and P. polonicum (Chessa and Bar- storage; on the other hand, its effectiveness was re-
bera, 1984; Rodriguez Felix et al., 1992; Granata and duced when it was applied at the end of cold storage
Sidoti, 2000; Swart et al., 2003; D’Aquino et al., 2015; (D’Aquino et al., 2015).
Faedda et al., 2015a). However, since fruit are generally • Hot water − either as dip treatment at 50−55° C for
not stored for long periods, the risk of decay does not 2−5 minutes (Schirra et al., 1996, 2002; Rodriguez et
represent a serious problem. al., 2005; D’Aquino et al., 2012) or by water brushing
at 60, 65 or 70° C for 30, 20 or 10 seconds, respec-
Post−harvest treatments tively (Dimitris et al., 2005) − inhibited the growth of
Despite limited treatments at commercial level and oc- pathogens naturally present on the fruit surface.
casional refrigeration for short periods, the increasing
• Curing (at 38° C for 24, 48 and 72 hours in a vapour−
worldwide demand for cactus pear − especially from
saturated environment) not only controlled decay, but
markets located far from producing areas − will pose
also improved fruit sensitivity to chilling temperatures
new challenges for the cactus pear industry in the fu-
(Schirra et al., 1997). Yet, curing the fruit at 38° C
ture. In order to extend the market window beyond the
with 75−80% relative humidity (RH) hastened detach-
harvest season and delay the decline in appearance,
ment of the piece of mother cladode left at harvest.
appropriate post−harvest treatments and handling
Additionally, the healing process of the stem−end scar
procedures are required to:
with respect to curing at the same temperature but
• reduce transpiration and respiration rates; with 100% RH decreased decay incited by pathogens
• increase fruit tolerance to chilling temperatures; and developing from the stem (D’Aquino et al., 2014).
• prevent microbiological spoilage. High temperature conditioning delayed fruit ageing
and weight loss, possibly due to melting and rear-
While refrigeration is undoubtedly the main means for rangement of the epicuticular wax layers with conse-
prolonging the post−harvest life of fresh fruit and vege- quent filling of microcracks separating wax platelets,
tables, the susceptibility of cactus pear to chilling injury the main fruit transpiration pathway (Schirra et al.,
poses limits to its use. On the other hand, as with other 1999a; López Castañeda et al., 2010).
commodities, as storage life increases, the natural de-
fence mechanism of tissue against pathogens declines Other treatments which can delay loss of freshness and
and fruit become progressively more susceptible to mi- increase tolerance to low temperature are: controlled
crobiological attacks, especially when moved from cold atmosphere storage (Testoni and Eccher Zerbini, 1990);
storage to a warm environment. intermittent warming (Chessa and Schirra, 1992); sali-
cylic acid dip treatments (Al Qurashi and Awad, 2012);
Experiments conducted with post−harvest fungicides and film wrapping (Piga et al., 1996, 1997; Shumye et
approved for other commodities revealed various de- al., 2014), although the latter may induce anaerobic
grees of effectiveness in reducing decay incidence in conditions and a build−up of undesirable volatiles (Piga
cold−stored cactus pears: et al., 1996) if film permeability to gases does not match
• Benomyl, captan or vinclozolin treatments were inef- the O2 requirement of packaged fruit.
fective in controlling post−harvest decay (Gorini et al.,
1993).
• Imazalil (IMZ) and thiabendazole (TBZ) treatments READY−TO−EAT CACTUS PEAR
prevented natural decay induced by Penicillium spp.,
Botrytis cinerea and Alternaria spp. in first−crop ‘Gial- Over the last two decades, the fresh−cut fruit market
la’ fruit over a 2−month storage period at 8° C fol- has seen steady growth due to consumers’ tendency to
lowed by 1 week of simulated marketing conditions eat healthy and convenient foods at any moment and
at 20° C; chilling injury symptoms were also reduced in any place. Ready−to−eat fruit and vegetables are at-
(D’Aquino et al., 1996). Sodium ortophenilphenate, tractive because the consumer requires no labour and
either alone or in combination with TBZ or IMZ, was generates no waste through peeling and coring (Rojas
phytotoxic, resulting in increased decay and weight Graü et al., 2011). This is particularly relevant in the case
loss (D’Aquino et al., 1996). TBZ efficiency was mark- of cactus pear, where the presence of glochids makes
edly increased when applied at 52° C even at a con- the fruit difficult to peel, especially for people not fa-
centration six times lower than that applied at 20° C miliar with it.
(Schirra et al., 2002). Fresh−cut fruit and vegetables are very perishable prod-
• Fludioxonil, a synthetic fungicide registered over the ucts; wounds caused by processing operations stimulate
last decade to control a wide range of decay−caus- respiration and ethylene production rates, hastening the
ing fungi in different commodities, very efficiently loss of respirable substrates, firmness and senescence.
Fruit production and post−harvest management 69

Damaged tissues are also subject to oxidative browning O2, coupled with increased concentration of CO2, may
due to the enzyme polyphenol oxidase (PPO) (Beaulieu lead to anaerobic conditions and in turn induce produc-
and Gorny, 2004). The increasing demand for fresh−cut tion of undesirable volatiles, compromising the sensory
cactus pear has had a marked impact on companies in- quality and altering the microbiological population both
volved in processing and distribution: more attention is quantitatively and qualitatively (Piga et al., 2000; Del
paid to hygienic requirements and new packing solu- Nobile et al., 2009).
tions are adopted to meet logistics and consumer re-
Coatings based on sodium alginate, agar and fish pro-
quirements (Timpanaro et al., 2015a).
tein gel did not affect yeasts and mesophilic bacteria but
The main factors affecting the quality of fresh−cut stimulated the load of lactic acid, psycrothrophic and
cactus pear are loss of acidity, firmness, juice leakage coliform bacteria (Del Nobile et al., 2009). When acetic
and, above all, microbiological spoilage. Spoilage is a acid was combined with chitosan, an overall decrease in
considerable hazard, especially when there is contami- the microbial population was achieved compared with
nation by pathogenic microorganisms with potentially the control fruit (Ochoa Velasco and Guerrero Beltrán,
harmful effects on consumer safety (Yahia and Sàenz, 2014). Coating fresh−cut cactus pear is not common
2011). When fruit are stored at the optimal temperature at commercial level and the literature indicates no clear
of 4−5°C, TSS are quite stable and sometimes increase; and consistent benefits from their application (Del Nob-
on the other hand, titratable acidity and juice pH are ile et al., 2009; Ochoa Velasco and Guerrero Beltrán,
quite stable but may decrease (Piga et al., 2000). High- 2014; Palma et al., 2015).
er temperatures reduce the potential storage life, while
Storage response may also be affected by the stage of
increased O2 requirements due to increased metabolic
ripeness and the ripening time of the fruit. The overall
activity may not be matched by the packaging perme-
quality of the summer crop declines faster than that of
ability to gases, leading to abnormal increases in titrat-
the scozzolatura crop, while fruit harvested at the com-
able acidity, ethanol and taste alterations as a result of
mercial stage maintains its quality longer than fruit har-
anaerobic respiration (Piga et al., 2000).
vested later, when fully ripe, especially in the case of
Its low acidity and high sugar content make cactus fruit coming from the scozzolatura crop (Allegra et al.,
pear − more than any other fruit − an ideal substrate 2015). Both aerobic mesophilic bacteria and mould pop-
for microbiological proliferation. In order to maintain ulation are affected more − albeit in different ways − by
bacteria and yeast populations below the legal limits of the stage of ripeness than by the time of ripening, with
107 and 108 CFU g−1 established in Spanish legislation fruit harvested at an advanced stage of maturity show-
(BOE, 2001), it is essential to use appropriate process- ing a higher load than fruit harvested at the commercial
ing equipment and adopt an effective sanitizing pro- stage (Allegra et al., 2015).
gramme, with personnel trained in hygienic processing
and maintenance of low storage temperatures. Surface
sterilization of the fruit surface prior to peeling is gen-
erally achieved by dip treatments in sodium hypochlo-
CONCLUSIONS AND FUTURE PERSPECTIVES
rite; however, new alternatives that are safer for work-
While there has been a general improvement in orchard
ers and consumers as well as environmentally friendly
practices over the past two decades, much remains to
and cheap, such as electrolyzed water (Pannitteri et al.,
be done to convince producers that cactus pear can
2015), are gaining in popularity.
achieve high yields and good quality if it receives appro-
Provided processing operations maintain low initial priate care and attention − just like any other crop. It is
microbiological load, storage temperature is the main hoped that by providing the latest technical and scientif-
factor affecting the microbiological population. The rec- ic information on the cultivation and post−harvest man-
ommended storage temperature range of 8−12° C to agement of the crop, productivity levels and especially
prevent chilling injury of the whole fruit is not optimal fruit quality standards worldwide will improve, enabling
for storage of ready−to−eat cactus pear; indeed, the the fruit to compete on an equal footing with other
best results in terms of maintenance of overall chemical, mainstream fruits on international markets. In order to
sensory and microbiological quality are at 4−5° C (Piga attract new consumers to cactus pear fruit and create
et al., 2000; Corbo et al., 2004; Del Nobile et al., 2007; higher demand, consistent high−quality fruit must be
Cefola et al., 2014). The permeability of the film used available on the market. Increased fruit productivity is
for packaging and in−package gas composition have easier to achieve than improved fruit quality; special
a minor impact on chemical composition and micro- attention should be therefore given to all horticultural
biological population when fruit are stored at 4−5° C. practices potentially affecting fruit quality, both pre−
However, temperature increments and reduced levels of and post−harvest.
 70 Crop ecology, cultivation and uses of cactus pear

Figure 1
Cumulative growth
cDespination of cactus
pear fruit by rotating
brushes: besides
removing glochids the
brushed make the fruit
shiny.

Figure 2
Sorting table used to
sort, grade and pack
the fruit

Figure 3
Chilling injury as scald
staining (a) and tissue
depression underneath
the peel around the
receptacle scar (b).

Figure 4
Chilling injury
symptoms in the form
pits (a) and brownish
scars (b).

Figure 5
Fruit splitting occurring
in fruit stored at high
humidity levels.

Figure 6
Soft rot on cactus pear
starting from the
stem end.
Fruit production and post−harvest management 71

Figure 7
Decay caused by
Penicillium spp. in cold
stored cactus pear.

Figure 8
Dry rot (Alternaria
spp.) develops slowly in
cactus pears stored for
long time.

Figure 9
Preparation of ready−
to−eat cactus pears:
fruit are first manually
peeled and placed in
plastic punnets (a) and
then sealed with a
polymeric film (b).
 07

Forage production
and supply for animal
nutrition
Jose C.B. Dubeux Jr.a, Hichem Ben Salemb and Ali Nefzaouic
a
University of Florida, North Florida Research and Education Center, Marianna,
United States of America
b
National Institute of Agricultural Research of Tunisia, University of Carthage, Ariana, Tunisia
c
International Center for Agricultural Research in the Dry Areas, Tunis, Tunisia
 07 Forage production and supply
for animal nutrition

INTRODUCTION The potential of cactus, however, remains under-

exploited. Major cactus species used as fodder
Livestock production remains the main source of in- include Opuntia ficus−indica Mill. and Nopalea
come for rural populations living in drylands. It is a cochenillifera Salm−Dyck., with several varieties
key component of resilient production systems and released in different countries.
is an indicator of wealth. However, the sector faces There are reports of the successful use of cactus
many challenges including feeding constraints and as a feed source in countries ranging from Brazil
climate change. Rangelands in semi−arid regions and Mexico to South Africa and Tunisia, usually
play an important role in livestock production sys- supported by strong research and extension
tems, although their contribution to animal feed- programmes. To achieve success, it is essential to
ing is decreasing. Rangeland productivity is usu- involve all stakeholders in the livestock production
ally low (< 5 tonnes dry matter [DM] ha−1 year−1) chain, including producers, suppliers, retail stores,
with low yields of consumable biomass (< 1 tonne research and extension institutions, and policy−
DM ha−1 year−1), leading to low carrying capacity makers. The livestock feed market is growing, and
(12−15 ha to sustain an adult cow) (Dubeux et al., it involves fewer risks than the fruit and vegeta-
2015a). Adapted perennial species are a potential ble market, representing, therefore, potential for
option for improv- entrepreneurial activity. In some countries, cactus
ing fodder availabil- cladodes from pruned fruit orchards can be used
ity in dry areas. The as livestock feed, complementing the farmer’s
global livestock pop- income. In general, the forage potential of cactus
ulation has stead- in semi−arid areas is underutilized. There are tre-
ily increased in re- mendous opportunities to develop cactus−based
cent decades, often livestock production systems, promoting human
resulting in range- livelihood and reducing the pressure on natural
land degradation. rangelands.
Water scarcity is an-
other important lim-
iting factor in dry-
CACTUS−BASED FOR AGE
lands, threatening
the sustainability of PRODUCTION SYSTEMS
livestock−based sys-
tems. Global pro- Given the lack of information on cactus surface
jections indicate that water use will increase in the areas in different agro−ecologies, it is not possible
Figure 1 coming decades; there will, therefore, be less water to make a full assessment of the importance of
Wild cactus in San Luis available for agriculture and livestock production. cactus within the various production systems. For
Potosí, Mexico example, data on native cactus or the role of cac-
(photo credit: In this scenario, cactus becomes one of the most tus in defensive hedges and copses are important,
Ali Nefzaoui) prominent crops for the twenty−first century. but rarely reported.
Cactus, a succulent and drought−tolerant species,
can produce > 20 tonnes DM ha−1 year−1 and pro- Native
vide 180 tonnes ha−1 year−1 of water stored in its
cladodes, representing a cost−effective option for The largest area of native cactus is located in Mex-
livestock watering (Dubeux et al., 2015b). At such ico and covers an estimated 3 million ha. Livestock
levels of productivity, it is possible to produce often go to graze in these areas and no special
sufficient forage to sustain five adult cows per treatment is applied. In some countries, cactus
year − at least a 60−fold increase over rangeland species may become invasive and plantations are
productivity. With small, intensively cropped cac- used in a similar way to native plantations − for
tus orchards, it is possible to produce feed and example, in Tigray, Ethiopia, livestock graze direct-
reduce the pressure on overstocked rangelands. ly the thorny cactus.
Forage production and supply for animal nutrition 75

Defensive hedges and copses the early 1930−1940s. Le Houérou (2002) reports
that fodder plantations were systematically devel-
Little is known about this form of cactus planta- oped, in particular in Tunisia, on the basis of the
tion, but it clearly has an important role in many research conducted by Griffiths and collabora-
countries, especially in North Africa and parts of tors in Texas, United States of America. In 1932,
Italy and Spain, where many farms are protected the Government of Tunisia invited Griffiths and
by biological fences using thorny cactus. applied his 30 years of experience on the use of
cactus as fodder in central Tunisia to mitigate the
In addition to their efficient defensive role, these
effects of drought on livestock. Furthermore, land
hedges have traditionally played an important
allotment was authorized in central Tunisia on
part in landscape organization and the local so-
condition that the contracting beneficiaries, inter
cio−economy as evidence of land rights and land
alia, planted 10% of the land allocated to spine-
ownership in countries or regions where collective
less cactus to act as an emergency buffer and
land ownership is prevalent, such as in North Af-
standing fodder crop reserve. This proved a wise
rica. Cactus hedges are often planted as proof of
decision, because the country faced 3 years of se-
land ownership. Moreover, they contribute to ero-
vere droughts during 1946−1948, when 70−75%
sion control, particularly when established along
of livestock were decimated – but not on farms
contours (Le Houérou, 2002).
with cactus plantations (Le Houérou, 2002).
Cactus plantations also take the form of a copse
From an agronomic view point, in order to reha-
around rural houses. This kind of “close−by” plan-
bilitate or improve rangeland, shrubland, bush-
tation is a source of feed for family livestock, as
land or poor farming areas characterized by
well as providing fruit for self−consumption and
shallow, stony, steep or sandy soil or where the
shelter for family poultry.
climate is too dry for practical farming, an appro-
priate planting strategy must be adopted: density
Mixed fruit−forage
of 1 000−2 000 single or double cladodes per ha,
with a spacing of 5−7 m between rows and 1−2
This is the most common and most widespread
m within rows. In general, no special treatments
system. It is prevalent in almost all countries where
(e.g. fertilizer application, pruning and treatment
environmental conditions allow cactus growing and
against pests and diseases) are applied. Supple-
cropping cactus is part of the local knowledge and
mental irrigation may be applied during establish-
tradition. There are two main types of plantation: in-
ment − but only if the first year is too dry. Plan-
tensive specialized fruit orchards, where the produc-
tations are exploitable after 3−4 years and fully
tion objective is to produce good quality fruits for
grown after 7−10 years; if rationally managed,
the local market or export; and orchards with low
they can remain productive for > 50 years.
inputs, where fruits are mainly for self−consumption
or the local market. In both cases, pruning provides The productivity of a rangeland planted with cactus
large amounts of cladodes that are sold and/or uti- can be increased by a factor of between 1 and 10
lized at farm level to feed the livestock. when very degraded and between 1 and 5 when in
good condition (Le Houérou, 2002); similar results
Rangeland rehabilitation are reported by Nefzaoui and El Mourid (2009).
Impressive results are obtained with fast−growing
Rangeland improvement using spineless cactus shrubs (Acacia cyanophylla) or cactus (Opuntia fi-
has been practised mainly in North Africa since cus−indica) in central Tunisia (Table 1).

TABLE 1 Productivity of natural and improved rangelands in Tunisia (Nefzaoui and El Mourid, 2009)
Productivity (forage unit per hectare
Rangeland type
and per year)a

Natural rangeland in Dhahar Tataouine, Tunisia (100 mm rainfall) 35−100

Private rangeland improved by cactus crop in Ouled Farhane, Tunisia

800−1 000
(250 mm rainfall)
Cooperative rangeland improved through Acacia cyanophylla, Guettis,
400−500
Tunisia (200 mm rainfall)
a
One forage unit is equivalent to 1 kg barley grain metabolizable energy (12.4 MJ kg−1 DM)
76 Crop ecology, cultivation and uses of cactus pear

Few plant species are able to increase land productivity ble, conservation−oriented manner. Alley designs can also
at such a high rate, especially on marginal lands charac- make better use of the space available between trees and
terized by low rain−use efficiency (RUE, measured in kg add protection and diversity to agricultural fields.
DM ha−1 year−1 mm−1), for example (Le Houérou, 1984):
There is a low adoption rate of cactus orchards as mon-
• degraded Mediterranean rangelands: RUE = 1−3 oculture for various reasons, from the technical design of
• rangelands in good condition: RUE = 4−6 plantations and mismanagement to competition for land
• desert rangelands: RUE = 0.1−0.5 often dedicated to cereal crops. However, alley cropping
overcomes some of these constraints because it:
Cacti, however, have a high RUE. Rangeland rehabili-
tated with O. ficus−indica exhibits a RUE of 10−20 kg • improves soil fertility;
of above−ground DM ha−1 year−1 mm−1 in arid areas • increases crop yield;
where rainfall is 200−400 mm year−1. • reduces weeds; and
• improves animal performance.
Alley cropping
Properly managed alley cropping allows diversification
Expansion of cereal cropping into rangelands and and farmers can benefit from several markets. It also
reduction in fallow practice are among the main rea- promotes sustainability in both crop and livestock pro-
sons for declining soil fertility and wind erosion. One duction by increasing land productivity and reducing
way of combating degradation resulting from cereal weather risks compared with cultivation of annual crops
monocropping is the introduction of adapted forage only. The benefits of cactus−barley alley cropping were
legumes, fodder shrubs/trees and cactus in cropping evaluated in Tunisia (Alary et al., 2007; Shideed et al.,
systems (Nefzaoui et al., 2011). 2007). Compared with barley alone, the total biomass
Alley cropping is an agroforestry practice where peren- (straw plus grain) of barley cultivated between the rows
nial crops are simultaneously grown with an arable crop of spineless cactus increased from 4.24 to 6.65 tonnes
(barley, oat etc.). Shrubs, trees or cacti are grown in ha−1 and of grain from 0.82 to 2.32 tonnes ha−1. These
wide rows (e.g. 10−15 m) with a crop in the interspace. results reflect the impact on the micro−environment
Alley cropping is a variation of hedgerow intercropping. created by alley cropping with cactus, in particular the
Leguminous and fast−growing tree or shrub species are beneficial “wind breaking” effect that reduces water
preferred (Saraiva, 2014). Legume trees improve soil at- loss and increases soil moisture. The barley crop stimu-
tributes (e.g. nutrient recycling, biological N2 −fixation), lated an increase in the number of cactus cladodes and
suppress weeds and control erosion on sloping land. fruits, while the cactus increased the amount of root
Alley cropping allows the farmer to continue cultivating material contributing to the soil organic matter.
the land while the trees or shrubs planted in intermit- Tree legumes are another option for alley cropping with
tent rows help maintain the quality of the soil. Cactus cactus. Tree legumes add N to the system, providing pro-
can act as a windbreak, resulting in improved grass and tein and fibre to cactus−based livestock diets. In Brazil,
cereal yields. Wide alleys allow animals to graze the Gliricidia sepium or Leucaena leucocephala were used
biomass strata or cereal stubbles in the summer, and with cactus and a comparison was made with cactus
cactus cladodes may be harvested, chopped and given planted in monoculture. Adding the legumes did not
directly to grazing animals as energy supplement to the
change the total biomass, but it did provide a diversi-
low−quality stubbles (Nefzaoui et al., 2011).
fied feed source (Table 3). Cactus and legumes can be
If properly managed, alley cropping can provide income at grown locally in semi−arid areas, reducing dependency
different time intervals for different markets in a sustaina- on foreign grain and increasing food security.

TABLE 2 Total biomass changes and barley crop yields in Sidi Bouzid (Tunisia)a

Alley cropping
Treatment Natural rangeland Barley crop (alone) Cactus crop (alone)
(cactus + barley)
Above ground biomass (tonnes ha−1) 0.51 0.53 1.87 7.11
Underground biomass (tonnes ha−1) 0.33 0.11 1.8 1.98
Barley grain yield, (tonnes ha−1) 0.82 2.32
Barley grain + straw + weeds
4.24 6.65
(tonnes ha−1)
a
Average rainfall in Sidi Bouzid is 250 mm year−1. No fertilizers were applied to all treatments. Source: Alary et al. (2007).
Forage production and supply for animal nutrition 77

TABLE 3 Biomass productivity in alley cropping with cactus and Gliricidia sepium or Leucaena leucocephala,
Pernambuco, Brazil

Cropping system Cactus biomass Legume biomass Total biomass

tonnes DM ha −1
year
−1

Cactus + Gliricidia 13.6a 4.2a 17.8a

Cactus + Leucaena 14.0 a
2.6 b
16.6a
Cactus 16.9a − 16.9a
Standard error 1.0 0.2 0.8
a,b
Means in the same column followed by similar small case letters do not differ by Tukey’s test at 5%. Source: Saraiva (2014).

Figure 2
Example of alley−
cropping technique
using O. ficus−indica
and maize (Photo
credit: Jaime Mena)

Figure 3
Alley cropping with
cactus and Gliricidia
sepium, Pernambuco
State, Brazil (tree
legumes planted in
rows, cactus planted
between rows) (Photo
credit: Jose Dubeux)
 78 Crop ecology, cultivation and uses of cactus pear

Intensive system
(with or without irrigation, high density)

The intensive system for cactus fodder production Recent experiences in Brazil indicate that drip−ir-
is restricted to a small number of countries, prev- rigating 10 mm month−1 (2.5 mm week−1) using
alent mainly in northeast Brazil and a few areas in water harvested through in situ water catchment
Mexico. Figure 4 illustrates an extreme example of increases productivity and enables the cultivation
intensification in Mexico, where cactus is cultivat- of cactus in areas where cactus growth was previ-
ed under drip irrigation and fertigation on a dairy ously limited because of warm night temperatures
cattle farm (Figure 4). and lack of soil moisture (Lima et al., 2015).

Figure 4
Cactus plantation for
fodder production
(top); details of irriga-
tion and fertigation
equipment (bottom),
Zacatecas, Mexico
Forage production and supply for animal nutrition 79

ferent locations and regions to select high−performing

AGRONOMIC PRACTICES ones. It is important to take into account the prevalence
of pests and diseases in some areas and choose the
Ecological adaptation best variety for the region. In Brazil, for example, com-
mon varieties include ‘Gigante’ and ‘Redonda’ (Opuntia
Opuntias are native in a range of environments: from ficus−indica Mill.), but an outbreak of cochineal (Dacty-
sea level in the Californian deserts to 4 700 m asl in lopius opuntiae Cockerell) led to a change in the variety
the Peruvian Andes; from tropical areas of Mexico with recommendation. Producers in the region are currently
temperatures always > 5° C to parts of Canada where planting varieties tolerant to this insect (‘Orelha de
temperatures reach as low as −40° C (Nobel, 1995). elefante Mexicana’ − Opuntia spp.; ‘Miuda’ − Nopal-
While cactus pear requires high soil fertility in order ea cochenillifera Salm−Dyck.). This is an example of
to reach its production potential, it also grows in poor an integrated approach, taking into consideration the
soils. However, it does need well−drained soils and different environmental limitations when choosing the
does not tolerate saline soils (Berry and Nobel, 1985). most suitable variety.
Opuntias have a crassulacean acid metabolism (CAM) Once the best variety has been identified, site selection
photosynthetic pathway: when exposed to water stress, is key for achieving success. The ecological character-
cactus opens its stomata only at night to reduce water istics must be taken into account, and the site should
loss. Therefore, cooler nights (15−20° C) are beneficial have well−drained soils and balanced soil fertility
for cactus development compared with warmer nights (Dubeux and Santos, 2005).
(> 25−30° C). Warmer temperatures increase the ca-
pacity of the air to hold water vapour, thus reducing Planting usually takes place in the last third of the
relative humidity. Conversely, cooler temperatures dry season in order to reduce the occurrence of plant
increase atmospheric relative humidity and reduce the pathogens, which are more common during the rainy
water loss from cactus. Nobel (1995) reported a reduc- season (Inglese, 1995; Farias et al., 2005). Cactus clado-
tion in atmospheric water−holding capacity from 39.7 des planted during the dry season are able to develop
g m−3 at 35° C to only 6.8 g m−3 at 5° C. Since night the initial roots after planting, thus reducing disease
temperatures are also related to elevation, cactus grows incidence in the wet season. When planting material is
better in highlands, because there is less water loss dur- harvested, it is recommended to leave it in the shade
ing the cooler nights. Rocha (1991) indicated that for to allow the cut portion to heal and reduce disease
every 100 m of elevation there is a reduction of 0.65° C incidence. Mondragón Jacobo and Pimienta Barrios
in the average temperature. (1995) recommend placing cladodes in the shade for
4−6 weeks before planting to heal the cut and reduce
Opuntia or Nopalea have the highest potential for fod-
der production compared with other species of cactus. disease incidence. In Brazil, it is common to place clado-
Nopaleas often require greater rainfall (> 600 mm des in the shade for 5−7 days to achieve good results
year−1) than opuntias (> 400 mm year−1). The interaction (Farias et al., 2005). Fungicide (e.g. Thiabendazol at
between rainfall and temperature at elevation affects 60% or “Bordeaux mixture”) can be used to reduce
the growth performance of cactus: at high elevation fungus disease (Mondragón Jacobo and Pimienta Bar-
with cool nights, cactus requires less rainfall than on a rios, 1995; Farias et al., 2005). Mature cladodes from
site with warm night temperatures. the intermediate portion of the plant should be used
for propagation (Tapia, 1983).
Establishment and varieties If mechanization is available, cactus can be planted in
furrows 15−20 cm deep. It is common, however, to
Cactus is vegetatively propagated. Cladodes are used as use a hoe and dig individual pits, particularly in small
propagative material and they preserve the genetic char- farming systems. In general, one cladode per pit (or
acteristics of the parent material. When establishing a furrow position) is recommended; planting more cla-
cactus orchard, various factors should be considered, in- dodes per position may result in higher yields, but the
cluding: cactus variety, type of cladode used (i.e. primary, higher establishment costs do not always justify the use
secondary or tertiary), number and position of cladodes, of two or three times more planting material (Mafra
timing of planting, plant spacing, planting method, weed et al., 1974). Likewise, a half cladode may be planted,
control, fertilization, and pest and disease occurrence. but productivity is usually reduced, particularly in the
Management practices usually interact and they should first harvest (Medeiros et al., 1997). Depending on the
be carefully analysed in a system approach.
cultivar, cladode mass varies, ranging from 0.5 to 1 kg
Varieties indicated for forage production and adapted for most Opuntia varieties, although heavier cladodes
to the specific environmental conditions should be are often found. Cladodes from Nopalea varieties are
used, and variety trials need to be carried out in dif- usually lighter (0.3−0.7 kg).
80 Crop ecology, cultivation and uses of cactus pear

Cladode position affects establishment. Mafra et al. (1974) average concentration (DM basis) of 0.9% N, 0.16% P,
tested three positions: horizontal, vertical or diagonal. 2.58% K and 2.35% Ca (Santos et al., 1990) and an
While no difference in biomass production was revealed, annual productivity of 20 tonnes DM ha−1, the nutrients
lodging was observed when cladodes were placed hori- exported (kg ha−1 year−1) via harvested material would
zontally at planting. Therefore, the authors recommend be 180 kg N, 32 kg P, 516 kg K and 470 kg Ca, without
planting in a vertical or diagonal position, with one−third considering other macro− and micronutrients (Dubeux
of the cladode buried. The orientation – north–south and Santos, 2005). Therefore, this nutrient demand
(N−S) or east–west (E−W) – of the cladodes may also be must be matched by fertilization in order for the system
important. In Mexico, Rodrigues et al. (1975) observed to be sustainable over time.
that cladodes facing N−S produced less than cladodes Cactus often responds better to organic fertilization
facing E−W, and they ascribed the results to the greater than to mineral fertilization (Méndez Gallegos and
light interception with E−W orientation. In Brazil, on the Martínez Hernandez, 1988; Santos et al., 1996). There
other hand, studies indicated no differences between N−S is usually interaction between fertilization level, plant
and E−W cladode position. Young cladodes actually take spacing and environmental conditions; the larger the
different directions, leading to a unique plant architecture plant population, the greater the necessary fertilizer in-
that optimizes light interception. It is important to follow puts. Cultivar ‘IPA−20’ (Opuntia ficus−indica Mill.) pre-
contour lines, particularly on steep slopes; cactus should sented linear responses up to 80 tonnes ha−1 of cattle
face the slope to reduce soil erosion. manure applied, with a plant population of 20 000 to
The recommended plant spacing varies according to 160 000 plants ha−1. Greater responses were observed
the production system and environment. Cactus can be with the combination of higher organic fertilization
planted as a sole crop or intercropped with cash crops. and higher plant population. Intensive systems like this
Farias et al. (1989) tested different plant spacings for may reach annual productivity of > 50 tonnes DM ha−1
cactus intercropped with sorghum (Sorghum bicolor (Silva, 2012). Producers in northeast Brazil, however,
L.). Cactus cladodes were spaced at 3 × 1 × 0.5 m or 7 usually apply 20–30 tonnes ha−1 of manure. Indeed,
× 1 × 0.5 m, and sorghum was planted between rows. smallholders may not have access to large quantities of
Cactus produced less biomass in the wider spacing, but manure and there may also be reduced availability of
sorghum compensated for the loss. If mechanization is both time and labour.
available, this too should be taken into account when Manure should be incorporated into the soil (Inglese,
choosing the optimal spacing, which may also vary 1995; Mondragón Jacobo and Pimienta Barrios, 1995)
with location. In dryer areas, spacing should be great- or surface−applied at planting and after each harvest.
er so that the individual plants can build greater root Saraiva (2014) evaluated different organic fertilizer
mass, reducing risks during drought periods (Dubeux et sources (cattle, sheep and goat manure, and broiler
al., 2011a, b). litter) on cactus (Nopalea cochenillifera Salm−Dyck) de-
Sites with better soil fertility and greater rainfall are velopment. Organic sources were applied at the same
more suited for hosting high−density cactus planta- rate (200 kg N ha−1) – although, given that N concen-
tions. Souza (2015) observed a linear increase in Nopal- tration differs among sources, the total amount actu-
ea cocheniliifera dry matter yield as population density ally varied – and the authors observed no difference in
was increased: from 10 417 plants ha−1 at 1.2 × 0.8 m cactus productivity. Berry and Nobel (1985) evaluated
to 83 333 plants ha−1 at 1.2 × 0.1 m. Linear responses mineral stress in two cactus species (O. ficus−indica
to plant spacing were also observed by Silva (2012) for and Ferocactus acanthodes) and observed that these
O. ficus−indica with the plant population ranging from species do not require high Ca levels in the soil solu-
20 000 to 160 000 plants ha−1. Even greater responses tion. In fact, their development was not affected when
were observed under higher organic fertilization (80 the soil pH was 4.5−8.5. High levels of Ca observed in
tonnes ha−1 of cattle manure). cactus cladodes reflect the high Ca−oxalates. Potassi-
um is the nutrient extracted in greatest quantities due
Fertilization to its high concentration in cactus cladodes. Therefore,
it is essential to replenish soil K after each harvest to
maintain cactus productivity.
Cactus (Opuntia and Nopalea) has a high productivity
potential, but it is only reached with appropriate fer- A combination of organic and mineral fertilizers might
tilization (Silva, 2012). Nutrient extraction from cactus be the best option when less manure is available.
fields dedicated to forage production is high because Dubeux and Santos (2005) estimated that a nutrient
cladodes are removed from the area in cut−and−carry deficit occurred in cactus orchards with a high plant
operations. The concentration of nutrients in cactus population (40 000 plants ha−1) and high productivity
cladodes varies depending on the cultivar, environ- (20 tonnes DM ha−1 year−1) if only 20 tonnes ha−1 of cat-
mental conditions and management. Considering an tle manure were applied. Nobel et al. (1987) observed
Forage production and supply for animal nutrition 81

increased productivity (Opuntia engelmannii ferent weed control methods on the development
Salm−Dyck) with 160 kg N ha−1 and 80 kg P ha−1. of cactus Opuntia ficus−indica Mill. cv. ‘IPA−20’.
The authors also observed that boron significantly When weeds were not controlled, the cactus
increased cactus yield. Nobel (1995) indicated that produced only 3 tonnes DM ha−1 after 2 years
N, P, K, B and Na are the nutrients that exert the of growth, compared with 12 tonnes DM ha−1
most influence on cactus productivity. Baca Cas- obtained using the best treatment (Tebuthiuron
tillo (1988) ranked P, N, K, Ca, B, Mg, Fe and Mn applied as pre−emergence herbicide at 2 litres
in decreasing order of importance, considering ha−1). Thus, weed control resulted in a fourfold
them to be the nutrients with the greatest effect increase in cactus yield. Mechanical control was
on cactus growth. In fact, Dubeux et al. (2006) not as efficient as the pre−emergence herbicides:
observed a linear increase in the productivity of Diuron, Tebuthiuron, or a combination of Simazine
Opuntia ficus−indica Mill. up to 300 kg N ha−1 and Ametryne. If the weeds are potential quality
year−1 on different sites in the semi−arid region of forage material, it is possible to avoid herbicide
Brazil. They observed that N fertilization increased application and instead make hay to use when
N concentration in cactus cladodes and improved feeding the cactus to livestock. This approach,
rainfall−use efficiency. Phosphorus fertilization however, results in reduced cactus productivity
produced positive responses in yield only when (Farias et al., 2005). Felker and Russel (1988)
soil P was < 10 mg kg−1. indicated that glyphosate can be used at 20 g
Saline soils are not suited for cactus (Opuntia and litre−1 for O. lindheimeri without causing damage.
Nopalea) cultivation, as they reduce shoot and root They also indicated that Hexazinone, Simazine
growth (Nerd et al., 1991c; Calderón Paniagua et and Tebuthiuron promote the best yields for that
al., 1997). Berry and Nobel (1985) showed that species. Plant spacing is important for mechanized
roots of O. ficus−indica Mill. exhibit saline stress weed control. In Brazil, plant spacings recently
when NaCl concentration in the nutrient solution used for cactus forage production are 1.6 × 0.2
reaches 25 mM, reducing shoot development. m for Opuntia and 1.2 × 0.2 m for Nopalea (Silva,
2012; Souza, 2015).
Weed management
Harvest management
Weed control is essential to increase cactus pro-
ductivity. Cactus has shallow roots that spread Cactus harvest management must take into con-
horizontally. In favourable conditions, roots can sideration harvest intensity, frequency and timing.
grow to a depth of 30 cm and spread horizontally Interaction between these factors affects cactus
in a radius of 4−8 m (Sudzuki Hills, 1995). There- regrowth. Several harvesting trials have been
fore, weeds compete with cactus for nutrients, carried out in northeast Brazil to determine the
moisture and light. Farias et al. (2005) tested dif- best harvesting strategy to maximize growth (Li-

Figure 5
Harvest, transportation
and utilization of cactus
in Pernambuco State,
Brazil (Photo credit:
Jose Dubeux)
82 Crop ecology, cultivation and uses of cactus pear

ma et al., 1974; Santos et al., 1996, 1998; Farias et al., 11.6 mm for O. ficus−indica and O. robusta, respective-
2000). In general, increased harvest frequency requires ly, was sufficient to fill the cladodes in plants subject to
reduced harvesting intensity, and these two factors water deficit. In rainfed systems, rainfall−use efficiency
interact with plant population. Residual photosynthetic (RUE) varies according to the environment and manage-
area after harvesting is critical to increase plant re- ment practices. Dubeux et al. (2006) report an average
growth, and this is the case also for cactus. Field trials RUE of 18 kg DM ha−1 mm−1, with values ranging from
often reveal a low cladode area index (CAI), which re- 5 to 35 kg DM ha−1 mm−1.
duces light interception and plant growth. Nobel (1995)
suggests a CAI of between 4 and 5 to increase cactus Productivity
productivity. Both faces of the cladodes are taken into
account when measuring the CAI. The larger the plant Cactus can reach high productivity in rainfed semi−arid
population, the higher the CAI, resulting in increased agro−ecosystems. A rainfed system in northeast Brazil
productivity when no other factors limit growth. Farias achieved productivity of > 50 tonnes DM ha−1 year−1 in
et al. (2000) observed that when cactus (O. ficus−in- intensive cultivation systems including the use of high
dica Mill.) was harvested every 4 years, there was no levels of manure (80 tonnes ha−1 year−1) and high plant
difference between preserving primary and secondary population (160 000 plants ha−1) (Silva, 2012). However,
cladodes. Harvesting every 2 years, on the other hand, the average productivity of small farming systems is
required a less intense cut with the preservation of all inferior, due to lower fertilization, smaller plant popu-
secondary cladodes. This trial, however, was done on a lations and absence of weed control. The data reported
cactus population of < 10 000 plants ha−1; with a more by Silva (2012) reflect the potential yield of cactus in
intense plant population, more frequent harvesting semi−arid regions. A cactus orchard productivity of 20
might be possible because of the greater residual CAI tonnes DM ha−1 year−1 (Santos et al., 2000) is sufficient
after cut. Souza (2015) observed that productivity in- to sustain 4−5 cows per year. In the same area, 15 ha
creased with greater plant population (≤ 83 333 plants of rangeland is needed to sustain just one animal unit
ha−1), harvesting every other year, and preserving pri- per year, meaning that cactus productivity is potentially
mary cladodes. This intensive system, however, requires 60−75 times higher. A small area of cactus can provide
higher inputs of organic fertilizer. enough forage to sustain the herds while reducing the
Cactus is usually harvested in the dry season, when pressure on the rangelands’ natural resources.
livestock feed is scarce. In more intensive production However, productivity in low input systems can be
systems, cactus is considered a component of the live- much lower. Farias et al. (2005) report productivity of
stock diet throughout the year, but year−round supply 1.5 tonnes DM ha−1 year−1 with no weed control in a
of cactus is not common in most regions. Producers cactus field. Cactus intercropped with other crops
usually take advantage of the rainy season for pastures often has diminished productivity. Farias et al. (2000)
and rangelands, leaving cactus for the dry season. report cactus productivity of 2.2−3.4 tonnes DM ha−1
During the rainy season, the moisture content of the year−1 with 5 000 plants ha−1 intercropped with Sor-
cladodes increases and there is more chance of patho- ghum bicolor (L.) Moench. In conclusion, productivity
gen incidence on the cut surface of cladodes, resulting varies with inputs and systems; the producer must take
in more disease problems. into account land availability and the economic value
of inputs and outputs when deciding what system best
Irrigation suits a particular condition.

Irrigation is not a common practice in cactus orchards

dedicated to forage production. In some regions where
FORAGE QUALITY
low rainfall associated with warm night temperatures
limits cactus development, the application of small
Nutrients
amounts of water has expanded the cactus planted area.
Dubeux et al. (2015b) report that the use of drip irriga-
Cactus (Opuntia spp.) cladodes are high in water, sugars,
tion (only 10 mm month−1) resulted in annual dry matter
ash and vitamins A and C, but they are low in crude pro-
yields of ≤ 19.6 tonnes ha−1 in a region where cactus (O.
tein (CP) and fibre (Ben Salem et al., 1996; Le Houérou,
ficus−indica Mill.) would otherwise not grow well be-
1996a; Batista et al., 2003a, b). They exhibit a high Ca−P
cause of the low rainfall and warm night temperatures.
ratio and are highly palatable (Tegegne, 2001; Nefzaoui
Salt concentration in irrigation water is a problem. Ap- and Ben Salem, 2001). However, numerous authors
plication of small amounts of water and use of organic (e.g. Le Houérou, 1996a; Azocar, 2001; Nefzaoui and
fertilizer reduce any potential salinization problem. Sny- Ben Salem, 2001; Tegegne, 2001) have demonstrated
man (2004) observed that an irrigation of only 13.6 and significant variability in the nutritive value of cladodes of
Forage production and supply for animal nutrition 83

different species and cultivars. Nutritive value also varies • Solid fermentation is recommended as an alternative
according to the season, agronomic conditions and the option for protein enrichment of cladodes (Araùjo et al.,
technical package adopted (soil texture, rainfall, fertiliza- 2005), although the technique is still at the laboratory
tion etc.). According to Le Houérou (1996a), 1− to 3−year stage of research. Cost–benefit analysis is required in
cladodes are high in water during winter and spring order to enhance adoption of this technique by farmers.
(85−90%), less in summer (75−85%), and the younger
the cladode, the higher its water content. Cladodes as Cactus cladodes are high in carbohydrates (approx. 600
forage can solve the problem of livestock watering, but g kg−1 DM), starch (approx. 75 g kg−1 DM) and β−caro-
attention should be paid to their low dry matter content tene (approx. 6.5 mg kg−1 DM) (Ayadi et al., 2009). Ac-
with regard to diet composition. To compensate for the cording to Abidi et al. (2009a), mucilage is high in the
low dry matter content, the ruminant consumes large cladodes of spineless (6−13 g kg−1 fresh material) and
quantities of cladodes, which may lead to diarrhoea. It is spiny (6−14 g kg−1 fresh material) cactus. Compared
therefore recommended to associate a fibrous feedstuff, with winter, mucilage concentration increases at least
as well as appropriate supplements, in particular those twofold in summer. It reduces salivation in ruminants,
rich in nitrogen. thus avoiding a rapid decrease in rumen pH. This phe-
nomenon was confirmed by Ben Salem et al. (1996),
Compared with conventional feedstuffs, Opuntia cla- who reported rumen pH values of 6.3−6.8 in sheep
dodes have a high ash content (Sawyer et al., 2001). fed on straw−based diets supplemented with increas-
Depending on the species and cultivar, the ash content ing levels of spineless cactus cladodes. These values fit
ranges from 100 to 250 g kg−1 DM, but often exceeds with the optimal pH (6.5−7) recommended for normal
200 g kg−1 DM. Ca − followed by K − is the most growth and activity of the microflora in the rumen.
abundant mineral in the cladodes, but the availabil- Other soluble carbohydrate−rich feedstuffs, such as
ity of Ca to rumen microflora and the host animal is molasses, cause acidosis in the ruminant, because they
compromised by the high content of oxalates and the are low in or free of mucilage. As a succulent plant,
extremely high Ca−P ratio. Cultivated in northern Brazil, cactus cladodes are low in fibre. In general, cell wall
the cladodes of Opuntia cv. ‘Gigante’, Nopalea cv. ‘Mi- neutral detergent fibre (NDF) ranges between 180 and
uda’ and Opuntia cv. ‘IPA−20’ contain 120−145 g kg−1 300 g kg−1 DM, although the cladodes of a spiny cactus
DM of ash, 29−42 g kg−1 DM of Ca and 3−4 g kg−1 DM (Opuntia imbricate) contain 400 g kg−1 of NDF. Ligno-
of P (Batista et al., 2003b). Data on trace minerals are cellulose (acid detergent fibre − ADF, 120−200 g kg−1
scarce; nevertheless, the concentrations of iron, copper DM) and lignin (sulphuric acid detergent lignin − ADL,
and manganese seem to be within the recommended 15−40 g kg−1 DM) are also low. Carotenes, titratable
range for ruminant diets (Abidi et al., 2009b). acidity and carbohydrates increase during development,
It is well documented that cladodes are low in CP. Under while protein and fibre decrease. It is worth noting that
Tunisian conditions, cladodes of O. ficus−indica f. inermis cladodes are high in malic acid and the content of this
contain 30−50 g kg−1 DM of CP. Most of total nitrogen acid oscillates due to a CAM−based diurnal rhythm.
(TN) is in soluble form (865 g kg−1, Ben Salem et al., Several authors (Lila et al., 2004; Mohammed et al.,
2002a). The older the cladode, the lower its CP content. 2004; Newbold et al., 2005) conducted in vitro and in
Therefore, nitrogen supplements in cactus−containing vivo studies to show that malic acid reduces methane
diets are mandatory to ensure normal microbial activity emissions. Therefore, it is expected that the integration
in the rumen and to improve livestock performance. The of cactus in livestock feeding could reduce methano-
literature suggests a range of options to solve the prob- genesis, contributing to a decrease in greenhouse gas
lem of the low N content of cactus cladodes: emissions. However, this hypothesis requires validation
by scientists.
• Inclusion of nitrogenous supplements (soybean meal,
urea, Atriplex nummularia foliage etc.) in cactus−con-
Anti−nutritive factors
taining diets is currently the most widely adopted op-
tion and encouraging responses have been observed
Like many plants, cactus cladodes contain phytochem-
in sheep. Examples are reported in Table 4.
icals with no apparent detrimental effects on livestock.
• Provision of chemical fertilizers (ammonite and super- Negesse et al. (2009) determined the content of some
phosphate) increased CP content of cladodes from 45 anti−nutritive factors in mature and young cladodes of
to 105 g kg−1 DM (Gonzalez, 1989). spineless cactus (O. ficus−indica) growing in Ethiopia
• Breeding successfully increased the CP content of cla- and reported their total tannin content as 21 and 42 g
dodes (Felker et al., 2006). Cladodes of the selected equivalent tannic acid kg−1 DM, respectively. However, a
clone of spineless cactus (clone TAMUK accession very low content of these tannins (1 g equivalent tannic
1270) reported by Peter Felker revealed higher (100 g acid kg−1 DM) in the same species of cactus but grow-
kg−1 DM) than normal (30–50 g kg−1 DM) CP content. ing in Tunisia was reported by Ben Salem et al. (2002a).
84 Crop ecology, cultivation and uses of cactus pear

TABLE 4 Intake, in vivo digestibility, and growth rate of different animal species receiving cactus−containing diets.

