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Introduction To Plant Nutrition

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Chapter 1
Introduction to Plant Nutrition

Keywords  Criteria essentiality · Nutrient accumulation · Nutritional requirement ·


Nutritional efficiency · Hydroponic cultivation · Silicon

The introduction to plant nutrition addresses basic and general topics on the impor-
tance of this area to meet nutritional requirements and promote crop growth, devel-
opment, and yield. We will address important topics, such as (1) concepts of plant
nutrition and its relationship with related disciplines; (2) the concept of nutrient and
criteria of essentiality; (3) relative composition of nutrients in plants; (4) nutrient
accumulation by crops and crop formation; (5) other chemical elements of interest
in plant nutrition, such as potentially toxic and beneficial elements, with emphasis
on silicon; and (6) hydroponic cultivation, preparation, and use of nutritional
solutions.

1.1  C
 oncepts of Plant Nutrition and Its Relationship
with Related Disciplines

Current knowledge of the concept of plant nutrition is historically recent. We will


briefly present some occurrences throughout the history of plant nutrition. The first
inference on some aspects of plant mineral nutrition started in antiquity, when
Aristotle (384–322 BC), the Greek philosopher and biologist, already made state-
ments about how plants fed. At that time, he indicated that plants were like inverted
animals, with their mouths on the ground. For him, food would be previously
digested by the soil, as vegetables did not show visible excretions, as animals do.
Investigations on the ways through which plants fed continued, and new discoveries
were found at each stage of human history.
In the nineteenth century, the Swiss researcher De Saussure (1804) made an
important publication establishing that the plant obtained C from CO2 of the atmo-
sphere, with hydrogen and oxygen being absorbed along with carbon. This publica-
tion also established that plant dry matter increased mainly due to the absorbed C,
H, and O, and that the soil supplied minerals indispensable to plant life. In that same

© Springer Nature Switzerland AG 2021 1


R. de Mello Prado, Mineral nutrition of tropical plants,
https://doi.org/10.1007/978-3-030-71262-4_1
2 1  Introduction to Plant Nutrition

century, the chemist Justus von Liebig (1803–1873), “father of plant mineral nutri-
tion,” established in Germany that foods of all green plants are inorganic or mineral
substances. This study was presented at an event of the British Association for the
Advancement of Science and resulted, in 1840, in the publication of the book
Organic Chemistry in its Application to Agriculture and Physiology. Liebig, with
his dominant vigor, convinced the scientific community of the time with his theory,
although being a compilation of studies by other authors (De Saussure, Sprengel,
and others) (Browne 1942). Sprengel (Professor of Agronomy) published a study in
1826 recognizing 20 elements as nutrients, with macronutrients among them.
According to Epstein (1975), Liebig’s main contribution to plant nutrition was to
end the humus theory that believed soil organic matter was the plant carbon source.
According to Liebig’s theory, plants lived on carbonic acid, ammonia (azotic acid),
water, phosphoric acid, sulfuric acid, silicic acid, magnesian lime, caustic potash,
and iron. Thus, throughout the end of the nineteenth century, the classic list of plant
nutrients consisted basically of N, P, S, K, Ca, Mg, and Fe, defining the requirement
of plants, especially regarding macronutrients and iron. At that time, Liebig contrib-
uted to the rise of fertilizer industries. The twentieth century saw the establishment
of the concept of micronutrients, nutrients equally essential, but required in smaller
quantities by plants. A new era for plant nutrition began in the 1930s–1950s with
Hoagland, who determined the ideal nutrient solution for plant growth. Afterward,
modern scientists wrote classic books on plant nutrition, such as Epstein (1972),
Mengel and Kirkby (1987), Marschner (1986), and Malavolta (1980) in Brazil. The
establishment of the first teaching and research institutions in Brazil took place in
the late 1800s (Federal University of Bahia (UFBA) in 1877, the Agronomic
Institute of Campinas (IAC) in 1887, and the Luiz de Queiroz College of Agriculture
(ESALQ) in 1901), establishing the basis for plant nutrition studies, which began in
the 1950s.
Although plant nutrition is a new science, with only 180 years old, it advanced
extraordinarily from the demystification of the humus theory, in 1840, to recent
discoveries regarding nutrient absorption through identification of genes that encode
proteins (carriers).
Thus, the study of plant nutrition establishes what are the essential elements for
the plant life cycle, how they are absorbed, transported, and accumulated, and how
the plant redistributes its functions, requirements, and disturbances when nutrients
are in deficient or excessive quantities.
It is clear that plant nutrition has links from nutrient acquisition by the roots,
related to soil science, to the functions nutrients perform in plants, and to aspects
studied by plant biochemistry and physiology. More broadly, plant nutrition and
agronomy are closely related, as it is known that the main objectives of agronomic
science are the production of food, fibers, and energy. For this, there are more than
50 factors of production that must be considered for maximum efficiency in agricul-
tural production systems. These factors of production are arranged in three major
systems, namely: soil, plant, and environment. The plant nutrition area centers on
the plant system, like other areas (plant physiology, molecular biology, plant breed-
ing, and phytotechnics). The areas of soil fertility, fertilizers/correctives, and
1.2  Concept of Nutrient and Criteria of Essentiality 3