Water TDM Initial Growth

Period Diet composition OMD CPD NDFD
Country Animal intake intake BWh rate Ref.i
(days) (g DM day−1) (%)h (%)h (%)h
(litres day ) (g W )
−1 −0.75 h
(kg) (g day−1)

Ethiopiaa Lamb 97 508 teff straw+58 NSC 1.02 65.0 46 48 50 17.0 23 a1

Lamb 97 273 teff straw+58 NSC+246 cactus 0.42 62.6 54 41 48 17.0 53 a1
Ethiopia Lamb 90 685 pasture hay 1.22 70.0 59 57 49 20.0 11 a2
Lamb 90 404 pasture hay+503 cactus 0.02 97.0 58 54 37 19.5 28 a2
Lamb 90 218 pasture hay+508 cactus 0.01 77.0 61 49 30 19.0 0.0 a2
Tunisiab Lamb 70 435 barley straw+178 barley+157 SBM 3.05 72.1 69 73 68 19.5 108 b1
Lamb 70 262 barley straw+157 SBM+577 cactus 1.30 85.2 71 71 69 19.5 119 b1
Tunisia Lamb 84 715 oat hay+300 concentrate 3.88 65.0 64 59 59 32.5 46 b2
Lamb 84 544 oat hay+295 cactus+67 concentrate 2.47 68.9 65 59 66 32.5 39 b2
Kid 84 454 oat hay+300 concentrate 3.20 74.9 65 63 59 19.3 46 b2
Kid 84 488 oat hay+346 cactus+67 concentrate 1.38 94.5 67 64 59 19.3 24 b2
400 tifton hay+280 ground corn+176 SBM+114
Brazilb Lamb 35 4.90 96.5 73 74 57 26.6 255 c1
wheat bran+15 salt+15 limestone
400 tifton hay+280 cactus+176 SBM+114 wheat
Lamb 35 2.30 102.1 83 87 77 27.9 231 c1
bran+15 salt+15 limestone
Brazilc Buck 21 250 Cynodon hay+600 cactus 0.04 84.4 74 77 48 13.7 c2
Buck 21 125 Cynodon hay+600 cactus+125 SBH 0.06 79.1 75 78 51 31.7 c2
Buck 21 0 Cynodon hay+600 cactus+250 SBH 0.03 66.5 78 80 59 31.7 c2
Brazil d
Heifer 395 sugar cane+93 corn+479 WB 103.6 70 45 c3
Heifer 390 sugar cane+92 corn+314 WB+163 cactus 107.4 76 47 c3
Heifer 385 sugar cane+91 corn+156 WB+320 cactus 111.8 77 40 c3
Heifers 381 sugar cane+92 corn+0 WB+469 cactus 103.4 80 37 c3
Indiae Ram 45 752 Cenchrus hay+191 concentrate 73.9 49 46 29.8 d
Ram 45 547 Cenchrus hay+243 cactus 61.2 31 45 31.7 d
Ram 45 672 Cenchrus hay+237 cactus+48 GNC 74.4 54 45 31.8 d
South 660 ground lucerne hay+300 maize meal+40
Wether 63 90.6 73 33.9 118 e
Africaf molasses meal
410 ground lucerne hay+240 dried cactus+300
Wether 63 89.2 76 33.9 116 e
maize meal+40 molasses meal+10 urea
285 ground lucerne hay+360 dried cactus+300
Wether 63 88.3 78 33.9 96 e
maize meal+40 molasses meal+15 urea
Tunisiaf Lamb 60 482 cactus+190 straw 0.38 72.6 58 59 49 19.9 8 f
Lamb 60 503 cactus+258 urea−treated straw 0.42 78.7 64 66 63 19.9 20 f
Lamb 60 513 cactus+198 straw+121 atriplex 0.44 85.0 65 72 65 19.9 31 f
Lamb 60 362 straw+393 concentrate 1.89 74.4 72 70 66 19.9 40 f
a
NSC: Noug (Guizotia abyssinica) seed cake.
b
SBM: soybean meal.
c
SBH: soybean hulls.
d
WB: Wheat bran. Values of dry matter intake (g kg−1 W0.75) were adapted from the data reported in the corresponding article.
e
GNC: Groundnut cake.
f
Values of dry matter intake (g kg−1 W0.75) were adapted from the data reported in the corresponding article.
g
Atriplex nummularia. Values of dry matter intake (g kg−1 W0.75) were adapted from the data reported in the corresponding article.
h
TDM: total dry matter. OMD: organic matter digestibility. CPD: crude protein digestibility. NDFD: total fibre (NDF) digestibility.
i
a1: Gebremariam et al. (2006). a2: Tegegne et al. (2007). b1: Ben Salem et al. (2004). b2: Abidi et al. (2009). c1: Costa et al. (2012). c2 : Souza et al. (2009).
c3: Monteiro et al. (2014). d: Misra et al. (2006). e: Einkamerer et al. (2009). f: Ben Salem et al. (2002b).
Forage production and supply for animal nutrition 85

The total oxalate content varies between 60 and 120 g flow, effluent bacteria and liquid and solid associated
kg−1 DM. Abidi et al. (2009b) reported that cladodes of bacteria isolated from fermenter flask contents. How-
spiny cactus are higher in oxalates (110−118 g kg−1 DM) ever, the spiny cactus−containing diet showed higher
than spineless cactus (102−105 g kg−1 DM) growing in acetate to propionate ratio (P = 0.016) and lower am-
an arid area of Tunisia. It is good that these oxalates are monia flow (P = 0.007).
insoluble, as they have no toxic effect. However, it is
known that insoluble oxalates form complexes with Ca The gas production technique was adopted by Negesse
and Mg rendering them unavailable for rumen micro- et al. (2009) to determine fermentation parameters
flora and the host animal. According to Ben Salem et and estimate the metabolizable energy (ME) content of
al. (2002a) and Abidi et al. (2009b), spiny and spineless some non−conventional feed resources, including ma-
cactus cladodes are low in saponins (2–5 g kg−1 DM), ture cladodes (MC) and young cladodes (YC) of spine-
total phenols (10–34 g kg−1 DM), total tannins (1 g less cactus growing in Ethiopia. The ideal ME content is
kg−1 DM) and condensed tannins (<1 g kg−1 DM). The 10−13.6 MJ kg−1 DM, but MC and YC revealed an ME
authors are not aware of the presence of any other of 7.5 and 8.5 MJ kg−1 DM, respectively, and were not
secondary compound in cactus cladodes with poten- considered good sources of energy.
tially negative effects on the nutritive value and animal
performance and health. Intake

Fermentation Considerable information on the response of different

animal species to cactus−containing diets is available.
Different in vitro techniques are used to evaluate the Feeding trials usually evaluate spineless cactus clado-
fermentation potential of the cladodes of different spe- des (henceforth referred to as cactus) as an alternative
cies and cultivars of cactus. fodder source for growing dairy and other ruminants.
Table 4 presents information generated by some
The gas production technique developed by Menke studies. The more fresh cactus is consumed, the less
and Steingass (1988) and using calibrated glass syringes water is drunk by the animal. The decrease in drinking
has been widely adopted in research laboratories in water consumption by the various categories of sheep
the last decade. The digestion of organic matter leads and other animal species (bucks, kids and heifers) is
to the production of volatile fatty acids (VFA), ammo- 40−98%, depending on the proportion of cactus in
nia and different gases, mainly dioxide carbon and the diet. Gebremariam et al. (2006) reported a 59%
methane. The technique is, therefore, based on the decrease in drinking water in lambs receiving a diet
following hypothesis: the more gas is produced in the composed of 43% fresh cactus. Furthermore, in a study
in vitro system, the more the substrate is digestible in conducted in Ethiopia, lambs stopped drinking water
the rumen. Equations have been developed to predict when there was access to a 55% fresh cactus−contain-
the digestibility and energy contents of a wide range of ing diet (Tegegne et al., 2007). Cactus plantations are,
feedstuffs. Batista et al. (2003a) measured the gas pro- therefore, a promising option for alleviating drinking
duction of three cultivars of spineless cactus growing in water scarcity in dry areas and during drought periods.
northern Brazil (‘Gigante’, ‘Miuda’ and ‘IPA−20’). They
reported high volumes of gas production from the 24− Cactus is often used to supplement low−quality forage,
hour fermentation of these three cultivars: ‘Gigante’ including straw and pasture vegetation. Its impact on
was highest (210 ml g−1 DM), followed by ‘Miuda’ (202 diet intake is improved when associated with a protein
ml g−1 DM) and ‘IPA−20’ (195 ml g−1 DM). In south- source. Total dry matter (TDM) intake in lamb receiving
ern Tunisia, higher values of potential gas production pasture hay and cactus was 26% higher than in lamb
were reported by Abidi et al. (2009b) for spineless (O. fed on pasture hay alone (Tegegne et al., 2007). The
ficus−indica f. inermis) and spiny (Opuntia amyclaea) replacement of energy sources, such as barley grain
cactus cladodes harvested in winter (138 and 140 ml (Abidi et al., 2009a) or ground corn (Costa et al., 2012),
g−1 organic matter [OM], respectively) and summer (140 by cactus in sheep diets increased TDM intake by 6%
and 145 ml g−1 OM, respectively). High gas production and 25%, respectively. Replacing barley in concentrate
in the early hours of incubation is a characteristic of with cactus increased TDM intake by 26% in growing
soluble carbohydrate−rich feedstuffs, including cactus. goat (Abidi et al., 2009a).
Single−flow continuous culture fermenters were used
Digestibility
by Abidi et al. (2009b) to compare the fermentation
traits of experimental diets composed of spineless or
spiny cactus cladodes (17 g), Atriplex nummularia foli- Soluble carbohydrate−rich feeds like molasses and
age (12 g) and wheat straw (24 g). Both diets (spineless/ cactus improve diet palatability and enhance rumen
spiny cactus) exhibited similar pH, total VFA, total N fermentation, usually leading to an increase in TDM
86 Crop ecology, cultivation and uses of cactus pear

intake and/or diet digestibility. Depending on the • buffer feed reserve during drought periods lasting
diet composition, feeding cactus results in increased 1−3 years.
apparent organic matter (OM) digestibility of the diet
or has no effect on this parameter. Studies conduct- In contrast to other fodder and forage crops which need
ed in Ethiopia (Gebremariam et al., 2006; Tegegne to be stored (e.g. hay or silage), cactus is a standing ev-
et al., 2007), Tunisia (Ben Salem et al., 2004; Abidi et ergreen crop and can be used year−round. The natural
al., 2009a) and Brazil (Costa et al., 2012) showed an and probably most efficient way of using cactus is to
increase of the OM digestibility of the diet by 2 to 10 cut the cladodes and feed them without any processing.
percentage units in cactus−receiving lambs and kids Moreover, cactus is rich in water and plays a crucial role
compared with cactus−free diets. This could be the in arid environments as a replacement for drinking water.
result of improved rumen fermentation. The provision Silage−making or drying is feasible, but entails additional
of cactus to Ethiopian lamb fed on teff straw or pasture costs in terms of handling, energy and labour.
hay resulted in a decrease in the apparent crude pro-
The different uses are outlined below, with a focus on
tein digestibility of the diet. Gebremariam et al. (2006)
direct browsing, cut−and−carry, drying, silage and, fi-
claims that the decreased CP apparent digestibility with
nally, feed blocks and their potential for making use of
increased cactus inclusion rate could be explained by
low−quality fruits.
the total tannin content of cactus. Indeed, tannins have
great affinity with proteins, making them unavailable
Direct browsing
for rumen microflora and the host animal. The authors
are not aware of any other study showing the effect of
Direct browsing of the stand occurs mainly in native plan-
cactus tannins on ruminal digestion in ruminants; the
tations with both thorny and spineless cacti. It is current-
interaction between cactus tannins and proteins has
ly practised in a small number of countries (e.g. Mexico),
not been investigated in either in vivo or in vitro studies.
where the wild cactus populations are directly browsed
Under Brazilian conditions, Costa et al. (2012) noted an
by livestock. In Ethiopia, where cactus is sometimes inva-
increase of 13 percentage units in the CP digestibility of
sive (e.g. in the Tigray region), it is also browsed directly.
lamb’s diet. Also in Brazil, Souza et al. (2009) reported
Camels and cattle are able to intensively browse spiny
high values of CP digestibility (77−80%) in bucks re-
cactus (Figures 6 and 7). In human−made plantations,
ceiving Cynodon hay, cactus and soybean hulls. Ben
direct browsing is not recommended because it usually
Salem et al. (2002b), Gebremariam et al. (2006) and
Misra et al. (2006) observed no effect of cactus on fibre results in rapid damage to the stands. The most effective
(NDF) digestibility in the diet. Costa et al. (2012) report and low−cost option is grazing with an electric fence,
a positive impact of cactus on NDF digestibility − an where all the biomass in a row must be completely con-
increase of 20 percentage units in NDF digestibility with sumed before the livestock can access a new row; the
a cactus−containing diet, compared with the control greatest danger of direct grazing (i.e. loss and wastage
(cactus−free) diet given to lamb. In brief, cactus supply because cladodes are only partially consumed) is thus
has no negative effect on diet digestibility and may avoided (Le Houérou, 2002).
even improve it. Spines in wild populations may be burnt directly prior
to grazing, as is practised in Texas. Maltsberger (1991)
conducted an extensive investigation on the economics
UTILIZATION of burning spines using propane and on the supple-
mentation of burnt cladodes to cattle. They found that
Cactus cladodes and waste fruits are a cost effective 15 litres of propane were required for 14 cows per day,
feed for ruminant animals. The benefits of using cactus and that 8 work−hours were required to prepare feed
as feed are well documented (Nefzaoui and Ben Salem, for 200 cattle per day. Pluenneke (1990) showed that
2001; Ben Salem and Abidi, 2009). labour and fuel costs to burn spines are significantly
reduced when the plantation is in rows. Nevertheless,
Plantations of cactus for fodder production (harvested)
there is still some wastage as cattle step on burnt por-
or as forage (directly browsed by livestock or wildlife)
tions of cactus; cut−and−carry is therefore preferred.
have been developed in Sicily and North Africa since
the mid−nineteenth century. Farmers wished to stabi-
Cut−and−carry technique
lize the fodder resource in arid and semi−arid zones,
where feed shortages in summer and autumn are a
Cut−and−carry is the most commonly used technique
quasi−permanent factor limiting livestock production
for cactus feeding. It prevents wastage and excessive
(Le Houérou, 2002). These plantations are still used as:
grazing. Both spiny and spineless cactus cladodes can be
• part of the common daily ration; harvested and transported to the barn, then chopped,
• supplementary feed in summer and autumn; and mixed with other feeds and put in troughs. Spines must
Forage production and supply for animal nutrition 87

Figure 6
Camel browsing thorny
cactus in Ethiopia

Figure 7
Cattle browsing cactus,
“fighting bull ranch” in
San Luis Potosí, Mexico
(Photo credit: Ali
Nefzaoui)

Figure 8
Hand−cutting of
spineless cactus
cladodes in Tunisia
(Photo credit: Ali
Nefzaoui)

Figure 9
Hand−propelled cactus
chopper in Tunisia

Figure 10
Hand−propelled
Ethiopian cactus
chopper (Photo credit:
Ali Nefzaoui)

Figure 11
Cactus processing
machines in Brazil

be burnt prior to chopping and feeding. Different Drying

types of choppers are available in the market, from
simple small choppers to more sophisticated ones. Under certain circumstances, cactus cladodes are
In North Africa, cactus chopping is done manually chopped into small slices then air−dried, for exam-
with knives and the whole family helps, especially ple, when the harvest period is short (pruning of
women and kids (Figure 8). In Tunisia and Ethiopia, cactus fruit plantations) or when the objective is
transportable, low−cost choppers are made locally to produce a commercial feed or mix the cladodes
(Figure 9 and 10) and are propelled using human with other ingredients. In all cases, sun−drying is
power. More sophisticated and motorized chop- recommended to avoid using expensive fuels that
pers are utilized in Brazil and Mexico (Figure 11). will increase production costs. Once dried, cactus
In addition to reducing wastage, slices of cactus are is ground to produce cactus meal − avoiding a
more convenient for incorporating in mixed diets. too−fine meal that could transit too quickly in the
88 Crop ecology, cultivation and uses of cactus pear

gut. Sometimes, cladodes are dried to reduce their high The TMR feeding strategy is recommended to reduce
water content (85–90%), on the basis that fresh cactus selectivity in the cows, which leads to an imbalance
consumption leads to very wet faeces. However, the between the diet offered and that consumed.
findings of De Waal et al. (2013) negate this assump-
tion: they show that, even when animals are fed with Silage
dried cactus, faeces remain wet due to the presence of
mucilage. Under certain circumstances, it may be necessary to
make silage from cactus cladodes: when the production
Cactus drying is mainly practised in Brazil and South Af-
rica. Several studies (Zeeman, 2005; Einkamerer, 2008; is concentrated in a short period (pruning); or when
Menezes, 2008) confirm that sun−dried and coarsely wet agro−industrial by−products cannot be stored for
ground Opuntia cladodes can replace a large proportion a long time and must be valorized. To make high−qual-
of lucerne hay in the diet of young Dorper lambs. De ity silage, good acid lactic fermentation is necessary,
Waal et al. (2013a) investigated the commercial poten- requiring appropriate levels of moisture (30−40%) and
tial of sun−dried cactus cladodes (O. ficus−indica) for sugar in a full anaerobic environment. Cactus cladodes
feedlot diets for Dorper wether Lambs (Table 5). They contain sufficient carbohydrates for good lactic acid
showed that there are no significant differences in car- fermentation, but their high water content means that
cass characteristics, and suggested that carcass quality is they need to be carefully mixed with other material
not affected by the inclusion of sun−dried and coarsely (e.g. chopped straw or bran).
ground cactus pear (O. ficus−indica) cladodes in diets or Abidi et al. (2013) conducted an experiment in Tunisia
by the type of nitrogen source. On the other hand, the where the olive industry is well developed. They tested
difference in average daily gain between the cactus− the ensiling potential of a mixture of chopped cladodes
pear−based diets shows the importance of high−quality (350 kg), olive cake (by−product of olive oil mills, 400
nitrogen sources for lambs. These results indicate good kg) and wheat bran (250 kg). The mixture was ensiled
prospects for the commercialization of sun−dried Opun- for 75 days and then distributed with concentrate sup-
tia cladodes in balanced diets for ruminants. plement (75% barley grain and 25% soybean meal) to
Two forms of cactus processing: knife−chopped (KC) lambs. Three diets were compared:
and fodder machine−sliced (FM), and two different • Diet 1 (control: oat hay ad libitum and 400 g concen-
feeding strategies: separate concentrate (SC) and total trate feed)
mixed ration (TMR), were investigated by Da Silva et • Diet 2 (silage and 400 g concentrate feed)
al. (2010). They monitored the feeding behaviour of
• Diet 3 (50% diet 1 and silage)
milking cows and concluded that cactus processing in
a fodder machine is recommended to maximize the dry The amounts of feed distributed were adjusted for
matter intake and avoid alterations in milk composition. iso−energetic and iso−nitrogenous diets. The silage

TABLE 5 Composition of diets (T0, T1 and T2) fed to young Dorper wether lambs (adapted from de Waal et al., 2013a)

Dietsa (g kg−1)
Feed ingredients (kg air−dried material)
T0 T1 T2
Sun−dried and coarsely ground Opuntia cladodes − 330 300
Coarsely ground lucerne hay 577 255 190
Yellow maize meal 358 340 275
Feed grade urea 10 20 −
Sunflower oilcake meal − − 180
Molasses meal (Enermol) 40 40 40
Feed lime 15 15 15
Average daily gain (ADG) in weight (g) 181a 125b 181a
Cost of diet head−1 day−1 (N$) 3.71a 2.73b 3.26b
a
T0: conventional feedlot diet. T1 and T2: Opuntia−based diets with different nitrogen sources (T1 – non−protein nitrogen; T2 – natural protein).
Means followed by the same letter (a/b), within the same row, do not differ (P > 0.05) in the Tukey test.
N$ = Namibia dollar (1 N$ ≈ 0.072 US$).
Forage production and supply for animal nutrition 89

obtained was of good quality with a pH of 4.5. The the growth rate of lamb on the cactus−containing diet
average daily weight gain of the lambs and the meat was considered tolerable, taking into account the low
quality were also similar across the three diets. production cost for smallholders and considering that
cactus is available year round (Einkamerer et al., 2009;
Feed blocks Costa et al., 2012). Although the association of cactus
with soybean meal produced an ADG in Barbarine
In many countries, and Tunisia in particular, large quan- lamb of 119 g day−1, Ben Salem et al. (2004) favoured
tities of cactus fruits are not harvested because of their a diet composed of locally available feed resources (i.e.
low quality or the labour costs. The over−ripened fruits cereal straw, cactus and Atriplex nummularia foliage)
attract the Ceratitis capitate fly, also called the “Med- for two reasons: i) it is less expensive than a diet con-
iterranean fruit fly” (“medfly” for short), which can taining barley grains and soybean meal; ii) the shrubs
cause extensive damage to a wide range of fruit crops. bring additional benefits (e.g. soil fixation and fruit
Therefore, farmers are encouraged to collect these production).
fruits and incorporate them in feed blocks for livestock.
Different formulas have been developed based on the To summarize, cactus can improve the nutritive value
replacement of molasses with cactus fruit. Chermiti of poor−quality diets (e.g. straw−based diets) due to its
(1998) investigated the potential of cactus fruit as a high content of soluble carbohydrates. It may also in-
feed block ingredient and evaluated the voluntary in- crease weight gain in small ruminants and heifers fed on
takes in heifers and ewes receiving an oat−vetch hay− crop residues or poor−quality pastures, provided that a
based diet supplemented by two kinds of feed blocks small amount of a nitrogen source is included in the diet.
(one containing cactus fruit, the other molasses). He
Most studies on the effects of cactus feeding on milk
concluded that the voluntary intakes by both animal
production and quality have been carried out in Brazil.
species were identical for both types of feed block. Ben
Cactus is highly valued in times of drought and water
Salem et al. (2003) mixed processed cactus fruits (90 g
scarcity. Spiny cactus is fed to cattle in northern Mex-
kg−1 DM) with olive cake (367 g kg−1 DM), wheat bran
ico and the southwestern United States of America
(243 g kg−1 DM), quicklime (154 g kg−1 DM), urea (73 g
(Maltsberger, 1991). It is a cheap source of cattle for-
kg−1 DM) and salt (73 g kg−1 DM) to make feed blocks.
age and is frequently used by ranchers and dairymen
They supplemented the feed blocks with oak (Quercus
in these areas. Spiny cactus is traditionally included in
coccifera, a tanniniferous shrub species) foliage to im-
ruminants’ diets, their thorns burnt from the standing
prove the nutritive value (total intake, organic matter
prickly pear plants using a propane−fuelled flame.
and fibre digestibility of the diet, nitrogen retention) of
Since spineless cactus have become widely disseminat-
the diet for goats. The administration of a small amount
ed in recent decades, many farmers now use cactus in
of polyethylene glycol in the feed blocks resulted in
dairy cattle feeding. According to farmers in northern
further significant improvements in the parameters, as
Brazil, Holstein cattle fed on a mixed diet of 60%
a result of the deactivation of the oak tannins with the
chopped fresh cactus cladodes, 20% fibrous feedstuff
re−agent.
(hay or straw) and 20% protein−rich concentrate
produce around 20 litres day−1 of milk (H. Ben Salem,
personal observation). Oliveira et al. (2007) studied the
ANIMAL PERFORMANCE replacement value of cactus (0, 12, 25, 38 and 51%) for
AND PRODUCT QUALITY cracked corn and Cynodon hay on dairy cattle raised
in northern Brazil. Based on their results, cactus can
Productive performances totally replace cracked corn and partially replace hay
(around 40%) without any significant effect on milk
In many articles, authors conclude that supplement- production (20.3−21.8 kg day−1).
ing low quality forage with fresh cactus increases the
average daily gain (ADG) of growing ruminants. The Many experiments in northeast Brazil show that meal
response is even more positive when a nitrogen source from dried cactus cladodes may replace corn in lamb
is provided (Table 5). Replacing around 50% of teff feeding (Véras et al., 2002). Four replacement levels
straw with cactus increased the ADG of Ethiopian lamb of corn (0, 25, 50 and 75%) were tested and results
from 23 to 53 g day−1 (Gebremariam et al., 2006). Un- show that there is no effect in the levels of replace-
der Tunisian conditions, the substitution of barley grain ment on the digestibility of dry matter, organic matter
with cactus increased the ADG of Barbarine lamb from or fibre. Véras et al. (2005) also tested the effect of
108 to 119 g day−1 (Ben Salem et al., 2004). It should replacement levels (0, 33, 67 and 100%) on the per-
be mentioned that in both studies the cactus−based formance of growing lambs in feedlots; they found
diet included protein supplements (noug seed cake that body weight gain and feed–gain ratio decreased,
and soybean meal, respectively). Any slight decrease in while intakes of NDF and ADF increased linearly with
 90 Crop ecology, cultivation and uses of cactus pear

Figure 12
Dairy cows in
Pernambuco State,
Brazil, feeding on
processed cactus
mixed with silage and
concentrate feed (Photo
credit: Djalma Santos)

corn replacement. The intakes of dry matter, SFA. Under northern Brazil conditions, Santos et
crude protein, organic matter and total carbohy- al. (2011) evaluated the muscularity and adiposity
drates and carcass yield were not affected by the of the carcass of Santa Ines lamb receiving cactus
replacement of corn with forage cactus meal. meal as a replacement of ground corn; there was
no effect on hot carcass yield, biological yield
Product quality (hot carcass weight to empty body weight ratio)
or on the proportion of fat required for ensuring
Taking into consideration consumer preference adequate carcass preservation.
and human health, some literature refers to the
Oliveira et al. (2007) conclude that cactus feed-
effect of cactus supply on meat quality. Abidi ing did not affect milk production in dairy cattle
et al. (2009) conclude that the replacement of receiving increasing levels of cactus. However,
barley with cactus as energy supplements in the it did significantly change milk composition,
local goat kid and Barbarine lamb diet did not in particular the profile of fatty acids. Indeed,
produce major changes on the intramuscular fat- cactus supply reduced the proportions of stearic
ty acid composition of meat. However, vaccenic and oleic acids, but did not affect the other long
acid was found to be higher in the cactus−con- chain fatty acids, such as linoleic (C18:2) and li-
taining diet. This fatty acid − like conjugated nolenic (C18:3) acids.
fatty acids − has a positive impact on the cardio-
vascular system. While gross forage (oat hay) was
used as basal diet in the research work reported
by Abidi et al. (2009), Atti et al. (2006) analysed FUTURE PERSPECTIVES AND
the meat quality of kids receiving cactus in RESEARCH NEEDS
concentrate−based diets supplemented with
a small amount of oat hay. The authors report Global warming, climate change, and increas-
that cactus feeding resulted in increased propor- ing human and livestock population all require
tions of linoleic, linolenic and conjugated linoleic more efficient use of dryland systems. Adapted
acid. They also obtained a higher proportion of perennial crops with greater productivity per
polyunsaturated fatty acids (PUFA) and higher unit area are required to protect natural range-
PUFA to saturated fatty acids (SFA) ratio. The two land systems from degradation. Cactus fits well
studies confirm that cactus included in the diets in this scenario, with productivity 60−75 times
of sheep and goats improves the quality of meat greater than natural rangelands. Small intensive
as per consumer preference: more PUFA and less cultivation of cactus can alleviate pressure on
Forage production and supply for animal nutrition 91

the natural resources of rangelands, reducing their The information currently available on cactus for fod-
degradation. At the same time, more animal products der production is adequate to implement successful
(e.g. meat and milk) produced by cactus−fed livestock systems in different areas. Further research is required
can alleviate hunger, increasing food security and re- to increase the knowledge of biotechnology, geno-
silience of populations living in arid regions. There are type × environment interactions, and locally adapted
numerous success stories from various countries and feeding systems utilizing cactus and local ingredients,
technologies are available to help other regions with and to enhance the efficiency of cactus use in different
similar environments. Policy−makers must be made livestock production systems. Caution is necessary to
aware of the benefits of cactus as a fodder as well as avoid the introduction of invasive species. The use of
its environmental benefits; they should establish poli- spineless cactus for forage production needs to be
cies to increase the planted area accordingly. Research encouraged. Insects and diseases are also a problem
and development work is needed in this area and in localized areas; it is necessary to generate and pro-
enhanced support from donors, policy−makers and mote varieties tolerant to diseases to deal with the
science managers can help achieve increased cultiva- problem. Therefore, conservation of genetic resources
tion of cactus in dry areas. It is also vital to increase and promotion of multilateral cooperation is essential
collaboration among international research teams. to address these challenges.
 08

Nopalitos or vegetable
cactus, production
and utilization
Candelario Mondragón Jacoboa and Santiago de Jesus Méndez Gallegosb
a
Faculty of Natural Sciences, Autonomous University of Querétaro
Juriquilla Querétaro, Mexico
b
Postgraduate College, San Luis Potosí Campus, Mexico
08 Nopalitos or vegetable cactus,
production and utilization

INTRODUCTION trend is destined to increase, encouraged by recent dis-

coveries of their functional properties (discussed in depth
One of the great challenges of developing and un- in this book). Nopalitos represent a unique example of
derdeveloped countries is how to achieve food security ancient Mexican horticultural wisdom: by utilizing the
without compromising the natural resource base, under vegetative structure, renewed by the continuous harvest,
continuous threat from the global trend of population farmers exploit the physiological effects of pruning, se-
increase and climatic change. Demographic growth also curing valuable greens year round, in a region dominat-
places additional pressure on water and land resources, ed by a climatic bimodal cycle of wet and dry seasons.
with competition for first quality land and sources of The introduction of vegetable nopalitos in other coun-
clean drinking water, as these assets are continuously tries and cultures has not been easy, despite the plant’s
removed from agricultural production and dedicated adaptability and high productivity in most locations.
to the needs of urban development. As a result, less The presence of thorns − even the so called “spineless”
land and water are available for agricultural production, cultivars present spines when young − the abundance
leading to the dilemma: incorporation of grasslands and of mucilage, and the lack of organoleptic appeal of
forests in agriculture versus undercut of development. cooked nopalitos have limited their adoption. Small
The search for alternative crops and efficient agricultural orchards of nopalitos are found in the United States of
technologies is clearly a reasonable approach. America, mainly in California, Texas and Arizona; they
Crops with built−in mechanisms conferring water efficien- produce limited quantities intended for local farmers’
cy, ability to grow in limiting soils and tolerance to cold markets, aiming at consumers of Hispanic origin.
and heat have been a beacon of agricultural science since This chapter describes the domestication of cactus pear
the second half of the last century, when the effects of and presents varieties and production systems − from
human population growth and poor resource planning open−field crop to greenhouse production − as well as
policies become more evident. Cactus pear (Opuntia fi- basic cooking techniques.
cus−indica), a plant native to the semi−arid highlands of
central Mexico and introduced to the rest of the world
since the sixteenth century as a curiosity, receives increas- EARLY UTILIZATION AND
ing attention from governments, non−governmental
organizations (NGOs) and private individuals. It comes
DOMESTICATION
up repeatedly as a serious alternative for the utilization of
In the earliest stages of utilization of cactus pear, the
semi−arid lands affected by natural and induced desertifi-
organ of interest was the fruit; indeed, the consump-
cation across the semi−arid tropics and subtropics.
tion of ripe fruits has been documented through copro-
Cactus pear is mostly known as a fruit crop and it is lites found in caves, demonstrating that the fruits were
slowly achieving the status of formal crop in world sta- part of the diet of ancient Mexican tribes (Hoffmann,
tistics. Estimated data indicate that it is present on a 1995). On the other hand, there is no record of the
commercial scale in over 20 countries, covering a plant- consumption of tender pads: due to their perishability,
ed area of around 150 000 ha (personal estimate). The there are no remnants on archaeological sites. It may
cultivation and utilization of forage cactus is destined be speculated that early Mesoamericans following the
to follow suit, since the intake of proteins of animal ori- gathering routes of cactus fruits, as the fruit became
gin continues to rise. Moreover, cultivation for forage is scarce, would turn to the succulent young cladodes
easier to adopt, both for commercial animal production as a source of water to quench their thirst. They were
and for pastoral production systems; cactus cultivation probably used as emergency foodstuff in times of fruit
may ease the depletion of the natural grasslands. scarcity − a frequent situation in the harsh semi−arid
climate predominant in central Mexico and exacerbated
The utilization of tender cladodes − known by Mexican
by the alternate bearing habit of the plant. They may
consumers as nopalitos − as a vegetable originated and
have roasted the cladodes to eliminate the spines − es-
evolved in central Mexico. Their consumption is deeply
sential to facilitate consumption.
rooted in the country’s gastronomic culture as cactus
pear is easy to cultivate and has high productivity. The Later on, an important part of the domestication process
Nopalitos or vegetable cactus, production and utilization 95

was the identification, nurturing and propagation ‘Milpa Alta’ Opuntia ficus−indica (L.) Mill.
of spineless plants, a mutant trait that limits the
survival of the plant in its natural environment (Col- An indigenous cultivar, it probably originated and
unga Garcia et al., 1986). All young cladodes, re- domesticated in the state of Hidalgo, Mexico
gardless of variety and species, present spines and (Reyes Agüero et al., 2004). The commercial name
glochids. Only mature cladodes may be spineless, ‘Milpa Alta’ refers to the region of the same name
allowing for easier manipulation. Modern vege- located in the outskirts of Mexico City where
table nopalito varieties are all spineless and were intensive production of nopalitos began. It has
originally obtained from family orchards. spread throughout Mexico, covering an estimated
7 500 ha. Its bright green, flat, thin, easy−to−peel
The new demands of the modern market, less de-
cladodes and its tenderness are appreciated by
pendent on local production and seasonal availabil-
merchants and consumers. It is planted in open−
ity, have led to increased interest in out−of−season
field plantations or under plastic covers, and may
production (ideally, year−round). Cultivated areas
be rainfed or irrigated in dry areas. The plant is
have, therefore, expanded to zones of mild winter
robust, erect, with oblong cladodes, highly pro-
or subtropical climate and, more recently, various
ductive but frost−sensitive. The fruit is medium
forms of greenhouse. Given its original status of
large, with yellow−orange skin and pulp, not too
emergent crop suited to growers with limited re-
juicy, with medium−sized seeds. It is well adapted
sources and sites of low productivity, greenhouse
to a semi−arid subtropical climate and to subtrop-
production systems for nopalitos have gradually
ical lowlands (Gallegos and Mondragón Jacobo,
evolved from basic to more sophisticated. Three
2011). This variety is similar to the spineless culti-
main types are in use in central and northern Mexi-
vars of yellow−orange fruit present in other parts
co and are described later in the chapter.
of the world: Italy, Morocco, Tunisia and South
Africa (Figure 1).

VEGETABLE NOPALITO VARIETIES ‘Atlixco’ O. ficus−indica (L.) Mill.

In contrast with the wide range available for This variety originated in the highlands of central
cactus pear production, the commercial varieties Mexico, but was named ‘Atlixco’ after the town
for cultivation as a vegetable are limited to ‘Milpa where its cultivation began. Nowadays, an esti-
Alta’, ‘Atlixco’ and ‘COPENA V1’, described below. mated 800 ha are planted, mostly in the neigh-

Figure 1
Nopalito varieties
(clockwise: ‘Milpa Alta’;
‘Atlixco’ or ‘Negrito’;
‘Blanco’; ‘Valtierrilla’)
96 Crop ecology, cultivation and uses of cactus pear

bouring states of Mexico City. The plant is vigorous, sustained some cottage size industries. Note that
erect, with rhombic spineless cladodes of intense green some local types of O. robusta are also used for brin-
colour and exceptional quality. The tender nopalitos are ing in San Luis Potosí (Figure 1).
easy to clean, with an ovoidal shape and thicker than • ‘Spineless 1308’ (Nopalea cochellinifera), selected
those of ‘Milpa Alta’. The fruit are large (reaching > 180 by P. Felker from accessions collected in the tropical
g) with orange skin and yellow pulp; eating quality is region of Tamaulipas, Mexico, cultivated in the Unit-
acceptable but inferior to that of standard fruit culti- ed States of America, planted in the coastal region of
vars. The mature cladodes are well accepted as forage.
Texas and Tamaulipas.
It is very productive under intensive cultivation, reach-
ing 400 tonnes ha−1 year−1 of fresh matter (Gallegos
and Mondragón Jacobo, 2011), and is well adapted to
field cultivation in the highlands under plastic tunnels AGRICULTURAL IMPORTANCE
(Figure 1). OF THE NOPALITO CROP

‘COPENA V1’ O. ficus−indica (L.) Mill. The cultivation of nopalitos reached the status of for-
mal crop in Mexico in the 1980s, when official statistics
In terms of commercial stock available for vegetable reported only two main production zones, Milpa Alta
production, ‘Copena V1’ is the only improved geno- and Tlalnepantla Morelos, both located near Mexico
type, obtained by the late Dr Facundo Barrientos at the City and covering about 2 000 ha. Mexico City and its
Postgraduate College in Chapingo, Mexico. The plant surrounding urban area still remain the largest market
is vigorous, fast−growing and with erect habit growth; for all fresh produce, including nopalitos, reaching al-
the cladodes are obovate, medium−sized, spineless. most 21 million people in the last census (INEGI, 2010)
Although selected for vegetable production, its fruits (Table 1). By 1990, urbanization and demographics,
are attractive: purple, large and sweet, of intermediate combined with new research on the health benefits
juiciness, and pulp with medium−sized seeds. While it of nopalito consumption, led to an increase in planted
is known that the breeder distributed planting material area; by 2010, it had expanded from 5 269 to 12 201
in several states, there is no accurate record of its acre- ha. The planted area currently ranks 12th compared
age (Figure 1). with other vegetable crops: an indication of its impor-
Other lesser known varieties useful for nopalito pro- tance (SIAP, 2015). Nowadays, plantations of vegetable
duction in tropical semi−arid areas are: nopalitos are scattered throughout the central, west,
north central and northeast parts of the country. A
• ‘Valtierrilla’, cultivated in a small area of the same
significant proportion of the production is destined
name located in central Mexico and its surroundings;
for the national market (90%); however, the share
characterized by densely spined cladodes; preferred
for its tender “baby” or “cambray” type nopalitos; for export to the United States of America has been
sensitive to frost damage; performs well when increasing steadily during the last decade, facilitated
brined. by the bilateral North American Free Trade Agreement
(NAFTA) and other similar treaties.
• ‘Blanco’ (Nopalea cochellinifera) (Figure 1), cultivat-
ed in Valtierrilla and in Michoacán and Tamaulipas Per capita consumption of nopalitos in Mexico in 2013
states; sensitive to frost damage, but performs better reached 6.4 kg, with demand concentrated in the cen-
than ‘Milpa Alta’ when brined − a feature that has tral and north central regions, where the fresh prod-

TABLE 1 Cultivated area of cactus plants in Mexico and product uses

Product Planted area (ha) Irrigated Rainfed

Fruit 55 254 720 54 534
Fodder 16 266 42 16 266
Vegetable 12 039 3 204 8 835
3 000 000
Wild stocks*
(1.5% of the national territory)

Source: SIAP (2015) and *Soberon et al. (2001).

Nopalitos or vegetable cactus, production and utilization 97

uct is available all year round. A decade ago, nopalito Family orchards
consumption was highest at Easter; today, however, as
more information becomes available about their func- Family orchards are a common sight in semi−arid rural
tional properties, demand is increasing, and nopalitos areas, characterized by the diversity of cactus pear spe-
have become a regular feature in supermarket vegeta- cies, the range of uses and the dominance of spineless
ble sections. types that can be fully utilized, providing tender clado-
des, fruits and sometimes mature pads for domestic
In spite of the growing popularity of nopalitos in Mex-
animals. Family orchards are found above all in the
ico, consumption has not taken off in other countries.
northern region. They functioned as gene reservoirs
There is a lack of information on the modes of prepa-
and as a breeding ground for local varieties, which in
ration and the beneficial effects attributed to its regular
turn evolved into the commercial varieties that sustain
consumption, and the cooked product lacks appeal.
the modern commercial market of cactus pears and
Consumers are deterred by the spines and glochids on
nopalitos (Pimienta Barrios, 1990).
the cladodes and the release of mucilage after cooking.
Despite the promotion of nopalito consumption in East
Traditional open−field production
and North Africa to utilize the naturalized stocks, there
has been little impact, and likewise in India and Brazil.
This production system began in Milpa Alta − the old-
As with other vegetables, nopalitos are subject to est nopalitos growing region in Mexico, located in the
sanitary regulations and specific norms are available suburbs of Mexico City − during the 1950s. It expand-
for Opuntia spp. and Nopalea spp. (NMX−FF−068−SC- ed to the neighbouring states of Morelos, Puebla and
Fl−2006 for the national market; CODEX STAN 185 −1 Hidalgo. The system relies on bushy plants (< 1.80 m),
993 for export). Both sets of norms relate to cladode started from single cladodes planted in rows 0.80–1.50
colour, absence of deformations, pest and disease dam- m apart. The individual plants are planted 50−75 cm
age and absence of contaminants. In addition, there are apart, resulting in planting densities of 10 000−40 000
official campaigns covering good agricultural practices plants ha−1 (Figure 2). The basal or mother cladodes
focused on the importance of offering the consumer a are trained to two or three branches; they fill the row
product that is hygienic, safe and free of contaminants. completely, but between the rows only partially so
Enforcement of these norms should help secure and as to allow for transit between all rows. Harvest can
increase the presence of nopalitos on international mar- begin just 2 months after planting, collecting excess
kets. cladodes once the primary branches are defined. Since
it is open−field production, the system is highly de-
pendent on soil moisture and air temperature. Tender
NOPALITO PRODUCTION SYSTEMS nopalitos are very sensitive to frost damage, and even
short episodes can damage the crown of the cladode,
Wild stocks affecting the final shape, productivity and appearance
of the cladode. Therefore, the harvest season is defined
Nopalito gathering is still a traditional activity in the cen- by the presence or absence of frosts; in other words,
tral and northern semi−arid regions of Mexico, which the length of the harvest season is determined by the
− in theory − cover 3 million ha (an overestimate, since duration of the frost−free period. In the highlands of
it includes extensive areas containing only a few plants). Mexico, this period extends from mid−March to late
Nopalitos are usually available at the end of winter or October or November, depending on the specific local
early spring, depending on the length and intensity of microclimate. The continuous emergence of new tender
the frost season. The most common species collected cladodes is also a response to timely harvest, because
wild are: O. streptacantha, O. robusta, O. leucotricha the continuous removal or harvest of tender cladodes
and O. xoconoxtle. The product is usually sold on local stimulates the emission of new ones. On the contrary, if
markets, and is characterized by variable quality, small market conditions are inconvenient, stalling the harvest
volume and irregular availability. Gathering is slowly delays new sprouts.
losing appeal as the density of wild stocks dwindle,
making way for expansion of rangeland, and agricultur- Mini plastic tunnels
al and urban development. Wild stocks are also prone
to the compounded effects of natural disasters: severe This system represents the earliest attempt to control
frosts, drought and natural fires, most likely fuelled by the effects of frost on nopalito production. It was first
climate change. Alternate bearing or cyclic growth also adopted during the 1970–1980s, and is still in use in
takes its toll on the volume of nopalitos and fruit col- small family orchards across the country. There is a wide
lected from wild plants. planting bed (120−150 cm), with 3−4 rows of cladodes;
 98 Crop ecology, cultivation and uses of cactus pear

row spacing is 30−40 cm, distance between plants of the beds will be 1.8 wide, and 45 m long, to
20−30 cm. The number of rows varies according to optimize usage of plastic rolls, which usually come
the width of the tunnel. With the above criteria, a as 100 m long roll. The tunnel can be built using
2−m long fraction of a bed can have 18−24 clado- 3/8” corrugated steel rods, cut into pieces 4m
des. The final planting density varies according to long − enough to build a tunnel 1.4 m high −,
bed width and corridor separation, which in turn shaped as arches of 1.8 m wide base. The arches
depends on the availability of labour and transport. will be inserted in pieces of white or clear colored
In large operations, removal of harvested produce, plastic hose (1/2 in thick) that will protect the plas-
application of fertilizer and manure, and spraying tic from sunburn due to direct contact with the
are performed with a tractor, and a blind bed metal rods. The tip of each rod needs to be buried
between 2−3 planted beds is therefore necessary. 30 cm deep to provide strength to the structure,
Plastic tunnels are built with arched steel rods (3/8 then they are placed 2 m apart. The plastic sheet
caliber [9.5 mm]) fixed to the ground, covered by is placed on top and the edge buried firmly into
transparent polyethylene sheet (caliber 600 mils) the ground on the east side. It is recommended to
and reinforced with polyethylene rope placed diag- use transparent plastic sheets 4m wide, and 600
onally over the tunnel to secure the plastic against mills caliber, with UV protection, special for green-
wind. The arches are usually < 2 m, designed to houses. The other edge is kept in place with some
cover a single bed of nopalitos. Ventilation is pas- burlap bags filled with soil, so they can be easily
sive, provided by manual lifting of the plastic cover removed to open the edge if temperature reaches
during the hottest hours of the day. more than 35 °C. Bury a short wooden stake on
each end of the tunnel 1.2 m away from the edge
This kind of structure provides protection from
of the bed; this stake will anchor seven strands
light frost (in the range of −1° C); its main func-
of polyethylene rope. Then stretch them and tied
tion is to reduce cold winds and increase the tem-
them around on top of each arch, placing them
perature during the day (Figure 2).
evenly on top and both sides along the tunnel, to
There are numerous variations of planting layout,
help the plastic keep the semi−cylindrical shape.
differing in bed width, cladode layout and height
of the plastic cover. Diaz and Maya (2014) describe
Low−tech greenhouses
a system useful for tropical regions:
The beds should be oriented N−S to reduce wind Mexico, like many other subtropical countries, is
force and maximize exposure to sunshine, the size experiencing a boom in protected agriculture; im-

Figure 2
Nopalito production
systems (clockwise:
open field in
Tlanepantla, Mor.
Mexico; mini−tunnels;
greenhouse;
low−rise tunnel)
Nopalitos or vegetable cactus, production and utilization 99

ported and national materials for greenhouses are for high−value crops, such as vegetable seedlings,
readily available for vegetable production. Market strawberries, raspberries, blueberries and peppers,
trends favouring out−of−season and year−round but are being slowly adapted to nopalito culti-
production result in increased product demand, vation in northern Mexico, especially in tropical
benefiting farmers and consumers alike. Given areas with low frost risk. The typical macrotunnel
the good response of nopal to irrigation, high does not have walls; it is built on 2−3 steel pipes
temperatures and fertilizers, protected cultivation secured in the ground, with steel arches and cov-
is the trend in both traditional and new cultiva- ered in UV−protected polyethylene. They are 4−5
tion areas. Several designs have been adapted to m wide and 2−3 m high; they are relatively easy
grow nopalitos, taking advantage of the rusticity to build and of low cost compared with regular
of the plant. Greenhouses for nopalito production greenhouses. The planting layout is designed to
are usually simple, providing little frost protection optimize the protected space available and is sim-
but maintaining a beneficial temperature range ilar to the wide bed system; irrigation is provided
(10−30° C). Attenuation of high temperatures is by dripping lines (Figure 3).
accomplished by means of manually operated win-
dows. Since the nopal plant is self−standing, the Hydroponic nopalitos
metal elements in the structure are reduced to a
minimum: 2 × 2 posts placed 2−3 m apart; arches The most advanced system for growing nopalitos
(usually made of galvanized steel) welded on site is in a hydroponic facility, for which there is an
and reinforced with galvanized steel poly latches; experimental model designed and promoted by
posts secured to the ground with concrete. Tun- the Regional Center for Arid and Semi−arid Zones
nel width varies from 6 to 9 m, depending on the (CREZAS) Postgraduate College. The system is
means of the grower, and planting layouts vary to based on a standard passive cooling plastic−cov-
optimize the covered space (Figure 2). ered greenhouse; it uses subirrigation and includes
a ground cover on alleys to ensure maximum
Macrotunnels water saving, optimum usage of fertilizers and
convenient weed control. Plants are established at
These structures are designed to provide cover high density and are kept bushy to reduce crowd-
and protection against high radiation, excessive ing and improve light interception by the nopal
rain, hail and wind damage, thereby improving canopy (Figure 4).
product quality. They were introduced in Mexico

Figure 3
Low−tech greenhouse
for nopalito production,
San Luis de la Paz, Gto.
Mexico (clockwise:
overall view; roof; foun-
dation; wall)
 100 Crop ecology, cultivation and uses of cactus pear

PRODUCTION PRACTICE of nopal or the synergic and antagonistic interac-

tions among nutrients for all different production
Crop nutrition systems. Nevertheless, Blanco Macías et al. (2006)
report positive interaction between Mg−Ca,
Nopal is generally considered a rustic crop, able Mg−K and K−P, compared with significant and
to survive in poor soils and dry areas. This is true; antagonistic interaction between Ca−N and
however, if the plant is exposed to good quality Mg−N. The optimal concentration ranges for five
soils, abundant fertilization and irrigation, produc- essential nutrients to attain a target yield of 46.7
tivity is significantly boosted. The beneficial ef- kg of fresh matter (considering only cladodes) are:
fects of the combination of manure and synthetic N = 1.29 ± 0.47%, P = 0.35 ± 0.08%, K = 4.24 ±
fertilizers for cactus pear under rainfed conditions 0.88%, Ca = 4.96 ± 1.73%, Mg = 1.61 ± 0.27%.
are reported by Mondragón Jacobo and Pimien- Precise information on the nutrition of vegetable
ta Barrios (1990). Higher responses are achieved nopalitos is also reported by Magallanes Quinta-
under irrigated conditions and using other organic nar et al. (2004), Valdez Cepeda et al. (2009) and
sources, such as vermicompost, composted ma- Blanco Macías et al. (2010), following several years
nure, solarized manure and mycorrhizal products. of field experimentation. According to the curve
The productivity of nopalitos or forage nopal can threshold line technique, nutrient requirements
be expressed as kg of fresh cladodes per plant; it depend on the quantity of nutrients available in
is a function of genotype, environment and crop the soil, expected yield and planting density. In
management practices, including fertilization. order to obtain 95% of the optimal yield (56−60
Little is known about the nutrient requirements kg plant−1, assuming dry matter content of cactus

Figure 4
Nopalito production
under macro−tunnels
(9.6 m wide × 4.3
m high); nopalitos
planted in wide beds
(1.40 m and 1.4 m
alley); hydroponic unit
for nopalito/forage
production (CREZAS−
CP, San Luis Potosí,
Mexico)
Nopalitos or vegetable cactus, production and utilization 101

pear is about 5%), the plant needs to acquire 23.7 g a bimodal pattern, starting at the end of June, with
N, 7.06 g P, 112.5 g K, 95.36 g Ca and 41.7 g Mg. For a slight depression in August and rising again in Sep-
a planting density of 10 000 plants ha−1, the soil must, tember. Frost risk increases from the end of October
therefore, provide 24 kg N, 71 kg P, 1124 kg K, 954 kg to March − when demand for nopalitos is at its peak.
Ca and 417 kg Mg ha−1, for a theoretical yield of 564 Under these conditions, manure and fertilizers should
tonnes ha−1 of fresh matter or 28 tonnes ha−1 of dry be applied at the end of the rainy season and early in
matter. Valdez Cepeda et al. (2009) warns of the high spring, if irrigation is available. In the case of fully rain-
response of nopal plants to nitrogen fertilization, and fed operations, fertilizers and manure can be incorpo-
the possibility of accumulation at toxic levels, as nitrate rated before the end of the rainy season. Fertilization in
concentration can build up in cladodes − potentially excess can cause cladode burning and rapid oxidation
dangerous if used as forage or vegetable. of nopalitos.