fertilization, among others, are centered on the soil, while irrigation, drainage, and
climatology are centered on the environment. Most factors of production can be
controlled in the field by the producer. However, some are difficult to control, such
as light and temperature.
Environmental factors are highlighted with climate change, especially due to
increasing air temperature, CO2, and water irregularity (flood and drought), affect-
ing plant nutrition and production (Viciedo et  al. 2019a, b; Barreto et  al. 2020;
Carvalho et  al. 2020a), and forage quality (Habermann et  al. 2019). However,
increased CO2 and temperature can increase nutritional efficiency and plant growth
in certain species, such as Stylosanthes capitata Vogel (Carvalho et al. 2020b). In
this species, warming combined with well-watered conditions increased leaf bio-
mass production by 38%, presumably due to a higher level of stoichiometric homeo-
stasis (Viciedo et al. 2021).
Plant nutrition is closely related with agronomy, specifically with the disciplines
of soil fertility, fertilizers/correctives, and crop fertilization. Fertilization = (QP –
QS). factor f; where QP = nutrient amount required by the plant (nutritional require-
ment); QS = nutrient amount in the soil; and f = fertilizer efficiency factor; which
can be reduced by losses in the soil (volatilization, adsorption, leaching, erosion,
etc.). In a cultivation system with soil tillage, efficiency factors of 50%, 30%, and
70% are admitted for N, P, and K, respectively, corresponding to the f values equal
to 0.50, 0.30, and 0.70, respectively. The use of twice as much N; 3.3 times more P;
and 1.4 times more K in fertilization ensures adequate plant nutrition.
In addition, nutrient cycling increases only in a consolidated no-tillage system
with live roots throughout the crop year. This improves efficiency factors for all
nutrients and, in the case of phosphorus, may double its efficiency.
There are other areas related to plant nutrition, such as microbiology, plant
breeding, and molecular biology, among others.

1.2  Concept of Nutrient and Criteria of Essentiality

There are many chemical elements in nature without considering isotopes, as stated
in the periodic table, with more than a hundred chemical elements. This number
may increase with new discoveries by science, which can occur even by synthesis in
laboratory.
However, when plant tissue is chemically analyzed, it is common to find approxi-
mately 50 chemical elements, and not all of them are considered plant nutrients.
Plants have the ability to absorb chemical elements in the soil or nutrient solution
with little restriction, which could be a nutrient, nonmineral nutrients from the
atmosphere, or a beneficial and/or toxic element (Kathpalia and Bhatla, 2018,
Fig. 1.1).
Considerations regarding the beneficial and/or toxic element are addressed in the
next item.
4 1  Introduction to Plant Nutrition

Fig. 1.1  Distribution of essential and beneficial mineral elements. All, except molybdenum, are
among the 30 lightest elements. Elements in bold letters are hyperaccumulators in plants

The nutrient is defined as a chemical element essential to plants, that is, without
it the plant cannot complete its life cycle. For a chemical element to be considered
a nutrient, it is necessary to meet the two criteria of essentiality, direct and indirect,
or both. The criteria were proposed by Arnon and Stout (1939), physiologists at the
University of California, thanks to the advancement of science regarding analytical
chemistry, allowing the determination of trace elements, and to the advancement of
cultivation techniques in nutritive solution. The essentiality criteria are
described below:
Direct:
• The element participates in some compound or reaction, without which the
plant does not live.
Indirect:
• In the absence of the element, the plant cannot complete its life cycle.
• The element cannot be replaced by any other element.
• The element, with its presence in the medium, must directly affect plant life
and not only play the role of neutralizing physical, chemical, or biological
effects unfavorable to the plant.
Epstein and Bloom (2006) proposed an adaptation of the criteria of essentiality,
that is, an element is essential if it meets one or both of the following criteria:
• The element is part of a molecule that is an intrinsic component of the plant
structure or metabolism.
• The plant is so severely deprived of the element that it causes abnormalities in
growth, development, or reproduction – that is, its performance – compared to
plants without its deprivation.
1.2  Concept of Nutrient and Criteria of Essentiality 5

The world literature considers 17 chemical elements as plant nutrients, namely:


C, H, O, N, P, K, Ca, Mg, S, Fe, Mn, Zn, Cu, B, Cl, Mo, and Ni. Nutrients are impor-
tant for life as they play significant roles in metabolism, either as substrate (organic
compound) or as enzymatic systems. Briefly, such functions can be classified as
follows (Malavolta et al. 1997):
• Structural (part of the structure of any vital organic compound for the plant).
• Constituents of enzymes (part of a specific structure, prosthetic group, or active
site of enzymes).
• Enzyme activator (not part of the structure). It should be noted that the nutrient
not only activates but also inhibits enzymatic systems, affecting the speed of
many reactions in plant metabolism.
Epstein and Bloom (2006) proposed another classification of nutrients with more
detail, organized by their functions in plants, as follows:
• Nutrients that are integral elements of organic compounds, for example, N and S.
• Nutrient for acquisition and use of energy and for the genome, for example, P.
• Nutrients structurally associated with the cell wall, for example, Ca and B (Si).
• Nutrients that are integral compounds of enzymes and other essential entities of
metabolism, for example, Mg, Fe, Mn, Zn, Cu, Mo, and Ni.
• Nutrients that activate or control enzyme activity, for example, K, Cl, Mg, Ca,
Mn, Fe, Zn, and Cu (Na).
• Nonspecific functions: nutrients that serve as counterions, for positive or nega-
tive charges, for example, K+, NO3-, Cl-, SO4-2, Ca+2, Mg+2, and (Na+).
• Nonspecific functions: nutrients that serve as osmotic agents of cells, for exam-
ple, K+, NO3-, Cl-, and (Na+).
The next chapters discuss the role of each nutrient after reaching the location
where they exercise their various functions, the cell (cell walls, cytoplasm, and
vacuole).
When a nutrient performs its function in the plant, that is, the integration of bio-
chemical functions, it affects one or several physiological processes (photosynthe-
sis, respiration, etc.), affecting crop growth and production.
Photosynthesis is the most important physicochemical reaction on the planet, as
all life forms depend on it. Organic compounds are synthesized with light (visible
from 400 to 740 nm) by photosynthetic pigments in plants (chlorophylls, carot-
enoids, and phycobilins). However, a small fraction of solar radiation (~5%) reach-
ing the Earth is converted into organic compounds by leaf photosynthesis.
In summary, the physicochemical reaction of photosynthesis occurs in two
phases. During the photochemical or light-dependent phase, sunlight is used to
break down the water molecule (H2O) into oxygen (O2) – conversion of light energy
into electrical energy – which generates chemical energy, with adenosine triphos-
phate (ATP) and nicotinamide adenine dinucleotide phosphate (NADPH) as pri-
mary products. Thus, light energy is captured to allow electron transfer by a series
of compounds that act as electron donors and receivers. Photolysis of the water
molecule and electron transport allow the creation of a proton gradient between the
6 1  Introduction to Plant Nutrition

thylakoid lumen and the chloroplast stroma. Ultimately, most electrons reduce
NADP+ to NADPH. Light energy also generates proton motive force across the
thylakoid membrane, which synthesizes ATP through the ATP synthase complex.
The light-independent phase or photosynthetic carbon reduction cycle is basically
an enzymatic phase, where light is not required, and the primary products from the
previous phase are used to obtain carbon hydrates (Cn(H2O)n), such as glucose,
from carbon dioxide (CO2). The free energy to reduce one mole of CO2 to glucose
level is 478 kJ mol−1.
The photosynthetic process occurs inside chloroplasts (Fig. 1.2), plastids located
in cells of palisade and spongy mesophylls. The number of chloroplasts per cell
ranges from one to more than one hundred, depending on the plant type and grow-
ing conditions. Chloroplasts are discoid shaped with diameter from 5 to 10 microns,
limited by a double membrane (outer and inner). The inner membrane acts as a bar-
rier controlling the flow of organic molecules and ions inside and outside the chlo-
roplast. Small molecules, such as CO2, O2, and H2O, pass freely through chloroplast
membranes. Internally, the chloroplast consists of a complex system of thylakoid
membranes containing most of the proteins required for the photochemical phase of
photosynthesis. The proteins required for CO2 fixation and reduction are located in
a colorless matrix called stroma. Thylakoid membranes form thylakoids, flat vesi-
cles with an aqueous inner space called lumen. Thylakoids, in certain regions, are
arranged in stacks called granum. Thus, the first phase of photosynthesis occurs in
the chloroplast inner membranes, the thylakoids, while the second phase occurs in
the chloroplast stroma, the aqueous region surrounding the thylakoids. Therefore,
the products formed in photosynthesis, carbon sources, are accumulated as sucrose