The addition of manure (mostly fresh or partially com- Crop irrigation

posted) is traditional in nopalito production, because
the high response of the nopal plant is well known Drip irrigation is an efficient method in nopalito
among Mexican growers – demonstrated by record orchards. The advantages of drip irrigation include
rates of 800 tonnes ha−1 of fresh cow manure, recorded (Vázquez−Alvarado et al., 2009):
in nopalito orchards in Milpa Alta (Fernández et al., • capacity to deliver soluble fertilizers;
1990; Aguilar, 2007; Flores, 2013). Such high rates are
• fertilizer savings of 30−50% (compared with furrow
justified, considering the long residual effect and the
and spray irrigation);
improvement of soil properties; however, they can also
result in burning if applied during the hottest months • reduction of weed competition; and
of the year. Field experiments conducted by Zuñiga • improvement of water and soil conservation.
Tarango et al. (2009) demonstrate that 100 tonnes ha−1
of fresh manure is sufficient to obtain profitable yields Orona Castillo et al. (2003) provide specific data on wa-
of nopalitos. ter use, following a study of superficial and subsuper-
ficial drip irrigation and three soil moisture levels – 30,
Manure application is the subject of increasing atten- 45 and 60% of daily evaporation (type A evaporation
tion, due to growing interest in organic products on the tank) – for several Opuntia species. They indicate a total
modern market and the continuous increase in prices of yield of 100.4 tonnes ha−1 and 18.8 kg of nopalitos per
synthetic fertilizers. Both the direct experience of grow- m3 of water. Total water use is 5 340 m3 ha−1, i.e. less
ers and the results of formal experimentation stress the than that required to produce corn, dry beans and for-
value of fertilization and irrigation at adequate rates in ages (which need 5 500−18 000 m3). Based on these
order to attain high yields (Flores, 2013). The sources data, the authors recommend for O. ficus−indica varie-
and rates applied depend on availability. The most ties: irrigation at 45% of daily evaporation; and annual
common source is fresh cow manure obtained from application of 161, 60.7 and 914 kg ha−1 of N, P and K,
dairies, followed by goat and chicken manure. respectively.

Due to rising transportation costs, there is growing Flores (2013) − on the basis of data obtained for an or-
interest in composts. In an evaluation for nopalito pro- chard planted in furrows, using a drip irrigation system
duction in Milpa Alta, Mexico, Aguilar (2007) reports − estimates water consumption at 391.9 mm year−1.
that similar yields (45−60 tonnes ha−1 year−1) were ob- Plants should be watered when the soil hydraulic ten-
tained with 3.3 tonnes ha−1 of vermicomposted dairy sion reaches 35 cb, associated with a daily consump-
manure instead of 800 tonnes ha−1 of fresh manure. tion of 1.1 mm, i.e. 2 321 m3 ha−1 of water year−1. Water
In this study, the cost of the composted manure was productivity is estimated at 111 kg m3 for O. ficus−indi-
just 22% of the usual rate of fresh manure. Given the ca plants fertilized with dairy manure and triple 17 solid
perennial nature of the nopal plant, manure application fertilizer, i.e. significantly higher than the productivity
may be repeated every 2−3 years based on the initial of the control treatment (76 kg m3). The recommended
rate; whether to apply a single dose or two fractions irrigation schedule is four waterings when the tensi-
depends on the local cost of manual labour. ometer reaches 35 and two when it reaches 70. Water
needs also vary according to location: Nobel (1998)
Fertilization, irrigation and pruning are associated with estimates 3.27 mm day−1 for California, while Fierro et
the rapid emergence of new vegetative buds. Careful al. (2003) recorded 1.65 mm day−1 in Milpa Alta, Mex-
management is important to reduce frost damage, ico. Finally, to improve the efficiency of irrigation, it is
optimize rainfall and, most important, take advantage necessary to consider the rainfall distribution and avoid
of specific market windows. The rainy season in central excess watering, using a single dripping line for two
Mexico presents some variations, but basically follows consecutive rows.
 102 Crop ecology, cultivation and uses of cactus pear

Training (Metamasius spinolae), shot hole weevil (Gerstae-

ckeria spp.) and areole weevil (Cylindrocopturus
Training the nopal plant for nopalito production biradiatus) are found. Earthworms (Phyllophaga
entails moulding it into a regular shape, avoiding sp.) are important in nopalito orchards planted
branching towards the alley, and maintaining plant in light volcanic soils. For detailed descriptions of
height < 1.8 m (in furrow or edge production sys- these pests, refer to Chapter 11.
tems) or below the fourth storey (for mini−tunnel
plantations). Training eliminates excess cladodes Harvesting nopalitos
produced on the second storey. Nopalito plants
are usually allowed to retain 2−3 main branches, Cladodes should be harvested 30−60 days after
configured as “rabbit ears” or in a “fan”; the sprouting, when they weigh 80−120 g and are
remainder are either harvested or discarded. In 15−20 cm long (Flores Valdez, 1995). Tender
practice, training is combined with harvesting; nopalitos present spines and glochids, and latex
however, when market demand stalls, the remov- gloves and knives are, therefore, used during
al of mature cladodes is programmed on the basis collection. The tender pads are carefully removed
of the expected date of market recovery. by inserting the knife between the joints of the
cladodes − an operation requiring skill to maintain
Pruning waste disposal the product intact. They are deposited in baskets
or plastic crates (Figure 5), and sorted according
A remarkable example of the utilization of pruning to market standards: small or baby, medium and
waste is the incorporation of fresh sliced cladodes large. Depending on the market destination, the
in the field from where they were collected. For grower may pack the nopalitos in cylindrical or
this operation the nopalito growers traditionally rectangular bales, wrapped in brown paper or
use machetes and knives. However, recently small polyethylene sheets.
specialized machines have been imported from
Brazil, originally designed to prepare forage nopal,
and similar machines of Mexican design are also BASIC PREPARATION TECHNIQUES
available. This practice increases the moisture and
organic matter levels in the soil and also suppress- Cleaning nopalitos
es weeds temporarily.
The most common method applied to nopalitos
Pests and diseases is still manual cleaning, using only a sharp knife
and neoprene gloves. Other tools have been tried:
Most of the pests present in cactus fruit orchards
are also found in nopalito orchards; however,
their incidence and damage are limited due to the
continuous pruning. The most serious pest in no-
palito production is the cochineal insect; the most
important Lepidopteran insect pests are the zebra
worm (Olycella nephelepsa), the white worm
(Megastes cyclades) and the cactus pad joint
borer (Metapleura potosi). Similarly, cactus weevil

Figure 5
Standard polyethylene
packaging of
nopalitos with 20−30
kg and the traditional
cylindrical bales
containing ≤ 300 kg
or ≤ 4 500 cladodes
Nopalitos or vegetable cactus, production and utilization 103

razor−blade−type knives, hollowed and sharpened On−the−spot cleaning of nopalitos offers the con-
spoons, potato peelers etc., but always with less sumer proof of freshness. This initiative has been
success. A laser machine was once proposed, but tried successfully in many Mexican supermarkets,
due to its high cost and low efficiency, it never where the freshly cleaned product is offered diced
passed the modelling phase. More recently, a or filleted as the customer chooses.
cleaning machine based on rotatory knives and
built from stainless steel has been introduced to Reducing the presence of mucilage
clean large volumes for the preserved nopalitos
industry. Developed by Mexican inventors, it is Mucilage is a common feature of all opuntias.
promoted by the brand Nopalli (http://www.agro- Mucilage is released in response to wounding, and
centro.org/#!services/c21r) and can be adjusted the quantity of mucilage released depends on the
for different shapes and sizes of nopalitos. The variety, as well as the cladode’s age and stage of
machine can process ≤ 40 pads minute−1 with an dehydration. Mucilage is also released when cook-
estimated wastage of < 15%. The same manu- ing, which normally involves boiling. Nevertheless,
facturer produces a second machine designed to mucilage cannot be completely removed from no-
slice clean nopalitos into different shapes: dices or palitos regardless of the cooking method adopted.
strips (Figure 6). Various practices have evolved to reduce or mask
its presence in the final preparation:
The complete removal by hand of thorns and
glochids takes skill and practice and a demonstra- • Addition of dry oregano leaves, bay leaves,
tion can be viewed in a public video (available at chopped onion stems, tomatillo husks, sea salt
https://www.youtube.com/watch?v=XfekDxpqB−I). grains, sodium bicarbonate, lemon juice, corn
husk or garlic cloves, depending on the nopalito
recipe used.
• Interrupting boiling to dip the nopalitos in cold
or chilled water.
• Addition of a few copper or silver coins during
boiling − although this particular method is the
least advisable given the high risk of contamina-
tion.
• Scalding with common salt and oregano for 7
minutes (Pensaben et al., 1995).

The most aggressive cooking methods have a

greater negative effect on the vitamin and mineral
content, reducing the functional properties and
beneficial effects of nopalito consumption. The
salt−oregano method (Pensaben et al., 1995) re-
duces the presence of mucilage without affecting
the green colour typical of the nopalitos.

Figure 6
Cleaning and dicing
nopalito machines,
Mad Industrias, Mon-
terrey N.L., Mexico
104 Crop ecology, cultivation and uses of cactus pear

Scalding and brining are the standard treatments therefore, recommends harvesting nopalitos 2
for large volumes of nopalitos used in the restau- hours after sunrise. In any case, harvest is typically
rant industry in Mexico and they led to the popu- early in the morning when humidity is highest and
larization of nopalitos in the 1990s. the exposure of workers to glochids is reduced,
making it the best time for work in the field.
Cladode acidity
Acidity seems to depend on the variety, according
Nopalitos are characterized by their acidity, which to Aguilar Sánchez et al. (2007) who studied 21
is due to the CAM photosynthetic pathway of cultivars utilized as vegetable: ‘Jalpa’ and ‘Morado
Opuntia and can affect acceptance by new con- Italiano’ around 0.43%; ‘Milpa Alta’ 0.68%; ‘Ore-
sumers. Acidity varies significantly (0.1−0.5% of ja de Elefante’, a forage cultivar, 0.69%. The same
titratable acidity), depending on the time of day. study revealed that ‘Jade’ and ‘Negrito’ present
Acidity decreases during the day and increases at low oxidation, another important trait for agro−
night, regardless of the time of harvest (Corrales industrial processing.
García, 2010). The acidity of several varieties of
nopalitos decreased during the day, in relation to Acidity can be modified by the temperature dur-
the exposure to light − a typical response of CAM ing storage. Chilled storage (5 °C) maintains or
plants (Flores Hernández et al., 2004). Even when slightly increases acidity, while storage at room
detached from the plant, the variations are no- temperature (20° C) decreases acidity. These
ticeable, since the cladode remains alive and pho- fluctuations modify the flavour of nopalitos, and
tosynthetically active. Cantwell et al. (1992) report according to Corrales García et al. (2004), the time
pH values of 0.94% in the morning, decreasing to of processing and consumption are more impor-
0.47% in the afternoon; Pimentel González (2013), tant than the harvest time.
 09

Cochineal breeding
Liberato Portillo and Ana Lilia Vigueras
Department of Botany and Zoology,
University Center for Biological and Agricultural Sciences,
University of Guadalajara, Jalisco, Mexico
 09 Cochineal breeding

INTRODUCTION For a long time, there was controversy over the ori-
gin of cochineal (D. coccus), with some authors cit-
Cochineal (Dactylopius spp) is an American group ing South America (Rodríguez and Niemeyer, 2000;
of Hemipteran insects (Spodek et al., 2014) thriv- Rodríguez et al., 2001). However, research has
ing on Opuntia cactus plants. Cochineal is impor- now led to acceptance that it originates in North
America, where its native ecological environment
tant to humankind for four reasons:
occurs and it has co−evolved with abiotic (Table 1)
1. The insects are a source of carminic acid, a pig- and biotic factors, mainly host plants (Table 2) and
ment (Figure 1) mostly extracted from Dacty- natural enemies (Table 3) (Griffith, 2004; Portillo,
lopius coccus, used in food colouring, cosmetics, 2005; Novoa, 2006). Based on mtDNA data and
drugs, fabrics and many other products (Cañam- climate niche modelling (Van Dam et al., 2015), its
ares et al., 2006; Chávez Moreno et al., 2009). origins were further specified to be in Mexico.
2. Some Dactylopius species have been used for
biological control against invasive cacti (Githure Cochineal has for centuries been one of the most
et al., 1999; Volchansky et al., 1999; De Felice, important pigments and it is still used today (Ser-
2004). rano et al., 2011, 2013). It was first used among
the ancient cultures of America (Donkin, 1977;
3. Dactylopius spp. can become invasive to Opun-
Piña, 1977), and when the Europeans arrived in
tia species where they are non−native (Van Dam
America, its use and breeding spread. In the nine-
and May, 2012).
teenth century, the first artificial dyes were syn-
4. There is ongoing research into the antioxidant thetized and the cochineal market declined and
and antimicrobial properties of carminic acid, almost disappeared. Today, however, the use of
to understand its potential applications in im- natural pigment sources is again on the increase
munology, wastewater treatment and solar cells thanks to their health benefits, and cochineal
(González et al., 2009; García Gil et al., 2007;
Bae and Huh, 2006; El Moselhy et al., 2011).

Figure 1 a b
a) Carminic acid
b, c, d) Diversity of
depredators

c d
Cochineal breeding 107

production is being re−established with a range Montiel (1995) states that since males undergo
of breeding technologies in many tropical and complete metamorphosis, their immature stages
subtropical areas of the world. must be termed larvae. Since characteristics of
dimorphism do not appear in the first immature
This chapter describes the various methods for
stages, and it is the bodies of the female that are
cochineal breeding and outlines harvest and
used to obtain the pigment, for the sake of con-
post−harvest management practices, with a focus
venience, all the immature stages are referred to
on both its biology and its ecology in native and
as nymphs.
non−native areas.
Life cycle
BIOLOGY Females and males go through the egg stage
(Figure 2a) and two immature stages. The
Dactylopius is the only genus of the Dactylopiidae biological cycle from egg to adult lasts 90−128
family (Hemiptera: Coccoidea). All 11 species are days, depending on temperature and other factors
parasitic of cactus plants (De Lotto, 1974; Spodek (Marín and Cisneros, 1977).
et al., 2014) and present a disjunct distribution
• Nymph I. The female in this immature stage
(Van Dam and May, 2012), native to:
has two substages: mobile and fixed. The first
• South America: D. austrinus De Lotto, D. con- substage is also known as crawler (Figure 2b),
fertus De Lotto, D. zimmermanni De Lotto, D. characterized by movement and absence of
salmianus De Lotto and D. ceylonicus Green; or white wax; in < 24 hours, it has to find a place
• North America: D. bassi (Tozzetti), D. coccus in the cactus pear to insert its mouthpart. Once
Costa, D. confusus (Cockerell), D. gracilipilus the nymph is fixed, it passes into the second
Van Dam and May, D. opuntiae (Cockerell) and substage, secreting large wax filaments over
D. tomentosus Lamarck. the body (Figure 2c); within days, the filaments
The species most used for commercial purpos- disappear and make way for a powdery wax.
es − due to its high pigment content (> 20% of • Nymph II. After a first moult (Figure 2d), this
carminic acid) − is the domesticated species, D. immature stage (Figure 2e) is bright red and
coccus (fine cochineal). In ancient times, it was within hours starts to be covered with wax.
known as nocheztli, a Nahuatl word meaning Nymph I and II stages present no ostensible
"blood of cactus", in reference to the insect and differences between males and females.
the dye it produces (Wright, 1963). In 1758, it was • Adult female. The female cochineal under-
called Coccus cacti by Linnaeus; in 1835, Costa goes another moult (Figure 2f), synchronized
classified it as Dactylopius coccus (Piña, 1977). with the adult males for copulation. During this
stage, the female increases in size (Figure 2g).
Sexual dimorphism Each female usually produces 420 eggs, but
about 10% of females show infertility (Vargas
Dactylopius males have wings on their adult instar; and Flores, 1986), which reduces the produc-
they are smaller than females and highly mobile. tion of new crawlers.
Females are about 6.24 mm, wingless, stationary, • Adult male. The male cochineal begins to devel-
oval−shaped and covered with a powder wax. op a cocoon (Figure 2h) to emerge as an adult

Figure 1
Two forms of
coccidoculture:
e) under cover
f) in the open

e f
 108 Crop ecology, cultivation and uses of cactus pear

male (Figure 2i). The male presents two pairs of both abiotic and biotic factors and it is, therefore,
wings, two tails, a non−functional mouthpart, vital to understand which factors are present in
and mobility to search for females for mating. a particular area. In the rainy season, it is recom-
mended to carry out propagation of cochineal on
isolated cladodes under protection. Indeed, in al-
COCHINEAL BREEDING most all of Mexico, coccidoculture takes this form
because of:
Abiotic and biotic factors • abiotic factors − unfavourable conditions in the
open (Table 1); and
Coccidoculture or cochineal breeding is an activity • biotic factors − essentially natural enemies (Ta-
that can be carried out in many places in tropical ble 3), with a wide diversity of depredators in
and subtropical regions, but it is conditioned by Mexico (Figures 1b, c and d).

Figure 2:
a - i) The biological
cycle from egg to adult
Cochineal breeding 109

TABLE 1 Abiotic factors limiting cochineal breeding

Factor Main effect Source

Altered length of biological cycle Méndez, 1992
Temperature
Dehydration of nymphs Flores, 1995
Elimination of 100% of cochineal population Flores, 1995
Rain Suffocation of cochineal Alzate and Ramírez, 1777
Hindered fixation and development of nymphs Flores, 1995
Limited fixation of cochineal Aquino, 1992
Wind
Increased dispersion and dragging of nymphs Flores, 1995
Movement of crawlers influenced Flores, 1995
Light
Growth and development of cochineal enabled Aquino, 1992
Hail Cochineal thrown to the ground Alzate and Ramírez, 1777
Frost Elimination of > 50% of cochineal Alzate and Ramírez, 1777

Cochineal breeding in Mexico has been reported on several Opuntia species (Table 2); the most impor-
tant cultivars for commercial production are spineless ‘Atlixco’, ‘Chicomostoc’ and ‘Milpa Alta’, which all
belong to O. ficus−indica.

TABLE 2 Opuntia species reported to host Dactylopius coccus in Mexico

Host species Source

Opuntia amyclaea Tenore Vigueras and Portillo, 2014
O. atropes Rose Rodríguez and Portillo, 1989
O. cochenillifera (L.) Salm−Dycka Piña, 1981
O. crassa Haworth Vigueras and Portillo, 2014
O. ficus−indica (L.) Miller Piña, 1981
O. fuliginosa Griffiths Portillo and Vigueras, 2003
O. incarnadilla Griffiths Vigueras and Portillo, 2014
O. jaliscana Bravo Portillo and Vigueras, 2014
O. megacantha Salm−Dyck Piña, 1981
O. pilifera Weber Portillo and Vigueras, 2003
O. sarca Griff. ex Scheinv. Portillo and Vigueras, 2003
O. streptacantha Lem. Piña, 1981
O. tomentosa Salm−Dyck Piña, 1981
O. undulata Griffiths Portillo and Vigueras, 2003
a
This species belongs to the subgenus Nopalea.

TABLE 3 Predators of Dactylopius coccus Costa recorded in Mexico

Species Order and family

Laetilia coccidivora Comstock (Lepidoptera: Pyralidae)
Eosalpingogaster cochenillivora Guerin−Meneville Diptera: Syrphidae
Sympherobius amiculus Fitch Neuroptera: Hemerobidae
Hyperaspis trifurcata Schaeffer Coleoptera: Coccinellidae
Chilocorus cacti Lineo Coleoptera: Coccinellidae
Leucopis bellula Diptera: Syrphidae

Source: Modified from Vigueras and Portillo (2014).

110 Crop ecology, cultivation and uses of cactus pear

Cactus pear plantations for cochineal breeding Infestation methods

Commercial plantations of cactus pear always use Regardless of the cochineal breeding method selected,
spineless cultivars of Opuntia ficus−indica for cochineal infestation is necessary in order to achieve harvest.
breeding, whether in the open or under cover. Cultivars Infestation methods require careful selection. Whether
with yellow fruits are preferred, but white, red and infesting whole plants or single mature cladodes (Llan-
purple cultivars may also be used. Plantations have an deral and Campos, 1999), it is highly recommended to
average of 20 000 plants ha−1, but the number may use different containers called “nests” to hold the adult
vary depending on soil quality (good drainage), altitude females in oviposition period (OP). New infestation
(800−2 300 m asl), temperature (13−27° C) and water methods are frequently introduced on open−air com-
availability (Salas, 2016). Plants receive around 30 tonnes mercial plantations to facilitate infestation and reduce
year−1 of manure (cow, goat or even chicken), and addi- labour:
tional fertilization as required (N100−P50−K10). • Oaxacan. Cylinders made of woven palm (length 10
cm, diameter 2 cm) carrying adult females in OP are
Cochineal breeding methods held by the spines on the host plants for ≤ 20 days for
infestation. Containers can be rotated to other plant
There are two forms for coccidoculture: under cover areas for better results. In Bolivia, the same concept
and in the open (Figures 1e and f). The choice of is applied using hollow cane sugar stems with a lot
breeding method largely depends on the local climatic of small holes (Tukuypaj, 1993), and recently in Peru
conditions. Structures for cochineal breeding under pro- using plastic hose pieces.
tection are traditionally made of wood, shade screens,
• Peruvian. The adult females in OP are introduced in
palm leaves, plastic (transparent or black) and other small tulle bags (10 × 10 cm), hung on the spines of
materials (Santibáñez, 1990; Méndez, 2013); in recent the host plants until infestation is observed (Quispe,
years, however, intensive production has used large 1983).
greenhouses (Escalante, 2013). The main methods are
described below: • Ricci. A small triangular incision (sides 2 × 2 × 2 cm,
depth 0.5 cm) is made in the centre of a cladode,
• Microtunnel. Structures (height approx. 1 m) are which becomes dry and serves as a support for the
made of greenhouse polyethylene plastic or shade adult females in OP to facilitate the infestation (Porti-
cloth (1 mm thick) (Figure 3a) to protect infested llo et al., 1992).
2−3−year host plants or single planted 1−year cla-
• Infested cladode. Infested cladodes with adult fe-
dodes (Vigueras and Portillo, 2014). This method is
males in OP are placed at the bottom or between two
useful in regions where heavy rains are common or
cladodes of the host plants to facilitate infestation
winter is cold (minimum 0−10° C).
(Tukuypaj, 1993).
• Tapesco. Shelters have a roof made from reeds,
• Cotton fabric. Small pieces of cotton fabric are
wood, logs, shade cloth, woven palm or plastic (Ro-
placed on adult females in OP, allowing the crawlers
dríguez and Portillo, 1989; Santibañez, 1990). These
to get on them; the pieces of fabric are then dropped
sheds are used to hang single mature cladodes for
on the host plants to be infested (Quispe, 1983).
small cochineal breeding or to increase a particular
cochineal colony while protecting it from undesirable • Gravity. As adult females start OP on hanging cla-
factors. dodes, the crawlers fall with the force of gravity on
new ones collocated at the bottom and yet to be
• Intensive production. This system (proposed by
infested. This method is adopted in new greenhouses
the Campo Carmín enterprise) was implemented for
in Mexico used for intensive production of cochineal
the first time in 2002. Greenhouses are built using
(Escalante, 2013). Gareca (1993) also describes a
modules. They cover 1 356 m2 and have a minimum
gravity−based system: a mesh tray with adult females
capacity of 432 000 hanged spineless cladodes
in OP is placed at the top, and twice daily horizontal
(Escalante, 2013). This method is currently the most
movements make the crawlers drop down to infest
common in Mexico (Figure 1e).
cladodes at the bottom.
• Irrigation. Single cladodes are established under
• Paper container. The simplest method: any piece
hydroponic systems in Peru and Mexico, using for-
of discarded paper is used to make a small container
mulations of macro− and micronutrients to promote
(length 8 cm) with adult females in OP inside (Téllez,
growth of the host plants and cochineal (Vigueras et
1911); it is attached by spines to the host plants allow-
al., 1993). This method is common on the south coast
ing the infestation (Figure 3b).
of Peru (Figure 1f), where high−density plantations
(around 9 000 or more plants ha−1) with a fertigation
system are infested in the open (Flores, 1995).
Cochineal breeding 111

Harvesting To dry cochineal, solar dryers and stoves are used, as

well as direct sunlight and in the shade; for shade−
Harvest is a delicate process that involves separation drying, the site must be completely dry. The selection
of the female insects from the host plant when they process involves the separation of the males, immature
are about to start oviposition, taking care to avoid in- cochineal, moults, wax and any other element that is not
juring them. There are several methods for harvesting the adult female cochineal. Selection also takes size into
cochineal, depending on the breeding method (open or account, as a bigger cochineal means more pigment;
under cover) and host plant (spineless or spiny). however, the end objective is to obtain cochineal with
a high carminic acid content. Currently the preferred
In the open cochineal has the following characteristics:
In the Andean region of Peru and Bolivia, there are • > 22% carminic acid content;
many instruments made of sticks (length 0.60−1.50 m), • 14−mesh size (> 1.295 cm); and
with spoons, brushes or metal strips attached at the end
• < 1% of impurities.
to gather cochineal (Figure 3c). Spoons and brushes are
used on spineless plants and metal strips on spiny ones. Cochineal meeting these specifications has an open
On the south coast of Peru and in Chile, cochineal is market; otherwise there is virtually no demand and the
harvested from spineless plants using brushes without price is low. The price of cochineal is directly influenced
the stick. In Chile and South Africa, another method by the carminic acid percentage (1% = 1 US$), although
with pressurized air has been used to harvest cochineal the price may vary depending on the market.
from spiny cladodes (Brutsch and Zimmermann, 1993).
In the Canary Islands, cochineal is harvested in Novem-
ber, 3 months after infestation, using a large spoon−like
ECOLOGY
instrument called a “milana”.
During harvest, the collected insects fall into bags made Any organism is in balanced interaction with the factors
of cotton, and are later distributed on wood, metal or prevailing in its native environment; however, if moved
clay trays, taking care to avoid clumping that may crush to a new range (non−native area), the equilibrium may
the fresh insects. be broken. This loss of equilibrium produces conse-
quences, one of which is that it may become invasive.
Under cover Biological control is the most common tool for reducing
In Mexico, the cochineal harvest takes place about 3 the effects of this phenomenon, controlling the alien
months after infestation; however, it may be carried out invasive species until it becomes non−invasive and natu-
at any time of year and using a wide range of instru- ralized (Shine et al., 2000).
ments (reeds, metal spoons, brushes, sticks and plastic Given their usefulness to humankind, species of Opun-
scrapers). Selection of the most suitable instrument tia and Dactylopius that are of particular interest for
depends on different factors: breeding method; extent commercial production, are frequently moved from
of cultivation area; and cultivar characteristics (spineless, their original distribution in the Neotropical and Ne-
spiny or even pubescent plants). Where cochineal breed- arctic regions, and they are now found in Afrotropical,
ing is a recent activity, new techniques and instruments Australasian, Palearctic and Oriental regions (Anderson,
may need to be developed for harvesting (Vigueras and 2001; Bravo Hollis, 1978; Majure et al., 2012b). These
Portillo, 2014). movements have led to the creation of new weed and
plague problems, with D. opuntiae on O. ficus−indica
Post−harvest plantations in Brazil, Spain and other countries of the
Mediterranean Basin (Portillo, 2009; Vasconcelos et al.,
Post−harvest activities involve the killing (in Mexico this 2009). Cochineal species may be considered invasive;
step is known as “sacrifice”) and drying of cochineal fine or domesticated cochineal (D. coccus) has been
(Santibáñez, 1990), as well as its selection. There are declared an invasive agent in Ethiopia (Belay, 2015)
several methods for killing cochineal: (Figure 3d). Nevertheless, the species is considered a
useful insect worldwide. Some species of Dactylopius
• immersion in boiling water (Alzate and Ramírez, 1777);
have been and still are used as biological control agents
• suffocation with steam; of several Opuntia species that are considered invasive
• freezing in temperatures below 0 °C (experimental); (Table 4). There is the risk that new issues will continue
• suffocation with hexane; to arise among cochineals and Opuntia cultivars.
• asphyxia in hermetic containers; and D. coccus shows adaptation and selectivity for particular
• heating with hot air, inside ovens, on clay posts (Téllez, Opuntia species or cultivars (Portillo et al., 1992), and
1911) or by solar radiation (temperatures around 40°C). similar behaviour is reported for D. opuntiae (Volchan-
 112 Crop ecology, cultivation and uses of cactus pear

Figure 3
a) Microtunnels
b) Infestation method a b
with paper container
c) Using instruments
made of sticks to
gather cochineal
d) Invasive cochineal
species

c d

TABLE 4 Dactylopius species used to control invasive Opuntia species

Cochineal species Opuntia species Country Source

D. austrinus O. aurantiaca Guilles ex Lindley Australia 1970; South Africa 1979 Moran and Cabby (1979)
D. ceylonicus O. vulgaris Miller Sri Lanka 1863 Volchansky et al. (1999)
D. opuntiae O. ficus−indica South Africa 1938 Moran and Zimmermann (1984a)
D. opuntiae O. stricta Haworth Australia 1921 Hoffmann et al. (2002)
D. opuntiae O. ficus−indica Brazil 2001 a
Batista et al. (2009)

a
In Brazil the introduction of D. opuntiae was not for biological control; it has now become an invasive species, destroying thousands
of hectares of cactus pear plantations.

sky et al., 1999; Githure et al., 1999). Cactus pear cochineal (D. opuntiae), cactus moth (Cactoblastis
cultivars have been introduced to many countries cactorum) and black spot (Pseudocercospora
outside Mexico and − from an ecological point of opuntiae) (Figure 3e). Plant sanitary borders must
view − cactus pear is often a foreign biotic factor be made stricter to counteract the new infestations
in a non−native area. Although it is sometimes of wild cochineal (D. opuntiae) that are constantly
considered invasive, it has now become a useful el- arising, most recently in Israel, Lebanon and Mo-
ement, known and appreciated by the local inhab- rocco. Wild cochineal is an insidious plague and
itants. It is utilized in many ways and ethnobotany highly aggressive to cactus pear, as was demon-
relations have been established in many communi- strated in Brazil (Figure 3f). Continuing efforts are
ties around the world. Nevertheless, the reality is needed to manage Dactylopius invasions and find
that new biotic factors may arise and these must a threshold; to this end, the authors are currently
be faced; potential threats to cactus pear are wild gathering more information.
Cochineal breeding 113

e
Figure 3
e) Black spot
(Pseudocercospora
opuntiae)
f) Wild cochineal

f
 10

Diseases of cactus pear

Giovanni Granata,a Roberto Faeddaa and María Judith Ochoab
a
Department of Agriculture, Food and Environment, University of Catania, Italy
b
Faculty of Agronomy, National University of Santiago del Estero, Argentina
10 Diseases of cactus pear

INTRODUCTION through to the other side of the cladode. The injuries

reach a diameter of 3−4 cm at the end of the infection.
Similar to other crops, cactus pear (Opuntia ficus−in- Different symptoms can appear simultaneously on the
dica [L.] Mill.) suffers from several biotic and abiotic same cladode (Ochoa et al., 2015b). Similar symptoms
diseases; how important they are depends on the des- have been associated with anthracnose caused by
tined use of the crop. Colletotrichum gloeosporioides (Penz.) Penz & Sacc.
However, on the basis of pathogenicity tests, Quezada
There is a shortage of accurate information on the Salinas et al. (2006) affirmed that C. gloeosporioides is
aetiology and epidemiology of the pathogens, and not the cause of black spot disease in cactus pear.
the scientific literature does not sufficiently cover the
geographic distribution and worldwide economic im- Disease management: Survey the cactus pear planta-
pact of many cactus pear diseases. Nevertheless, it is tions after the rainy season to detect and eliminate the
certain that most of the infectious diseases are caused cladodes at the first sign of disease symptoms. Adopt
by fungi, while a very small number of bacteria, phyto- preventive pruning to allow good ventilation. Apply
plasmas and viruses are reported as pathogens. copper−based fungicides just before the rainy season.

Due to the dry climate of the regions where cactus pear Cladode and fruit rot
is grown worldwide, these diseases become problem-
atic only in certain periods when conditions become Causal agent: Lasiodiplodia theobromae (Pat.) Giff. &
favourable. Nowadays, however, expansion of cactus Maubl. (teleomorph: Botryosphaeria rhodina Berk. &
pear cultivation, dense plantings and climate change, M.A. Curtis, Arx).
may contribute to a higher incidence and greater se- Syn.: Botryodiplodia theobromae (Pat.).
verity of diseases, as well as the possible emergence of Host range: More than 500 hosts, including humans.
new or unusual diseases. Early detection, precise iden- Geographical distribution: Brazil, Egypt, Italy, Mexi-
tification and strict monitoring of the diseases in fields co, South Africa, United States of America.
and on propagation material are essential to prevent Symptoms and signs: Black roundish areas (15–50
their diffusion. mm diameter) appear on cladodes leaking a black
This chapter provides concise descriptions of the most gum exudate from the margin of the lesion. Pycnidia
common diseases affecting cactus pear and recom- are often evident on the surface of the infected area.
mendations for their control. Probably, the same disease has been reported as gum-
mosis canker on cladodes (Figure 2) in Linosa (Italy)
and the causal agent attributed to Botryosphaeria ribis
(syn. Dothiorella ribis) (Somma et al., 1973).
FUNGAL DISEASES
Disease management: Avoid wounds, remove and
Black spot destroy diseased specimens. Apply fungicides (tiaben-
dazole and tiofanat−metil) proved effective for reduc-
Causal agent: Pseudocercospora opuntiae Ayala−Es- ing disease incidence.
cobar, Braun & Crous.
Host range: Opuntia spp., Nopalea spp. Alternaria dry rot
Geographical distribution: Bolivia, Brazil, Mexico, Peru.
Symptoms and signs: The first symptoms are cuticle Causal agent: Alternaria spp.
discoloration in the form of a circular spot; it becomes Host range: Highly polyphagous.
transparent and oily in appearance, and the brown Geographical distribution: Argentina, Italy, Brazil,
central area increases in size. Later, the cladode tissue Egypt, Mexico, South Africa.
presents a light brown colour at the margins of the Symptoms and signs: The first symptoms are circular
spot which changes to dark brown, presenting tissue chlorotic spots around the spines or wounds on the
collapse. The affected tissue turns black; conidiophores cladodes and fruits, which later become dark and
and conidia in the form of small grey protuberances necrotic (Figure 3). Penetration sites are generally the
can be seen (Figure 1). Finally the affected zone sinks spines, but may also be wounds in the cuticle caused
Diseases of cactus pear 117

Figure 1
Amaranthus cruentus

Figure 2
Gummosis canker
(Linosa Island, Italy

by hailstones. In South Africa, different species of Symptoms and signs: Root and stem rot is
Alternaria were isolated from cladodes with dry caused by Armillaria mellea, a basidiomycetous
rot symptoms (Swart and Kriel, 2002; Swart and fungus that colonizes the shoot and main roots of
Swart, 2002; Ammar et al., 2004; Souza et al., the plant and produces white mycelium and cor-
2010). In Italy, Alternaria alternata causes golden dons of hyphae, called rhizomorphs, that spread
spot (Figure 4) producing spots that are dark in from the infected tissues and contaminate neigh-
the centre and golden yellow around the edges bouring plants. Infected cactus pear plants suffer
(Granata and Sidoti, 1997). The same disease is from a decline in tissue turgor and chlorosis re-
known as mancha de oro or secamiento de la sulting from phloem disorganization and altered
penca in Mexico (Gutiérrez, 1992). A. alternata water and mineral nutrition. The fruit growing
also causes a post−harvest fruit decay. The symp- on infected plants do not reach full ripeness and
toms are a superficial dry rot of the peel, black remain mummified on the cladode. Basal stem
in colour (Figure 5). Generally, the lesions with rot symptoms can be observed with the release
irregular margins are localized in the central part of viscous exudate. Characteristic white mycelium
of the fruit and the decayed peel can be scraped fans of hyphae can be found beneath the surface
off (Faedda et al., 2015b). of the lesions. The rotten portion of the stem
has a bright red border which extends upwards
Disease management: Spray with a fungicide,
above the soil line. The disease may affect the
such as copper, mancozeb and iprodione, for ef-
main roots (Raabe and Alcon, 1968; Magnano di
fective control of the disease, especially after hail.
San Lio and Tirrò, 1983).
Apply hot water treatment for efficient control of
post−harvest disease. Disease management: Since Armillaria often
remains in the soil in residues from previous
Armillaria root and stem rot cultivations inevitably infecting new crops, it is
recommended not to cultivate soil infected by
Causal agent: Armillaria mellea (Vahl) P. Kumm. this fungus for 2–3 years and to remove sources
Host range: Highly polyphagous. of inoculum (roots of previous cultures). Effective
Geographical distribution: Italy, United States chemical control of the disease is not available yet.
of America.
118 Crop ecology, cultivation and uses of cactus pear

Figure 3
Alternaria dry rot
(Tunisia)

Figure 4
Golden spot (Italy)

Figure 5
Alternaria fruit rot
(Italy)

3 5

Figure 6 Cottony rot

Cottony rot
(Peru)
Causal agent: Sclerotinia sclerotiorum (Lib) de
Bary.
Host range: Polyphagous.
Geographical distribution of disease: Chile,
Mexico.
Symptoms and signs: The fungus causes a
cottony rot in the cladodes (Figure 6). Infected
cladodes initially become decoloured followed by
a slight softening of the cuticle. They later turn
dark, flake off and become covered with a white
wool. Numerous black sclerotia soon appear from
the decayed tissues (G. Granata, personal com-
munication).
Disease management: Remove and destroy in-
fected cladodes. It is of primary importance that
the sclerotia are prevented from touching the soil
6
where they may remain alive for many years.
Diseases of cactus pear 119

Figure 7
Phytophthora foot rot
(Italy)

Figure 8
Grey mould (Italy)

7 8

Rust approximately 90 different families and different

isolates have distinct host ranges.
Causal agent: Aecidium opuntiae Magn. Geographical distribution: Italy.
Syn.: Puccinia opuntiae (Magnus) Arthur & Holw. Symptoms and signs: Phytophthora nicotianae
Host range: Opuntia spp., Bouteloua simplex Lag. is the causal agent of foot rot in cactus pear in It-
Geographical distribution: Bolivia, Italy, Peru. aly (Cacciola and Magnano di San Lio, 1988). The
Symptoms and signs: Known as “Roya” in most typical symptom of foot rot is gum exudate
South America, the disease is caused by Puccinia oozing from the lower stem. Diseased plants pres-
opuntiae, which produces yellow–orange pustules ent water−soaked lesions with irregular brown
on the surface of cladodes or fruits. As the rust margins on the basal stem surface and a soft rot
spot dries, a hole develops on the cladode. Sim- of internal tissues that turn brown or reddish.
ilar symptomatology was described by Goidanich Affected plants become chlorotic, growth slows,
(1964) as rust scab and the causal agent attribut- and cladodes wilt, causing the plant to fall over
ed to Phyllosticta opuntiae Sacc. & Speg. (Figure 7).
Disease management: Remove and bury the Disease management: Clay soils where water
infected cladode. stagnates are prone to harbouring Phytophthora.
For this reason, cultivate cactus pear in well−
Scaly rot drained soil to prevent waterlogging.

Causal agent: Scytalidium lignicola Pesante. Pythium crown and stem rot
Host range: Auricularia polytricha, Citrus para-
disi, Mangifera indica, Manihot esculenta, Vanilla Causal agent: Pythium aphanidermatum Edson
fragrans, humans and animals. (Fitzp.).
Geographical distribution: Brazil. Host range: Widely polyphagous.
Symptoms and signs: The disease is widespread Geographical distribution: Mexico.
on cactus forage (palma−forrageira) in the north- Symptoms and signs: The first symptoms are
east region of Brazil. The symptoms begin with a soft, dark brown lesions on the cladode at the soil
dry rot that develops a scaly appearance at the line. The lesions gradually reach the upper part of
base of the cladodes (Souza et al., 2010). Plentiful the cladode and then extend upwards to the next
yellowish gum exudates gush from the infected cladode. Rotten cladodes are associated with root
tissues. This polyphagous fungus is a pathogen rots. The diseased plants finally collapse and large
causing skin, nail and hair infections in humans. plants collapse more rapidly than small plants
Disease management: To date, there has been (Rodríguez Alvarado et al., 2001).
no research to determine the optimum condi- Disease management: Avoid excessive water
tions for disease development. Remove infected and maintain good soil drainage.
plants to reduce the amount of inoculum infect-
ing nearby plants. Fusarium root rot

Phytophthora foot rot Causal agent: Fusarium oxysporum f. sp. opun-

tiarum (Pettinari) W.L. Gordon.
Causal agent: Phytophthora nicotianae Breda de Host range: Echinocactus grusoni, Schlum-
Haan. bergera truncate.
Host range: The pathogen infects plants from Geographical distribution: Brazil, Italy.
120 Crop ecology, cultivation and uses of cactus pear

Figure 9
Penicillium fruit rot
(Italy)

Figure 10
Macrophomina
soft rot (Italy)

9 10

Symptoms and signs: Diseased plants present infection by this fungus. Infected tissues become
root rot with intense red discoloration of affected brown−coloured, soft and watery. The lesions en-
tissues. If root rot becomes severe, infected plants large, a white mycelium emerges from the cracks
may develop other symptoms including stunting of decayed peel and a blue−green spore mass ap-
and wilting (Pettinari, 1951). pears on the surface of the peel along the lesions.
Disease management: Plant in well−drained soils Disease management: Control the disease by
and reduce soil compaction to make conditions less careful picking and handling of fruits to reduce
favourable to infection by Fusarium species. rind damage. In packing houses and storage fa-
cilities, sanitary practices may prevent sporulation
Grey mould of diseased fruits and reduce the amount of inoc-
ulum.
Causal agent: Botrytis cinerea Pers. (teleomorph:
Botryotinia fuckeliana [de Bary] Whetzel). Macrophomina soft rot
Host range: Highly polyphagous.
Geographical distribution: Italy. Causal agent: Macrophomina phaseolina (Tassi)
Symptoms and signs: The pathogen penetrates Goid.
through the wound that tends to occur when the Host range: Polyphagous with more than 500
fruit is detached from the cladodes. The infected plant hosts.
area of the fruits is dark grey and usually circular; Geographical distribution: Italy.
the internal tissues are decayed and soft (Figure Symptoms and signs: Symptoms begin with a
8). (G. Granata, personal communication). brown soft rot originating at the proximal portion
Disease management: Avoid wounds during and extending to the entire fruit at post−harvest
harvest and post−harvest processing. (Faedda et al., 2015b). In the advanced stage, the in-
fected area is covered by mycelial mass (Figure 10).
Penicillium fruit rot Macrophomina has also been reported in Brazil and
Mexico, causing charcoal rot on forage cactus pear
Causal agent: Penicillium spp. (Oliveira et al., 2003; Méndez Gallegos et al., 2009).
Host range: Highly polyphagous. Disease management: To date, there has been
Geographical distribution: Worldwide. no research on the control of this disease.
Symptoms and signs: Penicillium rot is regarded
as a major post−harvest disease of cactus pear Other fungi
fruits in many production regions of the world.
Penicillium italicum Wehmer and P. digitatum Besides those discussed above, several other fungi
(Pers.) Sacc. are reported as major causal agents have been found on cladodes and fruits of cactus
of this disease (Schirra et al., 1999b). Other Pen- pear plants. However, many of them are sap-
icillium species, including P. citrinum Thom, P. robes or secondary pathogens, often recovered
expansum Link (Scherm et al., 2003; Oliveri et al., from weakened plants, and their role as primary
2007) and P. polonicum K.M. Zalessky (Faedda pathogens has not been clearly proved. The most
et al., 2015a) are reported on cactus pear fruits commonly occurring fungi are outlined below.
in Italy (Figure 9). Physical injuries to the peel • Aspergillus niger − found in Egypt associated
during harvesting and handling as well as cold with rotted cladodes and fruits (Ammar et al.,
storage predispose the fruits of the cactus pear to 2004).
Diseases of cactus pear 121

• Candida boidimi − associated with soft rot of • Rhizopus sp. − found in Brazil and South Africa
cladodes and fruits in Italy (Granata and Varva- (Swart and Swart, 2002; Souza et al., 2010).
ro, 1990).
• Capnodium spp. and other fungi − cause sooty
mould in Mexico (Méndez Gallegos et al., 2009;
BACTERIAL DISEASES
Mondragón Jacobo et al., 2012).
• Cercospora sp. − reported in Bolivia and Peru Black soft rot
causing grey−coloured circular necrotic spots on
cladodes (Granata and Sidoti, 2002). Causal agent: Erwinia carotovora subsp. Caroto-
• Colletotrichum gloeosporioides − found in Bra- vora.
zil, Korea and Mexico, causing anthracnose of Host range: Polyphagous.
cladodes. The symptoms consist of dark brown Geographical distribution: Argentina, Italy.
spots usually starting at the pad edges and Symptoms: In spring, the cladodes display wa-
then extending over the whole pad (Osada and ter-soaked spots that become brown and coalesce
Càrcamo, 1991; Kim et al., 2000; Souza et al., (Figure 11). The external tissues become dry and
2010). frequently cracked; the internal tissues turn brown
• Fusarium species − found on cactus pear in to almost black. These symptoms can also be ob-
many parts of the world, including Brazil, Egypt, served on the fruits (Varvaro et al., 1993; Saad et
South Africa and Mexico (Swart and Kriel, 2002; al., 1998).
Ammar et al., 2004; Swart, 2009; Souza et al., Disease management: Remove and destroy the
2010): infected segments. Spray immediately with copper
− Fusarium oxysporum, F. proliferatum and F. compounds in high-risk situations, such as after a
sporotrichoides − associated with dry necrotic major wind or hail event.
lesions of cladodes. Figure 11
− F. solani − reported as causal agent of top rot. Black soft rot (Italy)
− F. lunatum − reported as causal agent of cir-
cular spots of cladodes.
• Cladosporium sp. − associated with pad spot in
Mississippi, United States of America (msucares.
com).
• Curvularia lunata − reported in Brazil and Mex-
ico causing symptoms of circular cladodes spot
(Souza et al., 2010; Flores Flores et al., 2013).
• Cylindrocarpon sp. − found in South Africa asso-
ciated with cladode necrosis (Swart and Swart,
2002).
• Hendersonia opuntiae − causes a disease known
as “scorch” or “sunscald”. The symptoms consist
of spots at first distinctly zoned, later enlarging
until entire cladodes turn a reddish brown. The
centre of the diseased area is greyish brown and
cracked (Horst, 2013).
• Mycosphaerella sp. − causes necrotic spots
on cladodes in South America. This disease is
known as mancha plateada in Mexico (Granata,
1995).
• Phoma sp. − found in Argentina, causing ne-
crotic spots on cladodes (Granata, 1995).
• Pleospora sp. − causes necrotic spots on clado-
des (Granata, 1995). Crown gall
• Pollaccia sp. − reported to cause cladode rot
on thornless cactus in the region of Alagoas Causal agent: Rhizobium radiobacter (previously
(Northeast Brazil) (Franco and Ponte, 1980). known as Agrobacterium tumefaciens).
122 Crop ecology, cultivation and uses of cactus pear

Figure 12 Host range: Highly polyphagous with more than

Phytoplasma 90 families of plants.
disease (Italy)
Geographical distribution: Mexico.
Symptoms: A tumorous overgrowth (gall) forms
at the base of the stem. The gall colour ranges
from brown to black and cracks at maturity. The
tumour at the basal stem varies in size with a di-
ameter of ≤ 15 cm (Gutiérrez, 1992).
Disease management: Do not plant susceptible
varieties in soils known to be infested with this
pathogen. Preventive treatment of cuttings with
the non-pathogenic biocontrol organism A. ra-
diobacter is a relatively inexpensive and effective
means of managing the development of crown
gall in commercial operations.

PHYTOPLASMA AND VIRUS DISEASES

One of the most serious diseases in cactus pear
orchards reported in Mexico is commonly known
as engrosamiento de cladodios (cladode enlarge-
ment) or macho (Pimienta Barrios, 1990). Typical
symptoms of this disease – assumed to be caused
by a phytoplasma – are severe stunting of clado-
des, flowers and fruits. The disease also occurs in
the United States of America (California), where
it appeared in the mid-1980s, and in South Africa
(Felker et al., 2010).
In Italy, symptoms of malformation, stunting and
lack of flowers, fruits and spine production on cactus
pear plants have been attributed to a phytoplasma
strain belonging to ribosomal subgroup 16Srll-C (Fig-
ure 12) (Granata et al., 2006; Tessitori et al., 2006).
Bertaccini et al. (2007) identified two different Felker et al. (2010) reported that in California
phytoplasmas (16SrI-B and 16SrV-A) associated cladode enlargement disease is caused by the to-
with cactus pear samples affected by cladode en- bacco bushy top virus, and it is possible that the
largement disease in California. symptoms observed in Mexico, South Africa and
Symptoms of cladode malformation, bud prolifer- Italy are induced by this same virus. Similar results
ation, thickening and heart-shaping of cladodes, were obtained by Suaste Dzul et al. (2012a), who
with arrested plant growth and yellowing of cla- demonstrated the presence of a virus in samples of
dodes were observed in the Saint Martin Pyramids cactus pear from Nopaltepec showing thickening
(northeastern State of Mexico) and associated and mosaic syndrome of the cladodes. Therefore,
with the 16SrII subgroup phytoplasma (Hernán- the co-infection between the phytoplasma and
dez Pérez et al., 2009a). Later, Fucikovsky Zak et the virus has been ascertained in this syndrome.
al. (2011) reported symptoms of phytoplasmosis The phytoplasma and viral diseases of this crop
on cactus pear plants in the region of Nopaltepec are difficult to control due to the lack of scientific
(State of Mexico) associated with the 16SrI phy- studies on cultivar susceptibility and insect vectors.
toplasma group (aster yellows strain, Candidatus
phytoplasma asteris). More recently, in the same
region, symptoms of distortion and thickening of ABIOTIC DISORDERS
the cladode, mosaic, yellowing, proliferation and
deformation of fruits were observed on cactus There is very limited scientific literature on non-in-
pear plants, but the causal agent was attributed to fectious diseases of cactus pear. The most common
the Mexican periwinkle virescence-16SrXIII phyto- abiotic disorders occasionally occurring in some
plasma (Suaste Dzul et al., 2012b). cactus-pear-growing regions are reported below.
Diseases of cactus pear 123

Frost injury ure 14). Since wounded tissues provide entry sites
for bacteria and fungi into the plant, protective
Frost damage due to temperatures < -5 °C may copper-based treatment soon after the event may
have a drastic effect upon the entire plant or affect prevent infection. Severely damaged cladodes and
only a small part of the plant tissue, reducing yield fruits should be pruned out as soon as possible.
or only product quality. Susceptibility to cold varies Herbicide injury
depending on the age and physiological state of
the plant and on the cultivar. Ripening fruits are Symptoms of herbicide toxicity vary according to
the most susceptible to frost injuries, which consist the kind of herbicide applied, the concentration of
of necrotic spots on the rind surface (Figure 13). the product and environmental factors (Figure 15).

Hail injury Fruit splitting

The degree of damage in cactus pear caused by Although the exact cause is unknown, fruit split-
hail depends on the duration and intensity of the ting occurs when plants take up water from rain
storm, as well as on the stage of development or irrigation after a long dry period and the fruit
of the plant. Hail injures result in wounds at the expands, bursting the peel in a crack across the
point of impact; even young cladodes and fruits bottom (Figure 16).
can be perforated by the impact of hailstones (Fig-

Figure 13
Frost injury (Italy)

Figure 14
13 14 Hail injury (Italy)

Figure 15
Glyphosate injury (Italy)

Figure 16
15 16 Fruit splitting (Italy)
 11

Insect pests
of cactus pear
Jaime Mena Covarrubias
National Institute of Forestry, Agriculture and Livestock Research, Mexico
 11 Insect pests of cactus pear

INTRODUCTION habits of the cactus plants, and plant bugs and

lepidopterans often lay their eggs on the spines
The Cactaceae family is a very distinct group of of the plant’s cladodes. No part of the cactus
plants with peculiar characters of growth and plant escapes from injury: insect species attack
habit and it has naturally evolved its own insect the roots, stems, cladodes and fruits (Figure 1).
complex. Indeed, more than 160 insect species Insect injury to cactus orchards affects the quan-
are harmful to cactus plants (Dodd, 1940; Mann, tity and the quality of the harvest, as well as the
1969; Zimmerman and Granata, 2002), and are duration of productive life. In general, insects
largely, but not solely cactus enemies. feeding on the fruits pose a higher risk of crop
loss than those feeding on the stems or cladodes.
Cactus insects have adjusted to the general con- Insect density also plays an important role in de-
ditions of the regions in which their host plants fining the status of an insect pest.
grow. For example, most of the adult stages of
beetle and lepidopteran insects are only active
at night, and their immature development takes INSECTS
place in the internal tissues of the cladodes,
branches and stems of the cactus plants; in fact, This chapter presents the main groups of insects
75% of all cactus−feeding insects are internal that feed on cactus plants worldwide, with a fo-
feeders (Zimmermann and Granata, 2002). These cus on identification, ecology and biology, as well
insects have also adapted to the structure and as on pest management tactics.

Figure 1
Insect damage to
cactus plants:
a) adult of Moneilema
on tender cladode,
b) shot hole symptoms,
c) wild cochineal
infestation on fruits,
d) hard gum exudates
on cactus main
stem from
Metamasius larvae,
e) grey bug damage,
and f) Metapleura
larvae inside a cactus
fruit.
Insect pests of cactus pear 127

Cottony cochineal insect (Dactylopius stage. On average, one female lays 131 eggs (the
opuntiae, Homoptera: Dactylopiidae) range is 62−617) (Flores Hernandez et al., 2006).
The adult male looks like a small fly with two
At present, the cottony cochineal (Dactylopius long tails. Males use flight to locate mates, and
opuntiae) is perhaps considered the most impor- are short−lived. The cottony cochineal insect is
tant insect pest in cactus orchards worldwide. In easily recognizable: large woolly masses of white
Brazil, it has infested 100 000 ha of cactus culti- wax cover its body and, when crushed, the bright
vated for forage production, lowering productivity crimson colour of the body fluid runs out and con-
by 80% (Lobos et al., 2013); in Mexico, it is the bi- trasts with the cottony colour (Figure 2).
otic factor that most limits cactus pear production Successful colonization of new cactus plants de-
(Mena Covarrubias, 2011); and in South Africa, it is pends on the female first−instar larvae, because
one of the two most important pests limiting for- they are the only ones with functional legs (Figure
age and fruit production (de Waal et al., 2013b). 2b). They also develop long waxy dorsal filaments,
D. opuntiae is characterized by sexual repro- which are well developed by the time the crawl-
duction and sexual dimorphism. The female has ers are 2−3 days old. In preparation for dispersal,
a four−stage life cycle (egg, first−instar larvae, female crawlers climb to the top of the host plant
second−instar larvae, adult), while the male has cladodes and “take off” with the help of the wind
a six−stage life cycle (egg, first−instar larvae, (Mow et al., 1982).
second−instar larvae, prepupa, pupa, adult). The Cochineal insects live on the surface of the cac-
life cycle duration depends on many factors, but tus plants, usually in aggregations or colonies
certain characteristics are common. The egg is composed of individuals in various stages of de-
oval and shiny red; hatching may occur inside the velopment, and settled at the base of the spines.
adult female; or the crawlers (first−instar larvae) All feeding damage is caused by the females and
may hatch within a few minutes of the eggs being nymphs as they suck sap from the cladodes and
laid (Flores Hernandez et al., 2006). For females, fruits. Within a few weeks of establishment on a
the larvae stage lasts 18.1 days; this is followed by plant, yellow areas appear on the cladode (Figure
a pre−oviposition phase lasting 18.8 days; and fi- 2e), which eventually drops off. Although new
nally the females spend 22 days in the oviposition cladodes may grow, these will also drop off if the

Figure 2
Wild cochineal,
Dactylopius opuntiae
life stages:
a) egg, b) crawler,
c) different stages of
immature development,
d) colony of several
adult females and some
crawlers on them, and
e) feeding damage on a
cladode.
 128 Crop ecology, cultivation and uses of cactus pear

cochineal remains on the plant. Depending on the The full−grown larvae of Cactoblastis, Olycella
severity of the infestation, the trunk may also die. and Metapleura all have bright body coloration.
Male cochineal insects never feed as adults. Cactoblastis larvae are uniform bright orange, or
For control, the first−instar larvae are the most orange−red with transversal rows of large black
susceptible and the easiest target. spots (Figure 3l); Olycella larvae are dark blue or
blue−black with white transversal bands (Figure
Lepidopteran insect pests 3i); the dull red−orange−yellow larvae of Meta-
pleura also have transversal rows of small black
The Pyralidae moth borers are the most important spots on a bright orange background (Figure 3f).
and abundant cactus insects; this family comprises Megastes larvae, on the other hand are creamy
about 50% more pest species than the cactus white in colour, with a light brown spot on each
longhorn beetles, and 66% more species than the side of the segments (Figure 3c). Cactoblastis
cactus weevils (Moran, 1980; Zimmermann and and Olycella females lay their eggs in chains called
Granata, 2002). The cactus moth (Cactoblastis “eggsticks” attached to the spines of the clado-
cactorum), the zebra cactus worm (Olycella nephe- des. There are 35−150 eggs on a single Cactoblas-
lepsa), the cactus white worm (Megastes cyclades) tis eggstick (Mann, 1969), and they are stacked
and the cactus pad joint borer (Metapleura potosi) one on another as if they were a part of the plant
are four of the most important lepidopteran in- spines (Figure 3k); the Olycella eggsticks have
sect pests feeding on cactus plants. Cactoblastis only 1−8 eggs and are attached at right angles to
poses the highest risk of damage since it is already the spine (Figure 3h), almost parallel to the cactus
present in South America and South Africa, and it pear cladode. The Megastes female lays masses of
threatens to invade commercial cactus pear pro- 40−90 eggs on the surface of the cladodes, with
duction areas in the United States of America and the edges of the eggs overlapping like roof tiles
Mexico (Zimmermann and Granata, 2002). These (Figure 3b). Metapleura eggs are laid singly on
four lepidopteran insects are all internal feeders the cracks, joints and surface of the cactus pads
of cladodes; Metapleura also invades the fruits, (Figure 3e). The adult stages of these four insects
boring from the cladode into the fruit, and leaving are moths with nocturnal activity (Figure 3a, 3d,
no trace of entry point (Mena Covarrubias, 2013). 3g, and 3j).

Figure 3
Lepidopteran insect
pests and their life
stages:
(a) Megastes cyclades
adult,
(b) egg mass and
(c) larvae
(d) Metapleura potosi
adult,
(e) eggs and
(f) larvae;
(g) Olycella nephelepsa
adult,
(h) egg mass and
(i) larvae;
(j) Cactoblastis
cactorum adult,
(k) egg mass and
(l) larvae.
Insect pests of cactus pear 129

Cactoblastis larvae can completely destroy a tunity varies from just one day (Cactoblastis) to
small−to−medium cactus plant in a single season: a few days (other species). The eggsticks of Cac-
they consume all the cladode internal tissues and toblastis are the easiest target: they are easy to
the larvae then leave the empty pad and move spot and they stay on the field for at least 3 weeks
in groups to the neighbouring untouched ones. before hatching (Mann, 1969).
Cactoblastis may have two or three generations
per year. Megastes larvae are not as mobile and Cactus weevils (Curculionidae)
destructive as Cactoblastis, but the galleries made
by their larvae every year cause the collapse of the The cactus weevil (Metamasius spinolae), the “shot
main branches and stems of the cactus plants. hole” weevil (Gerstaeckeria spp.) and the areole
Megastes has only one generation per year, but weevil (Cylindrocopturus biradiatus) are three of
it sometimes attacks the same plant for several the most important curculionid insect pests for
consecutive years. Metapleura poses a high risk cactus plants. Metamasius poses the highest risk,
for growers selling fresh cactus fruit, because it is since just a few larvae feeding on the base of the
difficult to detect an infested fruit; the consumer stem can knock out an entire plant; the other two
only realizes when it is too late, with negative con- genera feed mainly on the cladodes.
sequences for the consumer−farmer relationship.
Weevils are easily identified in the adult stage by
There can sometimes be as many as 15−20 larvae
their long snout; in Metamasius it is 22−25 mm
of Metapleura on the same cladode, inducing
long, in Gerstaeckeria 0.5−0.6 cm and in Cylindro-
partial rotting or, if the damage is in the joints,
copturus 3−3.5 mm. Metamasius is characterized
the cladodes fall to the ground; however, in most
by black elytra with two orange markings (Figure
cases, the damage is not significant. Finally, Olyce-
4a); Gerstaeckeria commonly has a black−brown
lla attacks young cactus orchards only in the first
body with white scales (Figure 4i); Cylindrocop-
3 years; damage is localized, and the single pests
turus is a silvery colour with a yellow double cross
are found in excavations beneath tumour−like
marking in the middle of its body (Figure 4e).
swellings in the cladodes.
The larvae of these curculionid beetles are legless
The target of control for all these pests are the white worms with brown heads (Figures 4c, 4g
newly hatched larvae, but the window of oppor- and 4j).

Figure 4
Cactus weevil insect
pests and their life
stages:
(a) Metamasius spinolae
adult,
(b) egg
(c) larvae and
(d) pupae;
(e) Cylindrocopturus
biradiatus adult,
(f) egg
(g) larvae and
(h) pupae;
(i) Gerstaeckeria spp
adult,
(j) larvae,
(k) cladode internal
damage by the larvae.
 130 Crop ecology, cultivation and uses of cactus pear

Metamasius and Gerstaeckeria are flightless The adult is the only stage that lives externally
weevils; the Gerstaeckeria adults are active only and for this reason is the main target of control.
at night. The cactus weevil is potentially present However, pruning could be an excellent option for
year−round, but emergence peaks at the begin- controlling Gerstaeckeria and Cylindrocopturus
ning of the rainy season; the shot hole and areole larvae and pupae during the winter months.
weevils are common from March to September.
The female of the cactus weevils burrows into Longhorn beetles (Cerambycidae)
the cladode tissue, lays an egg and seals it with a
mixture of food and regurgitated saliva. They have This group is represented by the genus Moneile-
only one generation per year, but the adults have ma (M. variolaris is one of the most common
a long life span: Metamasius ≥ 1 year, Gerstae- species in cactus orchards). They can inflict major
ckeria and Cylindrocopturus 4−5 months (Dodd, harm and just two or three larvae in the base of
1940; Mann, 1969; Mena Covarrubias, 2013). a full size cactus pear may cause the whole plant
to collapse. They also feed on the base of young
Most of the damage to cactus plants is caused
plants and pose the same risk of damage as the
by the larvae as they feed on the internal tissues
Metamasius weevil. These cerambycids are robust,
and set up conditions for rot, especially in the
shining black beetles; the elytra and prothorax
case of Metamasius and, to a lesser degree, Gers-
sometimes present irregular white marks. Their
taeckeria. Metamasius larvae are highly mobile,
length is 15−25 mm (males are smaller than fe-
travelling from the stem to the lower part of the
males), and the antennae are longer than half
main branches, in order to obtain a better supply
the insect body (Figures 1a and 5a). The larva is
of food. It is common to find single cladodes,
white and each segment is deeply creased along
branches and whole plants lying fallen on the
the body; it ranges in length from 15 to > 40 mm;
ground; scavenger insects greatly increase the rot-
the head is black (Figures 5a and 5c).
ting in affected areas (Dodd, 1940; Mann, 1969;
Mena Covarrubias, 2012). Adults prefer to feed Moneilema are nocturnal, flightless insects; how-
on young cladodes; Gerstaeckeria and Cylindro- ever, on cloudy days, these beetles are also active
copturus also use them for oviposition and larvae in the morning. The adults live for several months
development, while Metamasius prefers stems and are present from March to November. The
and older cladodes for those activities. eggs are laid singly and glued to the surface of ma-

Figure 5
Moneilema life stages:
(a) adult,
(b) egg,
(c) larvae and
(d) pupae.
Insect pests of cactus pear 131

ture segments, in cracks, against the stem or just as well as the mirid red bug (Hesperolabops) are
below ground level (Mann, 1969). There are one good examples of sucking bugs. These insects
or two generations each year; in the case of two pose less risk of damage to the cactus plants
generations, spring emergence is much bigger than compared with the lepidopterans, weevils, long-
autumn emergence (Dodd, 1940; Mann, 1969). horn beetles or cochineal insect pests described
above; very rarely will their feeding activities risk
The larvae infest the main stem and older joints of
destruction of a cactus plant. The most common
the cactus pear, constructing galleries filled with
damage from the grower’s point of view is the
copious exudations that soon become blackened.
reduced quality of the cactus fruit. Hesperolabops
Saprophytic insects are attracted to the feeding
has the greatest potential for causing this dam-
areas and help to make them spread. The larvae
age, because it reaches a wider population, feeds
move considerably inside the cactus plants, usu-
on both cladodes and fruits (Narnia also is a fruit
ally in a downwards direction, ending in the basal
feeder insect) and is harder to control.
stem of newly planted orchards. The adults, on
the other hand, feed superficially by gnawing at The adult stage of the cactus grey bug, Chelinidea
the edges of the newly formed pads and occa- tabulata (Burmeister), has a brown−green body;
sionally the fruits (Figures 1a and 5a). the newly hatched first instars are black, becom-
The adult stage is the target of control, because ing light green as they grow older; adults are
the larvae are difficult to locate and control inside 13−15 mm (Figures 6a and 6c). The cactus fruit
the plants. bug, Narnia femorata Stål is coloured deep purple
to black as an adult; it is slender and smaller than
Plant−sucking bugs the cactus grey bug; the immature stages are dark
grey with pale yellowing areas on the antenna,
There are very few cactus surface feeders with thorax and legs (Figures 6d and 6f). The adult
biting or chewing mouthparts − probably because stage of the cactus red bug (Hesperolabops gelas-
of the absence of true leaves on the cactus plant tops Kirkaldi) has a red head and prothorax, while
(Dodd, 1940). The most common are stinky bugs, the rest of the body is smoky with a pale band
cochineal insects, thrips and scale insects. The on the edge of the thorax and abdomen (Figure
coreid bugs of the genera Chelinidea and Narnia 6h); it is a small bug, length 6−8 mm. The newly

Figure 6
Cactus plant sucking
bugs life stages:
(a) Chelinidea tabulata
adult,
(b) eggs,
(c) and immature;
(d) Narnia femorata
adult,
(e) eggs,
(f) immature,
(g) and damage to
cactus fruits;
(h) Hesperolabops
gelastops adult,
(i) first instar,
(j) and older instar.
132 Crop ecology, cultivation and uses of cactus pear

hatched insects are completely red (Figure 6i) and they aspis echinocacti) and oyster scale (Lepidosaphes spp.)
maintain the colour as they grow, as well as the pale − feed externally on the cactus pads; their presence is
band around their bodies; only the wing pads are grey limited to a small number of plants per orchard and
(Figure 6j). Both Chelinidea and Narnia lay several eggs they only become important if their natural enemies are
end to end along a cladode spine (Figure 6) (Mena destroyed with the use of broad spectrum insecticides.
Covarrubias, 2013). Hesperolabops, on the other hand,
inserts its eggs under the epidermis layer of the cactus
cladode, with the operculum close to the epidermis INTEGRATED PEST MANAGEMENT
(Palomares Perez, 2011). Development periods for the FOR CACTUS INSECT PESTS
egg and immature stages are, respectively: 21 and 56
days for C. tabulata (Brailovsky et al., 1994); 13 and 67 Integrated pest management (IPM) is a strategy for
days for N. femorata (Vessels et al., 2013); and 269 and controlling insect pests using a full range of control
36 days for H. nigriceps Reuter (Palomares Perez, 2011). tactics. In order to maximize the reduction of the pest
Both the adults and the immature stages are gregarious population, a thorough knowledge is required of the
to different degrees and they are present in the cactus ecology and biology of both the insect pest and its
orchard year−round; they move at the same time on natural enemies. The key IPM management practices in
the cactus plant, searching for the opposite part, as if cactus orchards are described below:
they were trying to hide from the observer.
Adult and immature stages of the cactus plant−suck- Pruning
ing bugs feed by puncturing the cladodes, joints and
fruits; they leave a characteristic pale yellow circular Orchard maintenance includes yearly pruning, which
area around each feeding puncture, so the affected can affect an insect pest population in three significant
areas have a spotted appearance (Figure 1e). Dense ways (Mena Covarrubias, 2011, 2012):
populations can cause cladodes to look yellow and • A dense canopy offers abundant opportunities for
sickly, especially in newly planted orchards. Narnia fruit insect survival during winter and also reduces the ef-
damage produces a pulpy spongy area on each feeding ficiency of spraying. To optimize the benefits of prun-
site (Figure 6g); while the spot is less juicy and sweet, ing, cladode overlapping must be kept to a minimum.
it is small and superficial, and the average person does • Pruning provides an opportunity in winter to com-
not detect it. On the other hand, large populations of pletely eliminate from an orchard and destroy the
cactus red bug prevent the external tissue of the fruit areole weevil, the shot hole weevil and the cactus
from reaching its normal colour, and reduce the sugar cladode miner (Marmara opuntiella Busck), along
content, which affects the price of the product. In ad- with the infested first year cladodes. The efficiency of
dition, the cactus red bug is associated with “cacarizo”, this practice is improved if several farmers carry it out
a disease that affects the ‘Reyna’ cultivar in the Mexi- at the same time. Neighbouring wild Opuntia should
can Highlands and covers the epidermal tissues of the also be included in the campaign.
cladodes with blisters (Palomares Perez, 2011), reducing
• To optimize the positive effects of pruning, never
drastically the productivity of the orchard.
leave pruning waste on the orchard floor. The prun-
The targets of control for the cactus−sucking bugs are ings are a potential refuge for the development and
both the adults and immature individuals, because they reproduction of cochineal insects, cactus weevils,
are all external feeders; newly hatched insects are more longhorn beetles and cactus pad joint borers.
susceptible to control agents.
Targeting emergence
Other cactus insect pests
In order to control the newly hatched larvae of lepidopter-
There are a small number of species of fly that feed on an insect pests, it is essential to target them at emergence.
cactus plants. Asphondylia is the most important as it Once the larvae are inside the cactus tissue, external con-
feeds on the cactus seeds, while Mayetiola, Lonchaea trol is no longer possible. It is, therefore, critical to monitor
and Dasiops live in the cladodes and cause little dam- the adult stage (moths) and pinpoint population peaks. A
age (Mann, 1969). Fruit flies are occasionally reported pheromone has been developed to trap the cactus moth
as insect pests around the globe, particularly in the males (Heath et al., 2006). The other moth species can
Mediterranean area (Zimmermann and Granata, 2002). be sampled using light traps in September−October (cac-
Thrips are commonly found on cactus pear orchards, tus white worm), April−June and September−November
especially Neohydatothrips opuntiae; they damage the (cactus zebra worm) and year−round (cactus pad joint
skin of tender cladodes and young fruits early in their borer) (Mena Covarrubias, 2011, 2012). The adult stage of
development. Finally, scale insects − cactus scale (Di- Cactoblastis, Olycella and Melitara, on the other hand, do
Insect pests of cactus pear 133

not feed (Mann, 1969), and food traps cannot be adopted • provision of alternative food sources for the adult
for trapping or monitoring. stage, specifically pollen and nectar plants around
the orchard; and
Mass trapping • no spraying of broad spectrum insecticides (Mena
Covarrubias, 2014).
It is possible to carry out mass trappings of cactus weevils
because the males release an aggregation pheromone Biological control by augmentation can also be imple-
for both females and males, especially if the traps are mented using entomopathogenic fungi (e.g. Beauveria
set on cactus cladodes (Tafoya, 2006). Cactus weevils, bassiana) to control insects such as the cactus white
longhorn beetles and shot hole weevils are flightless worm (Lozano Gutierrez and Espania Luna, 2008). The
beetles and can be controlled by hand picking, especially challenge is how to place the agent of control in con-
if the infested area is flagged and the activity is concen- tact with the target insect, because most cactus insect
trated there. For longhorn beetles and shot hole weevils, pests are internal feeders and spend most of their life
control should be carried out at dawn and dusk because cycle inside the cactus tissues.
they are nocturnal insects; on the other hand, the adults
leave a typical feeding footprint (Figure 1a, 4m, 5a), Chemical control
and by day it is possible to pinpoint which cactus plants
were affected the previous night. Only a limited number of conventional insecticides are
permitted for the chemical control of cactus plants
Biological control However, organic insecticides − such as pyrethrum,
diatomaceous earth, Bacillus thuringiensis, neem and
Biological control is a fundamental element of insect insecticidal soaps − are available and can potentially
pest management. In the case of cactus insect pests, increase the spectrum of chemical alternatives for in-
biological control entails the conservation of the natural sect control. Further research is required on spraying
enemies present in the orchards, which in turn requires: coverage and the impact speed of the droplet in order
• identification of the beneficial insects; to optimize pest control on cactus plants.
 12
Processing and utilization
of fruit, cladodes
and seeds
Carmen Sáenz
Department of Agro−industry and Oenology,
Faculty of Agronomic Sciences, University of Chile, Santiago, Chile
 12 Processing and utilization of fruit,
cladodes and seeds

INTRODUCTION of the bioactive compounds of these cactus parts,

their activities and the relative health benefits.
A Sicilian writer once called cactus pear "a treas- There has been much research on the chemical
ure under the thorns" because of its immense composition of the edible parts of the fruit from
benefits, some of which were little known before plants grown in different parts of the world, includ-
today. ing Egypt, Saudi Arabia, Mexico, Chile and Argenti-
In many countries − Argentina, Chile, Peru, Bo- na. There is extensive information available regard-
livia, South Africa, Egypt, Turkey, Ethiopia, Eritrea ing the composition and uses of seeds as a source
and other countries in South America and the of oil, fibre and protein, in particular the detailed
Mediterranean Basin − it is traditionally the cactus analysis of Sáenz et al., eds (2006). This chapter
pear fruit that is eaten. In Mexico, on the other presents the latest information on the subject.
hand, in addition to the fruit, the tender cladodes
(nopalitos) are consumed. However, both fruit and Within the genus Opuntia, the most cultivated
species is Opuntia ficus−indica, characterized by
nopalitos are perishable and processing technolo-
sweet and juicy fruit and pulp of different colours:
gies are needed to increase their shelf−life. More-
white–green, yellow, orange, red or purple. The
over, both fruits and cladodes contain numerous
proportion of pulp in the fruit varies and the peel
bioactive compounds which must be preserved
is generally thin. Coloured ecotypes (Figure 1)
during processing if consumers are to reap the
have a dual application: production of natural
full benefits. Cactus pear is a multipurpose fruit
colourants (betalains); and provision of health
and a wide range of products and by−products
benefits thanks to their antioxidant properties
can be derived from it. The same applies for the
(Butera et al., 2002; Galati et al., 2003a; Kuti,
cladodes. Sáenz et al. (eds, 2006, 2013) present
2004; Tesoriere et al., 2005a; Stinzing et al., 2005;
numerous alternatives for processing fruits and
Azeredo, 2009; Fernández López et al., 2010).
cladodes. The recent advances in this field are
presented in this chapter. Other bioactive compounds present in the fruits
are vitamin C, carotenoids and dietary fibre (Mo-
rales et al., 2009; Sáenz et al., 2009, 2012a). Cac-
CHEMICAL COMPOSITION AND tus cladodes have a high content of water, dietary
fibre and minerals (Pimienta Barrios, 1990; Sáenz
BIOACTIVE COMPOUNDS
et al., eds, 2006). The seeds are rich in polyun-
Figure 1
saturated essential fatty acids such as linoleic acid
Cactus pear fruits Before any raw material is processed, it is essential
coloured ecotypes (Ennouri et al., 2005; Özcan and Al Juhaimi, 2011).
to understand its chemical and phytochemical
(Opuntia ficus−indica)
composition, as well as any technological charac- There are minor variations in the chemical com-
(garden varieties, An-
tumapu Experimental teristics regarding the industrial process. For cactus position of coloured cactus pear fruits; major
Station, University of pear, information is required about the chemical differences are related to the pigment content.
Chile, Santiago) composition of the fruits, seeds and cladodes. Table 1 summarizes the ranges of certain chemi-
(Photos: C. Sáenz and Furthermore, a thorough understanding is required cal compounds and technological characteristics,
A.M. Fabry).
Processing and utilization of fruit, cladodes and seeds 137

TABLE 1 Chemical and technological characteristics of cactus pear pulp from coloured fruits

Parameters Green cactus pear Purple cactus pear Yellow−orange cactus pear
pH 5.3–7.1 5.9–6.2 6.2–6.3
Acidity (% citric acid) 0.01–0.18 0.03–0.04 0.55–0.57
Soluble solids (°Brix) 12–17 12.8–13.2 13.5–14.5
Vitamin C (mg 100 g−1) 4.6–41.0 20.0–31.5 24.1–28.0
β−carotene (mg 100 g ) −1
0.53 − 0.85–2.28
Lutein (µg g−1) 26.0 0.15 0.04
Betacyanins (mg kg ) −1
0.1–0.8 111.0–431.0 2.4–11.0
Betaxanthins (mg kg−1) 0.4–3.1 89.4–195.8 16.0–76.3

Source: Askar and El Samahy, 1981; Pimienta Barrios, 1990; Sawaya et al., 1983; Sepúlveda and Sáenz, 1990; Sáenz and Sepúlveda,
2001a; Sáenz et al., eds, 2006; Stintzing et al., 2005; Hernández Perez et al., 2009b; Morales et al., 2009; El Gharras et al., 2006; Coria
Cayupan et al., 2011; Sáenz and Fabry (unpublished data).

based on values reported by several authors. Red, pur- counts to a safe level compared with the treatment of
ple and yellow−orange cactus pears contain betalains, acidic foods (pH ≤ 4.5). Such high temperatures (gen-
while red and purple contain betacyanins and yellow− erally > 121° C) can negatively influence traits such
orange betaxanthins (Stintzing et al., 2005; Sáenz et al., as taste, colour and aroma. The pH and high soluble
2012b). solids content of the pulp favour the growth of micro-
organisms (Sáenz and Sepúlveda, 1999; Sáenz, 2000);
The chemical composition may vary depending on dif-
it is, therefore, important to control heat treatments in
ferent factors:
preservation processes.
• origin of the plants (i.e. the climate where it is culti-
vated); From a sensory point of view, the green fruit in some
countries (e.g. Chile) has a better texture, taste (sweet-
• agronomic factors, such as cultivation, fertilization
er) and flavour than purple and orange ecotypes, which
and irrigation; or
tend to be floury. Nevertheless, purple, red and orange
• genetic differences (Muñoz de Chavez et al., 1995; fruits have great potential for processing, because the
Ochoa, 2008). betalains contained in the colour ecotypes are more
stable than the chlorophylls, with regard to both pH
Cactus pear is rich in calcium, although McConn and
and heat (Merin et al., 1987; Montefiori, 1990; Castel-
Nakata (2004) report that calcium bioavailability might
lar et al., 2003; Sáenz and Sepúlveda, 2001a; Sáenz et
nevertheless be low, because it is present as calcium oxa-
al., 2012b).
late, which is not absorbed. The high potassium content
and low sodium content offer clear nutritional benefits
for people with kidney problems and hypertension.
Cladodes

The presence of mucilage and pectin in the cladodes

influences the viscosity of some products, such as
TECHNOLOGICAL CHARACTERISTICS powder preparations mixed with water or juice before
consumption. Both compounds are part of dietary fibre
Fruit and are hydrocolloids, known for their ability to absorb
and retain water. They can also be extracted and used
Besides the chemical composition and bioactive com-
as thickeners in foodstuffs (Sáenz et al., 2003, 2004;
pounds, there are other characteristics to consider dur-
Sepúlveda et al., 2003a, 2007).
ing processing. In general, cactus pear fruits have a high
pH (5.3−7.1) and are therefore classified as low acid The chemical composition of nopalitos has been re-
foods (pH ≥ 4.5); an exception is O. xoconostle, which ported by Pimienta Barrios (1990) and Maki Díaz et al.
has higher acidity (pH > 3.5) (Mayorga et al., 1990). It is (2015). As with other vegetables, there is a high con-
well known that heat treatment temperatures depend tent of water and fibre. Polyphenols are present and
on the pH (Casp and Abril, 1999). For this reason, when have antioxidant activity important in the diet; during
non−acidic foods are pasteurized or canned, unless the preservation, however, they can cause browning due
pH is reduced (by citric acid addition, for example), a to oxidation (Rodríguez Felix, 2002). Furthermore, the
higher temperature is needed to reduce the microbial acidity of nopalitos varies during the day − as a result
138 Crop ecology, cultivation and uses of cactus pear

of crassulacean acid metabolism (CAM) (Cantwell et Dehydrated cactus pear products

al., 1992) − and the optimal harvesting time must be
chosen depending on the process to be applied. Dehydration is one of the oldest food preservation pro-
cesses. It can be done naturally − solar drying − or with
Seeds equipment, such as dehydration tunnels, rollers, dryers
and spray dryers. Highly controlled processes have re-
The seeds represent about 15% of the edible part of the cently been developed to produce more homogeneous
fruit and they have a variable oil content (on average, 9.8 and better quality dried products.
g 100 g−1 of seed) (Ramadan and Mörsel, 2003a).
With regard to cactus pear, there have been various
Seed oil is rich in unsaturated fatty acids (Sepúlveda and studies about dehydrating thin layers of pulp to prepare
Sáenz, 1988; Ennouri et al., 2005; Ghazi et al., 2013); it chewable, natural products. They are known as “fruit
is, therefore, of interest to the pharmaceutical and cos- sheets”, “fruit leather”, "fruit bars" or “fruit rolls”
metic industries, for example, in Morocco and Tunisia. and differ in thickness and moisture content: sheets
Given the low yield of oil from the seed, it is neither are thinner and low in moisture, while bars have a
economical nor attractive as an edible oil. The presence high moisture content (≤ 20%). The University of Chile
of tocopherols, recognized as natural antioxidants, developed a process whereby a blend of cactus pear
ranges from 3.9 to 50%. Matthäus and Özcan (2011) pulp and quince or apple pulp is used to prepare fruit
and Özcan and Al Juhaimi (2011) report that fibre and rolls. Coloured ecotypes are also used to make prod-
minerals are also important components in the seeds, ucts with a pleasant flavour and texture and attractive
with 12.5% crude fibre and high amounts of calcium, appearance (Sepúlveda et al., 2000, 2003b) (Figure 3).
potassium and phosphorus, as well as other minerals. Table 2 shows the characteristics of certain products.
The relatively high protein content (approx. 6%) means All treatments contain:
that cactus pear seed is a source of protein for human • 75% cactus pear pulp;
consumption (Tlili et al., 2011).
• 25% apple pulp;
• varying amounts of sucrose (T1 and T2 = 6% and T3
= 0% sucrose and 0.01% sucralose); and
PROCESSING TECHNOLOGIES
• varying amounts of flaxseed (T1 = 0% and T2 and
A wide range of traditional preservation technologies T3 = 1%).
can be applied to cactus pear fruit, cactus cladodes The total dietary fibre content of the treatments is
and seeds. Some technologies are described in Sáenz 14.1−43.9%; the purple ecotype has a high fibre
et al., eds (2006) and some of the most innovative are content due to its high pulp content, and sucralose
described here. (instead of sugar) is added.
The flow chart in Figure 2 shows the various steps
Dehydrated products
involved in the production of cactus pear sheets and
Figure 3 shows the cactus pear and apple pulps, as
Water activity (aw) is a measure of the "available wa-
well as an oven used for drying to make the sheets.
ter" in a food. Availability of water in plant tissues is
variable, and a distinction is made between "free wa- El Sahamy et al. (2007a) prepared orange–yellow cac-
ter" and "bound water". The proportions of free and tus pear sheets; they tested different drying tempera-
bound water depend mainly on the food composition, tures (60 and 70 °C) and a range of ratios of sucrose
as compounds such as hydrocolloids have higher water (0, 1, 2, 3, 4, 5 and 10%). The prepared pulps were
retention capacity. The mucilage present in Opuntia spread to a thickness of 10 mm and dehydrated in an
pads and fruits is an example of a hydrocolloid. air convection oven for 44 hours. The preferred sheets
Microbial growth can be controlled by lowering aw. The were those prepared with 2 and 3% sucrose.
minimum aw for microbial growth is variable. Accord- Dehydrated products do not usually contain additives;
ing to Roos (2007), it is > 0.90 for bacteria, 0.87−0.90 they are, therefore, “natural products”, accepted by
for yeast, 0.80−0.87 for moulds and 0.60−0.65 for consumers because they are considered safe.
osmophilic yeasts.
This simple technology, to the best of the author’s
The technologies used to decrease aw to preserve food knowledge, has not been used at commercial level.
include dehydration, concentration and freeze−drying While there are companies offering apple, strawberry,
(drying under frozen conditions); the latter combines cherry and apricot rolls, these products are not usually
cold with decreasing aw to control the growth of mi- 100% natural and are typically made from pear puree
croorganisms. concentrate and artificial flavourings and colourants.
Processing and utilization of fruit, cladodes and seeds 139

TABLE 2 Dietary fibre and total phenolics compounds in coloured cactus pear rolls mixed with apple
pulp and flaxseed

Parameter/Treatment Rolls from yellow−orange cactus pear pulp

T1 T2 T3
Total dietary fibre (g 100 g−1) 14.1 a 24.3 b 38.8 c
Total phenolics (mg GAE kg−1) 1 445.3 a 1 365.0 a 1 640.1 b
Rolls from purple cactus pear pulp
T1 T2 T3
Total dietary fibre (g 100 g−1) 20.2 a 28.9 b 43.9 c
Total phenolics (mg GAE kg−1) 1 404.7 a 1 438.0 b 1 846.0 b
a
GAE = Gallic acid equivalent.
Means following different letters differ at P < 0.05.
Source: Sáenz et al., unpublished data

Figure 2
Wash Preparation of cactus
Manual peeling Selection Harvest and
(chlorinated water) pear fruits rolls
spine removal
(adapted from Sáenz
et al., eds)

Mix ingredients
Pulp/sieve (cactus pear Mix Place on trays
(1 mm) and other fruit (stir) (2.5 mm)
pulps, sugar etc.)