Fig. 1.2  Scheme of the photosynthetic process responsible for crop growth, development, and
production
1.2  Concept of Nutrient and Criteria of Essentiality 7

in vacuoles and starch in chloroplasts, in order to be subsequently used in photosyn-


thesis, such as in respiration and synthesis of reserves and structural materials.
The photosynthetic process depends on some nutrients acting with structural or
enzymatic function. Moreover, the products formed by photosynthesis also depend
on nutrients to produce other vital organic compounds for plant development and
production, which are detailed in the next chapters.
Thus, we perceive the importance of nutrients for plant life. According to the
history of plant nutrition, these chemical elements, which are currently considered
plant nutrients, were discovered and had their essentiality demonstrated from 1804
until recently, with the last inclusion in 1987 (Table 1.1).
There are disagreements on the author who demonstrated the essentiality of
some nutrients, often due to problems in the scientific rigor of the research, for
example boron, whose authorship of essentiality is attributed to Warington (1923).
Although these nutrients are equally important for plant production, they are
classified based on the proportion in which they are found in the dry matter of veg-
etables. Therefore, there are two major groups of plant nutrients (not considering C,
H, and O), namely:
• Macronutrients – Nutrients absorbed or required by plants in larger quantities: N,
P, K, Ca, Mg, and S (expressed in g kg−1 of dry matter). Macronutrients can be
further divided into primary macronutrients (N, P, and K) and secondary macro-
nutrients (Ca, Mg, and S).
• Micronutrients – Nutrients absorbed or required by plants in smaller quantities:
Fe, Mn, Zn, Cu, B, Cl, Mo, and Ni (expressed in mg kg−1 of dry matter).

Table 1.1  Discovery and demonstration of essentiality of plant nutrients


Nutrients Atomic mass Discoverer Year Demonstration Year
C 12.01 – – De Saussure 1804
H 1.01 Cavendish 1774 De Saussure 1804
O 16.00 Priestley 1774 De Saussure 1804
N 14.01 Rutherford 1772 De Saussure 1804
P 30.98 Brand 1772 Ville 1860
S 32.07 – – Von Sachs, Knop 1865
K 39.10 Davy 1807 Von Sachs, Knop 1860
Ca 40.08 Davy 1807 Von Sachs, Knop 1860
Mg 24.32 Davy 1808 Von Sachs, Knop 1860
Fe 55.85 – – Von Sachs, Knop 1860
Mn 54.94 Scheele 1744 McHargue 1922
Cu 63.54 – – Sommer 1931
Zn 65.38 – – Sommer & Lipman 1926
B 10.82 Gay Lussac and Thenard 1808 Sommer & Lipmana 1939
Mo 95.95 Hzelm 1782 Arnon & Stout 1939
Cl 35.46 Schell 1774 Broyer et al. 1954
Ni 58.69 – – Brown et al. 1987
Researchers defined the essentiality of B for nonleguminous plants
a
8 1  Introduction to Plant Nutrition

In some cases, crops that accumulate certain micronutrients may have higher
contents of this nutrient than of a macronutrient. Prado (2003) found for Averrhoa
carambola that the leaf content of Mn (1.7 g kg−1) exceeded that of the macronutri-
ent S (1.4 g kg−1). Thus, other classification systems for nutrients emerged based not
on the amount accumulated by the plant, but on the (biochemical) role they play in
the plant life. Mengel and Kirkby (1987) classified nutrients into four groups. The
first group consists of C, H, O, N, and S, structural nutrients in the constitution of
organic matter, which also participate in the enzymatic system and in assimilation
in redox reactions. The second group consists of P, B, and Si in some crops, nutri-
ents that easily form connections of ester type (energy transferors). The third group
consists of K, Mg, Ca, Mn, Cl, and (Na), nutrients responsible for enzymatic activ-
ity, maintenance of the osmotic potential, ion balance, and electrical potential, espe-
cially K and Mg. Finally, the last group consists of Fe, Cu, Zn, and Mo, which act
as prosthetic groups of enzymatic systems and also participate in electron transport
(Fe and Cu) to different biochemical systems.
The author highlights that the list of 17 chemical elements considered essential
may increase with the progress of research. There are isolated studies whose authors
indicate certain elements as essential to plants, such as silicon in tomato plants
(Miyake and Takahashi 1978), sodium in Atriplex vesicaria (Brownell and Wood
1957), cobalt in lettuce (Delwiche et al. 1961) and alfalfa (Loué 1993), and Se (Wen
et al. 1988). However, for a chemical element to be included in this list (if it occurs,
it is more likely to be a micronutrient), additional studies are necessary in order to
satisfy the criteria of essentiality for a considerable number of plant species and to
convince the international scientific community. There are strong candidates for
inclusion in the list of nutrients, such as Si, Na (Malavolta et al. 1997), Se, and Co
(Malavolta 2006).