Packaging Cooling Dehydration

Cutting/rolling
(film HDPE) (20–25°C) (58–60°C, 78 h)

Figure 3
Cactus pear and apple
pulps; electric tray drier;
rolls from coloured
ecotype
(Photos: C. Sáenz and
A.M. Fabry)
140 Crop ecology, cultivation and uses of cactus pear

The quality of such products could be easily result in a herbaceous aroma and flavour, and the
improved by using coloured cactus pear pulp to mucilage present in the cladodes produces tex-
prepare a more natural and healthier product. ture defects (Sáenz et al., 2012a). The research of
Sáenz et al. (2012a) resulted in a purified natural
Dehydrated cladode products dietary fibre with > 80 g 100 g−1 of total dietary
fibre and 20−22 g 100 g−1 of soluble dietary fibre,
The dehydration options for cladodes are differ- one of the scarcest types of fibre present in veg-
ent from those for cactus pear fruits. The clado- etables. Purification results in an increase in total
des are not dehydrated for direct consumption, dietary fibre, a decrease in the green colour of the
but are transformed into powders (Sáenz et al., powder and a decrease in the total phenolic com-
2010) with a high content of dietary fibre. This pounds, in particular when cladodes are washed
powder can be used to make cookies (blended at high temperatures. This purification process is
with wheat flour), puddings and − in some coun- promising and could result in more widespread
tries, in particular Mexico − breakfast cereals or use of cactus cladode powder as a food additive,
tortillas. It can also be used in the production of especially in markets where consumers are not
food supplements (capsules, tablets etc.). familiar with the herbaceous flavour of cactus
cladodes and therefore are less accepting. Further
Rodríguez García et al. (2007) observed that research is required to obtain a powder rich in
during the development of young cladodes, in- dietary fibre, low in flavour and colour and with
soluble fibre increases (from 29.87% in a 60−g high antioxidant capacity for use as an ingredient
cladode to 41.65% in a 200−g cladode), while in new food formulas.
soluble fibre decreases (from 25.22 to 14.91%,
respectively). Ayadi et al. (2009) prepared powder Cactus pear concentrates
using 2−3−year cactus pads; the total dietary fi-
bre was 51.24%, of which 34.58% insoluble fibre The range of concentrated products derived from
and 12.98% soluble fibre. cactus pear includes syrups, jams and concen-
This powder can make a nutritional contribution trated juices (Sáenz, 2000). Morales et al. (2009)
developed dessert sauces (toppings) from colour-
to several food products by increasing the daily
ed ecotypes with excellent results, preserving
dietary fibre intake. Inclusion of the powder in
both their attractive colour and their functional
Figure 4
foods, however, implies certain technological
compounds. Vacuum concentration was used
Coloured cactus challenges, as some aspects of taste and texture
on mixtures of cactus pear pulp with sugar
pear toppings on a require improvement (Sáenz et al., 2002b, c;
milk dessert (22.0−30.25%), fructose syrup (13.75−22.0%),
Ayadi et al., 2009). For example, heat treatments
glucose (11.0−19.25%), citric acid (0.14%) and
modified starch (1.5%). These attractive products
may be used in a range of different dishes (Fig-
ure 4). Table 3 lists bioactive compounds present
in two coloured toppings.
A wide variety of cactus−pear−based foods are
manufactured (Sáenz et al., eds, 2006). Compa-
nies use the Internet to advertise different ways
of eating and enjoying cactus pear fruit products,
offering concentrated products, such as jams,
syrups and candies. The information available on-
TABLE 3 Bioactive compounds in coloured cactus pear toppings

Yellow−orange cactus pear

Bioactive compound Purple cactus pear topping
topping
Carotenoids (μg g−1) 0.186 ± 0.001 0.021 ± 0.001
Total phenolics (mg GAE litre )−1 a
350.50 ± 15.25 131.48 ± 5.72
Betalains 81.06 ± 1.83 63.80 ± 1.86
Betacyanins (BE mg kg ) −1 b
66.09 ± 1.03 0.92 ± 0.00
Betaxanthins (IE mg kg−1)c 14.97 ± 1.53 62.88 ± 1.86
a
GAE: gallic acid equivalent; b BE: betanin equivalent; c IE: indicaxanthin equivalent (Morales et al., 2009).
Processing and utilization of fruit, cladodes and seeds 141

line suggests that these are mainly artisanal and from cactus pear.
small−scale companies. Cactus pear sweet gels
These cactus pear sweet gels are made with:
Research by the author’s group at the University • seedless cactus pear pulp (1 000 g);
of Chile has recently led to the development of
• sugar (760 g)
cactus pear sweet gels using the pulp of differ-
ent coloured ecotypes to exploit the attractive • pectin (52 g) − another gelling agent (e.g. car-
pigments (Sáenz and Fabry, unpublished data). boxymethyl cellulose − CMC) could be tried,
These products – obtained evaporating the pulp or a mixture of fruit pulps rich in pectin (e.g.
and adding gelling agents such as pectin – have quince); and
proved very popular with small farmers in arid • citric acid (16 g) − or substituted with lemon
zones of Chile, where there are shortages not only juice.
of water, but of electric power. These sweet gels
are made with available technologies, enabling Figure 6 shows cactus pear sweet gels made by
small farmers to add value to their cactus pear small farmers in the village of Codpa (Arica and
production. The flow chart in Figure 5 shows the Parinacota, Chile) in a workshop led by our group.
different stages in the production of sweet gels The village is located in Camarones, in the middle

Figure 5
Pulp/sieve Preparation of cactus
Fruit harvest Wash Manual (1 mm)
and spine pear sweet gels
(chlorinated water) peeling/selection Washing
removal (chlorinated water)

Boll/stir (until 70° Brix) Mix Blend powder ingredients

Mix ingredients (cactus pear (cactus pear pulp Pulp/sieve (1 mm)
and other fruits pulps) and powder ingredients) (sugar/pectin/citric acid)
sugar or sucralose etc.

Fill hot (glass) Remaval from trays Storage

until gelling dehydration Packaging (20–22° C)
(58–60° C, 7– 8 h) in HDPE film

Figure 6
Cactus pear coloured
sweets made in a
workshop in the north
of Chile (Photos: C.
Sáenz and A.M. Fabry)
 142 Crop ecology, cultivation and uses of cactus pear

of the Tarapacá Desert in the north of Chile − an 31.3 and hab = 16.6, corresponding to a dark
area with water and electric energy restrictions and purple colour. Despite the soluble solids content
where cactus pear grows as an alternative crop. of 56.5° Brix, the product shows a high aw (0.92).
For this reason, to ensure a good shelf−life, the
Recently, Sáenz and Fabry (unpublished data)
use of preservatives (e.g. sodium benzoate and
developed cactus pear pulp gummy confections,
potassium sorbate) is recommended.
mixing a fine screened pulp (2.5 litres) with unfla-
voured gelatin (0.45 kg), water (2 litres) and sugar Nopal jams
(1.5 kg) (Figure 7).
The preparation of jams combines heat treat-
The final product has a brilliant and attractive ments with a decrease of aw (and sometimes also
appearance (Figure 8) and a high sensory accept- of pH to enable less severe thermal treatment).
ance. The colour parameters of purple gummy There are different jams and syrups on the market
confections are L = 5.6, a* 30.0, b* = 9.0, C* = in several countries (Sáenz et al., eds, 2006). One

Figure 7
Wash
Preparation of Pulp/sieve Fine screen
(chlorinated water)
cactus pear gummy (1 mm) (thin cloth)
and peel
confections

Blend water Weight ingredients

Add cactus
and unflavoured gelatin (gelatin, sugar,
pear pulp
and stir preservatives, etc.)

Mix with sugar Boil Measure

and lemon juice (100° C, 10–22 min) in a jar
(optional)

Cooling
Storage/packaging/ (4–5° C or <20–25° C) Fill silicone
distribution until solid moulds

Figure 8 innovative product, for which technology was

Gummy confections recently transferred to small farmers in the north
made from purple of Chile, is cladode and lemon jam: nopal jam. To
cactus pear, remove the excess mucilage affecting the texture,
yellow–orange cactus
the cladodes are pre−treated with Ca(OH)2. This
pear and a mixture of
both (Photo: C. Sáenz pre−treatment may be avoided if the cladodes
and A.M.Fabry) have a low mucilage content. The flow chart in
Figure 9 describes the steps involved in the
preparation of nopal jam.

Nopal jam represents a new alternative for pro-

cessing cladodes. The jam can be eaten with
crackers and baked goods, or served with meat
and other dishes (Figure 10).
Processing and utilization of fruit, cladodes and seeds 143

Chop Wash Nopal harvest

(cubes 0.5 x 0.5 cm) brush and peel cladodes (1–3 year)

Pre-treatment with Wash Blanch

Ca (OH)2 at 2% (lh) in profuse water (100° C water, 10 min)

Figure 9
Preparation of nopal
Mix ingredients
Concentration Drain jam (cladode and
(Nopal, lemon juice
(65–67° Brix) lemon) (Modified from
and peel, sugar, gelling agent)
Sáenz et al., eds, 2006)

Filling/sealing Labelling/packaging Storage/distribution

Figure 10
Cladode and lemon jam
(Photos: C. Sáenz and
A.M.Fabry)

Cactus pear juices was assessed after 40 days at 5° C. The authors

observed an increase in sedimentation and turbid-
The pasteurization of cactus pear juice made with ity (Nephelometric Turbidity Unit − NTU) during
green fruit has produced unsatisfactory results, storage, and a decrease in colour, probably due to
due to the unpleasant colour and aroma changes the Maillard reaction.
that occur during the process. On the other hand,
De Wit et al. (2014) made juices from eight cactus
purple cactus pear juice blended with other juices
pear cultivars, with different fruit colours: one O.
(e.g. pineapple) might represent an interesting al-
robusta cultivar (‘Robusta’, purple) and seven O.
ternative (Sáenz and Sepúlveda, 2001a). However,
ficus−indica cultivars (‘Berg’ × ‘Mexican’, pink;
acidity is a potential issue: the addition of organic
‘Fusicaulis’, green−white; ‘Meyers’, red; ‘Algerian’,
acid to increase acidity facilitates the heating pro-
red; ‘Santa Rosa’, orange; ‘Skinners Court’, green–
cess but can modify the taste, while consumers
white; ‘Morado’, white). The juices were pasteur-
generally prefer cactus pear juices with the origi-
ized (60 °C for 10 minutes). The authors reported
nal acidity of the fruit, similar to that of the fresh
that the heat treatment had an unfavourable
fruit (Sáenz and Sepúlveda, 1999). El Samahy et al.
effect on the flavour of the juice and, for some
(2007a) studied the production of pasteurized and
of the cultivars (i.e. ‘Santa Rosa’), pasteurization
sterilized cactus pear juices, adjusting the pH and
caused the cactus pear and melon taste of the
adding preservatives in some treatments (sodium
fresh juice to turn bitter and astringent.
benzoate). Colour changes were observed at high
temperatures, but microbiological stability was An example of a commercial product is a drink
achieved in all the treatments. Baccouche et al. made from coloured cactus pear concentrate, wa-
(2013) produced different formulations of a cactus ter and natural flavouring. This product is available
pear beverage made with whey; it was pasteurized in the United States market, packaged in 1−litre
at 80° C for 20 minutes, and the physical stability Tetra Pak and sold for approximately US$6 litre−1.
144 Crop ecology, cultivation and uses of cactus pear

Cladode juices and nectars (−196° C), are widely used in the food industry
to improve the quality of the final product. The
Rodríguez (1999) reported various formulations of faster the freezing, the smaller the ice crystals
a drink made from tender cladodes, which were formed and the better the product quality
blanched at 95° C, liquefied and filtered. For the achieved. Freezing combines the effects of low
best results, the liquid was diluted 30% with wa- temperature (microorganisms cannot grow, chem-
ter, the pH was adjusted with citric acid (to reach ical reactions are reduced, and cellular metabolic
pH 3.5) and aspartame was used as a sweetener reactions are delayed) with a decrease in water
(1 g 335 ml−1). The product was pasteurized at 76° C activity (aw) (Casp and Abril, 1999; Vieira, 1996;
for 15.2 minutes, with mild loss of heat−sensitive Delgado and Sun, 2000). However, experiments
nutrients and other compounds. on cactus pear have not yet yielded good results
(Sáenz et al., eds, 2006). Several studies indicate
Recent years have seen an increase in the supply of
that freezing technologies could achieve better
cladode juices and nectars, mainly sold in Mexico.
results with cactus pear pulps, rather than whole,
The process for obtaining cactus pad juice is as half or sliced fruits. Defrosting − regardless of the
follows: type of cut (whole, halves or slices) − presents
• Remove spines from the cladodes. some problems, with excessive mucilage drip after
• Cut into pieces. defrost, resulting in an unpleasant appearance.
• Grind using industrial or domestic equipment This happens even when individually quick frozen
(blender). (IQF) technologies are adopted, involving freezing
temperatures close to −40° C.
• Add water to facilitate the process.
• Filter the liquid to separate the solids. Ice creams made from coloured cactus pear pulps
can be found in very few countries and may be
Cladode juice is produced by several companies an interesting alternative for the use of pulps or
in Mexico, and mixed juices are also made, com- concentrates. El Samahy et al. (2009) studied ice
bining cladodes with orange, pineapple, guava cream made with the addition of concentrated
or celery; the juices are available in the domestic red cactus pear pulp (30°Brix); a 5% addition of
market and for export. Another product sold in pulp was found to be the most acceptable. The
the Mexican market is “nopal water", a drink author’s research group is currently conducting
made with cactus cladodes juice and sugar, usu- experiments with this attractive product.
ally produced on a small scale. Other products
include cladode syrups, made from a base of
sucrose syrup (55−75° Brix) with the addition of OTHER TECHNOLOGIES
cladode juice. In Texas (United States of America),
a company produces blackberry and blueberry Fermentation
syrups with added cactus mucilage.
Fermentation is one of the oldest food preserva-
Canned nopalitos (tender cladodes) tion techniques and has been used with cactus
pear to obtain various products. O. streptacantha
Canning and pasteurization technologies are (cactus pear ‘Cardona’) has been used to produce
widely adopted in the manufacture of various alcoholic beverages in Mexico since the pre−His-
products made from nopalitos. In Mexico, for panic period; the most traditional drink made
example, pickled or salted tender cladodes (nopal- with the juice of this species is colonche (Corrales
itos) have been available on the market for many García and Flores Valdez, 2003; Diaz, 2003).
years. (Corrales García and Flores Valdez, 2003).
Detailed descriptions of the preparation of pickled Flores (1992) experimented with O. streptacantha
and O. robusta to make a wine and a distilled al-
nopalitos and nopalitos in brine, among others,
cohol. A wine of 11.6 ºGL was obtained from juice
are given in Sáenz et al., eds (2006).
concentrate (20 °Brix) using O. streptacantha; the
distilled alcohol reached 56.2 ºGL. The two spe-
Frozen products
cies, O. streptacantha and O. robusta, produced
alcohol with similar characteristics and a pleasant
Freezing is widely adopted for food preservation.
fruity aroma.
More than any other method, freezing allows
the taste, texture, and nutritional and functional Another product with interesting potential is
features to be preserved. Techniques, such as cold vinegar and some manufacturing experiences
air tunnels (−40 °C) or spraying liquid nitrogen are worthy of note. Pérez et al. (1999) prepared
Processing and utilization of fruit, cladodes and seeds 145

vinegar from orange cactus pear using two types seawater. Initial research on cactus pear juice
of substrate for acetic fermentation: must with focused on the application of MF and UF − the
previous alcoholic fermentation (13.5º GL) and methods normally used in juice clarification. Cas-
cactus pear juice with added sugar (22° Brix). Ace- sano et al. (2007) experimented with cactus pear
tobacter pasteurianus was used in the first case, cv. ‘Gialla’ (yellow−orange) and combined mem-
Acetobacter xylinum in the second. Both vinegars brane technologies with concentration by osmotic
obtained presented an intense yellow−amber col- distillation (OD); they used low temperatures and
our, clean and bright, with a fresh and acetic acid preserved the organoleptic, nutritional and sen-
smell. sory characteristics of cactus pear juice. The con-
centration process (OD) resulted in 61° Brix and a
Prieto et al. (2009) studied the development of
good balance of betalains.
balsamic−type vinegars from coloured cactus pear
juices; the resulting products had an attractive Cassano et al. (2010)
colour, pleasant aroma and good sensory accepta- later compared the
bility (Figure 12). performance of MF
and UF in the phys-
Membrane technologies icochemical com-
position of yellow
Membrane separation technologies have been in- cactus pear juice. In
creasingly used in the food industry in the last 25 both processes, the
years. Today their use is widespread and adopted solids suspended
for a range of purposes, including cold pasteuri- in fresh juice were
zation, juice clarification and bioactive compound completely removed
concentration (Cissé et al., 2011; Rai et al., 2006; and a clarified juice
Todisco et al., 2002). Membrane technologies obtained, and the
have advantages over other separation technolo- betacyanins were Figure 12
gies (traditional filtration and concentration): retained. This retentate, rich in betalains, could Balsamic−type vinegars
from coloured cactus
• Operation at low temperatures (15−35° C) be pasteurized and added to a juice concentrate
pear juices
− consequently affordable (low energy con- to prepare, for example, pulpy juices, ice cream, (Photo: C. Sáenz)
sumption) and degradation of heat−sensitive jellies or infant formula (Cassano et al., 2010).
compounds is avoided. The research group of the authors of this chapter
• No use of chemicals (filter aids or enzymes) − recently used this technology to separate or con-
unlike in filtration or traditional separation (Cas- centrate betalains from purple cactus pear MF, UF
sano et al., 2010). and NF (Cancino, Robert and Sáenz, unpublished
data). Thanks to the avoidance of high tempera-
In membrane separation technologies, the mem-
tures, there was no degradation of the pigments,
brane (ceramic or polymer) acts as a barrier: it al-
and the betalain extracts obtained were free of
lows only certain components in a mixture to pass
mucilage and had a reduced content of low mo-
and retains others. The flow of these substances
lecular sugars. Table 4 shows the characteristics
is determined by various driving forces, including:
of purple cactus pear pulp: prediluted (P), ultrafil-
pressure, concentration and electric potential. This
tered (UF) and nanofiltered (NF). The ultrafiltered
selectivity means, for example, that it is possible to
and nanofiltered extracts were, as expected, fully
enrich a flow with one or more substances present
clarified solutions (0 NTU), in comparison with
in the feed. Two flows emerge from the feed: per-
the prediluted pulp (2 453 NTU), which contained
meate (or filtrate) and retentate (or concentrate).
mucilage. In the UF, the betacyanin content (247.9
The retentate contains substances that do not
mg BE litre−1) was similar to that of P; however, in
cross the membrane, while the permeate is rich
the NF, the betacyanin values (216.3 mg BE litre−1)
in substances that pass through (Raventós, 2005).
were lower than those in P and UF, taking into
The selectivity depends on the size of the pores in
consideration that the pulp (feed) is diluted.
the membrane and the chemical affinity between
the membrane and the substances (Cheryan, Cassano et al. (2010) obtained the highest betalain
1998; Raventós, 2005). values with UF (32.8 mg BE litre−1). The polyphenol
content was concentrated in both membrane pro-
Membrane technology processes used in the food
cesses (UF and NF). Cassano et al. (2010) − apply-
industry include: microfiltration (MF), ultrafiltration
ing UF to cactus pear cv. ‘Gialla’ and using other
(UF), nanofiltration (NF) and reverse osmosis (RO).
membranes and process conditions − reported
RO is known for its effectiveness in desalinating
lower values of total polyphenols (552.17 mg
146 Crop ecology, cultivation and uses of cactus pear

TABLE 4 Physical and chemical characteristics of prediluted pulp, ultrafiltered and nanofiltered extract
from purple cactus pear

Parameter P UF NF
Total sugars (%) 13.2 ± 0.0 b 9.2 ± 0.3 c 17.5 ± 0.1 a
Turbidity (NTU) 2453 ± 64 a 0.00 ± 0.00 b 0.00 ± 0.00 b
Betacyanins (mg BE litre ) −1
254.0 ± 0.2 a 247.9 ± 4.3 a 216.3 ± 7.0 b
Betaxanthins (mg IB litre ) −1
85.4 ± 0.1 b 88.6 ± 1.2 a 79.1 ± 2.6 c
Total phenolics (mg GAE litre−1) 534.8 ± 4.4 b 659.7 ± 5.0 a 673.5 ± 13.5 a
Colour
L 12.5 ± 0.9 c 17.8 ± 0.1 b 19.3 ± 0.03 a
C* 48.0 ± 1.7 c 59.3 ± 0.1 b 62.4 ± 0.1 a
Hab 25.9 ± 1.0 c 30.3 ± 0.1 b 31.4 ± 0.1 a

P = pulp; UF = ultrafiltered pulp; NF = nanofiltered pulp; BE = betanin equivalent; IE = indicaxanthin equivalent; GAE = gallic acid equivalent.
Different letters in rows: means statistical differences (Tukey p < 0.05).
Source: Cancino, Robert and Sáenz (unpublished data).

GAE litre−1). The results obtained for the different sausages. However, it has some disadvantages:
extracts (P, UF and NF) are due to the membrane an earth−like flavour, imparted by geosmin and
pore size, as well as the chemical affinity between 3−sec−butyl−2 methoxypyrazine, and high nitrate
the membrane and the interaction between the levels. Red or purple cactus pears are, therefore,
different compounds and the membrane. an interesting alternative as a source of betanin
for the production of food colourants (Sáenz et
These kinds of products open new possibilities for
al., 2009; Castellar et al., 2003) (Figure 13).
the production of colourants from cactus pear.
However, further research is required to improve The use of a purple concentrate cactus pear juice
the pigment concentration and other properties as food colouring for dairy products (e.g. yogurt)
of the extracts. was studied some years ago (Sáenz et al., 2002d;
Sepúlveda et al., 2002, 2003c). Topics including
pigment purification and stability, crucial for appli-
OTHER PRODUCTS cation in industry, are currently being addressed.
Castellar et al. (2008) obtained a betalain concen-
Cactus pear colourants trate by fermentation (Saccharomyces cerevisiae
var. Bayanus AWRI 796) of O. stricta juice. The
Natural colourants − in particular, red and pur- final product had 9.65 g betanin litre−1, pH 3.41,
ple − are highly appreciated by consumers since 5.2° Brix and a viscosity of 52.5 cP.
synthetic red colourants used as additives in food
Other studies focused on dehydration of cactus
have been restricted by official regulations in the
pear pulp by spray−drying or freeze−drying to ob-
European Union and the United States of America
tain colourant powders. Mosshammer et al. (2006)
because of the possible adverse effects on hu-
adopted lyophilization and spray−drying with
man health (Tsuda et al., 2001). Therefore, there
maltodextrin as carrier (18−20 dextrose equivalent
is a growing interest in new, natural red pigment
[DE]) to dehydrate O. ficus−indica cv. ‘Gialla’ juice;
sources and their potential application in foods.
they reported high betalain retention (≤ 90%) for
Red and purple cactus pear ecotypes have a var- both kinds of powder. Similarly, various species of
iable betalain content in both the pulp and peel purple cactus pear (O. stricta, O. streptacantha, O.
(Odoux and Domínguez López, 1996; Sepúlveda lasiacantha) were spray−dried, using maltodextrin
et al., 2003c). This pigment is commercially ob- as drying carrier (10, 20 DE). Diaz et al. (2006) ob-
tained from red beetroot and is widely used in served that after 24 weeks at 25° C, the powder
the food industry. Its use in food is permitted by retained 86% of the original content of betanin.
both US and EU legislation (Sáenz et al., 2009). Obón et al. (2009) used glucose syrup (29 DE)
The red beet extract (rich in betalains) is used and reported that after 1 month stored at room
mainly to colour food, such as dairy products, temperature, the colour powder retained 98%.
confectionery, ice cream, desserts, beverages and Rodríguez Hernández et al. (2005) observed that
Processing and utilization of fruit, cladodes and seeds 147

Figure 13
Purple cactus
pear fruits and
microparticles
(Photos: C. Sáenz)

cactus pear powder, when reconstituted, presented Extruded products

a slightly different colour from the fresh juice.
Extrusion cooking is a high−temperature short−time
Betalain stability is affected by several factors: pH,
(HTST) technology. While there has been little relative
water activity, exposure to light, oxygen, enzyme
study in cactus pear and cladodes, this technology is
activity and, above all, temperature (Azeredo,
2009; Herbach et al., 2006; Castellar et al., 2003). widely applied in food processes, such as breakfast
A technology − microencapsulation − is availa- cereals, salty and sweet snacks, baby foods and snack
ble to stabilize the pigments (Sáenz et al., 2009, foods. Food materials are plasticized and cooked,
2012b; Gandía Herrero et al., 2010; Vergara et al., combining moisture, pressure, temperature and
2014; Robert et al., 2015). shear. The factors affecting product quality include
extruder type, screw configuration, speed, tempera-
Microencapsulation is a technique involving the ture and feed rate, as well as the composition of the
introduction of bioactive compounds (solid, liquid raw material and the type of food ingredients used
or gaseous) into a polymeric matrix to protect (Singh et al., 2007; El Samahy et al., 2007b).
them from the environment, interaction with oth-
er food components or controlled release (Yáñez El Samahy et al. (2007b) studied cactus pear rice−
Fernández et al., 2002). Microencapsulation by based extruded products − an innovative option
spray−drying is the most widely used technique: for production of a new value−added snack based
it obtains a powder of low water activity, enabling on cactus pear pulp concentrates. In this study,
easier transportation, handling and storage and orange−yellow and red cactus pear pulps were
ensuring microbiological quality (Hayashi, 1989). concentrated (40º Brix), added to rice flour and
the blend put in a single−screw extruder. Different
Vergara et al. (2014) combined membrane tech- formulations (varying the ratio of rice flour and
nology (to separate betalains) and microencap- cactus pear pulp concentrate) were tested. Substi-
sulation (to protect them), obtaining a healthy tution levels of 5% and 10% of concentrated cac-
colourant that can be used in the food industry. tus pulps gave the best results for extruded prod-
Gomez (2013) studied the microparticle betalains ucts with good functional, nutritional and sensory
stability in soft drinks, comparing microparticles characteristics. The poor sensory attributes of the
from cactus pear pulp, and ultrafiltered and na-
formula without cactus pear pulp were significant-
nofiltered extracts; he concluded that the stability
ly improved by adding cactus pear concentrates.
of betalains is affected both by their source (pulp
or extract) and by the encapsulating agent used. Sarkar et al. (2011) extruded cactus pear pulp
Betalains from cactus pear pulp are more stable (yellow variety) with rice flour. The authors tested
in soft drinks, probably thanks to the mucilage. different solids ratios (rice flour solids : puree sol-
Alfaro (2014) applied purple cactus pear pulp mi- ids − 6 : 1, 8 : 1 and 10 : 1). The blends were dried
croparticles in yogurt, and reported that after 45 in a twin−screw extruder. The results revealed that
days’ storage, 60% of betalains were retained. some characteristics, such as porosity, decreased
when the fruit solid level increased.
There has been relatively little research to date on
the encapsulation of betalains from cactus pear Cactus pear pulp has potential for the production
(O. ficus−indica) (Sáenz et al., 2009; Vergara et of extruded products; further study is required to
al., 2014; Pitalua et al., 2010; Gandía Herrero et better understand the behaviour of cactus pulp in
al., 2010; Robert et al., 2015; Otárola et al., 2015). this process and to improve characteristics of the
Further investigation is required. products obtained.
148 Crop ecology, cultivation and uses of cactus pear

Hydrocolloids (mucilages) from cladodes Seed oil

Hydrocolloids are polysaccharides with variable com- Cactus pear fruits contain variable amounts of seeds, but
plexity, generally used in the food industry as additives they are usually present in a high proportion (10−15 g
to provide viscosity to, for example, beverages, puddings 100 g−1). In most cactus pear fruit processes, the seeds
and salad dressings. They include cactus mucilage, a pol- are separated from the pulp, resulting in large quantities
ysaccharide−type arabinogalactan present both in cactus of discarded seeds that become an environmental waste
pads and in cactus pear fruits (Sáenz et al., 2004; Matsu- problem. For this reason, in recent decades, researchers
hiro et al., 2006). The mucilage has an important physio- in different countries have studied the seed composition
logical role in the Opuntia species as it has a high water and sought different possible uses for the seeds.
retention capacity (Nobel et al., 1992). It can be extracted
Sawaya et al. (1983) studied the seed composition and
from the matrix (cladodes or fruit peel) with water, and
its potential utilization in animal feed. They reported
precipitated with ethanol; or other techniques can be
adopted, such as pressing (Sepúlveda et al., 2007). In 16.6% protein content, 17.2% fat, 49.6% fibre and
general, the extraction yields are low (≤ 2% fresh weight 3.0% ash. The mineral content is high in sodium (67.6
[FW]), but they nevertheless offer an interesting prospect, mg 100 g−1), potassium (163.0 mg 100 g−1) and phos-
considering that the cladodes (pads) are usually pruning phorus (152.0 mg 100 g−1).
residues and are available throughout the year. Various Cactus pear seed oil is edible; it could be another nutritive
extraction methods have been reported using different and functional product of potential interest to the food
solvents to precipitate the mucilage, such as ethanol, industry, but perhaps not for direct consumption (as
isopropyl alcohol and acetone (Rodríguez González et al., indicated below). The oil is usually extracted in research
2014; Cai et al., 2008; Sepúlveda et al., 2007; Yahia et using organic solvent (4.4−14.10%) (Sawaya and Khan,
al., 2009; Medina Torres et al., 2000). Some studies have 1982; Sepúlveda and Sáenz, 1988; Ennouri et al., 2005;
researched the application of this hydrocolloid in fruit Becerril, 1997; Tlili et al., 2011; Ouerghemmi et al., 2013;
nectars to replace other thickeners normally used in the Chougui et al., 2013), depending on factors such as grow-
food industry (e.g. CMC) (Sepúlveda et al., 2003a, 2004). ing conditions, variety and fruit maturity (Özcan and Al
The mucilage has also been tested as an edible coating Juhaimi, 2011). Use of a cold press to obtain the seed oil
to protect fresh fruits. Del Valle et al. (2005) used muci- was only reported by Gharby et al. (2015) from Moroc-
lage as edible film to increase the shelf−life of strawber- co, with a yield of 6−7%. This type of extraction is more
ries stored at 5° C, reporting that the fruits maintained environmentally friendly as it avoids the use of organic
their texture and flavour and no deterioration occurred solvents.
after 9 days of storage. This edible film could provide an Yields of edible oils are 6−17%, which, in terms of
alternative for the preservation of different fresh fruits, waste stream, compares reasonably with other com-
such as berries. However, application to different prod- monly used seed oils. The production of cactus pear
ucts is a challenge, because the respiratory rate of each seed oil as edible oil is only viable with integrated pro-
type of fruit must be taken into account. Aquino et al. cessing, using all parts of the plant (Sáenz et al., eds,
(2009) used a mucilage solution blended with different 2006).
citric acid and sodium bisulphite concentrations to inhibit
the browning of banana slices during the drying process. Cactus pear seed oil is rich in unsaturated fatty acids
The authors reported that a combination of 500 ppm so- and has a high linoleic acid content (57.7−73.4%) and
dium bisulphite and citric acid (1%), following treatment low linolenic content. Table 5 shows the percentages
in the mucilage solution (35 mPas), inhibited browning of the main seed oil fatty acids.
and made the banana slices shinier. The oil has a high unsaturated fatty acid content, as
Recently, Medina Torres et al. (2013) and Otárola et al. well as other healthy compounds, such as sterols,
(2015) reported that mucilage can be used as an en- tocopherols, vitamin E, β−carotene and vitamin K
capsulating agent for bioactive compounds, gallic acid (Ramadan and Mörsel, 2003a; Kouba et al., 2015). Phe-
and betalains; this points to new opportunities in the nolic compounds were reported by Tlili et al. (2011) and
industrial sector. Sáenz et al. (2009) also reported this Chougui et al. (2013): respectively, 61 mg GAE 100 g−1
behaviour in a study of the encapsulation of betalains and 268 mg 100 g−1, expressed as rutin equivalent. The
from purple cactus pear fruits. researchers in Mexico and Taiwan reported higher fig-
ures (Cardador Martínez et al., 2011). Further research
Lira Ortiz et al. (2014) extracted low methoxyl pectins
is required before definitive conclusions can be drawn.
from O. albicarpa Scheinvar skin, and discovered its po-
tential for use in the food industry as a thickening and These and other physical and chemical properties,
gelling agent − the latter when calcium ions were added. including refractive index, iodine index and saponifi-
The authors obtained a yield of 98 g kg−1 of dry matter. cation number, highlight the similarity between cactus
Processing and utilization of fruit, cladodes and seeds 149

TABLE 5 Fatty acid content (%) in cactus pear seed oil (Opuntia ficus−indica) from different countries

Fatty acid Countries

Morocco a
Turkey b
South Africa a
Tunisia a,c,g Germany d Chile e Algeria f
Palmitic (C16 : 0) 11.9 10.6−12.8 13.7 12.2−12.7 23.1 16.2 13.1
Stearic (C18 : 0) 3.4 3.3−5.4 3.38 3.2−3.9 2.67 3.3 3.5
Oleic (C18 : 1w9) 21.3 13−23.5 15.7 16.4−22.3 24.1 19.9 16.3
Vaccenic (C18 : 1n−7) − 5.1−6.3 − 4.8 − − 5.3
Linoleic (C18 : 2w6) 60.8 49.3−62.1 64.38 53.5−60.6 32.3 57.7 61.8
a
Gharby et al. (2015); b Mattháus and Özcan (2011); c Tlili et al. (2011); d Ramadan and Mörsel (2003a); e Sepúlveda
and Sáenz (1988); f Chougui et al. (2013); g Ouerghemmi et al., 2013.

pear seed oil and other edible vegetable oils, such to diversify their production. When, on the other
as corn oil or grape seed oil. Oil extraction yields hand, the aim is the creation of new agro−indus-
are low compared with other common edible tries for cactus pear processing, investment is re-
seed oils; cactus pear seed oil could nevertheless quired and must come from governments, NGOs
be used in the food industry as a fat replacer in or other sources. In some countries, a cooperative
special confectionery products. However, pharma- model could be promoted. The immense variety of
ceutical and cosmetics uses offer more and better cactus pear products and by−products can bring
alternatives. In this context, cactus pear seeds also huge benefits to many people, in particular those
contribute with essential oils, a group of com- living in arid and semi−arid zones of the world.
pounds used mainly in the pharmaceutical field.
Ouerghemmi et al. (2013) reported that essential
Figure 14
oils comprise, among others, terpenes, esters and (a) Seed separator
aldehydes, and the yield for cactus pear seeds is
close to 4%.
In recent years, other applications have emerged,
in particular for cosmetics; this industry has ex-
ploited the oil’s characteristics (polyunsaturated
fatty acid content, tocopherols, sterols, phenolics)
and in countries such as Morocco a promising
industry has emerged. There are several cooper-
atives and private companies that extract this oil
for cosmetic purposes, and at least 20 producers
of cactus oil exist (Abderrahman Ait Hamou, AN-
ADEC, Morocco, personal communication). The
industry extracts the oil by cold press − an envi-
ronmentally friendly process that avoids the use
of solvents (Berraaouan et al., 2015; Gharby et al.,
2015). Before the oil is extracted by cold press, the
seeds must be separated using a special machine
(Figure 14).

CONCLUSION
There are a vast array of alternatives for process-
ing the fruits, cladodes and seeds from Opuntia
plants. In general, the technologies used are
available to many small−scale agro−industries that
could take advantage of this new raw material
 13
Nutritional properties
and medicinal derivatives
of fruits and cladodes
Mónica Azucena Nazareno
Research and Transfer Center of Santiago del Estero
National Scientific and Technical Research Council
National University of Santiago del Estero, Argentina
 13 Nutritional properties and medicinal
derivatives of fruits and cladodes

INTRODUCTION be used as natural medicines to prevent and cure

serious diseases. These beneficial properties of cac-
Scientific investigation has confirmed that fruits tus, already known to ancient civilizations, are now
and cladodes of cacti may be efficiently used as generating interest among the scientific communi-
a source of nutrients and phytochemicals (e.g. ty. The scientific literature concerning the medicinal
sugars, mucilage, fibres, vitamins and pigments) properties of cactus is in constant expansion, with
of nutritional and functional importance. Cactus new discoveries reported about the plant constitu-
products show promising functional character- ents responsible for such activities.
istics due to their health−promoting properties.
This chapter presents a detailed description of the
major active components of the different cactus NUTRITIONAL ASPECTS
species studied worldwide.
Since ancient times, cactus plants have been The nutrient composition of fruits and cladodes
used to cure diseases and heal wounds. Cacti are depends on many factors: species, cultivar or va-
traditionally used as natural medicines in several riety; environmental factors, such as the climatic
countries for the treatment of many diseases. and edaphic conditions; crop management, in-
Moreover, cladodes and cactus pear fruits are cluding fertilization and post−harvest treatment;
still used today in folk medicine as therapeutic and maturity status.
agents. Remarkable progress has been made in
recent decades to characterize plant constituents Fruits
and explain the role of natural molecules in rela-
tion to disease prevention. In this context, there Many cactus species produce edible fruits. Cur-
are strong recommendations to incorporate fruit rently, most of the cactus fruits in the global
and vegetables in the diet. Several manufactured market belong to the Opuntia ficus−indica species
products are currently available on the global mar- and the fruits are known as “cactus pear”. The
ket that exploit the medicinal properties of cactus typical cactus pear fruit is an oval−shaped berry
plants; there is increasing interest in the industrial with an average weight of 100−200 g. The juicy
Figure 1 use of cactus products as nutraceuticals. pulp constitutes 60−70% of the total fruit weight
Cactus pear chemical and contains numerous small hard seeds, varying
composition (% in Cactus plants are multipurpose crops: not only do from 100 to > 400 per fruit. Figure 1 shows the
fresh matter) and they provide food and feed, they contain bioac- fruit composition. Cactus pear is a non−climac-
fruit constitution as
tive phytochemicals. Fruits and cladodes already teric fruit, it has a low respiration rate (Cantwell,
a percentage of fruit
weight, fw (El Kossori provide energy and nutrients for both humans 1995) and low ethylene production. Hence, the
et al., 1998; Duru and and livestock. However, cactus plants are also rich nutrient concentrations remain in the fruit without
Turker, 2005) sources of health−promoting substances that can significant changes throughout storage; however,
weight and firmness may change. Furthermore,
the fruit’s pH, acidity and total soluble solids do
Peel not vary during post−harvest storage; although
or pericarp
30-40% some changes in vitamin C content have been
fw
Fruit reported, depending on the storage conditions
water (85) (Coria Cayupán et al., 2009).
Seeds
2-10%
sugars (6-14)
fw The chemical and nutritional composition of cac-
pectin (1) tus pears has been extensively studied (El Kossori
protein (0.3) Pulp
et al., 1998; Butera et al., 2002; Feugang et al.,
lipids 60-70% 2006). Sugars, fibre, mucilage and pectins are the
fw
major fruit constituents; proteins, amino acids,
vitamins and minerals are the minor components
(Tesoriere et al., 2005b). Secondary metabolites
with antioxidant properties have been described
Nutritional properties and medicinal derivatives of fruits and cladodes 153

(Kuti, 2004; Yahia and Mondragón, 2011; Coria Cay- rently under development by the cosmetics industry.
upán et al., 2011). Cactus pear is characterized by high The seed endosperm is composed of arabinan−rich pol-
sugar content (12−17%) and low acidity (0.03−0.12%) ysaccharides, while the major seed−coat component
with pH values between 5 and 7. Furthermore, total is D−xylan (Habibi et al., 2002). Xylan has been used as
soluble solid content varies from 11.6 to 15.3° Brix, an adhesive, a thickener and as an additive in plastics;
increasing with fruit maturity (Yahia and Mondragón, it is of growing interest to the food industry due to its
2011). The relative ratio of glucose and fructose varies potential for packaging films and coating food; it also
among different species and within fruit tissues: 53% acts as an emulsifier and protein foam stabilizer during
glucose and 47% fructose of total sugar content (Kuti heating. Nowadays, it may play an important role in
and Galloway, 1994). The fruit pulp is also rich in min- the development of new biomedical products for novel
erals, such as calcium (59 mg 100 g−1) and magnesium drug delivery systems, especially for controlled drug
(98.4 mg 100 g−1) (Stintzing et al., 2001). Fruit peel and release.
pulp constituents contain mucilage in the dietary fibre;
it is a hydrocolloid and has a high capacity to absorb Cladodes
water. The total amino acid content (257.24 mg 100
g−1) is higher than the average of other fruits; the prin- Cladodes are modified flattened stems with a charac-
cipal amino acids are proline, taurine and serine. Signif- teristic ovoid or elongated shape; they are able to exert
icant amounts of ascorbic acid are present in Opuntia a photosynthetic function. Young tender cladodes,
ficus−indica, ranging from 180 to 300 mg kg−1 of fresh called nopalitos, are consumed as fresh vegetables;
fruit (Piga, 2004). they are used as an ingredient in a wide range of dishes,
Tocopherols, the fat−soluble vitamin E, are found in the including sauces, salads, soups, snacks, pickles, bever-
lipid fraction of both the cactus fruit seed and pulp. The ages, confectionery and desserts (Saenz et al., 2002a).
vitamin E homologues isoforms γ− and δ−tocopherols The major components of cladodes are carbohydrate−
are the main components in seed and pulp oils, respec- containing polymers, comprising a mixture of mucilage
tively, constituting up to 80% of the total vitamin E and pectin. The chemical composition of fresh young
content (Ramadan and Morsel, 2003a). Lipids are dis- cladodes is reported by Saenz et al. (2002a):
tributed in the fruit peel, pulp and seeds. The fruit peel • moisture 91% (weight of water [w w])
contains appreciable amounts of polyunsaturated fatty • total carbohydrates 4.5%
acids, in particular linoleic acid. It also contains other
• protein 1.5% (w x in dry basis [DB])
fat−soluble compounds, such as sterols, beta−carotene
and vitamin K1; the main sterol is β−sitosterol (Ram- • fat 0.2% (w w DB)
adan and Mörsel, 2003b). Peel polysaccharides and • ash 1.3% (w w DB), of which 90% calcium
pectins have been characterized by Habibi et al. (2004).
Guevara Figueroa et al. (2010) analysed the cladode
proximate composition of wild Opuntia spp. (Figure 2).
Seed and seed oil In addition, 100 g of cladodes contain 11 mg vitamin C
and 30 µg of carotenoids.
Cactus pear contains many hard−coated seeds that
represent 10−15% of the pulp weight. The fruits The mucilage is a polysaccharide; it can be found in
contain a large number of seeds although their oil specialized storage cells or free within cells or intracel-
content is relatively low. Oil content is 7−15% of the lular spaces of the chlorenchymatic and parenchymatic
whole seed weight. Essential fatty acids (mainly linoleic tissue of the cladodes. The dried mucilage comprises
acid) form a significant percentage of the unsaturated on average: 5.6% moisture; 7.3% protein; 37.3% ash;
fatty acids of seed oil extract. Unsaturated fatty acids 1.14% nitrogen; 9.86% calcium; 1.55% potassium
account for about 80% of all fatty acids (Ennouri et (Sepúlveda et al., 2007). Based on its chemical compo-
al., 2005). The linoleic acid content varies between sition, the mucilage is considered a polymer (similar to
61.4 and 68.9%. The α−linolenic acid concentration in pectin), composed of arabinose, galactose, xylose and
all cultivars is < 1%. The oleic acid content varies be- rhamnose as neutral sugars, and of a small amount of
tween 12.4 and 16.5%. Therefore, although the seed galacturonic acid (Medina Torres et al., 2000; Mad-
oil content is relatively low, its fatty acid composition jdoub et al., 2001). Besides the direct consumption of
indicates that it has potential for use in the health and tender shoots, mature cladodes are ground to make
cosmetic industries (Labuschagne and Hugo, 2010). flour and other products. Nopal flour is a rich source
Seed oil destined for cosmetics sells at a very high price of dietary fibre reaching up to 43% DB (Saenz et al.,
as organic oil for use in the production of anti−ageing 2002a); it can be used to fortify food recipes containing
and anti−wrinkle products. New applications are cur- flours from other sources.
 154 Crop ecology, cultivation and uses of cactus pear

BIOACTIVE PHYTOCHEMICALS sible for the ripe fruit colours and, therefore, they
IN CACTUS PLANT are a major factor in consumer acceptance. The
concentration of pigments depends on several
factors, in particular fruit ripening status (Coria
Cactus plants are also important sources of bio-
Cayupán et al., 2011). Stintzing et al. (2005) char-
active substances and excellent candidates for nu-
acterized the betalain composition of cactus pear
traceutical and functional food preparation. The
fruit. These pigments do not only provide colour:
fruits, cladodes, seeds and flowers all have a high
their antioxidant properties are greater than those
content of chemical constituents, adding value to
of ascorbic acid (Butera et al., 2002; Stintzing et
cactus products.
al., 2005). In contrast with anthocyanins (anoth-
er group of natural red pigments), betalains are
Fruits
stable in a wider pH range and are thus more
appropriate for use as food colourants in low−ac-
Cactus pear fruits present high antioxidant ac-
id products (Stintzing et al., 2001). Due to their
tivity, ascribed to the presence of vitamin C, fla-
wide structural variety, and hence colour diversity,
vonoids and betalains (Galati et al., 2003a; Kuti,
betalains constitute a very promising source of
2004). Antioxidant activity in the fruit is twice that
natural colourants to be used as functional colour-
of pear, apple, tomato, banana and white grapes,
ants. Compared with red beets, cactus pear not
and is similar to that of red grapes and grapefruit
only offers a wider range of colours, it also has
(Butera et al., 2002).
technological advantages: absence of geosmin (an
Betalains are nitrogen−containing pigments and undesirable earth−like flavour); low nitrate levels;
can be classified as red betacyanins (e.g. beta- and lack of soil microbial contamination.
nin) and yellow betaxanthins (e.g. indicaxanthin).
The presence of phenolics was also detected in
Betacyanins are ammonium conjugates of bet-
cactus fruit pulp. Kuti (1992) reported an antiox-
alamic acid with cyclo−DOPA; betaxanthins are
idative effect due to the major flavonoids found
conjugates of betalamic acid with amino acids or
in cactus fruits (e.g. quercetin, kaempferol and
amines. Betalains are the main pigments respon-
isorhamnetin derivatives). Flavonol derivatives de-
tected in Opuntia spp. were reviewed by Stintzing
and Carle (2005). The peel has a higher phenolic
content than the pulp (Stintzing et al., 2005).
Consequently, from a functional point of view, it
is advantageous to process both peel and pulp.
Lee et al. (2002a) found that cactus flavonoids are
Cladodes Cladode dry matter also effective in protecting plasmid DNA against
water (88-95) carbohydrates (42-81)
the damage induced by hydroxyl radicals.
ash (19-23) ash (5-20)

fibres (18) fibres (5-20)

Cladodes
protein (4-10) protein (6-19)

lipids (1-4) lipids (0.1-2)

Other functional components, such as chlorophyll
derivatives, amino acids and flavonoids, are found
in cactus pads (Figure 3). Guevara Figueroa et
al. (2010) analysed cladodes of commercial and

Figure 2
Cladode proximal
composition, %
(Stintzing and Carle,
2005; Guevara Figueroa
et al. 2010)

Figure 3
Main phytochemicals
in cactus fruits and
cladodes
Nutritional properties and medicinal derivatives of fruits and cladodes 155

wild Opuntia spp. from Mexico and evaluated the tance (El Mostafa et al., 2014). The cactus plant
polyphenolic and flavonoid profiles of both fresh and can be fully exploited since its bioactive compo-
processed nopal products. The presence of five ma- nents can be extracted from its various parts:
jor flavonoids (isoquercitrin, isorhamnetin−3−O−glu- flowers, fruit, cladodes, roots and seeds (Nazare-
coside, nicotiflorin, rutin and narcissin) was observed no, 2014). Figure 4 summarizes the most relevant
in all varieties and nicotiflorin was predominant. medicinal properties ascribed to cactus products.

Flowers Fruits

Cactus flowers accumulate betalains as well as col- Diet supplementation with cactus pear fruits in
ourless phenolic compounds (Ahmed et al., 2005). a healthy human decreases oxidative stress, by
The chemical composition of O. ficus−indica and improving the overall antioxidant status. The
O. stricta flowers extracted at four−flowering stage effect of cactus betalains on oxidative stress in
was studied by Ammar et al. (2012). The antiradical, humans was studied by Tesoriere et al. (2003,
antibacterial and antifungal activities in ethanol, 2004, 2005a), who reported that fruit ingestion
as well as hexane extracts of flower, were also produced a decrease in oxidative stress markers,
analysed. Phenolic content varies markedly with inhibited LDL oxidation and increased resistance
flowering stage, and these active constituents are to oxidative haemolysis red blood cells in ex vivo
at their peak during post−flowering stage. De Léo experiments. Budinsky et al. (2001) showed that
et al. (2010) reported the chemical profile of meth- regular consumption of O. robusta reduces oxida-
anol extract of O. ficus−indica flowers. The volatile tion−mediated damage.
fraction of three Opuntia species obtained from
Zou et al. (2005) studied the ability of cactus pear
aqueous distillation was characterized and assayed
fruit extract to suppress carcinogenesis in cultured
as antifungal agents (Bergaoui et al., 2007).
human cancer cell lines in vitro and an animal
model in vivo. The results showed that the cactus
extract inhibited the growth in vitro of ovarian,
MEDICINAL PROPERTIES bladder and cervical cancer cells and suppressed
tumour growth in the animal inoculated with
Cactus plants were used by ancient civilizations to ovarian cancer cells. The antiproliferative effects
cure diseases and heal wounds for thousands of of betanin on human chronic myeloid leukaemia Figure 4
years. The origins and history of cactus as med- cell line−K562 were reported by Sreekanth et al. Medicinal properties
icine are, therefore, closely linked to the ancient (2007). O. humifusa fruit extracts were assayed of cactus
Mesoamerican civilizations. For over 12 000 years,
fresh cactus has been consumed by the natives
for its nutritional qualities and healing properties.
Cactus cladodes, fruits, seeds and flowers have ANTI
ANTI OBESITY ANTI
been used as folk medicines in several countries HYPERLIPIDEMIC INFLAMMATORY
for centuries.
There are numerous reports indicating that a diet ANTI
ULCEROGENIC CICATRIZANT
rich in fruit and vegetables is linked to lower inci-
dence of coronary heart disease and some types
of cancer; this suggests that this kind of diet has
positive effects on health (Bazzano et al., 2002).
HYPOGLYCEMIANT ANTIVIRAL
These beneficial effects are associated, not only
with the nutrient and vitamin content of foods,
but with the action of certain bioactive compo-
nents. Phytochemicals with antioxidant properties HEPATO Cactus CHOLESTEROL
promote good health by protecting against the PROTECTOR Medicinal REDUCING
ACTION
oxidative damage induced by reactive oxygen spe- Properties
cies (Prakash and Gupta, 2009).
Several studies demonstrate that both cactus NEUROPROTECTOR ANTICANCER
fruits and cladodes have high levels of important ANTI DNA
GENOTOXIC PROTECTOR
nutrients, minerals and vitamins, as well as antiox-
idants. The cactus plant appears to be an excellent
source of phytochemicals of nutraceutical impor-
156 Crop ecology, cultivation and uses of cactus pear

against breast cancer and glioblastoma human cell lines Moreover, consumption of young cladodes reduces obe-
(Hahm et al., 2010). Emerging studies indicate remark- sity and blood glucose. The anti−obesity factor, hypogly-
able anticancer activities displayed by cactus pear ex- cemic action and antidiabetic effects were all observed in
tracts. The chemopreventive and anticancer activities of rats and humans (Bwititi et al., 2000; Frati Munari et al.,
crude extracts from plants belonging to the Cactaceae 2004; Yang et al., 2008).
family, as well as their main active constituents are well
Galati et al. (2001) proposed that O. ficus−indica cladodes
reviewed by Harlev et al. (2013).
stimulate a protective response in the gastric mucosa,
Studies in vitro of the interaction between purified preventing the development of ethanol−induced ulcers
betalains and hypochlorous acid and human myelop- (preventive treatment) and promoting a faster recovery
eroxidase revealed the anti−inflammatory action of (curative treatment). The cytoprotective effect of O.
these fruit pigments (Allegra et al., 2005). Recently, ficus−indica cladodes is ascribed to the physicochemical
remarkable anti−inflammatory effects of indicaxan- properties of the mucilage (Galati et al., 2001). O. ficus−
thin were reported in an animal model (Allegra et al., indica cladodes produce cytoprotection by increasing
2014). The anti−ulcerogenic and antigastritis effects of mucus secretion in the gastric mucosa of rats affected by
cactus fruits were studied in rats by Lee et al. (2001) ethanol−induced ulcers (Galati et al., 2002). Cactus cla-
and Galati et al. (2003a). Hepatoprotection of O. dode administration is recommended for both preventive
ficus−indica fruit juice and extract was demonstrated and curative treatment of gastric ulcer (Lee et al., 2002a).
in rats (Galati et al., 2005; Alimi et al., 2012). Kim et Opuntia ficus−indica plant extract can alleviate alcohol
al. (2006) demonstrated the neuroprotective action of hangover symptoms in humans (Wiese et al., 2004).
O. ficus−indica fruit extracts against neuronal oxidative
Furthermore, the protective effect of Opuntia ficus−ind-
injuries induced by excitoxins in mice cortical cells. They
ica cladode juice against Ni−induced toxicity is reported
also reported in vivo experiments where methanolic
by Hfaiedh et al. (2008). Experimental exposure to Ni
fruit extracts reduced neuronal damage produced by
was found to generate reactive oxygen species (ROS),
global ischemia in gerbils by 36% and ascribed these
leading to increased lipid peroxidation, loss of mem-
effects to bioflavonoid antioxidant action. Wolfram et
brane integrity and alterations of the cellular antioxidant
al. (2002) reported that the ingestion of 250g day−1
system. There is indication that cladode juice can prevent
of O. robusta fruits produced an anti−hyperlipidemic
oxidative stress and decrease the related parameters in
effect and cholesterol−reducing action in non−diabetic
rats; in addition, regular ingestion of cladode juice can
hyperlipidemic humans; there are also reports of the ef-
counteract the peroxidative effects of nickel. Similar
fects on glucose metabolism. The hypocholesterolemic
protective effects against oxidative damage induced by
action can be partially explained by the fibre (pectin)
various toxics are also reported by Ncibi et al. (2008) and
content of fruits. Although the hypoglycaemic mech-
Zourgui et al. (2008). The neuroprotective action against
anism is still unclear, investigations indicate that this is
neuronal oxidative injuries of O. ficus−indica flavonoids
because fruits seem to promote − in both diabetics and
extracts has also been assessed; they were found to be
non−diabetics − a faster and better glucose entry into
effective in cultured mouse cortical cells and against
the cell, and the enhancement of cellular sensitivity to
global ischemia in gerbils (Kim et al., 2006).
insulin is suspected. Platelet function improvement due
to regular consumption of cactus pear (250 g day−1) in Cladode extracts might have a hepatoprotective ef-
healthy volunteers and in patients with mild familial fect against aflatoxicosis in mice; they probably act by
heterozygous hypercholesterolemia was described by promoting the antioxidant defence systems (Brahmi et
Wolfram et al. (2003). Fruit ingestion induces benefi- al., 2011). Experiments concerning the antiviral action
cial actions on the cardiovascular system by decreasing of cactus extracts have been conducted. A cladode
platelet activity and improving haemostatic balance. extract from Opuntia streptacantha was reported to
exhibit antiviral properties towards DNA viruses, such
Cladodes as herpes, and ribonucleic acid (RNA) viruses, such as
influenza type A and human immunodeficiency virus
When O. ficus−indica cladodes were fed to hypercho- HIV−1. The active principle was located in the outer
lesterolemic rats, a marked decrease in cholesterol and non−cuticular tissue and ascribed to a protein with
triglycerides levels was found in plasma samples. An unknown mechanisms of action (Ahmad et al., 1996).
anti−hyperlipidemic effect and cholesterol reduction Replication of both DNA and RNA viruses was inhib-
were observed in guinea pigs, rats and mice (Galati et ited. An International Patent1 claims that nopal cactus
al., 2003b; Oh and Lim, 2006). Experiments in diabetes has an effect against herpes simplex and influenza A
mellitus non−insulin−dependent patients confirmed the viruses. Chlorophyll derivatives are proposed as the
hypoglycemic effects of Opuntia streptacantha cladodes. active compounds.
1
International Patent (1993): Skinner and Ezra (GB, Israel). Nopal cactus effect on herpes simplex and influenza A viruses.
Nutritional properties and medicinal derivatives of fruits and cladodes 157

TABLE 1 Medicinal properties of cactus products

Cactus species, parts and activities Studied system and reference

Antiviral action
O. streptacantha cladode extract Intracellular virus replication inhibition and extracellular virus
inactivation (Ahmad et al., 1996)
Opuntia sp. cladodes Guinea pigs (Fernández et al., 1994)
Anti−hyperlipidemic effect and cholesterol level reduction
O. robusta fruits Non−diabetic hyperlipidemic humans (Wolfram et al., 2002)
O. ficus−indica cladode Rats (Galati et al., 2003b)
O. ficus−indica seeds and seed oil Rats (Ennouri et al., 2006a, b, 2007)
O. ficus−indica var. saboten Mice (Oh and Lim, 2006)
Anti−obesity factor
Opuntia sp. cladode Humans (Frati Munari et al., 2004)
O. megacantha Diabetic rats (Bwititi et al., 2000)
O. lindheimeri Diabetic pigs (Laurenz et al., 2003)
Hypoglycemic and antidiabetic effects
O. ficus−indica, O. lindheimeri and O. robusta fruits Diabetic rats (Enigbocan et al., 1996)
O. streptacantha Humans (Meckes Lozyoa, 1986)
O. monacantha cladode extract Diabetic rats (Yang et al., 2008)
O. ficus−indica seeds and seed oil Rats (Ennouri et al., 2006a, b)
O. streptacantha Humans (Frati Munari et al., 1992)
O. filiginosa fruit extract Rats (Trejo González et al., 1996)
Anti−inflammatory actions
O. humifusa extracts O. ficus−indica indicaxanthin Nitric oxide−producing macrophage cells (Cho et al., 2006); rat
pleurisy (Allegraetet al., 2014)
Healing properties
O. ficus−indica cladodes Human (Hegwood, 1990)
Neuroprotection
O. ficus−indica var. Saboten extract Primary cultured cortical cells (Dok Go et al., 2003)
O. ficus−indica fruit extracts In vitro studies in cultured mouse cortical cells e in vivo studies
in gerbils (Kim et al., 2006)
Antiulcerogenic effects and antigastritis
O. ficus−indica cladodes Rats (Galati et al., 2001, 2002)
O. ficus−indica fruit juice Rats (Galati et al., 2003a)
O. ficus−indica var. Saboten stems Rats (Lee et al., 2002a)
O. ficus−indica fruit Rats (Lee et al., 2001)
Decreasing effect on the oxidative stress in humans
O. ficus−indica fruits Humans (Tesorieri et al., 2004); in vitro human LDL (Tesorieri et
al., 2003) ex vivo human cells (Tesorieri et al., 2005b)
O. robusta fruits Humans (Budinsky et al., 2001)
Alcohol hangover symptoms alleviation
O. ficus−indica plant extract Humans (Wiese et al., 2004)
Protection upon nickel−induced toxicity
O. ficus−indica cladode extract Rats (Hfaiedh et al., 2008)
Protection against oxidative damage induced by zearalenone
O. ficus−indica cladode Mice (Zourgui et al., 2008)
158 Crop ecology, cultivation and uses of cactus pear

(continued)

Cactus species, parts and activities Studied system and reference

Diuretic effect
O. ficus−indica cladodes, flowers and Rats (Galati et al., 2002)
non−commerciable fruits
DNA damage reduction
O. ficus−indica fruit extract Human peripheral lymphocytes (Siriwardhana et al., 2006)
Cancer preventive properties
O. ficus−indica fruits aqueous extracts Ovarian and cervical epithelial cells, as well as ovarian, cervical,
and bladder cancer cells (Zou et al., 2005), ovarian cancer cells
(Feugang et al., 2010); leukaemia cell lines (Sreekanth et al.,
2007)
O. humifusa fruit extracts Breast cancer and glioblastoma human cell lines (Hahm et al.,
2010; Harlev et al., 2013)
Opuntia spp. fruit juice Prostate, colon, mammary and hepatic cancer cells (Chavez
Santoscoy et al., 2009)
Hylocereus spp. extracts In vitro antiproliferative action (Kim et al., 2011; Wu et al.; Jaya-
kumar and Kanthimathi, 2011)
O. humifusa cactus fruit powder Decreased number of papillomas and epidermal hyperplasia in
mice (Lee et al., 2012)
Liver protection
O. ficus−indica fruit juice and extract Liver (Galati et al., 2005; Alimi et al., 2012; Brahmi et al., 2011;
Ncibi et al. 2008)
Anticlastogenic capacity
Cactus fruit juice Mice (Madrigal Santillán et al., 2013)
Bone density increase
O. humifusa freeze−dried cladode Rats (Kang et al., 2012)
Insulin sensitivity improvement
O. humifusa freeze−dried cladode Rats (Kang et al., 2013)

PERSPECTIVES There is scope to further exploit the functional proper-

ties of cactus products in the food, cosmetic and phar-
Cacti can be considered an important source of bioac- maceutical industries, but more scientific research is
tive substances and excellent candidates for nutraceu- required in these fields. Although the progress made to
tical and functional food preparation. Scientific data date is significant, much remains to be explored. Cactus
reveal a high content of some chemical constituents in cladodes have health−promoting properties and they
fruits, cladodes, seeds and flowers, which can add val- are dehydrated or powdered to prepare nopal pills and
ue to cactus products. Additionally, some constituents capsules. Seed oil, on the other hand, is used in cosmet-
show promising characteristics as health−promoting ics. The global growing demand for nutraceuticals and
substances. Several manufactured products are current- healthy products is accompanied by the development
ly available in the nutraceutical market benefiting from of natural products for the treatment and prevention of
the medicinal properties of cactus plants. Fruits are pro- human diseases.
cessed to prepare confectionery, syrups, spreads, jams More study of cactus species is required to find new ac-
and jellies. Fruit juice can be used to cure hangovers. tive compounds and their pharmaceutical and industrial
Natural cactus pear juice is promoted as a healthy thirst applications. Moreover, antioxidant formulations must
quencher rich in vitamin C, flavonoids and antioxidants, be tested in the search for possible synergistic effects
and as an anti−ageing and anti−inflammatory agent. between components. Market demand should be stim-
It is also considered to promote optimal cellular health ulated by publicizing these cactus properties. Although
and detoxify the body. the beneficial properties of cactus have been known
since ancient times, only recently have they been scien-
tifically proven.
 14
Cactus ecosystem
goods and services
Mounir Louhaichia, Ali Nefzaouib and Juan Carlos Guevarac
a
International Center for Agricultural Research in the Dry Areas, Amman, Jordan
b
International Center for Agricultural Research in the Dry Areas, Tunis, Tunisia
c
Argentinean Institute for Arid Land Research, Mendoza, Argentina
14 Cactus ecosystem
goods and services

INTRODUCTION plus the wide range of economic uses, makes them

ideal for agricultural development in areas affected
The Cactaceae family includes about 1 600 species by the world’s two biggest environmental problems:
native to America but disseminated worldwide. Opun- desertification and climate change (Nefzaoui and El
tia is the most widely known genus in this family and Mourid, 2008).
O. ficus−indica (L.) Mill. is cultivated in more than 20 “Humble”, “aggressive”, “gold green”, “green jewel”,
countries (Nefzaoui et al., 2014). Cacti are cultivated “fruit of the poor”, “fruit of thorns and delights”,
on 2.6 million ha across the world, and mostly used “priceless treasure”, “treasure under the thorns”,
for forage or fodder: in Tunisia (600 000 ha), Mexico “dromedary of the vegetal world”, “plant of the fu-
(230 000 ha) and Algeria (150 000 ha) (Nefzaoui and ture” and “monstrous tree” are just some of the many
Ben Salem, 2006); in South Africa (525 000 ha) and epithets used to describe the plant and fruit of cactus
Ethiopia (355 000 ha) (Reveles Hernández et al., 2010); pear (Opuntia spp.). They reflect what “tuna” means
in Brazil (> 600 000 ha) (Torres Sales, 2010); and in to those who work or live with a plant appreciated and
southern Morocco (90 000 ha) (Anegay and Boutoba, loved by many, but feared and hated by others. In any
2010). In Argentina, the cultivated area of cactus is case, this “humble” plant continues quietly but firmly,
estimated at 10 000 ha for forage and fruit produc- to earn a leading place in programmes aimed at the
tion, with syrup as a secondary product (Dubeux et al., agricultural development of arid and semi−arid areas
2013); the area cultivated with Opuntia solely for fruit in many countries.
production was 2 000 ha in 2003 (Ochoa, 2006).
Cacti, endemic to America, were incorporated into
Cold winter temperatures are the principal limitation to many Native American cultures. For example, the
cactus cultivation in parts of Argentina and northern nopal (Opuntia spp.) is one of the most important re-
Mexico (Borrego Escalante et al., 1990), the Mediter- sources in arid and semi−arid areas of Mexico, where
ranean Basin (Le Houérou, 1996a), the arid highland it has influenced culture, history and traditions. In the
steppes of western Asia (Le Houérou, 1996b) and the social, economic and religious life of the Aztecs, the
southwestern United States of America (Parish and
plant played a very important role and the symbol of
Felker, 1997). Under a range of climatic conditions,
Great Tenochtitlan (now Mexico City) was an eagle on
the thermal limit for frost−sensitive species such as
a cactus devouring a snake. This symbol is now incor-
O. ficus−indica is indicated by a mean daily minimum
porated in the Mexican national emblem. Opuntia is
temperature in the coldest month of 1.5−2.0 °C (Le
also linked with feelings of national unity in Mexico
Houérou, 1995). Cactus and other drought−tolerant
and is frequently associated with the Virgin of Guada-
and water−efficient fodder shrubs can survive with as
lupe and the Indian Saint Juan Diego.
little as 50 mm rainfall in a given year, but with nei-
ther growth nor production. Mean annual rainfall of Tuna is now part of the natural environment, and
100−150 mm is the minimum requirement for the suc- has been integrated into the culture of many coun-
cessful establishment of rainfed cactus plantations (Le tries where it is well adapted to zones characterized
Houérou, 1994), provided soils are sandy and deep (Le by drought, erratic rainfall and poor soils exposed to
Houérou, 1996a). Plantations of drought−tolerant and erosion. It is naturalized in the landscape as part of
water−efficient fodder shrubs, especially Opuntia spe- the local flora, as can be seen in postcards and tourist
cies, have been established as buffer feed reserves as advertisements in Italy, Spain, Morocco, Israel, Kenya,
a strategy to mitigate the effects of drought in animal Yemen and Saudi Arabia. Notably, Lorenzo Bernini
production systems in various arid and semi−arid are- (1598−1680) included the cactus in the Fountain of the
as of the world (Le Houérou, 1991). Cacti have good Four Rivers in Piazza Navona in Rome where the Río de
water−use efficiency, thanks to the crassulacean acid la Plata is represented.
metabolism (CAM) photosynthetic pathway (Han and
The controversy over whether it is a helpful or harmful
Felker, 1997; Nobel, 1991, 1994); for this reason, they
plant depends on the species, where, when and how it
are especially suited for forage production in arid land.
is grown, and to whom it applies. There are contradic-
Opuntia spp. can withstand prolonged drought, high tory assessments, true in one case and not in another,
temperatures, and wind and water erosion. This ability, based on experiences under different ecological, eco-
Cactus ecosystem goods and services 161

nomic and social conditions. Different situations evolve landscape and nature conservation, mitigation of soil
depending on the individual reality. For example, in erosion, water protection and cultural heritage. How-
Australia and South Africa, biological control was used ever, these public services do not have a market price,
to stop its expansion and even exterminate it in cer- are difficult to disaggregate, are highly interrelated in
tain areas. In Ethiopia and Eritrea, on the other hand, complex dynamic ways and are difficult to measure.
where the climate is suitable and no natural enemies The strong links between cactus pear production and
exist, nopal effectively invaded thousands of hectares the provision of diverse ecosystem goods and services,
after it was introduced > 150 years ago. In any case, especially in marginal areas, need to be considered and
while it may affect local plant genetic resources, the integrated into a standard evaluation framework for
current reality is that after so many years, people have environmental impacts of agricultural production. The
an economic dependency on cactus food and prod- main objective of this chapter is to highlight the
ucts, regardless of whether the introduction of nopal benefits generated from cactus pear.
was a curse or a blessing. Its potential for adaptability
and rapid expansion in wilderness areas or in areas
previously disturbed by human intervention is anoth- RANGELAND IMPROVEMENT
er matter requiring examination. Cactus pear is more
than useful: it is a vital plant that has been called “a Rangeland improvement using spineless cactus has
crop that saves lives of humans and animals”, especial- been practised since the early 1930−1940s, principal-
ly in times of severe drought (Arias Jimenez, 2013b). ly in North Africa. Le Houérou (2002) reported that
There are numerous reasons behind the diffusion of fodder plantations were systematically developed,
Opuntia spp. around the world, particularly of O. ficus− especially in Tunisia, following the research conducted
indica, including: by Griffiths and collaborators in Texas, United States of
America. Griffiths was invited by the Government of
• simple cultivation practices required to grow the crop;
Tunisia in 1932 and his 30 years’ experience with the
• rapid establishment soon after introduction in a new use of cactus as fodder has since been applied in central
area; Tunisia to mitigate the effects of drought on livestock.
• easy multiplication practices that favour rapid diffu- The development of cactus for fodder was strongly
sion and exchange of material among users; supported by the Government. Conditional land allot-
• ability to grow in very harsh conditions characterized ments were authorized in central Tunisia on condition
by high temperature, lack of water and poor soil; that the contracting beneficiaries planted 10% of the
• generation of income from the sale of much−valued allocated land with spineless cactus. This was to serve
and appreciated fruits; as an emergency standing fodder crop reserve, which
would stand as a buffer in times of fodder shortages.
• use of stems in the human diet and as forage for live-
This was a strategic move as the country faced 3 years
stock;
of severe droughts from 1946−1948, during which
• useful deployment of plants for fencing farms; livestock were decimated by 70−75%. Livestock losses
• nutritional value of juicy fruits; were lower for those who had cactus plantations (Le
• long shelf−life of fruit; and Houérou, 2002).
• production of a wide range of industrial derivatives The plantation of cactus is applicable in many settings,
from fruit. particularly where the environment is too limited or
challenging for traditional agricultural crops or where
These and other factors have contributed to such a
the land is in need of rehabilitation. Cactus is recom-
wide distribution, from the regions of origin in Latin
mended wherever soils are too shallow, too stony, too
America to remote areas, spanning continents, cultures
steep or too sandy, or when the climate is too dry for
and traditions.
practical farming. As a result, they have a key role in re-
habilitation strategies to improve rangeland, shrubland,
Main objective of this chapter bushland or poor farming areas. For rehabilitation, the
planting density is 1 000−2 000 single or double pads
Despite its ecological, economic and social importance, per ha, with a spacing of 5−7 m between rows and 1–2
Opuntia (cactus pear) continues to receive limited sci- m within rows. Fertilization, pruning and pest/disease
entific, political and media attention. Well−maintained treatment are not generally applied, but they can be in
cactus plantations generate positive externalities and order to improve productivity. Occasionally, if the first
environmental goods and services: they can play a year is too dry, supplemental irrigation is applied during
major role, not only in terms of biodiversity enhance- establishment. Plantations are exploitable after 3−4
ment and carbon sequestration, but with regard to years and fully grown after 7−10 years; they can remain
162 Crop ecology, cultivation and uses of cactus pear

productive for > 50 years when managed well. Since (Winckler, 2002). Water and soil are the most precious
Opuntia can survive with minimal management, it is renewable natural resources. Drought avoidance and
recommended in rehabilitation programmes. coping strategies are imperative: for example, choose
drought−tolerant crops, maintain low plant densities,
Intensive management is not a prerequisite for cactus
and apply water conservation and water harvesting.
survival, but a plantation can reach high productivity
However, only a small fraction of rainfall becomes us-
levels if appropriation practices are applied. The pro-
able soil moisture: 1−10% of the rain that falls in the
ductivity of rangelands planted with cactus can be
drylands ends up in the tissue of natural vegetation
increased by a factor of 1 to 10 when rangelands are
and crops of economic significance (El Beltagy, 1999).
very degraded and by 1 to 5 when rangelands are in
Water erosion is accelerated by tillage on slopes and
good condition (Le Houérou et al., 1991). Nefzaoui and
gully margins. Soil productivity is rapidly declining.
El Mourid (2009) compared the productivity of range-
lands in central Tunisia when they were rehabilitated Soils of arid and semi−arid zones are very susceptible
with cactus (Opuntia ficus−indica) and fast−growing to water erosion (Cornelis, 2006), mostly due to the
shrubs (Acacia cyanophylla). Rehabilitation with cactus scarce vegetation cover, low organic matter content
(O. ficus−indica) yielded higher productivity rates than and poor resistance to erosion forces. The magnitude
rehabilitation with fast−growing shrubs (A. cyanophyl- of water erosion also depends on texture, water con-
la) (Table 1). tent, evaporation, percolation and leaching. These soil
Few plant species are able to increase land productivity characteristics are not favourable to the resistance
at the high rate mentioned above, particularly in the of soil to water erosion (D'Odorico and Porporato,
case of marginal lands. Cacti can − because of their 2006). In arid and semi−arid areas, soils with little or
rain−use efficiency (RUE). Indeed, degraded Mediterra- no vegetation cover are exposed to torrential precipi-
nean rangelands have a RUE of 1−3 kg DM ha−1 year−1 tation events, characterized by short duration and high
mm−1, rangelands in good condition exhibit a RUE of intensity, and are prone to physical and chemical pro-
4−6, and degraded rangelands may have a RUE as low cesses that change the surface layer conditions, such
as 0.1–0.5 (Le Houérou, 1984). In contrast, rangelands as surface sealing and crusting. When the surface is
rehabilitated with Opuntia ficus−indica exhibit a RUE dry, a hard layer is formed (crust). Crusting soils are
of 10−20 kg of above−ground DM ha−1 year−1 mm−1 in typical of dry areas, where soil degradation is induced
arid areas with annual rainfall of 200−400 mm. by diminishing infiltration rates and increasing runoff
and erosion rates (Ries and Hirt, 2008). Arid and semi−
arid areas are fragile environments where vegetation
cover is scarce and soil erosion processes occur rapidly
ALLEVIATION OF SOIL EROSION and severely after rainfall events. However, even under
such conditions, native vegetation has a very important
Land degradation occurs in all continents and affects
role in the regulation of surface hydrological processes
the livelihoods of millions of people, including a large
(Vásquez Méndez et al., 2011).
proportion of the poor in the drylands (Nefzaoui et
al., 2014). Dry zones with annual moisture deficits of Erosion control is another important use of cactus
> 50% cover approximately 40% of the earth’s land pear (Opuntia spp.), as it grows quickly and has small
surface. More than 70% of all dry areas suffer from roots that regrow each year from the main root during
desertification, currently accounting for 36 million km² times of rain. In dry periods, the small roots die, add-

TABLE 1 Productivity (forage units per hectare) of natural and improved rangelands in Tunisia (Nefzaoui and El
Mourid, 2009)

Rangeland type Productivity (forage unit per hectare)a

Natural rangeland in Dhahar Tataouine, Tunisia
(100 mm rainfall) 35−100

Private rangeland improved by cactus crop in Ouled Farhane, Tunisia

(250 mm rainfall) 00−1 000

Cooperative rangeland improved through Acacia cyanophylla, Guettis,

Tunisia (200 mm rainfall) 400−500

a
One forage unit is equivalent to 1 kg barley grain metabolizable energy.
Cactus ecosystem goods and services 163

ing organic matter to the soil. With an increased

organic matter content, it is easier for the soil
to absorb rainwater. Opuntia spp. are utilized in
programmes to prevent soil erosion and combat
desertification; they are very adaptable, growing
in severely degraded soil which is inadequate
for other crops, and are ideal for responding to
increases in atmospheric carbon dioxide (CO2)
levels. Opuntia is also important as cover in arid
and semi−arid areas, because it can survive and
spread under conditions of scarce and erratic rain-
fall and high temperatures (Reynolds and Arias,
2001).
Tests involving the planting of cactus for water
harvesting strips, contour ridges, gully check
structures and biological control of rills and small
gullies have had good results. Contour ridges
consist of parallel stone ridges built 5−10 m apart
to stop runoff water (and the soil it carries) from
damaging downstream areas. Each ridge collects
runoff from the area immediately upstream/ ed at low cost in Tunisia and Algeria by contour Figure 1
uphill, which is channelled to a small plantation planting of cacti (Nefzaoui et al., 2011). Management in Tunisia
for the protection of
of fodder shrubs or cactus. With a combination In a comparison of different cultivation systems, watershed
of well−designed ridges and cactus, farmers can such as downhill planting, contour planting, re-
meet a large proportion of their fodder require- duced weeding, and intercropping with contour
ments. In North Africa, particularly Tunisia, cactus hedges, it was found that soil losses (0.13−0.26
is successfully associated with water−harvesting tonnes ha−1 year−1) are lowest with contour hedg-
structures. Planted according to contour lines, es. Cactus planting in contour hedges may help
cactus hedges play a major role in erosion control retain ≤ 100 tonnes of soil ha−1 year−1. Experiments
(Figure 1). Soil physical properties and organic conducted in Brazil and Tunisia show clearly that
matter content are considerably improved under planting cactus in an agroforestry system is more
these hedges and in the immediate adjacent efficient for soil and water conservation than con-
areas, with an improvement in organic matter ventional land use (Table 2) (Margolis et al., 1985).
and nitrogen compared with non−treated fields.
There have been reports of rates of 40–200% The eruption of massive dust storms in the Sahara
increase in organic matter and nitrogen. Topsoil can move 66–220 million tonnes of fine sediment
structural stability is enhanced, sensitivity to sur- each year. Wind erosion is a major cause of soil
face crusting, runoff and erosion are reduced, degradation on agricultural land in arid and semi−
and permeability and water storage capability are arid areas throughout the world. Wind damages
increased. Marginal lands have been rehabilitat- soil by removing the lighter, more fertile and less

TABLE 2 Comparison of soil loss (tonnes ha−1 year−1) under different crops in semi−arid northeast
Brazil (Margolis et al., 1985)

Harvest until
Soil prepara- Cultivation Total soil
Crop type next growing C factor
tion phase phase losses
season
Bare soil 7.19 8.2 13.71 29.1 1
Cotton 2.42 1.77 6.72 10.91 0.392
Maize 1.51 0.68 3.75 5.94 0.199
Maize + beans 1.36 0.55 2.02 3.93 0.119
Opuntia ficus−indica 0.48 0.02 1.48 1.98 0.072
Perennial grass 0 0.02 0.01 0.03 0.001
164 Crop ecology, cultivation and uses of cactus pear

dense soil components, such as organic matter, clay activities with a sound research initiative to investigate
and silt. Reduced soil productivity is not the only ag- all possible benefits and the efficiency of new technol-
ricultural impact of wind erosion. Blowing sediment ogies using cacti as a keystone species to help control
cuts and abrades plants, reduces seedling survival and desertification and adapt to global warming (Nefzaoui
growth, lowers crop yields, and can increase suscep- and El Mourid, 2010).
tibility to diseases and the spread of plant pathogens
(Northcutt, 2001). In arid lands subject to wind erosion,
cactus planted alone as a biological barrier or together BIOLOGICAL FENCING/VEGETATIVE
with physical barriers is an easy, cheap and efficient BARRIERS
way to prevent and control topsoil loss, and it facilitates
the accumulation of wind−borne deposits.
Cactus pear can be grown as hedges and fences by
The somewhat scattered results obtained to date are planting them around 30 cm apart. Within several
testimony to the lack of research in this domain. Devel- years, the plants grow together to form a wall of spiny
opment actions are mainly based on assumptions and pads protruding at all angles. Plantings can also be
observations gathered by practitioners (Nefzaoui and El established for erosion control in deforested areas. In
Mourid, 2010). time, cacti such as O. ficus−indica may grow into freely
branching plants 3–6 m high (University of California
Many Cactaceae species live in arid environments and
Cooperative Extension, 1989). The use of various spe-
are extremely drought−tolerant. One survival technique
cies of cactus for fencing has great benefits which reach
of O. microdasys (Lehm.) Pfeiff., which originates in the
well beyond the savings made by not having to use
Chihuahua Desert, lies in its unique and efficient fog
expensive resources such as iron. Once created, a living
collection system, attributed to the integration of its
fence provides fruit and excellent security for crops and
multilevel surface structures (Ju et al., 2012; Bai et al.,
homes, while providing home or habitat for wild fauna
2015). This system comprises well−distributed clusters
species. Deer and its main predators can jump over it or
of conical spines and trichomes on the cactus stem;
crawl through natural tunnels. Also, if given reasonable
each spine contains three integrated parts, each with
amounts of water, cactus can grow quickly and – since
a different role in fog collection depending on their
most farms and homes have water – a fence can be
surface structural features. The gradient of Laplace
established in as little as 1 year. The closer pads are
pressure, the gradient of surface−free energy and mul-
planted, the quicker the fence fills in, but a difficult
tifunction integration provide efficient fog collection.
pruning job later becomes necessary. Once established,
There is also evidence that some cactus species can
a cactus border provides security, beautiful flowers
harvest dew on their stems and spines (Malik et al.,
(bees and other insects are attracted to them) and fruit.
2015).
Each variety should be sampled for ease of handling
Opuntia can tolerate drought in open spaces by in- and flavour, as some are too seedy. A spiny variety of
creasing and moving chloroplasts and avoiding drastic standard cactus pear Opuntia makes a good fence; and
decreases in their osmotic potential (Delgado Sánchez spineless varieties will not keep humans away if the
et al., 2013). O. ellisiana Griffiths is a CAM plant, and its lack of spines can be seen from a distance. The bluish
conversion efficiency of water to dry matter is several O. violacea Engelm var. santa−rita (Griffiths & Hare) L.
times greater than either C3 or C4 plants. A significant D. Benson makes an excellent fence – the fruit is beet
quantity of water (17 mm = 170 000 kg ha –1) can be red, packed with seeds and has almost no edible pulp;
stored in this cactus and used for animal drinking water it also has thick spines clustered on the fruit. To make a
(Han and Felker, 1997). Cactus can take advantage of fence, cut the thickest pads from a plant, allow the scar
the lightest rainfall, because its roots are close to the to heal for about a week and then plant the pads at a
soil surface. The water is quickly collected by the roots 1.3 m spacing. However, if there is little time available,
and stored in thick, expandable stems. The fleshy stems place the pads flat on the ground on the same day and
of the barrel cactus (Ferocactus wislizeni Britton & Rose) leave for a year or two; they form a cup−shape and
are pleated like an accordion and they shrink as mois- retain rainwater. If this latter method is adopted, very
ture is used up. The green stems produce the plant’s thick or woody pads are best (White Dove Farm, 2015).
food, but they lose less water than leaves do, thanks to
Thorny cacti O. ficus−indica var. amyclaea (Ten.) A.
their sunken pores and a waxy coating on the surface
Berger and var. elongata Shelle are often used as de-
of the stem. The pores close during the day and open
fensive hedges for the protection of gardens, orchards
at night and, therefore, only release a small amount of
and olive groves throughout North Africa and in parts
moisture. The price paid by cactus for these water−sav-
of Italy and Spain. These hedges demarcate boundaries
ing adaptations is slow growth (Zemon, 2015).
while helping to control erosion. However, in regions
There is an urgent need to enhance ongoing research where winters are temperate to mild (mean daily min-
Cactus ecosystem goods and services 165

imum temperature in January > 3° C), the fruits of al., 2002; Root et al., 2003). Research has focused on
Opuntia hedges may host the fruit fly, Ceratitis capitata evaluating the potential of biological CO2 sequestration
L. Fruit−bearing cacti may, therefore, need to be treat- for various types of plants. In comparison with C3 and
ed against fruit fly or eliminated from other fruit crop C4 plants, CAM plants (agaves and cacti) can use water
areas. In the case of fodder cacti, however, if harvested much more efficiently with regard to CO2 uptake and
every 2−3 years, they do not produce fruit and cannot, productivity (Nobel, 2009). Biomass generation per unit
therefore, host fruit fly. Not only do these hedges have of water is on an average 5−10 times greater than in C4
a very efficient defensive role (particularly when estab- and C3 plants (Table 3). The potential of CAM systems
lished in double rows), they also play an important part to accumulate high biomass depends on their capacity
in landscape organization and the local socio−economy, to partition more carbohydrates to growth than to noc-
defining land rights and land ownership in countries or turnal acid metabolism (Borland and Dodd, 2002).
regions where no land registry exists. Cactus hedges are
The role of cactus plantations in the carbon cycle is
often planted as testimony of land ownership, because
extremely important. They help complete the cycle of
in some countries, tradition dictates that tribal land may
life by recycling building block nutrients to the plants
become the property of whoever among the rightful
and carbon (CO2) to the atmosphere (Doran, 2002).
users has established a permanent crop on it. This is a
This is important in the soil decomposition process and
strong motivation for planting cactus hedges (and olive
is often mediated by organisms in the soil. Various ex-
groves) on communal lands, and explains their popular-
periments in different regions have been carried out to
ity in countries such as Tunisia. Cactus hedges also play
quantify the carbon sequestration potential of Opun-
a major role in erosion control and land−slope partition-
tia. Measurements of gas exchange in O. ficus−indica
ing, particularly when established along contours. Soil began in the early 1980s, when Nobel and Hartsock
physical properties and organic matter content are con- (1983) measured CO2 uptake on single cladodes, using
siderably improved under hedges and in the immediately portable infrared gas analysers with cuvettes adapted
adjacent areas (Monjauze and Le Houérou, 1965b). The to fit cladode morphology. At optimal temperature and
aggregates in the topsoil become more stable and less intercepted radiation, instantaneous values of net CO2
sensitive to surface crusting, runoff and erosion; perme- uptake of 1−year cladodes may reach 18 mmol m –2 s–1,
ability and water storage capacity increase. Moreover, with a total daily CO2 uptake of 680 mmol m –2 (Nobel
hedges are a physical obstacle to runoff: they help accu- and Bobich, 2002). In a similar study − evaluating the
mulate temporary local runoff and silting and prevent re- effects of seasonal variations in temperature, irradiation
gressive erosion. Some badlands, developed in outcrops and soil moisture content on the photosynthetic rates
of shale and stony/rocky slopes, have been rehabilitated of O. ficus−indica − the total daily net CO2 uptake was
at low cost in Tunisia and Algeria with contour planting 393 mmol m –2 averaged over five measurement dates,
of cacti. In arid lands subject to wind erosion, cactus and annual CO2 uptake was 144 mol m –2 (Pimienta Bar-
hedges are an easy, cheap and efficient way to prevent rios et al., 2000).
and control topsoil loss and aid accumulation of wind−
borne deposits (Le Houérou, 1996a). Opuntia has greater water−use efficiency than C4 or
C3 plants due to the CAM photosynthetic pathway,
The costs of establishing a fence depends on the mate- which is more efficient in converting water and CO2 to
rial used. A metallic fence (four strands of barbed wire) plant dry matter (Nobel, 1991, 1994; Han and Felker,
costs about US$1 m−1, i.e. US$150 ha−1 in Tunisia for 1997; Nobel, 2009). As stated by Nobel (2009), the
plantations of about 10 ha. A fence made of a double consequences of nocturnal gas exchange depend on
row of spiny cactus costs less than US$60 ha−1, but temperature. Temperatures are lower at night, which
needs to be established for at least 2 years before it reduces the internal water vapour concentrations in
begins to function (Le Houérou, 1989). CAM plants, and results in better water−use efficiency.
This is the key reason for which CAM species are the
most suited plants for arid and semi−arid habitats. The
importance of nocturnal opening and diurnal closure
CARBON SEQUESTRATION POTENTIAL of stomata in CAM species for water conservation has
long been recognized (Black and Osmond, 2003). In
Over the last four decades, it has become evident that addition to the advantages of being a CAM species,
rising atmospheric CO2 from fossil fuel consumption Opuntia plants are also known for their ability to regen-
is causing increased climate variability. This leads to erate and grow easily. They act as carbon sinks and can
important issues associated with global warming and be grown on a large scale in areas where precipitation
modified continental patterns of precipitation that are is inadequate or unreliable. They can grow where evap-
already having significant effects on species distribu- oration is so great that rainfall is ineffective for crop
tion and function in the plant biosphere (Walther et growth (Osmond et al., 2008). C3 and C4 plants suffer
166 Crop ecology, cultivation and uses of cactus pear

irreparable damage once they lose 30% of their wa- species (Rascher et al., 2001; Nelson et al., 2005; Grif-
ter content, while many cacti can survive an 80−90% fiths et al., 2007). Wang and Nobel (1996) found that
loss of their hydrated water content and still survive. the growth of O. ficus−indica in elevated CO2 for 3
This is due to the ability of CAM plants to store large months showed little evidence for downregulation of
quantities of water; to shift water around among cells photosynthesis commonly found in herbaceous plants.
and keep crucial metabolism active; and to tolerate ex- Herbaceous plants usually experience sink limitations
treme cellular dehydration (Nobel, 2009). These abili- and feedback effects of sugar on CO2 assimilation and
ties, in turn, stem from the cactus characteristics: extra gene expression. However, in O. ficus−indica, higher
thickness of the cuticles providing an efficient barrier CO2 assimilation (source capacity) was found with
for water loss; presence of mucilage; and daytime greater sucrose transport in the phloem and stronger
stomatal closing. In addition, cacti are characterized by sink strength. Drennan and Nobel (2000) concluded
asynchronous development of various plant organs, so that high−biomass CAM communities offer potential
that even under the worst conditions, some part of the as a low−input system for atmospheric CO2 seques-
plant remains unaffected. It is well known that cacti tration in arid habitats. Although more research is
grow in the desert where temperatures are extreme- needed, long−lived CAM plants in arid ecosystems
ly high. Many authors (e.g. Nobel, 2009) report that may present effective regional carbon sequestration
many agaves and cacti can tolerate high temperatures systems on time scales of decades to centuries.
of 60−70° C.
Limited data are available on cladode net CO2 uptake
This aspect is covered in detail by Nobel (2009). In view according to cladode age. Liguori et al. (2013a) used an
of the specific phenological, physiological and struc- open gas exchange chamber to measure whole plant
tural adaptations of cacti described above, they may be or single organ net CO2 uptake in cactus pear, particu-
considered well positioned to cope with future global larly to understand the response of the whole plant to
climate change. Opuntia ficus−indica, for example, can environmental stress. Unlike with single cladode meas-
generate a carbon sequestration of 20 tonnes of dry urements, after 60 days of drought the whole plant
matter (equivalent to 30 tonnes of CO2) per ha and per maintained the same level of net CO2 uptake, although
year under suboptimal growing conditions similar to there was substantial water loss in the parenchyma of
those in the arid regions of North Africa. the most photosynthetically active cladodes. Future
One Opuntia species is known to occupy open and research on individual cladode CO2 uptake is required
abandoned farmland and to invade open scrubland to understand the best pruning practices needed to
and forest. This occurs especially in the unpredictable increase CO2 uptake, particularly for cultivated fields of
but frequently wet habitats of central eastern Australia O. ficus−indica.
that have non−effective rainfall for the growth of agri-
cultural crops (Leeper, 1960; Osmond et al., 1979). The
plant has succeeded in its adopted habitat for multiple ALLEY CROPPING
reasons: in part, because it is a CAM plant with ex-
ceptional water−conserving potential; in part, because Expansion of cereal cropping into rangelands, com-
its extraordinarily low root−to−shoot ratio, dominated bined with not allowing lands to go fallow, is one of
by above−ground cladode biomass, can focus on the major reasons for declining soil fertility and wind
photosynthetic activity; but most of all, because it is erosion. One way of combating degradation result-
characterized by extraordinary vegetative and sexual ing from cereal monocropping is the introduction of
reproductive activities. adapted forage legumes, shrubs/fodder trees and cac-
Increased atmospheric CO2 stimulates further growth tus in the cropping system (Nefzaoui et al., 2011). Alley
and carbon sequestration of O. ficus−indica (Gomez cropping is an agroforestry practice where perennial
Casanovas et al., 2007). Drennan and Nobel (2000) crops are grown simultaneously with an arable crop.
reported that doubling atmospheric CO2 stimulated Shrubs, trees or cactus are grown in wide rows and
the total CO2 uptake by an average of 31% for six large the crop is grown in the interspace. Alley cropping is
cacti, and stimulated growth and biomass by 33%. a form of hedgerow intercropping. Leguminous and
These responses were unexpected because elevated fast−growing tree/shrub species are preferred because
CO2 was not expected to stimulate CO2 assimilation of their soil−improving attributes, i.e. their capacity to
in the presence of closed stomata in the light; further, recycle nutrients, suppress weeds and control erosion
it was expected that CO2 assimilation in the dark by on sloping land. This technology enables the farmer
phosphoenolpyruvate carboxylase would be saturated to continue cultivating the land while the trees/shrubs
at internal CO2. However, these responses are in good planted in intermittent rows help maintain the quali-
agreement with what is now known of the diffusion ty of the soil. Cactus can function in this system as a
limitations of CO2 fixation in all growth states of CAM windbreak, resulting in improved grass/cereal yields.
Cactus ecosystem goods and services 167

The wide alleys allow animals to graze biomass

strata or cereal stubble during summer; cactus
pads can be harvested, chopped and given direct-
ly to grazing animals as an energy supplement to
low−quality stubbles (Nefzaoui et al., 2011).
Although cacti are well known as the best plants
for reforestation of arid and semi−arid areas
because of their resistance to scarce and erratic
rainfall and high temperatures, alley−cropping
systems in Tunisia are a largely new phenomenon.
When properly managed, alley cropping can pro-
vide income at different time intervals for differ-
ent markets in a sustainable, conservation−orien-
tated manner. Alley designs can also optimize the
space available between trees, adding protection
and diversity to agricultural fields.
The practice of planting only shrubs is not widely
adopted for various reasons, including the tech-
nical design of the plantation, mismanagement, creates a beneficial “windbreak”, reducing water Figure 2
and competition for land often dedicated to ce- loss and increasing soil moisture. The barley crop Alley cropping using
real crops. Some of these disadvantages can be stimulated an increase in the number of cactus Opuntia ficus−indica
overcome by alley cropping, which: and barley crop
cladodes and fruits, while the cactus increased
• improves soil; the amount of root material contributing to the
• increases crop yield; soil organic matter.
• reduces weeds; and
• improves animal performance.
BIODIVERSITY CONSERVATION
Properly managed alley cropping allows diversi-
fication and the farmer can benefit from several
The intensification of agricultural practices in a
markets. It also promotes sustainability in both
context of climate change is cause for concern,
crop and livestock production. The benefits of
as it dramatically alters soil characteristics and
cactus−barley alley cropping were evaluated in
Tunisia (Alary et al., 2007; Shideed et al., 2007). affects the local flora and fauna communities
Compared with barley alone, the total biomass (Ouled Belgacem and Louhaichi, 2013). These dis-
(straw plus grain) of barley cultivated between turbances affect biodiversity, the most important
rows of spineless cactus increased from 4.24 to factor affecting the stability of ecosystems and
6.65 tonnes ha−1, while the grain yield increased agro−ecosystems (Fontaine et al., 2011). Stopping
from 0.82 to 2.32 tonnes ha−1 (Table 3, Figure 2). or reversing the decline in biodiversity is a major
These figures are the result of the micro−environ- challenge for the maintenance of biodiversity and
ment created by alley cropping with cactus, which wider ecosystem services.