1.3  Relative Composition of Nutrients in Plants

In a freshly harvested plant, depending on the species, we observe that the largest
proportion of its mass, from 70% to 95%, consists of water (H2O). After drying this
plant in a forced air circulation oven (±70 °C for 24–48 h), the water evaporates and
we obtain the dry matter or dry mass; and when the plant is subjected to mineraliza-
tion, either in a muffle furnace (300 °C) or in strong acid, the organic and mineral
components (nutrients) are separated. After analyzing this dry plant material, we
generally observe predominance of C, H, and O, composing 92% of plant dry matter
(Table 1.2).
The results of chemical analysis of plant material are expressed based on the dry
matter for being more stable than fresh matter, which varies according to the
medium, that is, the time of day, soil water availability, and temperature,
among others.
It is noteworthy that C comes from atmospheric air in the form of carbon dioxide
(CO2); H and O come from water (H2O); and minerals (macronutrients and
1.4  Nutrient Accumulation by Crops and Crop Formation 9

Table 1.2  Relative composition of nutrients in plant dry matter


Participation Total
Classification Nutrient (element)a %
Organic macronutrients C 42
O 44
H 6
92
Macronutrients N 2.0
P 0.4
K 2.5
Ca 1.3
Mg 0.4
S 0.4 7
Micronutrients Cl, Fe, Mn, Zn, B, Cu, Mo, Ni 1
Total 100
The element form of nutrients is not always the chemical form that plants absorb
a

micronutrients) come from the soil, directly or indirectly. Therefore, we notice that
plant nutrients come from the three following systems: air, water, and soil, and
approximately 92% of plant dry matter comes from air and water systems, with only
8% coming from the soil. However, although the latter is less important quantita-
tively in relation to the others, it is the most discussed topic in plant nutrition stud-
ies. In addition, it is the most expensive material in agricultural production systems,
especially if we consider that air and rainwater have no cost (in nonirrigated produc-
tion systems).

1.4  Nutrient Accumulation by Crops and Crop Formation

Nutrient accumulation in plants reflects their nutritional requirement, ranging


according to several factors, such as the level of production, species, cultivar, soil
fertility and/or fertilization, climate, and crop treatments.
In general, crops have nutritional needs representing the amount of macronutri-
ents and micronutrients that plants remove from the soil during cultivation to meet
all development stages, resulting in adequate harvests.
Crops in general, such as sugarcane, soybean, and wheat, have high demand of
nitrogen and/or potassium and low demand of copper and molybdenum (Table 1.3).
However, the order of requirements for other nutrients may vary between crops and
even between cultivar/hybrid.
The standard, decreasing order of extraction of crops is generally as follows:
Macronutrients: N > K > Ca > Mg > P ↔ S
Micronutrients: Cl > Fe > Mn > Zn > B > Cu > Mo > Ni
10 1  Introduction to Plant Nutrition

Table 1.3  Total extraction (shoot) and export (stems/grains) of nutrients by commercial crops.
Sugarcane
Nutrient (100 t ha−1) Soybean (5.6 t ha−1) Wheat (3.0 t ha−1)
Crop Crop
Stems Leaves Total Grain residue Total Grain residue Total
kg ha−1
Macronutrient N 90 60 150 152 29 181 75 50 125
P 10 10 20 11 2 13 15 7 22
K 65 90 155 43 34 77 12 80 92
Ca 60 40 100 8 43 51 3 13 16
Mg 35 17 52 6 20 26 9 5 14
S 25 20 45 4 2 6 5 9 14
g ha−1
Micronutrient B 200 100 300 58 131 189 100 200 300
Cu 180 90 270 34 30 64 17 14 31
Fe 2500 6400 8900 275 840 1115 190 500 690
Mn 1200 4500 5700 102 210 312 140 320 460
Mo – – – 11 2 13 – – –
Zn 500 220 720 102 43 145 120 80 200

Considering the crops shown in Table 1.3, we note that the order of total extrac-
tion of nutrients changed. For macronutrients, sugarcane requires more K compared
to N, while S is the third most required nutrient in wheat. For micronutrients, we
note that Cl was the most extracted (not mentioned). The same change in the stan-
dard order occurs especially between Zn and B, for example, sugarcane demands
more Zn, while soybeans and wheat demand more B.
Regarding the export of nutrients from the agricultural area, a significant amount
of elements are mobilized to the harvest product (stem or grain) (Table 1.3). We note
that a significant part of N, S, P, and Zn, among others, is mobilized in the grains.
Thus, nutrients are stored in seeds in the form of specific organic compounds, for
example, N and S, accumulated in specific storage proteins (Müntz 1998), and P and
various cations, accumulated in the form of phytates (Raboy 2001). Each phytate
molecule contains six phosphate groups that form complexes with cations, and most
of K, Mg, Mn, Ca, Fe, and Zn in seeds are associated to the phytate (Epstein and
Bloom 2006).
Consequently, for living beings (humans and animals), seeds are more nutritious
than the rest of the plant. Thus, higher nutrient levels in the seeds benefit food qual-
ity, reducing human malnutrition, which is high in several regions of the world
(Stein 2010). In addition, this quality would influence the initial growth of the new
crop in seed production fields. Many plants can live on the P in the seed for approxi-
mately two weeks (Grant et al. 2001).
In practice, crops that export a large part of the absorbed nutrients in harvest, or
those in which the harvested product is the whole shoot (sugarcane, maize silage,
pasture), leave little crop residue. These crops require more attention regarding the
need to replace these nutrients through maintenance fertilization.
1.4  Nutrient Accumulation by Crops and Crop Formation 11