TABLE 3 Total biomass changes and barley crop yields (tonnes ha−1) in Sidi Bouzid (Tunisia)a (Alary et
al., 2007)

Natural Barley crop Cactus crop Alley cropping

Treatment
rangeland (alone) (alone) (cactus + barley)
Above−ground biomass (tonnes ha−1) 0.51 0.53 1.87 7.11
Underground biomass (tonnes ha−1) 0.33 0.11 1.8 1.98
Barley grain yield (tonnes ha−1) 1.51 0.82 2.32
Barley grain + straw + weeds (tonnes ha )−1
1.36 4.24 6.65

a
Average rainfall in Sidi Bouzid is 250 mm year−1. All treatments were without fertilizers.
168 Crop ecology, cultivation and uses of cactus pear

Cactus pears are prominent in many arid and semi−ar- assassin, Apiomerus crassipes Fabricius, lies in wait in
id habitats. They have an important role in the ecology cactus flowers and preys upon its bee or ant victims by
and are important for the fauna and flora sharing their injecting a paralyzing enzyme with its hypodermic−like
habitat. Padilla and Pugnaire (2006) report that some beak. The cochineal insect, Dactylopius coccus Cos-
plants benefit from closely associated neighbours, a ta, is a cryptic species (a taxon that uses anatomy or
phenomenon known as facilitation. Cacti often act behaviour to elude predators) and the females spend
as “nurse plants” in hot climates: their shade, and their entire lives in colonies on the stems of prickly
sometimes nutrients associated with their presence, pear. This insect species is noted for the secretion of
help seedlings of other species to become established, carminic acid, used by the ancient Aztecs to produce a
which they might otherwise not be able to do in hot or crimson dye. Hundreds of butterfly, moth and skipper
poor soil. Cacti can be an attractive source of shelter species are known pollinators of Cactaceae (Hogan,
for wildlife and their shade is very important for ani- 2015). The flowers contain large amounts of nectar
mals, as well as for other plant species. Cactus species and the fruits are rich in water.
provide significant nesting sites for birds, rodents and
Cacti are also important for many desert animals. Nec-
other animals. Birds perch on their branches to exam-
tar−feeding bats, Leptonycteris curasoae curasoae Mill.
ine their surroundings. Bird droppings often contain
and Glossophaga longirostris elongata Petit and Pors,
seeds of other plants and the shade of the cacti can
are valuable for the pollination of many important
provide a microclimate that promotes other plant life.
plants in Curacao, and they depend on columnar cacti
Cacti provide fruits and flowers for a range of ani- for their survival. Their diet consists mainly of cactus
mals – many species of bird, bat and insect, including nectar, pollen and fruits when available. Larger mam-
bees. Cacti survive in the natural environment without mals, such as white−tailed deer (Odocoileus virginianus
artificial watering and they produce most of their flow- Zimmerman), actually consume the pads of prickly
ers and fruits during the dry season, when very few pears, despite the formidable spiny armour (Ramawat,
other resources are available for wildlife; moreover, 2010). The cactus pads are filled with water, used by
their flowers attract butterflies and other pollinating several animals including prikichi (brown−throated
insects. Some Opuntia species produce juicy fruit in parakeet, Aratinga pertinax L.) and the white−tailed
summer, appreciated by many birds. The succulent deer. Coyotes (Canis latrans Say) feed in a semi−de-
fruits containing many seeds are particularly attractive sert grass shrub habitat and consume a variety of
to passerine birds; for this reason, it is not uncommon foodstuffs throughout the year. Many cactus species
to find plants developing to maturity in rocky outcrops, in the Sonoran Desert bear fruit during summer – an
on wooden fence posts or along wire fence lines. important food source in drier months. Coyotes feed
Prickly pear cactus offer good escape cover for birds. on cactus fruits when they are ripe, as shown by the
For instance, the cactus wren (Campylorhynchus brun- analysis of scats collected in autumn (Short, 1979). In
neicapillus Lafresnaye) is native in the southwestern Arizona, packrats, also known as the American rat, live
United States of America and southwards to central primarily in the desert beneath fallen cactus and debris
Mexico. It is found in deserts and arid foothills char- piles. They burrow under a cactus (usually cactus pear),
acterized by cactus, mesquite, yucca and other types killing the roots and causing the cactus to collapse over
of desert scrub. It nests in cactus plants – sometimes them. This creates a prickly armoured home relatively
in a hole in a saguaro and or where it is protected by safe from mammal and bird predators. Some species
the prickly leaves of a cholla or yucca. Building the nest use the base of a prickly cactus pear as the site for
in cactus provides some protection for the young, but their home, utilizing cactus spines for protection from
the wrens also use these nests throughout the year predators.
to roost. They eat mainly insects, occasionally seeds
Unfortunately, the habitat provided by cactus is not
or fruits, but they rarely drink water, getting moisture
always beneficial or agreeable to humans. As a result,
from their food.
during the summer months many ranchers burn cactus
Hundreds of species of ants use cacti for food and – to- to eliminate them from their fields. While cacti are
gether with other insects – are important cactus polli- often considered invasive weeds in much of the south-
nators. While other predators find the spines daunting, ern United States of America, the presence of many
ants forage along cactus stems, capturing small fauna Opuntia species was actually documented by the first
and feeding from extra−floral nectaries, which provide settlers to the area. Also in Mexico, the plant is often
a sugar mixture often high in amino acid content. regarded as a weed, but it is an important food source
Furthermore, ants and other insects feed on cactus in Mexican culture. Cactus pads have recently become
seeds and play a role in seed dispersal. Cacti provide a health trend and they are sold as a vegetable in
a habitat for many different types of insects. The bee many supermarkets in the United States of America. It
Cactus ecosystem goods and services 169

is hoped that its use as a food will highlight the impor- receive limited scientific and media attention and there
tance of the plant; however, many have yet to realize is insufficient focus on their conservation merits. This is
its ecological importance for biodiversity. largely due to limited knowledge and a narrow vision
focusing on cacti for forage and fruit production. A
major shift in the role of cactus and its production is re-
quired, towards a much broader concept of ecosystem
CONCLUSION AND RECOMMENDATIONS goods and services. A holistic approach is needed,
with a balance among environmental conservation,
Human societies derive many essential goods and ser- farming system production and socio−economic devel-
vices from natural ecosystems. This chapter focuses on opment. It would be of great benefit to promote the
the importance of cactus pear and its significant role ecological, economic and social benefits of cacti and
in the livelihood of farmers. The goods and services strengthen the technical capacity of human resources
provided by cactus pear include: soil and water erosion dedicated to these species. In addition to the analysis
control; regulation of climate through carbon seques- of current trends in research, it is vital to present new
tration; biodiversity conservation; habitat for wildlife; discoveries and plans for future research in all areas
pharmaceutical and industrial benefits – in addition concerning cacti. Public policies and credit are essential
to their aesthetic beauty as evergreen plants. Despite in order to increase cultivation of this important plant
their ecological, economic and social importance, cacti in the arid and semi−arid regions of the world.
 15

Global invasions
of cacti (Opuntia sp.):
control, management
and conflicts of interest
Helmuth Zimmermann
Helmuth Zimmermann and Associates, Pretoria, South Africa
15 Global invasions of cacti (Opuntia sp.):
control, management and conflicts of interest

INTRODUCTION conflicts of interest based on true or false perceptions

of cactus/prickly pear as invasive or beneficial, resulting
The value of cactus pear, Opuntia ficus−indica (L.) Mill., is in delays in the implementation of biological control
indisputable (Barbera et al., eds, 1995; Mondragón Jaco- programmes. In view of the increased importance of
bo and Pérez González, 2001; Nobel, ed., 2002; Sáenz et commercially cultivated Opuntiae, permission for the
al., eds, 2006) and there have been considerable efforts release of the relevant biological control agents would
to introduce this emerging crop to several developing not have been granted today, with direct consequences
countries with some success (De Waal et al., 2015). The for future projects on biological control of new emerg-
value of cactus pear as a source of fodder has almost ing species in both subfamilies.
surpassed its value as a fruit crop. Nevertheless, several This chapter provides a historical overview of cactus in-
countries – including developing countries in sub−Saha- vaders in the three subfamilies Opuntioideae, Cactoideae
ran Africa – have yet to seriously adopt this new crop and Pereskioideae − according to the taxonomy of Ander-
plant, for example, Botswana, Namibia, Angola, Zimba- son (2001) and Hunt (2006) − and the control of these
bwe, Mozambique, the United Republic of Tanzania and invaders, focusing on new approaches to overcome the
Zambia. Cactus pear remains an underutilized, emerging increasing problem of conflicts of interest. Cactus densifi-
crop with many potential uses. Other recently commer- ers, which have become weeds within their native ranges
cialized cultivated cacti are also gaining in popularity, for in the Americas, are not included in these discussions.
example: Nopalea cochenillifera (L.) S−D, O. undulata
Griffiths (Brazil) (Cordeiro dos Santos and Gonzaga de
Albuquerque, 2001), O. monacantha Haw. (Kaufmann,
CACTI AS A PROBLEM
2001), Hylocereus undatus (Haworth) Britt. & Rose and
Cereus hildmannianus KSch (= C. peruvianus) (Israel)
Despite the limited information available, an attempt
(Nerd et al., 2002). Unfortunately, commercially valuable
has been made to identify all the major invasive cactus
cactus species (including O. ficus−indica and O. mona-
species, the extent of invasion, their origins, utilization
cantha) have become invasive in several countries (Novoa
and management (if any). This information is summa-
et al., 2016), resulting in well−publicized attempts to
rized in Tables 2 and 3. The perceptions of cultivated
manage and control them (Kaufmann, 2001; Zimmer-
and invasive Opuntiae, and how these relate to the
mann and Moran, 1982).
management of problematic species and the accept-
Over the past 50 years, there has been an increase in ance of cactus pear as a new emerging crop, are also
the number of invasive species, particularly within the presented. Solutions are then proposed for achieving
subfamily Opuntioideae, but several new emerging in- acceptable management options.
vaders from the Cactoideae and Pereskioideae subfam-
Within the Opuntioideae subfamily, the genera Cylin-
ilies have also become invasive, bringing the total num-
dropuntia and Opuntia contain most of the invaders
ber of reported invasive species to about 57 (Novoa
outside the Americas; they are confined mainly to Aus-
et al., 2016). The number of invasive species in South
tralia and sub−Saharan Africa (Table 1).
Africa increased from 13 (all Opuntiae) in 1947 to 35 in
2014, including at least eight Cactoideae species. Some Within the Opuntioideae, the genus Opuntia counts
invasive Opuntia species have required drastic control by far the most invaders; they are detailed in Table 2.
measures e.g. chemical control, usually followed by At least 37 countries have actually reported invasive
biological control when conventional control methods Opuntiae, but small invasions are emerging in many
proved unsuccessful (Pettey, 1948; Annecke and Mo- more countries, e.g. numerous islands and developing
ran, 1978; Moran and Annecke, 1979; Zimmermann countries in Africa. Table 3 lists the invasive cacti in the
and Moran, 1982). The spiny form of O. ficus−indica, Cactoideae and Pereskioidea subfamilies; compared
known as prickly pear, and two newly recorded Opun- with Opuntioideae, there are fewer, but the numbers
tia invaders, namely O. robusta Pf and O. engelman- of reported invasive species have nevertheless steadily
nii S−D, are currently being considered, mainly as a increased in the past decade. Only eight Cactoideae
source of fodder, partly in order to reduce infestation, species were recorded in 1984 (Moran and Zimmer-
but mostly to derive benefits from a large untapped mann, 1984b), compared with 18 in 2015. New serious
resource (Felker, 1995). In the past there have been invasions can be expected from this group.
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 173

TABLE 1 Invasive cacti by genera in three subfamilies and in countries where serious invasions have been recorded

Total no. species No. invasive

Genera in Opuntioideae Countries where invasive
in genus species
Austrocylindropuntia Australia, Kenya, Namibia, South Africa, Spain, Italy, France 11 2
Corynopuntia Australia 14 1
Cylindropuntia Australia, Botswana, Israel, Kenya, Namibia, Spain,
33 8
Zimbabwe
Opuntia Angola, Algeria, Australia, Austria, Canary Islands, China,
Croatia, Czech Republic, Eritrea, Ethiopia, France, Germany,
Ghana, India, Kenya, Madagascar, Mauritius, Morocco,
Namibia, New Zealand, Pacific Islands, Portugal, Reunion, 181 27
Saudi Arabia, Somalia, Spain, Sri Lanka, South Africa,
Swaziland, Switzerland, United Republic of Tanzania,
Tunisia, Uganda, Yemen, Zimbabwe
Tephrocactus Australia, South Africa 6 1
Total no. species No. invasive
Genera in Cactoideae Countries where invasive
in genus species
Acanthocereus Australia 6 1
Cereus Australia, South Africa, Namibia 34 3
Harrisia Australia, South Africa, Namibia 20 4
Hylocereus Australia, China, New Zealand, Portugal, Pacific Islands,
18 4
Spain, South Africa
Echinopsis Australia, South Africa 128 3
Myrtillocactus South Africa 4 1
Peniocereus South Africa 18 1
Selinicereus Pacific Islands 28 1
Total no. species No. invasive
Genera in Pereskioideae Countries where invasive
in genus species
Pereskia Australia, South Africa 17 1
TOTALS 518 58

TABLE 2 Major alien Opuntioideae invaders: origin, countries of invasion, utilization, status and management

Countries
Species and where
Utilization Status as invader Management Key references
origin introduced
and invasive
Genus:
Austrocylindropuntia
A. cylindrica (Lam.) Australia Not utilized
Backeb. South Africa Popular garden Emerging invader Chemical control Chinnock, 2015
Ecuador, Peru ornamental
A. subulata Australia
(Muehlenpf.) Kenya Serious in Chinnock 2015
Widely
Peru, Bolivia Namibia Kenya Emerging invader Chemical control Walters et al., 2011
utilized as a living fence
South Africa in other countries
Spain
Genus: Opuntia
Australia Not utilized Serious invader Fairly good biocontrol Hosking et al., 1988
O. aurantiaca Lindl.
South Africa Not utilized Serious invader Fairly good biocontrol Zimmermann and Moran,
Argentina, Uruguay
1982
Zimbabwe Not utilized Status unknown None
174 Crop ecology, cultivation and uses of cactus pear

(continued)

Countries
Species and where
Utilization Status as invader Management Key references
origin introduced
and invasive
O. cordobensis Speg.
(= O. ficus−indica) Hawaii Not utilized Insignificant Good biocontrol
Argentina
Australia Good biocontrol
O. elatior P. Miller Only problematic in Rao et al., 1971
Indonesia Utilized as a living except in Kenya
Columbia, Venezuela, Kenya with increasing Hosking et al., 1988
Kenya fence in Kenya where it is still a
Panama populations
weed
O. elata Link & Otto Australia Some biocontrol
Becoming increasingly
ex S−D Not utilized Walters et al., 2011
South Africa invasive in South Africa No control
Argentina, Uruguay
Australia
Inadequate biocontrol
Caribbean Indies
in South Africa
O. engelmannii S−D Ethiopia? Not utilized in countries Moran and Zimmermann,
Management in other
ex Eng. (six varieties South Africa invaded Utilized in Serious invader in some 1984b
countries unknown
recognized) (Varieties Mexico and Texas as countries Winston et al., eds, 2014
C. cactorum having
USA, Mexico engelmannii and stock feed
an impact in
lindheimeria are
Caribbean
invasive
Hawaii Biocontrol effective
South Africa Extensively utilized in Perceptions on weediness in South Africa,
O. ficus−indica (L.)
Australia South Africa, Ethiopia, varies Hawaii and Australia Zimmermann et al., 2009
Mill. (spiny prickly
Ethiopia Eritrea Conflicts of interest only Serious weed Haile et al., 2002
pear variety)
Eritrea Yemen prevail in other countries
Mexico
Saudi Arabia where limited control
Yemen is done
O. humifusa (Raf.) Australia
Serious invader in South Biocontrol successful Majure et al., 2012b,c
Raf. South Africa Not utilized
Africa in South Africa Walters et al., 2011
Eastern USA
O. leucotricha (DC) Australia Serious invader in South Biocontrol successful Majure et al.., 2012b,c
Not utilized
Mexico South Africa Africa in South Africa Walters et al., 2011
O. microdasys Australia Utilized as ornamental Becoming increasingly No control Smith et al., 2011
(Lehm.) Pf South Africa only invasive
Mexico
Australia
O. monacantha India
(Wild.) Haw. Kenya Intensively utilized in Low status after Successful biocontrol
Zimmermann et al., 2009
Argentina, Uruguay Sri Lanka Mauritius Madagascar only successful biocontrol everywhere
Brazil Madagascar
South Africa
Australia Seriously invasive De Kock and Aucamp,
O. robusta Wndl Utilized as fodder in Biocontrol partially
Spiny form becoming 1970
(spiny varieties) South Africa South Africa successful
invasive Walters et al., 2011

Australia
Angola
Ghana
Moran and Zimmermann,
O. salmiana (Haw.) Kenya Fruit is consumed Biocontrol effective 1984b
Haw. (including Ethiopia fresh by some but where implemented Mann, 1969
stricta and dillenii Madagascar is mainly utilized for Very serious invader in Other control
Julien and Griffiths, eds,
subspecies) Namibia confectioneries, juice many countries measures
uneconomical 1998
Mexico, USA, India and colourant Cladodes
Zimmermann et al., 2009
Caribbean Sri Lanka not utilized
Saudi Arabia
South Africa
Yemen
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 175

(continued)

Countries
Species and where
Utilization Status as invader Management Key references
origin introduced
and invasive
O. streptacantha Small remnant
Australia Not utilized Good biocontrol Mann, 1970
Lem. Mexico populations left
O. tomentos
Australia Small remnant
Salm−Dyck Not utilized Good biocontrol Mann, 1970
South Africa populations left
Mexico
Genus: Corynopuntia
Corynopuntia sp. Western Australia Unknown origin Emerging weed Chemical control Chinnock, 2015
Genus: Cylindropuntia
C. fulgida (Engelm.)
South Africa
F.M. Knuth var.
Zimbabwe Living fences Serious invader Good biocontrol Paterson et al., 2011
fulgida
USA, Mexico
C. fulgida var. Australia
mamillata South Africa Popular ornamental Serious invader Good biocontrol Paterson et al., 2011
USA Namibia
Australia
C. imbricata Botswana
Moderately good
(Haw.) F.M. Knuth South Africa Not utilized Serious invader Zimmermann et al., 2009
biocontrol
USA, Mexico Zimbabwe
Namibia

C. pallida Australia Introduced as

South Africa ornamental Chemical control
(Rose) F.M. Knuth Walters et al., 2011
Serious invader Biocontrol under
(= C. rosea) Namibia Laguna et al., 2013
Not utilized investigation
Mexico Spain
C. kleiniae (DC)
Southern USA, Australia Not utilized Emerging invader Biocontrol Chinnock, 2015
northern Mexico
C. leptocaulis (DC)
F.M. Knuth Australia
Not utilized Limited invasions Good biocontrol Zimmermann et al., 2009
Southern USA, South Africa
northern Mexico
C. prolifera (Eng.)
F.M. Knuth
Australia Not utilized Emerging invader Unknown Chinnock, 2015
Southern USA,
Baja California
C. spinosior (Eng.) Chemical control
Australia Serious invader in
F.M. Knuth Not utilized Biocontrol under Chinnock, 2015
South Africa Australia only
Southern USA investigation
C. tunicata (Leh.)
F.M. Knuth Australia
Not utilized Emerging invader Chemical control Chinnock, 2015
Southern USA, South Africa ?
northern Mexico
Genus: Tephrocactus
C. articulatus (Pf)
South Africa Popular garden
Bkbg Emerging invader Chemical control Walters et al., 2011
Namibia ornamental
Argentina
 176 Crop ecology, cultivation and uses of cactus pear

sions reported from China, Italy, Spain, Portugal,

Saudi Arabia, Ethiopia, Yemen and France (Novoa
et al., 2014). Many notorious cactus invaders are
cultivated or have been naturalized in Madagas-
car, but are not yet recognized as invasive (Kull et
al., 2012). In 2010, a brief survey of invasive cacti
in Namibia revealed 25 species seriously invading,
but with no incentives to manage them or prevent
new invasions (Zimmermann, 2010). Only two
introduced cactus species have been recorded for
Angola (Figueiredo and Smith, 2008). In general,
cactus invader lists are grossly underestimated
on account of poor herbarium specimens and
inadequate surveys. It is difficult to prepare good
herbarium specimens of succulents; in addition,
According to Moran and Zimmermann (1984b), many samples, including type specimens, become
Figure 1 46 cactus weeds were recorded globally in 1983 lost due to decay or poor processing (Anderson,
Opuntia engelmannii – amounting to 42 if the latest taxonomic classi-
invasion in South 2001; Chinnock, 2015).
fications of Hunt (2006) and Anderson (2001) are
Africa: origin USA
adopted. In 2014, 57 of the approximate 1 922 Nowadays aggressive cactus invaders are com-
recognized cactus species were recorded as inva- monplace in private and public rockeries as far
sive (Novoa et al., 2014). The figure now stands at afield as Greece, Turkey and Croatia. Easy access
58, but it remains a gross underestimate as only to known cactus weeds in nurseries (Figure 1) and
a small number of developing countries have an on the Internet (Humair et al., 2015) will inevitably
inventory of invasive cacti. South Africa and Aus- increase the establishment of, and subsequent in-
tralia are the invasion hotspots with, respectively, vasion by, such weeds. Fortunately, cactus weeds
35 and 27 species recorded as invasive (Novoa tend to have a long lag period before they are rec-
et al., 2014). Between 1947 and 2014, 24 cactus ognized as invaders; this provides opportunities
invaders were listed in South Africa (Pettey, 1948; for early detection and rapid response initiatives.
Anon., 2014). Even the established invaders have An exponential increase in cactus invasions can be
drastically expanded their range and are now in- expected.
vasive in many more countries, with serious inva-

TABLE 3 Major alien Cactoideae and Pereskioideae invaders: origin, countries of invasion, utilization, status and
management

Countries
Genera, species, where
Utilization Status as invader Management Key references
origin introduced
and invasive
Subfamily: Cactoideae
Genus: Acanthocereus
A. tetragonus (L.)
Hmlk Australia Ornamental Not serious and localized None Mann, 1970
Central America
Genus: Cereus
C. jamacaru (DC) South Africa Biocontrol reasonably
Ornamental Serious Paterson et al., 2011
South America Namibia effective but slow
C. hildemanianus
Schum (= C. South Africa Ornamental Not serious None Walters et al., 2011
uruguayanus)
C. hexagonus (L.) Mill.
Northern South Australia Ornamental Not serious None Hosking et al., 1988
America
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 177

TABLE 3 (continued) Major alien Cactoideae and Pereskioideae invaders: origin, countries of invasion, utiliza-
tion, status and management

Countries
Genera, species, where
Utilization Status as invader Management Key references
origin introduced
and invasive
Genus: Echinopsis
E. oxygona (Link.)
Zucc. ex Pf Australia Ornamental Not serious None Novoa et al., 2014
Northwest Argentina
E. chamaecereus
Australia Ornamental Not serious Novoa et al., 2014
FrdH & Glae
E. schickendantzii
Limited chemical
Weber (E. spachiana) South Africa Ornamental Serious Walters et al., 2011
control
Northwest Argentina
Genus: Harrisia
H. balansae (K. Scum) Rapid response
Not serious and very
Tayl. & Zappi South Africa Ornamental and hedges programme aimed at Walters et al., 2011
localized
Argentina eradication
H. martinii (Lab.) Biocontrol with
Australia Not serious after good
Britt. Ornamental Hypogeococcus Klein, 1999
South Africa biocontrol
Argentina festerianus
H. pomanensis (Web.) Rapid response
Britt. & Rose Australia programme in South
Ornamental Very localized Novoa et al., 2014
Chaco of South South Africa Africa aimed at
America eradication
Genus: Hylocereus
H. undatus (Haw.)
Commercial cultivations
Britt. & Rose South Africa Not serious and localized None Walters et al., 2011
for fruit
Central America

H. polyrhizus (Web.)
Pacific Islands Ornamental Unknown status None Novoa et al., 2014
Britt. & Rose
H. costaricensis
Pacific Islands Ornamental Emerging weed None Novoa et al., 2014
(Web.) Britt. & Rose
H. triangularis (L.)
Spain Ornamental Emerging weed None Novoa et al., 2014
Britt. & Rose
Genus: Myrtillocactus
M. geometrizans (Pf)
Not serious and very
Cons South Africa Ornamental None Walters et al., 2011
localized
Central America
Genus: Peniocereus
P. serpentinus
Australia
(Lag. & Rodr.) Tayl Ornamental Not serious None Walters et al., 2011
South Africa
Mexico
Genus: Selenicereus
S. macdonaldiae
(Hk) Britt. & Rose Australia Ornamental Not serious None Randall, 2002
Honduras
Subfamily: Pereskioideae
Genus: Pereskia
First attempt at
South Africa Hedge and fruit was Serious in South Africa biocontrol using two
P. aculeata Mill. Paterson et al., 2011
Australia used for confectioneries Localized in Australia insects
Chemical control
 178 Crop ecology, cultivation and uses of cactus pear

MANAGEMENT OF INVASIVE Biological control

OPUNTIOIDEAE SPECIES
The first reports of a deliberately planned and
successful biological control project on cactus
Wide−ranging options exist for the manage-
date back to 1913 when the cochineal, Dacty-
ment and control of invasive Opuntioideae:
lopius ceylonicus (Green), was introduced from
• prevention (pre−border control); India to control O. monacantha in South Africa
• early detection and rapid response; (Lounsbury, 1915; Zimmermann et al., 2009). This
• risk analysis; success was repeated in Reunion, Mauritius and
Australia and it encouraged scientists in Australia
• chemical, biological and mechanical control;
to embark on one of history’s largest and most
and
dramatic projects of biocontrol of weeds, namely,
• utilization. the successful biocontrol of O. stricta (Haw.) Haw.
Control is often hampered by conflicts of inter- in Queensland and New South Wales in the 1920s
est: a cactus weed may be perceived by some and 1930s (Dodd, 1940; Mann, 1970; Zimmermann
as beneficial, but by others as problematic and et al., 2009). Since then, biocontrol has been de-
necessitating control. This can become the big- ployed against 28 other cactus invaders, involving ≥
gest stumbling block in the search for solutions, 22 successfully established cactus−feeding insects
particularly among Opuntiae and the Hylocereeae and mites (Zimmermann et al., 2009). Almost 42%
and Cereeae tribes. of these projects have been completely successful;
another 18% have achieved substantial control
Figure 2 (Zimmermann et al., 2009; Klein, 2011; Hosking,
Opuntia stricta iinvasion 2012; Winston et al., eds, 2014). Most recently, Cy-
in Saudi Arabia:
lindropuntia fulgida var. fulgida (Eng.) Knuth and C.
Origin USA
fulgida var. mamillata (Figure 2) were successfully
1
For a more detailed controlled in South Africa (Paterson et al., 2011).
update of the biolo-
gical control of cactus Table 4 lists the most important cactus invaders
invaders, consult that have been subjected to biocontrol. The list
Winston et al., eds does not include species that have become invasive
(2014) or in their native habitats, and it does not include bio-
www.ibiocontrol.org/
logical control agents that have neither established
catalog
nor had an impact on their cactus host1.

TABLE 4 Invasive alien cactus invaders: countries of introduction and outcome of deliberately introduced natural
enemies for biological control

Country where biocontrol

Cactus invader Utilization Status as invader
was introduced
Subfamily: Opuntioideae
Genus: Opuntia
Australia Moderate variable
Cactoblastis cactorum (Berg)
South Africa Moderate
O. aurantiaca Lindl.
Australia Zophodia tapiacola (Dyar) Moderate
(Argentina, Uruguay)
Australia Considerable
Dactylopius austrinus De Lotto
South Africa Considerable
O. elata Link & Otto ex S−D Australia Dactylopius ceylonicus (Greeen) Trivial
Indonesia Extensive
O. elatior Mill. Dactylopius opuntiae (Cockerell)
Kenya ?
(Central America) “ficus−indica” biotype
India Extensive
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 179

(continued)

Country where biocontrol

Cactus invader Utilization Status as invader
was introduced
O. engelmannii S−D ex Eng. South Africa Moderate to trivial
Australia Dactylopius opuntiae (Cockerell) depending on the variety
3 varieties
“ficus−indica” biotype
(USA) Kenya Unknown
Australia Moderate
Dactylopius opuntiae (Cockerell)
South Africa Considerable
“ficus−indica” biotype
Hawaii Considerable
Australia Considerable
O. ficus−indica (L.) Mill. (spiny) South Africa Considerable
(Mexico) Cactoblastis cactorum
Hawaii Considerable
Mauritius Considerable
South Africa Metamasius spinolae (Gyllenhall) Considerable (localized)
Hawaii Considerable
Archlagocheirus funestus (Thomson)
South Africa Trivial
Dactylopius opuntiae (Cockerell)
Extensive
O. humifusa (Raf.) Raf. (USA) South Africa “stricta” biotype
Cactoblastis cactorum (Berg) Trivial
Dactylopius opuntiae (Cockerell)
O. leucotricha (DC.) Moderate
South Africa “ficus−indica” biotype
(Mexico)
Cactoblastis cactorum (Berg) Trivial
Australia Extensive
India Extensive
Kenya Moderate
Sri Lanka Extensive
Madagascar Dactylopius ceylonicus (Green) Extensive
O. monacantha Haw.
Mauritius Extensive
(South America)
Tanzania Moderate
South Africa Extensive
Ascension Island ? Moderate
Mauritius Considerable
Cactoblastis cactorum (Berg)
South Africa Trivial
Australia Dactylopius opuntiae (Cockerell) Considerable

O. robusta (spiny) South Africa “ficus−indica” biotype Moderate

(USA) Australia Considerable
Cactoblastis cactorum (Berg)
South Africa Considerable
O. salmiana (Parm.) ex Pf
South Africa Cactoblastis cactorum (Berg) Trivial
(Argentina)
Australia Initially extensive
India Extensive
Kenya Extensive
O. strictab (Haw.)
Dactylopius opuntiae (Cockerell)
Haw. (two varieties) India Extensive
“stricta” biotype
(USA, Mexico) Namibia Moderate
Saudi Arabia Probably extensive
Sri Lanka Considerable
180 Crop ecology, cultivation and uses of cactus pear

(continued)

Country where biocontrol

Cactus invader Utilization Status as invader
was introduced
Australia Extensive
Kenya Trivial
O. strictab (Haw.) Namibia Cactoblastis cactorum (Berg) Moderate
Haw. (two varieties)
(USA, Mexico) New Caledonia Considerable
South Africa Moderate
Australia Moneilema blapsides Newman Trivial
Dactylopius opuntiae (Cockerell)
Considerable
“ficus−indica” biotype

O. streptacantha Lem. Cactoblastis cactorum (Berg) Moderate on young plants

Australia
(Mexico) Archlagocheirus funestus (Thomson) Trivial
Moneilema blapsides Newman Trivial
Lagocheirus funestus Thomson Trivial
Australia Dactylopius opuntiae (Cockerell) Moderate
O. tomentosa S−D. “ficus−indica” biotype
South Africa Considerable
(Mexico)
Australia Cactoblastis cactorum (Berg) Trivial
Dactylopius opuntiae (Cockerell)
O. tuna a (L.) Mill. Initially moderate
Mauritius “ficus−indica” biotype
(Caribbean)
Cactoblastis cactorum (Berg) Extensive
Genus: Cylindropuntia

C. fulgida (Eng.) Knuth var. South Africa Dactylopius tomentosus (Lamarck) Extensive
fulgida (Sonora, Mexico, USA) Zimbabwe “cholla” biotype Extensive
Australia Good progress
C. fulgida (Eng.) Knuth var. Namibia Dactylopius tomentosus (Lamarck) Extensive
mamillata (USA) South Africa “cholla” biotype Extensive
Zimbabwe Extensive
Australia Extensive
Namibia Dactylopius tomentosus (Lamarck) Considerable
C. imbricata (Haw.) Knuth “imbricate” biotype
Botswana Considerable
(Mexico, USA)
South Africa Considerable
South Africa Cactoblastis cactorum (Berg) Trivial

C. leptocaulis (DC) Knuth Australia Dactylopius tomentosus (Lamarck) Considerable

(USA) South Africa “imbricate” biotype Extensive
C. kleiniei (DC) Knuth Dactylopius tomentosus (Lamarck)
Australia Considerable
(Mexico, USA) “imbricate” biotype
C. rosea (DC) Bkbg Dactylopius tomentosus (Lamarck)
Australia Trivial
(Mexico) “imbricate” biotype
Subfamily: Cactoideae
Genus: Acanthocereus
A. tetragonus (L.) Hmlk Hypogeococcus festerianus
Australia Moderate
(Argentina) (Lizer & Trellis)
Genus: Cereus
Considerable where
Nealcidion cereicola (Fisher)
C. jamacaru (DC) established
South Africa
(Argentina, Brazil) Hypogeococcus festerianus (Lizer &
Considerable
Trellis)
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 181

(continued)

Country where biocontrol

Cactus invader Utilization Status as invader
was introduced
Genus: Harrisia
H. balansae (KSch) Taylor & Hypogeococcus festerianus
South Africa Considerable
Zappi (Argentina) (Lizer & Trellis)
Australia Hypogeococcus festerianus Considerable
H. martinii (Lab.) Britt. (Lizer & Trellis)
South Africa Considerable
(Argentina)
South Africa Nealcidion cereicola (Fisher) Trivial to moderate
H. pomanensis (Web) Britt. & Hypogeococcus festerianus
South Africa Considerable
Rose (Argentina) (Lizer & Trellis)
H. regelii (Wngt) Borg Hypogeococcus festerianus
Australia Considerable
(Argentina) (Lizer & Trellis)
H. tortuosa (Otto & Dietr.) Hypogeococcus festerianus
Australia Considerable
Britt. & Rose (Lizer & Trellis)
Subfamily: Pereskioideae
Genus: Pereskia
P. aculeata Mill.
South Africa Phenrica guerini Bechyne Trivial
(Argentina, Brazil)

a
O. tuna is no longer recognized as a species (Hunt et al., 2006).
b
O. dillenii is now regarded as a spiny form of O. stricta and is sometimes cited as a subspecies.

Biological control of cactus weeds has generally been 1999; Volchansky et al., 1999; Mathenge et al., 2009;
easier than of taxa in other plant families. This is due Jones et al., 2014). They provide excellent control of
to the absence of native cactaceous species outside O. stricta var. stricta and C. fulgida var. fulgida and C.
the Americas, with the exception of Rhipsalis baccifera fulgida var. mamillata resp., not only in South Africa but
(Mill.) Stearn, which arrived in South Africa, Mada- also in Australia, Kenya and Saudi Arabia (Paterson et
gascar and Sri Lanka probably with migrating birds al., 2011). Further research is focusing on finding new
(Rebman and Pinkava, 2001). This set of circumstances cochineal biotypes for O. engelmannii, O. elata S−D,
has permitted the safe use of phytophagous insects C. pallida (Rose) Knuth and C. spinosior (Eng.) Knuth.
and mites that are less host−specific. However, as The existence of host−adapted biotypes within the
cactus pear (O. ficus−indica) is becoming increasingly mealybug species, Hypogeococcus festerianus Granara
important worldwide, the choice of biocontrol agents de Willink, to control invaders in the genera, Harrisia
for new cactus weeds is increasingly restricted, particu- and Cereus, should not be ruled out. The prospects of
larly in the case of opuntias, as the safety of this new biological control of new invaders from the Cactoideae
world crop cannot be allowed to be compromised. It is are generally good.
increasingly difficult to find natural enemies outside the
Dactylopiidae that are sufficiently host−specific to be The risks of biological control are minimal. There are
considered for biocontrol of Opuntia invaders. no known cases of unpredicted non−target effects
of cactus biological control agents. Cactophagous
More recently, the successful biological control of arthropods are known to be host−specific to Cactace-
cactus invaders may be attributed to new association ae species; as the family is endemic to the Americas,
effects (Hokkanen and Pimentel, 1989). This applies it is very unlikely that an unpredicted host−switch
primarily to cochineal insects and entails the use of to another plant family will occur outside its native
biocontrol agents that do not have a recent history range. The cactus moth, Cactoblastis cactorum (Berg),
of intensive interaction in evolutionary time with their and the cochineal, D. opuntiae, were known to have
host. The objective is to take advantage of the lack of many hosts within the genus Opuntia; it was predicted
evolved commensalism common to many predator– that both insects would establish on cactus pear and
prey systems. Two cochineal biotypes (=genotypes) invasive Opuntiae. The use of these two insects for
have recently been identified and evaluated: Dacty- biological control was based on calculated risks, given
lopius opuntiae (Cockerell) (“stricta” biotype) and D. the seriousness of cactus invasions at that time. On the
tomentosa (Lamarck) (“cholla” biotype) (Githure et al., other hand, the deliberate introduction of the cactus
182 Crop ecology, cultivation and uses of cactus pear

moth to some Caribbean islands in 1957 was most Prevention

unfortunate and ill−conceived (Zimmermann et al.,
2001). It was introduced to control the indigenous O. Research is underway to prevent the introduction and
triacantha (Willd.) Sw and it was known that the moth establishment of new cactus invaders. Initiatives include
would also develop on most other species in the genus strict pre−and post−border control, risk assessments
Opuntia in the West Indies. In time, the moth spread before species are introduced, and identification and
to the American mainland through the nursery trade regulation of the potential pathways of introduction
(Pemberton, 1995), via weather events or by multiple (Novoa et al., 2015). Legislation can play an important
introductions directly from South America (Marsico et role in preventing the establishment of new invaders
al., 2011). It is now threatening native Opuntia species or in controlling invasive cacti. This may include the
in the United States of America and Mexico. compilation of lists of prohibitive species and of species
with prescriptive control measures. In South Africa, 34
The cochineal, D. opuntiae, widely used as a biological cactus species are listed in several categories of invasive
control agent outside its native range (United States alien plants in terms of the National Environment Man-
of America and Mexico), may have been carried as a agement Authority (NEMA) regulations, which make
contaminant on vegetative plant material to several control mandatory (Anon., 2014).
countries, including the Mediterranean region. It has
also been introduced inadvertently, mistaking it for An analysis of the 57 cactus invaders on a global scale
the carmine cochineal, D. coccus. In addition, since it shows that species in some genera are more likely to
is wind dispersed, there is also a remote chance that it become invasive than species in others. In the genus
reached cactus pear plantations through air currents, Opuntia alone, 26 species (of a total of 181) are already
especially where plantations are close to invasive invasive somewhere in the world. Similarly, in the genus
Opuntia populations infested with the insect. This can Harrisia, five species (of a total of 20) are now inva-
only occur where D. opuntiae is used for biocontrol, sive. It might, therefore, be justifiable for a particular
for example, in South Africa, India, Sri Lanka, Kenya, country to declare an entire genus as unwanted. This
Indonesia, Australia, Hawaii and Mauritius. It is vital to approach is already applied in South Africa, where it is
ensure that only contaminant−free vegetative material prohibited to introduce all new species in the genera
is used for new cactus pear plantings. Cylindropuntia, Harrisia and Opuntia (Anon., 2014),
while those species that are already naturalized (except
for the commercially valuable species) are subject to
Chemical and mechanical control compulsory control (Anon., 2014). With the exception
of spineless O. ficus−indica, all species in the genera
Mechanical control is seldom a viable option for dealing Opuntia, Austrocylindropuntia and Cylindropuntia are
with cactus invasions, because vegetative reproduction declared “weeds of national significance” in Australia
is possible from small fragments that remain after phys- (the genus Corynopuntia may be added to the list)
ical clearing. Chemical control is recommended where (Chinnock, 2015). It is highly unlikely that permits will
small and new invasions are to be confined, controlled be granted for the introduction of any species in these
or even eradicated, but it is seldom viable when large genera. Other countries may have to consider similar
and extended populations need to be controlled. Sev- legislation.
eral chemical campaigns in the history of cactus control
have failed owing to the costs involved and the rapid re- There are more than 800 species in the ornamental
covery of populations: chemical control is never−ending cactus trade, including 25 known invaders (Novoa et
(Dodd, 1940; Annecke and Moran, 1978; Moran and al., 2016). At least 266 of these species are introduced
Annecke, 1979). to other countries primarily as seeds. Screening such
large numbers of introduced species for prohibited or
The thick protective wax layers and the crassulacean potentially invasive species is a challenge, especially
acid metabolism (CAM) photosynthetic pathway (closed when they are imported as seeds (Humair et al., 2015),
stomata during daytime) in most cacti severely restrict and New Zealand is the only country that screens all
the uptake of foliar applied herbicides, unless effective incoming postage for seeds. However, a general rule
wetting additives are used. Most cactus invaders, how- of the thumb is that species with large and heavy seeds
ever, lend themselves to stem injection treatments with are more likely to be invasive than species with small
systemic herbicides due to their succulent nature. The seeds (Novoa et al., 2016) (exceptions are the genera
advantages are rapid translocation to all parts of the Echinopsis, Cereus and Harrisia). The best option for
plant, minimal damage to non−target species and low prohibiting the introduction of harmful species may be
costs. Access to the stems in certain thicket−forming a risk assessments at genus level, carried out in consul-
and spiny cacti is a potential problem when using the tation with importers and the horticultural cactus trade.
stem injection method (Grobler, 2005). One recommendation exists that all genera containing
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 183

invasive species should be prohibited. For now, South There were many years of debate and negotiations
Africa has only prohibited imports of species not already before it was decided to embark on a biological control
in the country in the genera Opuntia, Cylindropuntia programme against prickly pear (O. ficus−indica) to halt
and Harrisia (Anon., 2014). Most cactus species in the further invasions in South Africa (Annecke and Moran,
ornamental trade belong to genera with no recorded 1978). Two natural enemies, the cactus moth (C. cacto-
invasive species. To date, no invasive species have been rum) and the cochineal (D. opuntiae), were released in
reported in the large genus Mamillaria; this very popu- 1933 and 1938, respectively. Following successful bio-
lar group of ornamentals could thus be excluded from control (Pettey, 1948), an effort was made to utilize the
regulation. small infestations remaining in order to further reduce
its status as an invader (Brutsch and Zimmermann, 1993,
Other characteristics that can be useful in risk assessments
1995). Today these remaining infestations are well used,
are reproduction strategies and vectors of dispersal.
mainly as a source of fruit; control is no longer needed,
except in particular situations, such as in conservation
areas (Beinart, 2003). In contrast, Ethiopia (Tigray prov-
UTILIZATION AND CONFLICTS OF INTEREST
ince) and Eritrea have not implemented biocontrol and
are unlikely to do so in the future; consequently, dense
A culture of utilization of cactus species (mainly Opuntiae)
infestations of > 300 000 ha in Ethiopia alone persist
has developed over thousands of years, mainly in Mexico
and some countries of Central and South America. Over (Behailu and Tegegne, 1997; Haile et al., 2002). Local
the past 400 years, when a plant has been introduced to farmers have become very dependent on prickly pear as
another country, the culture of utilization has not always there is little natural vegetation left. They are reluctant
transferred with it. In countries where prickly pear (spiny to destroy even part of the cultivation through biological
O. ficus−indica) has become invasive, there has been a control, because they fear losing the resource on which
clear shift from “useful” during establishment to “trou- they have become dependent. Despite an effort by FAO
blesome” in the subsequent expansion phase until it is to promote new uses and to improve the utilization of
eventually perceived as a national disaster. More recent- fruit and fodder, there is no evidence of reduction of the
ly, however, there has been a significant revival in the infestations (Portillo, personal communication). In other
multiple uses of cactus pear (spineless O. ficus−indica), countries with large invasions, such as Saudi Arabia and
and these can also be applied to prickly pear. There is a Yemen, active utilization programmes have yet to begin
wide range of literature on these uses, including many and prickly pear populations continue to increase.
recipes and by−products of “tuna” (fruit) and nopalitos
Madagascar is a special case: it was not prickly pear (O.
(tender cladodes used as a green vegetable) and medic-
ficus−indica), rather O. monacantha (Raketa), that in-
inal applications (De Waal et al., 2015; Sáenz et al., eds,
vaded large parts of south Madagascar. The cactus was
2006). Use of the vegetative parts of O. ficus−indica, O.
intensively utilized − mainly as a source of fodder and
robusta and more recently N. cochenillifera as a source
fruit − until the local population became fully dependent
of fodder is increasingly practised; in some countries,
on it (Middleton, 1999; Kaufmann, 2001). Nevertheless,
such as Brazil, it has developed into a substantial in-
some locals saw the cactus as a problem (Kaufmann,
dustry (Mondragón Jacobo and Pérez González, 2001;
2001). Similar large−scale invasions occurred in India,
Dubeux et al., 2015a). Opuntia ficus−indica is also the
Australia, Mauritius, Sri Lanka and South Africa, but the
principal host for the production of the carmine cochi-
weed was not actually utilized, except for a brief period
neal, D. coccus (Costa), which is a major industry in Peru
in India when it was used to rear the carmine cochineal,
(Flores Flores and Tekelenburg, 1995). There have been
D. coccus (Zimmermann et al., 2009; Winston et al., eds,
considerable efforts to apply these many uses to invasive
2014). The introduction of the cochineal, D. ceylonicus,
populations of prickly pear, in an attempt to reverse the
to Madagascar in 1924 (apparently mistakenly identified
status of the invader and turn it into an asset. Utilization
as D. coccus, with the intention of starting a cochineal
alone is not sufficient, however, to curtail the densifi-
cation and spread of prickly pear in Eritrea, Yemen and industry) resulted in the destruction of practically all
Ethiopia, where many thousands of hectares are invaded cactus populations within 4 years (Middleton, 1999).
and expanding. This led to severe starvation among the local popula-
tion. According to Middleton (1999), the rehabilitation
There is much to be learned from countries where of the pastures following the demise of the cactus was
cacti in all three subfamilies have been widely utilized too slow to be able to support the same level of pasto-
by ancient civilizations for thousands of years, such as ralism, resulting in the death of thousands of cattle. The
Mexico (Hoffmann, 1995; Anderson, 2001). In Mexico, conflicts of interest between pastoralists, local commu-
well−known aggressive invaders, for example, Cyl. nities, authorities and politicians are well documented
imbricata (Haw) Knuth and O. engelmannii, are widely by Middleton (1999) and Kaufmann (2001, 2004). It
utilized for forage (Benson, 1982; Nobel, 1994). seems that the Malagasy people learned to live with this
184 Crop ecology, cultivation and uses of cactus pear

transformer species, involving intense utilization of this southwards to countries that depend on cochineal for
foreign new resource (Kull et al., 2012, 2014), in part the biological control of invasive cacti.
because they had no other choice. The O. ficus−indica
invasions in Tigray (Ethiopia) offer parallels.