Studies on nutrient extraction can identify the requirement for a given nutrient in
crops, enabling to meet its demand and increase crop production.
In Brazilian agriculture, fertilizer application often may not meet the nutritional
requirements of crops, consequently limiting agricultural production. This fact is
verified by comparing plant nutritional requirements with the average use of fertil-
izers in the respective crops (Table 1.4).
In addition to fertilizers, there are other nutrient sources, such as steel slag (Ca,
Mg, and micronutrients) (Prado et al. 2002a, b), calcium silicates (Ca) (Prado and
Natale 2005), and biomass ash, among others, which were not considered in
Table 1.4.
We obtained crop requirements for yield levels close to the Brazilian average.
Based on these average results, we infer that soil depletion (in a soil already poorly
fertile) may be occurring. However, there are many areas in Brazil with adequate
fertilizer application, reaching the highest yields in the world, such as for soybean.
Researchers at the International Fertilizer Development Center (IFDC) stated
that most of the agricultural soils in the world are undergoing depletion of some
nutrients, except for North America, Western Europe, and Australia/New Zealand.
The authors conclude that we need to maintain this agricultural technology; other-
wise, we will not achieve the food production required for the future. The growth of
the world economy and the use of agricultural raw materials for fuel production
increase the world demand for food, which will only be ensured with the increase of
agricultural production by meeting crop nutritional requirements.
Still regarding nutritional requirements, nutrient extraction from the soil does not
occur constantly throughout the crop production cycle. The curve of nutrient extrac-
tion or accumulation over the period of cultivation (uptake rate) follows that of plant
growth, explained by a sigmoid curve. It is characterized by an initial phase of
decreased growth and nutrient uptake and, in the next phase, fast (almost linear)

Table 1.4  Nutritional requirement and apparent consumption of fertilizers (N + P2O5 + K2O) of


some crops in Brazil
Crop Nutritional requirementb Fertilizer consumption
N + P + K N + P2O5 + K2Oc N + P2O5 + K2O
Soybeana (2.8 t ha−1) 90(54) + 7 + 38 152 (97) 145
Sugarcane (73.0 t ha−1) 73 + 9.7 + 76 186 206
Citrus (26 t ha−1) (fresh fruit) 66.5 + 8.3 + 52 192 122
Maize (3.7 t ha−1) 176 + 32 + 149 430 110
Rice (3.2 t ha−1) 82 + 8 + 47 157 77
Bean (1 t ha−1) 102 + 9 + 93 235 31
Cassava (16,600 plants) 187 + 15 + 98 339 8
a
For soybean, the author estimates that 60% of N requirements come from biological fixation, and
the rest from the soil (54 kg ha−1 of N)
b
Fertilization needs are higher than nutritional requirements due to nutrient loss in the soil of 50,
70, and 30%, on average, for N, P, and K, respectively
c
P × 2.29136 = P2O5; K × 1.20458 = K2O
12 1  Introduction to Plant Nutrition

plant growth and increased nutrient uptake/accumulation. In the last phase, plant
growth/development and nutrient uptake stabilize until completing the production
cycle. However, at the end of the latter phase, accumulation of certain nutrients (K,
N) can stabilize or even decrease due to the loss of senescent leaves and rain wash-
ing out the nutrient (K) from the leaf. The sigmoid curve explains nutrient uptake
during the crop life cycle (perennial or annual). P accumulation for the formation of
sugarcane seedlings is high during a period from 65 to 110 days after transplanting,
requiring fertilization (Fig. 1.3).
In soybeans, Bataglia and Mascarenhas (1977) verified a period of maximum
demand for each nutrient, which would correspond to the maximum speed of nutri-
ent accumulation by the plant. The authors found that most nutrients reach the
period of maximum uptake along with the period of maximum accumulation of dry
mass (60–90 days), except for K, P, Cl, Mn, Mo, and Zn, which reached this period
at 30–60 days. In maize, the maximum speed of accumulation would be in the
period of 60–90 days for most nutrients, also corresponding to the maximum accu-
mulation of dry matter. For P and K, the maximum speed of accumulation occurred
earlier, at 30–60 days. However, nutrient application, for example, K, should prefer-
ably occur until 30 days, corresponding to the beginning of the period of maximum
requirement by the crop.
Although most studies in the literature established the nutrient uptake rate using
chronological data (in days), plants develop as thermal units and accumulate above
a base temperature, while growth stops below this temperature. Through thermal
accumulation, also known as degree-days, we obtain excellent correlations with the
duration of the crop cycle or with the stages of phenological development of a given
cultivar. Therefore, new studies on nutrient uptake rates or accumulation curves as
a function of degree-days accumulated during the crop cycle are relevant.

0.3 Accumulation of P 300


Shoot dry matter
250
Accumulation of P (g per plant)

0.2
Shoot dry matter (g)

200
0.2
150
0.1
100

0.1
50

0.0 0
30 40 50 60 70 80 90 100 110 120 130 140 150
Days after transplant (DAT)

Fig. 1.3  Phosphorus accumulation by sugarcane seedlings as a function of plant age


1.4  Nutrient Accumulation by Crops and Crop Formation 13

Studies on uptake rates are important to detect at which development stage the
crop demands more of a given nutrient, that is, at which stage the nutrient has the
highest uptake rate. Through this information, it is possible to predict the moment
of nutrient application to satisfy the nutritional requirement at the respective stage
of crop development, avoiding nutritional deficit and decreased production.
Thus, the best period for fertilizer application depends on the evaluation of the
nutrient uptake rate by the crop, besides depending on the nutrient and its release
dynamics in the soil.
The author notes that crop nutritional requirements are specific to the species and
to the cultivar/variety of the same species. Thus, nutritional differences between
crops will be discussed for each nutrient in the following chapters. In order to ensure
maximum efficiency of fertilization, that is, the most appropriate moment for nutri-
ent/fertilizer application, it is necessary to know the plant (i.e., its nutrient uptake
rate) and environmental factors, such as soil (texture), water (irrigated or nonirri-
gated), and cultivation system (conventional or no-till).
Modern agriculture requires maximum economic benefit regarding the environ-
ment. Thus, the Brazilian producer has a great challenge in face of the low fertility
of tropical soils and the high cost of fertilizers, which are the nutrient source.
A rational solution for sustainable agriculture would be to adapt the plant to the
soil by using crops/cultivars that are efficient in the process of crop formation, doing
more with less. In recent decades, especially in the 1990s, agricultural production
increased, although fertilizer application decreased. This could be explained by
increased efficiency of nutrient use by crops (Epstein and Bloom 2006).
As nutrient uptake, transport, and use can be genetically controlled, it is possible
to improve and/or select more efficient cultivars regarding nutrient use (Gabelman
and Gerloff 1983). For the plant to use nutrients with increased efficiency, it is nec-
essary to optimize several physiological and biochemical processes of crop forma-
tion. The possible mechanisms to control the nutritional needs of plants include
nutrient acquisition from the environment (soil or nutrient solution), their move-
ment through the roots and release to the xylem, their distribution in the organs, and
use in metabolism and growth (Marschner 1986).
There may be a crop with the same nutritional requirement, but more efficient
agronomically. For example, a hybrid wheat A can accumulate the same nitrogen
amount as a hybrid B. However, hybrid B uses N for increased grain production,
compared to hybrid A that prioritizes increased production of vegetative organs
(crop residues). Thus, for the same dry matter production (6 t ha−1), hybrid B pro-
duces 40% more grains than hybrid A, although both accumulate the same N amount
in the shoot (90 kg ha−1).
These results illustrate the current trend in agriculture, that is, it is better for the
plant breeder to select plants with increased grain production (commercial product)
and harvest index than total biomass. The harvest index is the ratio between the
harvested dry matter (grains) and the total dry matter of the plant. Therefore, photo-
assimilate partitioning by plants is also an important production factor for modern
agriculture.
14 1  Introduction to Plant Nutrition