In South Africa, on the other hand, conflict raged CONCLUSIONS

for years over whether to start a biological control
programme against O. ficus−indica (Beinart, 2003). Cactus pear is a multipurpose dryland crop destined to
A project was finally launched in 1932 and biological become more important as global warming and desert-
control reduced infestations by about 80%, leaving ification increase. It offers great potential for developing
enough plants to be widely utilized by the rural popula- countries in arid regions. At the same time, however,
tion (Annecke and Moran, 1978). The pastoralists were these countries face the escalating problem of invading
never as dependent on prickly pear as in Madagascar, cacti. Innovative utilization of these cacti, mainly from
and the resulting deficiencies were partly made up for the genus Opuntia, may be one way to partly alleviate
by cultivated spineless cactus pear and spineless O. the threat. A more feasible option is to prevent the
robusta – a project initiated by the South African Gov- introduction and establishment of new invaders in the
ernment to compensate for the loss of cactus fodder first place. Research is required to find innovative and
and fruit (Beinart, 2003). The resistance to biological integrated methods to manage these invasions. It is
control of prickly pear invasions in some countries is recommended that research focus on utilizing the large
understandable; this resistance is exacerbated when the biomasses of many invasive species. In some cases, it
population’s dependency on the plant increases, as in may be possible to find new species−specific biocontrol
Ethiopia, Eritrea and Yemen. agents, including new cochineal biotypes, which would
solve some of the problems (Zimmermann et al., 2009).
More recently, severe invasions of O. stricta, O. dillenii Invasive experts in South Africa and elsewhere have
(Ker−Gawler) Haw. (= O. stricta var. dillenii), O. elatior undertaken valuable research to predict new invaders,
Mill. and O. engelmannii are emerging at an alarming particularly from within the succulent nursery trade.
rate in Madagascar, Ethiopia, Yemen, Somalia, Angola, Conflict of interest has until now been confined to
Namibia and Kenya. Since most effective biocontrol Opuntiae, but new emerging crops in the Cereus and
agents are not species−specific, and due to conflicts Hylocereus genera (Nerd et al., 2002), both of which
of interest (past or present) in some of these countries, contain invasive species, could cause new conflicts to
there is little chance of implementing biological control arise. Natural enemies (e.g. H. festerianus) that have
on these species. Risk–benefit analysis will eventually been released for the control of invasive Harrisia and
show that biological control can be considered part of Cereus species in Australia and South Africa are known
a management plan, even if it may entail slight damage to have the potential to cause damage to several species
to cultivated species. Most cactus invasions are already in the Cereeae and Trichocereeae tribes (McFadyen,
beyond the stage where chemical control can be imple- 1979; McFadyen and Tomley, 1981).
mented. In time, the option of biological control will be-
Compared with 20 years ago, there are many new in-
come more compelling until it might eventually become
vaders in the Cactoideae subfamily, mostly originating
part of a broader integrated management programme.
from the nursery trade. This trend is likely to continue.
Many new cactus invaders lie in waiting, ready to move
The “ficus” biotype of the cochineal, D. opuntiae, has
into the exponential phase of invasion. In Madagascar
become a pest on cultivated cactus pear, not only where
alone, of 52 cactus species that are cultivated or have
biocontrol was deliberately implemented, but in other
become naturalized, only two are currently regarded
countries, such as Brazil, Morocco, Spain, Egypt and
as invasive (Kull et al., 2012, 2014). Of the remaining
Lebanon, where the cochineal was accidentally intro-
50 species, 24 have been recorded as invasive in other
duced. Another level of conflict of interest may develop
countries (Tables 2 and 3), including some of the worst
as some countries contemplate the introduction of co-
cactus weeds known: O. aurantiaca, O. stricta, O. dil-
chineal predators from Mexico for the biological control
lenii, C. leptocaulis, C. tunicata and Harrisia spp.
of cochineal in cactus pear cultivations. These predators
are not host−specific and may control all Dactylopius With the exception of the insect enemies associated
species and biotypes, including those that are vital for with the genus Harrisia (McFadyen, 1979) in South
the control of several invasive cacti. This may severely America, the cactophagous insects associated with the
reduce the efficacy of biological control. For this reason, Cactoideae subfamily have not yet been surveyed ex-
South Africa has a moratorium on the importation and tensively, unlike those associated with the genera of the
release of any predator that may negatively impact on Opuntioideae (Mann, 1969). There are good prospects
natural enemies of cochineal. Unfortunately, any releas- for finding effective and host−specific natural enemies,
es of predators in North Africa will eventually migrate should biological control become necessary.
Global invasions of cacti (Opuntia sp): control, management and conflicts of interest 185

Figure 3
Opuntia ficus−indica
invasion in South
Africa: Origin Mexico

Figure 4
Opuntia robusta
invasions in Australia:
origin Mexico

3 4

Figure 5
Opuntia humifusa
invasion before
biological control in
South Africa: origin
USA.

Figure 6
Opuntia humifusa after
biological control in
South Africa.
Helmuth Zimmermann
5 6

Figure 7
Opuntia stricta
invasions in Ethiopia

Figure 8
Opuntia aurantiaca
invasions in South
Africa and Australia:
origin Argentina

7 8
Figure 9
Excellent biological
control of Opuntia
stricta in South Africa,
Kenya, Saudi
Arabia and Australia
using a host−specific
cochineal, Dactylopius
opuntiae "stricta"
biotype.

Figure10
9 10 Opuntia stricta invasion
in the Tsavo National
Park, Kenya, before
biological control.,
2012) (photo A. Witt)

Figure11
The same locality after
successful biological
control using a host−
specific biotype of
Dactylopius opuntiae
(2015) (Photo A. Witt)
11
 16

Biogas production
Maria Teresa Varnero and Ian Homer
Department of Engineering and Soils, Faculty of Agricultural Sciences,
University of Chile

LAMP

TV

PRESSURE
GAUGE
DESULFRIZER

NOPAL CLADODES
ANIMAL
MANURE
STOVE

BIOGAS STORAGE
INLET OUTLET POOL

BAG
 16 Biogas production

INTRODUCTION USING CACTUS WASTE IN

BIOGAS PRODUCTION
Non−conventional renewable energy (NCRE) is
increasingly prominent, providing an inexhaust- Dry climate areas have reduced availability of or-
ible energy source compatible with human and ganic waste − an obvious disadvantage for biogas
environmental sustainability. The various forms production. This obstacle can be overcome by de-
of NCRE include wind, solar, small hydro, tidal, veloping energy crops well adapted to arid areas.
geothermal and biomass. Biomass uses biological, In this context, opuntias − among them, Opuntia
chemical and physical processes to generate liquid ficus−indica (L.) Mill. − characterized by crassu-
or gaseous biofuels, such as biodiesel, bioethanol lacean acid metabolism (CAM), are recommended
and biogas. as an alternative energy source as they have a
Biogas is a viable and essential form of energy in high potential for biomass production (García de
agricultural and rural areas, obtained from the Cortázar and Nobel, 1992; García de Cortázar
processing of organic waste through anaerobic and Varnero, 1995). Farmers can thus reduce
digestion. In addition to biogas (comprising mainly their electricity and gas bills (liquefied petroleum
methane and carbon dioxide, plus other trace gas- gas, LPG) by producing their own energy, while
es), the process also produces a stabilized organic improving the quality and conditions of the soil by
waste, digestate (also known as biol or bioferti- applying the digestate to the fields.
lizer), which can be used as a soil conditioner or In the Faculty of Agricultural Sciences of the
biofertilizer (Varnero, 1991, 2001). University of Chile, experiments with Opuntia fi-
The biodegradation rate of organic residues is cus−indica (Uribe et al., 1992; Varnero et al., 1992;
related to the microbial activity in the anaerobic Varnero and López, 1996; Varnero and García de
system. This activity depends on the type of raw Cortázar, 1998) indicate that the cladodes are not
material, the pH of the medium, the total level of a good methanogenic material. The quality of
solids, the temperature of the process and other the starting material in the digesters, particularly
Figure 1
parameters that determine the digestion period when batch−loaded (Hilbert, 2009), is vital for
Family size
biogas system for the production of biogas and biofertilizer. the process. It is, therefore, necessary to include

LAMP

Family size biogas system TV

PRESSURE
GAUGE
DESULFRIZER
GENERATOR
ORGANIC NOPAL CLADODES
TRASH
AND ANIMAL CROP
TOILET MANURE
STOVE

BIOGAS STORAGE
INLET OUTLET POOL

BAG
Biogas production 189

a particular material derived from another digest- accelerating the methanogenic process and reduc-
er and enriched with methanogenic bacteria, or ing the time required for this activity (Varnero and
to incorporate a percentage of animal manure. García de Cortázar, 2013).
Such adjustments advance the starting time of the
methanogenic phase in the digester and increase
the production of biogas. Moreover, the pH of the OPUNTIA SPP. PLANTATIONS
pulp is very low, and this too affects the production
FOR BIOGAS PRODUCTION
of biogas; for this reason it is preferable to mix with
other raw materials, mostly animal manure.
Opuntias can grow successfully in areas with a
The fermentation efficiency of mixtures containing range of climates and soils; therefore formal plan-
different proportions of cladodes and animal ma- tations can be established to optimize biomass
nure showed that it is crucial to maintain the pH of production. Its economic evaluation is still pending.
the mixture close to pH = 6 in order to obtain biogas Studies have shown that 1 ha of Opuntia over 5
with a methane content of > 60%. The composition years old can produce up to 100 tonnes of fresh
of the biogas produced by methanogenic fermenta- cladodes per year in areas with little rainfall (≤
tion is closely related to the pH of the raw materials 300 mm) (García de Cortázar and Nobel, 1992).
biodigested. At pH < 5.5, biogas is predominantly In some semi−arid parts of Mexico, cladodes are
carbon dioxide, with reduced combustibility and traditionally collected from wild cactus plants as a
energy content; conversely, with a neutral or basic source of forage; regular pruning boosts yield and
pH, the biogas is methane−enriched. It is, therefore, improves fruit or nopalitos quality.
important to increase the proportion of animal ma-
nure in the mixture and use cladodes older than 1 Pruning can yield approximately 10 tonnes of dry
year. The particle size of the chopped material has matter (DM) ha−1 year−1, and the prunings can be
no significant influence on the efficiency of the used for biogas, compost or animal feed (García de
fermentation process (Varnero and López, 1996; Cortázar and Varnero, 1995). Pruning can also pro-
Varnero and García de Cortázar, 1998). vide the raw material to feed digesters, combined
with animal manure. Mature cladodes (1 year old)
During the anaerobic digestion of animal manure, can be cut, chopped and fed directly into the digest-
the addition of cactus cladodes promotes the meth- ers. It is important to use them as soon as they have
anogenic fermentation, provided that the pH of the been chopped, in order to reduce biodegradation
mixtures of these raw materials remains within a and improve the efficiency of biogas and biofertiliz-
neutral or slightly acidic range. Furthermore, adding er production. If the capacity of the digester is not
an appropriate percentage of cladodes to the animal sufficient for immediate use, the cladodes can be
manure helps the fermentation process start earlier stored in a shaded, cool, dry place for several days
(Uribe et al., 1992; Varnero et al., 1992): the energy (Varnero and García de Cortázar, 2013).
and carbon content of the cladodes favours the
development of acidogenic bacteria, which generate As the plantation matures, the growth of the
the substrate required by methanobacteria, thereby cladodes slows, because the net photosynthetic

Gases
CH4 CO2 Figure 2
(%)
Biogas composition
80 as a function of pH
of manure−cladode
mixture (Varnero and
60 Arellano, 1991)

40

20

0
7,5 7,4 6,7 6,1 5,3 4,8
pH of manure-cladodes mixture
190 Crop ecology, cultivation and uses of cactus pear

rate decreases due to the shading effect of the • Scenario 3: optimum production. A commer-
upper cladodes (Acevedo and Doussoulin, 1984). cial plantation with irrigation and fertilization
The dry matter content is not affected, howev- can produce ≤ 30−40 tonnes DM ha−1 (García
er, as growth continues throughout the year. In de Cortázar and Nobel, 1992; Franck, 2006).
Chile, the maximum commercial fresh fruit yield Production of 30 tonnes year−1 is equivalent to
is estimated at 16 tonnes ha−1 for plants 16−20 82.2 kg day−1, which can be used as raw mate-
years old under good management. This begins to rial for biogas production with a potential of 29
decline between 21 and 35 years of age, reaching m3 day−1 (82.2 × 0.360 = 29 m3 biogas day−1),
8 tonnes ha−1 (Acevedo and Doussoulin, 1984; or 10 885 m3 ha−1 year−1 biogas − comparable to
Pimienta Barrios, 1990). During the January−April 6.4 tonnes of oil (Varnero, 1991).
harvest season, yield is 5−16 tonnes ha−1, while
There is a significant difference in biogas produc-
and in June−September, it is just 0.5 tonnes ha−1
tion between the best and worst scenario as a
(Sudzuki et al., 1993).
result of the constraints involved.
Tohá (1999) indicates that 3 kg of dried cladodes
produce 1 m3 of biogas, which is equivalent to an Based on the family biogas system for small oper-
output of 10 kWh. Moreover, Baeza (1995) indi- ations illustrated in Figure 1, organic waste can
cates that the calorific value of biogas from cactus be collected by connecting the bathroom to the
is 7 058 kcal m−3 (range of 6 800−7 200 kcal m−3) digester, and/or by accumulating kitchen waste.
and the biogas potential of Opuntia is equivalent Furthermore, if there are animals (e.g. one cow
to 0.360 m3 kg−1 DM. and two pigs), they too provide organic matter
(Table 2). This would give a biogas potential of
• Scenario 1: low production. With a yield of 1.05 m3, leaving a further 2.6 m3 to achieve the
10 tonnes DM ha−1 year−1, the potential biogas 3.61 m3 needed. In order to determine the min-
production is equivalent to 9.86 m3 biogas day−1 imum cactus plantation area required to supply
(27.40 kg DM day−1, with a potential estimated enough raw materials to achieve this amount, it
0.36 m3 biogas kg−1 (27.40 × 0.360 = 9.86 m3 should be considered that a maximum of 3 kg of
biogas day−1). cactus is necessary to produce 1 m3 of biogas.
• Scenario 2: intermediate conditions. Prun- The digester must, therefore, be provided with 7.1
ing waste production of 18 tonnes ha−1 year−1 kg day−1; this is obtained in an area of 0.28 ha,
generates 17.75 m3 biogas day−1 and 49.3 kg assuming an availability of 10 tonnes of cladodes
DM day−1. ha−1 year−1, equivalent to 27.47 kg ha−1 day−1.

Practical example
TABLE 1 Average consumption of biogas energy in a family of 5 people

Average biogas consumption Cactus biogas consumption

Calorific value 5 000 kcal m −3
Calorific value 7 058 kcal m−3 (75% of CH4)
Kitchen (5 hours) 0.30 m3 h−1 × 5 h = 1.50 m3 day−1 0.21 m3 h1 × 5 h = 1.05 m3 day1
3 lamps (3 hours) 0.15 m h × 3 h × 3 = 1.35 m day
3 1 3 −1
0.11 m3 h1 × 3 h × 3 = 0.99 m3 day1
Cooling medium 2.20 m3 h1 × 1 = 2.20 m3 day1 1.57 m3 h−1 × 1 = 1.57 m3 day−1
Total 5.05 m day
3 −1
3.61 m3 day−1

Source: Baeza, 1995.

TABLE 2 Summary of the calculations

Potential biogas
Quantity (units) kg unit−1 kg Biogas (m3)
(m3 biogas kg−1)
Kitchen waste 5 0.56 2.8 0.092 0.26
Human faeces 5 0.13 0.65 0.092 0.06
Cow manure 1 10 10 0.04 0.40
Pig manure 2 2.8 5.6 0.06 0.336
subtotal 1.053
Cladodes 0.28 27.47a 7.7 0.3 2.60
total 26.64 3.62
a
10 tonnes ha−1 year−1 (364 days).
Biogas production 191

On the basis of the biogas production described • Continuous. Material loading is frequent (daily
above, 0.45 m3 of gas is obtained per m3 of digest- or weekly), each load replacing approximately
er; the minimum size of the digester is, therefore, 5−15% of the total volume. The solids concentra-
8 m3. In addition, a daily load of 26.24 kg must tion is low (6−8% of the volume), and once the
be incorporated, combined with sufficient water digestion process starts, the biogas production
for a solid concentration of 7%: equivalent of 221 rate is relatively constant (this is mainly depend-
litres (also equivalent to the digester’s volume, di- ent on temperature). Continuous digesters are
vided by 35 days, i.e. the time required to degrade best suited to situations where there is a constant
organic matter). When loading the digester with production of material for biodigestion, i.e. if
221 litres, the same amount of biofertilizer is pro- cladodes are collected throughout the year. They
duced, which can be used for irrigation, fertiliza- are also suited to small properties where house-
tion and organic matter input (5.20 g N kg−1, 3.90 hold waste can be added as a raw material − for
g P kg−1, 3.60 g K kg−1 and 561 g Mo kg−1 DM). example, incorporating faeces produced by the
farm animals or through a connection between
the bathroom and digester (Varnero and García
BIODIGESTERS DESIGN de Cortázar, 2006; FAO, 2011). Three models of
AND OPERATION continuous digester are available:
− Taiwan type, made of plastic sleeves (polyeth-
The biodigester must have certain characteristics: ylene) (Figure 4a);
• Airtight − to prevent both output of undesirable − Indian type (Figure 3a) − gasometer included
gas and intake of unwanted air. in the digester in the form of a floating bell;
• Thermally insulated − to avoid major tempera- and
ture changes. − Chinese type (Figure 3b) − closed, with gas
• Fitted with safety valve. accumulation at the top, while the Indian
digester.
• Easily accessible − for loading and unloading
of the system with raw material, removal of • Batch. Discontinuous digesters (Figure 3c)
digester scum and maintenance of the digester comprise a sealed battery of tanks or deposits,
(Varnero, 1991, 2001). with a gas outlet connected to a floating gas-
ometer, where the biogas is stored. With multi-
There is extensive information available in various
ple digesters, one is always loading or unloading
countries, including India, China and Germany,
while the others are in biogas production. Feed-
on the design of biodigesters (García de Cortázar
ing or charging the digester with the raw mate-
and Varnero, 1995). While most production and
rial, which has a higher concentration of solids
use of biogas is obtained from family biodigesters
(40−60%), is done only once, since there is no
(Figure 1), community digesters may also be fea-
recharging during the fermentation process.
sible in some situations, in particular when large
The stabilized organic material is discharged
amounts of raw material and technological exper-
once the biogas production is complete. Biogas
tise are available.
production has an initial waiting period, during
There are two types of digester: continuous and which fermentative hydrolysis, organic acid
batch (discontinuous). formation and methane formation take place.
Most of the biogas production then occurs, be-

a) Biogas Figure 3
b) c)
a) Indian digester;
b) Chinese digester;
c) Batch digester

Biogas
 192 Crop ecology, cultivation and uses of cactus pear

fore slowing down and eventually decreasing to ECONOMIC ASPECTS

almost zero, as the batch−loaded materials run
out. The total duration of the process depends The initial cost of biogas production in rural
on the temperature. The discontinuous system households is around US$50 per biodigester
is suited to certain situations, for example, (Bui Xuan An et al., 1999). This cost is recovered
when: raw materials exhibit handling problems within 9−18 months through savings in fuel costs.
in continuous systems; materials are difficult to In rural areas where the main fuel is wood, the
digest by methanogenic fermentation; or raw use of biogas reduces ecosystem damage (less
materials are available intermittently. Raw ma- deforestation and contamination) and leads to
terial from the cladode harvest is available once time savings of up to 5 hours a day per house-
or twice a year (Varnero and García de Cortázar, hold − time which can be used for other more
2006; FAO, 2011). productive tasks (Rutamu, 1999). To calculate the
economics of using biogas, it is assumed that one
Under optimal conditions and for the same vol- pound (0.45 kg) equals 1 m3 of biogas; therefore a
ume of dry matter, the two types of digester theoretical calculus of 3.61 m3 day−1 would corre-
produce the same amount of biogas. Therefore, spond to 3.61 pounds (1.63 kg) of gas daily, with
the choice should be based on the frequency of a value of approximately US$2.98 day−1, or US$1
waste production (in this case, cladodes) and the 078 year−1.
availability of water.
The residue obtained from digestion processes
For small and medium−sized producers, a wide (Figure 5b) has a high nutrient content; it is,
range of materials may be used for constructing therefore, a valuable fertilizer and allows to save
a biogas digester. The most economical continu- on the expense of commercial fertilizers. Accord-
ous types are made from low−cost polyethylene ing to Varnero (1991), 1 tonne of biofertilizer is
tube (or EPDM, PVC, HDPE) as shown in Figure equivalent to 40 kg of urea, 50 kg of potassium
4a. Known as the Taiwan type, it is widespread nitrate and 94 kg of triple superphosphate. In-
in Asia and some Latin American countries. The ternational fertilizer prices vary from US$255 to
material costs for this type are US$7 m−3. The Indi- US$380 tonne−1 (Indexmundi, 2015). Assuming
an or Chinese models can be made with different an average price of US$0.32 kg−1 of fertilizer, each
materials (Figures 4b−f). tonne of biofertilizer saves US$58.8 on fertilizers

Figure 4
Different materials used
for the construction of
biodigesters:
a) plastic sleeves;
b) brick;
c) concrete;
d) recycled plastic
drums;
e) recycled metal
drums;
f) prefabricated
a b c

d e f
Biogas production 193

Figure 5
a) Solid digestate;
b) biofertilizer (biol
or liquid digestate)

a b

costs; this saving is in addition to the important fore adapts better to a larger scale, given the high
contribution in terms of microorganisms and or- investment costs, and produces concentrations of
ganic material, as well as the possibility of obtain- > 98% ethanol. At fermentation, specific yeast is
ing solid materials when emptying the digester required to maximize the alcohol production. The
(Figure 5a). ethanol concentration at fermentation is 8−12%
(García de Cortázar and Varnero, 1995), attain-
able only by distillation to achieve the required
OTHER BIOENERGY USES concentration of ethanol as fuel.

Cactus cladodes have other bioenergy uses, such Estimates indicate that cactus mucilage can be used
as biodiesel or ethanol production. With an an- to produce small amounts of ethanol: about 20 ml
nual production of 40 tonnes ha−1 in crops grown kg−1 of mucilage. On the other hand, 8.6 litres were
specifically for energy use or 10 tonnes ha−1 in produced from 100 kg of dried cladodes, and 24.7
pruning waste from fruit plantations, energy can litres from 100 kg of dried fruits, and so it is not
be obtained through direct burning. The cladodes considered competitive compared with production
are harvested, sun−dried and crushed, then used
from fermented fruits. With a density of 635−5
in direct burning or cogeneration mix coal−fired;
000 plants ha−1, if only the cladodes are used (Re-
the calorific value is 3 850−4 200 kcal kg−1.
tamal et al., 1987), an average of 300−3 000 litres
The technology for ethanol production is more of ethanol can be obtained from non−irrigated and
complex than that for biogas production; it there- irrigated plantations, respectively.
 17

Marketing
and communication
constraints and strategies
Marcos Mora
Department of Agrarian Economy, Faculty of Agronomy
University of Chile
17 Marketing and communication
constraints and strategies

INTRODUCTION importance of cactus pear: natural cactus pear stands

give subsistence to an entire population in months or
The aim of this chapter is to analyse agribusiness strat- years of food scarcity.
egies applied to cactus pear and derived products and In most producing countries, cactus pear has de-
to examine business issues affecting their develop- veloped in arid and semi−arid zones (Inglese et al.,
ment. The records reviewed are from various sources, 1995b; Russell and Felker, 1987b). It is a marginal crop
but are mainly of Mexican, Italian and Chilean origin. in the fruit market, but in Mexico it produces employ-
However, it should be noted that the product informa- ment and income in areas where few other crops can
tion is scarce, and moreover, it is not up to date, as in- be produced (Timpanaro et al., 2015b). The crop is
dicated by Inglese et al. (2002a). Consequently, Chile often located in areas of high "rurality", cultivated by
is the main focus of this chapter, and the analysis is small and micro−farmers; this makes it attractive from
supplemented and discussed with literature associated a strategic point of view and it should be seriously
with the major producer countries, Mexico and Italy. considered in public policy development actions.
According to the Foundation for Agrarian Innovation Moreover, with regard to sales and marketing, the
(FIA, 2010), many countries throughout the world fresh fruit is marketed mainly in the domestic fresh
produce cactus pear, and Mexico is the world's largest market with small and isolated incursions into export
producer (currently close to 70 000 ha). Other impor- markets. Italy, the second producer worldwide, is the
tant cactus−pear−producing countries are Italy, South leading exporter of cactus pear. In the Italian island
Africa, Argentina, Chile, Bolivia, Peru, Colombia, of Sicily, numerous festivals take place around harvest
United States of America, Morocco, Algeria, Libya, time (October−November) in villages, including San
Tunisia, Egypt, Jordan, Pakistan, Israel, Greece, Spain Cono, Biancavilla, S. Margherita Belice and Rocca-
and Portugal. However, in a significant number of palumba (Sáenz et al., eds, 2006).
these countries, especially in Africa, cactus pear fruits
are considered a by−product, as the plant is mainly
used for the conservation of degraded soils. There is PRODUCTION AND MARKETING
potential for development of cactus pear through a BACKGROUND
wide range of applications, including:
• cultivation as forage complement, of which Brazil is Mexico is the largest producer, accounting for about
the main producer (Callejas et al., 2009); 80% of world production of cactus pear, estimated
• consumption as vegetable (cladodes) and fresh fruit; at about 500 000 tonnes. Italy (12.2%) is the second
largest producer, and South Africa (3.7%) ranks third.
• medicinal uses;
These three countries account for approximately 96%
• industrialization for processed food (e.g. cladode of world production. Average yields of cactus pear
flour, jams and juices); vary enormously: from 6.5 tonnes ha−1 (García et al.,
• non−food industrialization (e.g. bioenergy and cos- 2003) in Mexico to 20 tonnes ha−1 in Italy and 25
metics); and tonnes ha−1 in the United States of America and Israel.
• carmine production. In Chile, cactus pear is traditionally cultivated between
Arica and Parinacota and the Bío−Bío regions; how-
Cactus pear also plays an important role in subsist- ever, more recently it has also been grown between
ence agriculture in many areas of the world. Whether Atacama and Maule, mainly to meet domestic con-
cultivated on a small scale or in natural stands, cactus sumption demand. There is a strong concentration
pear is a crucial source of food (fruits and its deriva- of the crop in the central zone of Chile. The area
tives) for the rural poor, as well as fodder or forage covered by cactus pear in Chile has declined to 800
for their livestock. This is the case, not only in North ha and according to the Office of Agricultural Studies
Africa and the Horn of Africa (Eritrea, Ethiopia), but and Policies (ODEPA, 2015a) and Franck (2010), it is
in Mexico, the Near East and the Americas. While it concentrated in regions close to large cities, such as
is not possible to give an economic dimension to this Santiago, Viña del Mar and Valparaíso (Table 1), and
phenomenon, northern Tigray (Ethiopia) conveys the in Atacama and Maule. Sicily accounts for over 96%
Marketing and communication constraints and strategies 197

of the total Italian cactus pear harvest, with 8 300 ha Sicily, 15−20% exported within Europe and a very small
producing about 87 000 tonnes annually (Timpanaro fraction exported to Canada (Timpanaro et al., 2015b).
et al., 2015b). North Africa fails to reach the European markets in ear-
ly summer (June−July), when no fruits are available in
The value addition and creation of derivatives for com-
Italy and high temperatures during the summer harvest
mercial purposes is minimal. Nevertheless, there has
affect post−harvest management and exports. Mexico
been some important research with the development
produces fruits from June to October, with very modest
of agro−industrial products from cactus pear (Sáenz
exports to the United States of America and Canada. In
et al., eds, 2006). On the other hand, a quite different
the Southern Hemisphere, Argentina and South Africa
dynamic is present in Italy with cultivation increasing
produce from January until March and only for local
over the past 20 years. Nevertheless, development is
markets. Rarely do they adopt the Italian practice of
limited for various reasons, including limited supply,
scozzolatura (removal of the first flush of flowers and
lack of coordination throughout the value chain supply
cladodes) to obtain a second, out−of−season harvest.
and scarcity of resources to boost research, innovation
Chile, on the other hand, produces two crops with no
and cultivation techniques (Timpanaro et al., 2015b).
technical help, the second one in June−September.
The average yield of a cactus pear plantation in Chile
Overall, cactus fruit comes on the market seasonally;
is around 10 tonnes ha−1. However, with modern culti-
however, with better coordination among the produc-
vation practices, yields could more than double.
ing countries and accurate post−harvest management,
The most popular cactus pear ecotype is a local se- cactus fruit could be supplied to markets at almost any
lection with yellow−green flesh (CEZA, 2011). There time during the year, as is the case with kiwi fruit.
are, however, other coloured Opuntia ficus−indica
In Chile, the main market for cactus pear is represent-
genotypes with purple, orange, yellow, red and white
ed by wholesale traders (Lo Valledor and Mapocho
fruits (Aquino et al., 2012). These ecotypes are only
Fair), who then sell to retailers. According to Mora et
produced at experimental level in Chile (Sáenz and
al. (2013), wholesalers account for about 60% of cac-
Sepúlveda, 2001b). Marketing of cactus pear is atom-
tus fruit sales. Another important outlet, accounting
ized at producer level and oligopsonic/oligopolistic
for approximately 18% of produce, are informal shops
at wholesale level; this generates information asym-
located on street corners and at traffic lights. Finally,
metries between producers and buyers and increases
supermarkets account for a mere 2%. Prices have a
the market power of buyers, hindering the develop-
slightly upwards trend generally. During the year,
ment of producers in Chile. In many cases, it means
prices rise when smaller volumes reach the wholesale
producers cannot sell directly or establish business
markets and vice versa. There is also a direct relation-
contacts with other buyers. Consequently, producers
ship between size and price. It should be noted that
wait for the same buyers to come every year, bringing
during winter (June−September in the Southern Hemi-
their own workers for the harvest. A decade ago, the
sphere), a small quantity of cactus pear tends to reach
same situation existed in Italy (Inglese et al., 2002a):
prices well above the seasonal average (Figure 1).
most of the crop was sold by the farmer in the field,
with limited participation of the producer in the value Table 2 shows the unit value per kg of cactus pears
chain. Nowadays, farmers’ organizations are more for export by air – around US$1.9 in 2011. If export
sophisticated; farmers get higher incomes and make costs (approximately US$0.6 kg−1) are subtracted (free
a greater impact in the market. One of the most ef- on board, FOB), the unit value is around US$1.3 kg−1
ficient examples is Euroagrumi s.c.c., a consortium (based on the September 2015 exchange rate of 685.6
operating in Sicily and trading 1 500 tonnes year−1 of Chilean pesos = US$1). While this is an attractive fig-
cactus pears to a value of €2.5 million. ure, it is difficult to bring together large volumes of
export quality cactus pears, since only a very small
Fruit production is from late July to late November in the
fraction reaches this quality. In contrast, the domestic
Northern Hemisphere, plus a small winter production in
market pays an equivalent of about US$0.6 kg−1, i.e.
Israel in January−March, mostly destined to the home
almost half the export price.
market. Italy concentrates its production in October−
November, with > 60% of the market concentrated in
198 Crop ecology, cultivation and uses of cactus pear

TABLE 1 Chile: Producer number and cactus pear area

Region/Variety Number Total (ha)

Atacama Region

‘Chilena’ 2 5.18

Total 2 5.18

Coquimbo Region

‘Blanca’ 1 14.09

‘Chilena’ 7 69.01

No information 2 2.57

‘Til−Til’ 1 3.91

Total 11 89.58

Valparaíso Region

‘Blanca’ 3 1.73

‘Chilena’ 58 65.75

No information 6 6.32

‘Til−Til’ 2 8.72

Total 69 82.52

Metropolitana Region

‘Chilena’ 22 228.11

‘Nopal De Castilla’ 4 29.30

No information 6 20.60

‘Til−Til’ 50 306.22

‘Undulatta Griffiths’ 1 2.84

Total 587.07

L.B. O'Higgins Region

‘Chilena’ 1 0.20

No information 1 7.00

‘Til−Til’ 1 18.00

Total 3 25.20

Maule Region

‘Chilena’ 1 5.00

‘Til−Til’ 1 5.00

Total 2 10.00

Total country 799.55

Source: ODEPA−CIREN, 2015a, b, c; ODEPA−CIREN, 2014, 2013.

 Marketing and communication constraints and strategies 199

Price
in chilean Trade
currency volume
(Box 20 kg) Average Price in Chilean Currency Kg (kg)
25.000 9.000 Figure 1
Price and trade of
8.000 cactus pear (high
quality) in principal
20.000 7.000 wholesale markets,
Chile
6.000 Source:
15.000 ODEPA (2015b).
5.000

4.000
10.000
3.000

2.000
5.000
1.000

0 0
/2010
/2010
/2010
/2010
/2011
/2011
/2011
/2012
/2012
/2012
/2013
/2013
/2013
/2014
/2014
/2014
/2014
/2014
/2014
/2015
/2015
/2015
/2015
/2015
/2015
/2015
02
03
05
09
03
08
11
03
08
09
01
04
08
01
02
03
04
05
09
01
02
03
04
05
07
08
Time (month/year)

TABLE 2 Fresh cactus pear exports by air from Chile

Year FOB Volume (kg) FOB/US$ Country for export and transport type

2002 53 795 23 641 2.3 Canada, USA

2003 37 829 18 766 2.0 Japan, USA

2004 24 252 12 423 2.0 Saudi Arabia, USA

2005 29 497 17 152 1.7 Canada, USA

2006 54 626 17 168 3.2 Saudi Arabia, USA rest

2007 75 963 20 172 3.8 Spain, USA rest

2008 42 173 12 138 3.5 Spain, UK, USA rest

2009 28 779 11 201 2.6 Spain, USA rest

2010 0 0 −

2011 21 738 11 362 1.9 USA

2012 0 0 −

2013 0 0 −

2014 0 0 −

Source: ODEPA (2015b).

200 Crop ecology, cultivation and uses of cactus pear

PERCEIVED QUALITY IN CACTUS fibre, hydrocolloids (mucilage), pigments (betalains and

carotenoid), minerals (calcium, potassium) and vitamins
PEARS: FROM INTRINSIC TO (e.g. vitamin C), which are present in the fruit and the
EXTRINSIC ATTRIBUTES cladodes (Piga, 2004; CEZA, 2011).

According to Olson and Jacoby (1972), quality attributes Most consumers of fresh cactus pear fruit focus on the
may be described as: either extrinsic − related to the intrinsic attributes, because there has been practically
product (e.g. price, colour, labelling and packaging); or no development of the extrinsic attributes. Intrinsic
intrinsic − tied to the physical attributes of the product signals are related to the physical aspects of a product,
(e.g. nutrient composition and flavour). As Grunert et al. such as colour, size, texture, shape and appearance
(1996) asserts, quality is a multidimensional phenome- (Mora et al., 2011; Sulé et al., 2002).
non, described by a set of attributes that are subjective- Extrinsic signals are related to the product, but are not
ly perceived by consumers before (quality expectation) part of its essence, for example: brand, packaging, seal
and after (quality experience) the purchase. Credence of quality, price, PDO, shop, sales force and production
attributes, however, increasingly influence consumer information. Migliore et al. (2015a) report that important
preferences in response to rising concerns about safety, attributes of credibility for the Italian market are environ-
health, convenience, locality, ethical factors and the mental sustainability, healthiness and Italian origin.
environmental. (Migliore et al., 2012, 2015a; Bernués et
al., 2003).
The marketing of cactus pears in Chile and the main VALUE ATTRIBUTES AND
producing countries has focused on the fresh fruit, CONSUMER PREFERENCES
highlighting its essentially intrinsic characteristics. Chile
has seen the commercial development of containers, la- In Chile, there are 17 ecotypes of the genus Opuntia,
bels, brands or denominations of origin, but none have probably introduced from Mexico by the Spaniards;
succeeded as formal commercial agents. In Italy, two the green variety (Opuntia ficus−indica) is the most
protected designation of origin (PDO) labels are active: commercialized at present (Sudzuki et al., 1993). In this
• “Ficodindia dell’Etna” refers to production in the area context, a study of cactus pear preferences developed
of the Etna volcano, where the white flesh ‘Trunzara by our group (Matamala et al., 2015) focused on the
Bianca’ is the most important Italian cultivar, because study of six ecotypes of different colours, and evaluated
of its firm flesh and early ripening time in summer intrinsic attributes, such as weight, number of seeds,
(July−August). peel thickness, internal and external colour, and sweet-
• “Ficodindia di San Cono” refers to the most important ness. According to Mokoboki et al. (2009), a weight of
cactus pear cultivation area in Italy where the three around 120 g is a positive attribute on the market. The
main cultivars grow (‘Gialla’, ‘Rossa’ and ‘Bianca’). thickness of the shell and the seeds also have commer-
cial impact (FIA, 2010). Green fruits are preferred by
According to FIA (2010), the agro−industrial potential the Chilean population, while yellow cactus pear is the
of countries such as Mexico and Italy has enabled the most consumed worldwide and red is the most attrac-
development of a variety of alternatives for the agrifood tive to first−time consumers (Migliore et al., 2015a).
sector, for example, flour, tortillas, jams, supplements,
confectionery, snacks, frozen products and juice. In It- Matamala et al. (2015) identify three preliminary market
aly, however, the sector is limited to some liqueurs and segments: i) traditional, regular consumption of cactus
− on a very small scale − jams, while the interest in and pear; ii) sporadic, occasional consumption; and iii) skep-
commerce of by−products for nutraceutical and medici- tical about the characteristics of the product. Esparza
nal uses is undergoing rapid expansion. Sáenz et al., eds (2015), on the other hand, identifies three consumer
(2006) point out that there are a significant number of market segments: i) "practical" (48.8% of the surveyed
products that can be obtained from the various parts population); ii) "essential" (18.5%); and iii) "healthy"
of the plant, and even more by exploiting the colour- (13.5%). In general, price was the most important at-
ed fruit ecotypes. The fruit can be used to prepare a tribute, followed by place of origin and lastly, by the
large number of products, including jams, liqueurs colour of the flesh. There also exists a potential market
and vinegars, sauces, juice concentrates and canned segment attracted to the purchase of minimally pro-
products. Cladode powders, on the other hand, can cessed cactus pear.
be used as functional foods and colouring ingredients, Migliore et al. (2015a, b) report similar findings in the
while other encapsulated bioactive compounds act as Italian market. Inglese et al. (1995b) indicate that the
colouring agents (Sáenz et al., 2009). More specifically, challenge for researchers is related to reduced fruit size
many functional ingredients are characterized by their and the high number of seeds, together with poor pro-
potential beneficial effects on health, thanks to the motion and issues of quality standardization. Moreover,
Marketing and communication constraints and strategies 201

there are other attributes related on an emotional level: particularly in public catering and in the refrigerated
moods and emotions are factors that constitute the section of supermarkets. Research is required to explore
motivational bases of the purchasing process. With re- the market potential of other cactus pear derivatives,
gard to cactus pear, Chironi and Ingrassia (2015) report such as jams, flour, juices, concentrates and functional
three consumer profiles. The first analyses cactus pear, products. Finally, for a quality product preferred by con-
not only for the qualitative and sensory elements, but sumers, innovation of the primary activities in the value
also in relation to new hedonistic needs. The second chain is required: it is necessary to improve cultivation
associates the image of the fruit with the territory. The techniques, with particular attention to organic practic-
third allows consumers to develop secondary needs, es (Timpanaro et al., 2015b; Migliore et al., 2015a).
increasing the chances of overcoming the limitations
related to this fruit. Promotion and advertising

In general, the cactus pear is a product known and

consumed by a specific part of the market. A commu-
PROPOSED COMMERCIAL DEVELOPMENT nications strategy must, therefore, present the intrinsic
STRATEGY FOR FRESH CACTUS PEAR attributes − functional ingredients with beneficial ef-
AND ITS PRODUCTS fects on health, such as fibre, pigments (betalains and
carotenoids), minerals (calcium, potassium) and vitamins
Callejas et al. (2009) provide guidelines for a better (e.g. vitamin C) − and make them known to a wider
product in terms of aspects, quality, added value and public. Communication actions associated with public
competitive price. policy are essential and are recommended for cactus
pear and its derived products.
Price

This variable depends a lot on doing things the right CONCLUSIONS

way; however, in a competitive value chain, cactus pear
prices are potentially attractive. Increasing competitive- Cactus pear producers throughout the world face simi-
ness through improved performance quality (based on lar challenges with regard to improving the value chain.
a yield of around 12 tonnes ha−1, i.e. 600 boxes − 20% The main issues are outlined below:
more than current production in a plantation in central
Chile) would result in an estimated average price of 6 • Irrigation systems are required at production level, as
000 Chilean pesos (US$8.75) per box premium (lowest cactus pear grows in places characterized by water
wholesale price offer is projected). This would generate shortages.
an income of 3.6 million Chilean pesos (US$5 251) ha−1. • Fruit quality must be improved, with particular attention
to the intrinsic attributes associated with homogeneity.
The Italian experience indicates that fruit of the best
size (classified as A or AA) may reach prices of €1.5−3.0 • Marketing strategies are needed. As cactus pear
at farm level, while smaller sizes show a sharp decrease producers are generally small and lack the resources
in price, down to 30−40% for class B fruits and even to access modern technologies, it is recommended
less for class C fruits. Therefore, only with proper crop that they develop associations to improve organiza-
management is it possible to reach the best results in tion, optimize resources and deal with the significant
the market, by cultivating class A size cactus pear fruits, asymmetries and failures inherent in markets.
totally without glochids and defects, harvested at the
• More up−to−date information needs to be made
appropriate ripening stage and with an adequate pro-
available, especially with regard to markets, costs,
portion (55−65%) of flesh (Inglese et al., 2002a).
prices and quality. Major producing countries could
develop a joint project for the generation of a data-
Product
base with standardized information and economic
and business updates.
Quality as perceived by consumers depends mainly
on the intrinsic attributes of the fruit. Certain aspects • The social role played by cactus pear in its production
require improvement to achieve, for example, uniform areas must be recognized. Public policies should be
size, thin epidermis and fewer seeds. For fresh cactus implemented to help associations improve productive
pear fruit in Chile, the preference is for green and and economic results and to generate innovation
yellow cultivars. Yellow (Inglese et al., 2002a) and red throughout the value chain, in both primary and sec-
cactus pears (Migliore el al., 2015a) are the most appre- ondary activities. The aim is to improve the level of
ciated in the Italian market. Moreover, there is a grow- competitiveness and welfare of the people living in
ing market space for minimally processed cactus pears, those territories.
202 Crop ecology, cultivation and uses of cactus pear

• Market research is required to understand the po- trial), bring them to prototype stage and continue
tential interest in products prepared from cactus testing and market validation.
pear and cladodes. Many interesting attributes could − Communicate the intrinsic attributes that create val-
be well received in the market. Quality assurance ue but which the average consumer is not aware of.
systems must be installed for processed products, as
they are essential tools for dealers selling such prod- − Explore new commercialization channels, such as
ucts. speciality shops, catering businesses, pharmaceuti-
cal companies and the Internet.
• Development of the intrinsic attributes of cactus
pear is at the centre of any marketing strategy: − Continue research to decrease the number of
seeds.
− Improve current systems of production (productivi-
ty and quality). − Develop minimally processed cactus pears.
− Carry out economic and technical feasibility studies − Elaborate and communicate credence attributes
to assess the production and trade potential of with labelling of food safety, quality, nutritional
both domestic and international markets. and environmental aspects.
− Explore consumer receptivity for new products • Development of the extrinsic attributes (packaging
developed by research centres (mainly agro−indus- and labelling) is also essential for added value.
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