In addition, modern cultivars develop shallower root systems (Rublo et al. 2003),
with increased recovery rates for fertilizers or organic matter mineralization.
The term nutrient use efficiency emerged in this context, which is the ability of a
species or genotype to provide high yields even in soils with deficit of the nutrient
under study (Graham 1984). It is also the relationship between production and con-
centration of the nutrient in the tissue (Lauchli 1987). Therefore, a species or culti-
var with superior nutritional efficiency develops adequately in low fertility soils due
to an increased ability to absorb the necessary nutrients in less quantity and/or to
distribute them more efficiently in the various plant components, ensuring adequate
plant metabolism with high conversion into dry matter.
Simple ways to increase nutritional efficiency are to reduce fertilizer doses to
levels that are still economical to produce or by genetic improvement through selec-
tion of plants with better nutritional indexes.
From dry matter and nutrient content in the plant, we can calculate nutritional
indexes comprising the efficiency of absorption, translocation/transport, and nutri-
ent use for conversion into dry matter. These indexes are shown below:
Uptake efficiency = (total nutrient content in the plant)/(root dry matter) (Swiader
et al. 1994). This index indicates the ability of the plant to extract nutrients from
the growing medium (soil). Mechanisms developed for high uptake efficiency
differ between plant species. Some species grow an extensive root system and
others have high uptake rates per unit of root length, that is, increased nutrient
influx (Föhse et al. 1988).
Translocation efficiency = ((shoot nutrient content)/(total nutrient content in the
plant)) ×100 (Li et al. 1991). This index indicates the ability of the plant to trans-
port nutrients from root to shoot. Samonte et al. (2006) observed the correlation
between N translocation index and protein content in rice grains.
Utilization efficiency (UE) (coefficient of utilization) = (total dry matter)2/(total
nutrient content in the plant) (Siddiqi and Glass 1981). This index indicates the
ability of the plant to convert the absorbed nutrient into total dry matter. According
to Gabelman and Gerloff (1983), the ability of a plant to redistribute and reuse
minerals from an older and senescent organ characterizes utilization efficiency in
the metabolism of the growth process.
Usually, the efficiencies aforementioned are more used in pot assays due to easi-
ness to work with plant root systems compared to field conditions. This line of
research in plant nutrition is very important, as adequate nutrient use is essential to
increase or sustain agricultural production.
Such efficiencies will be increasingly correlated with agricultural yield if the
cultivars have increased assimilate partitioning for organs of interest (such as
grains).
Thus, through field experimentation, other nutritional indexes similar to the pre-
vious ones emerged, although concerned to indicate nutritional efficiencies regard-
ing the dry matter of the commercial part (such as grains).
In this sense, Fageria (2000) observed increased correlation with rice production,
agronomic efficiency (AE), agrophysiological efficiency (APE), and physiological
1.4  Nutrient Accumulation by Crops and Crop Formation 15

efficiency (PE) (similar to the aforementioned utilization efficiency). In this same


crop, Samonte et al. (2006), who studied N application, observed that grain yield
correlated with N utilization efficiency and N content. In addition, the authors added
that it is interesting to select plants that not only have increased yields, but also use
the nutrient efficiently and with quality (grains with high protein content). Svecnjak
and Rengel (2006) observed differences in N utilization efficiency in canola culti-
vars, although they absorbed the nutrient similarly, as certain cultivars considered
efficient produced more dry matter with decreased N content in different plant
organs, except for the root. In maize plants, N uptake efficiency was more important
than utilization efficiency (Erying et al. 2020).
Thus, the forms of calculating nutritional efficiencies (agronomic, physiological,
agrophysiological, recovery, and utilization) will be presented, being used in field
assays, as indicated by Fageria et al. (1997):
Agronomic efficiency (AE) = (GPwf –GPnf)/(NAa), given in mg mg−1, where:
GPwf = grain production with fertilizer, GPnf = grain production without fertil-
izer, and NAa = nutrient amount applied. This index indicates the grain produc-
tion capacity per unit of fertilizer applied in the soil.
Physiological efficiency (PE) = (TOPwf–TOPnf)/(NACwf–NACnf), given in mg
mg−1, where: TOPwf = total organic production (shoot and grains) with fertilizer;
TOPnf = total organic production without fertilizer; NACwf = nutrient accumu-
lation with fertilizer, and NACnf = nutrient accumulation without fertilizer. This
index indicates the shoot production capacity per unit of nutrient accumulated in
the plant.
Agrophysiological efficiency (APE) = (GPwf – GPnf)/(NACwf – NACnf), given in
mg mg−1, where: GPwf = grain production with fertilizer, GPnf = grain produc-
tion without fertilizer, NACwf = nutrient accumulation with fertilizer, and
NACnf = nutrient accumulation without fertilizer. This index is similar to the
previous one, although indicating the specific capacity of grain production per
unit of nutrient accumulated in the plant.
Recovery efficiency (RE) = (NACwf–NACnf)/100(NAa), given in percentage,
where: NACwf = nutrient accumulation with fertilizer, NACnf = nutrient accu-
mulation without fertilizer, and NAa = nutrient amount applied. This index indi-
cates the amount of nutrient applied in the soil absorbed by the plant.
Utilization efficiency (UE) = physiological efficiency (PE) x recovery efficiency
(RE). This index indicates the total production capacity of the shoot per unit of
nutrient applied. This index differs from the index (c), as it computes the ­fertilizer
recovery efficiency, that is, the plant capacity for nutrient uptake/acquisition
from the soil.
In these experiments, plants are grown in soils with low and high content of the
given nutrient. Plants with better nutritional efficiency in soils with low nutrient
content are considered efficient, that is, they produce more under stress conditions,
while plants with the best nutritional indexes when submitted to soils with high
content of the given nutrient are considered responsive.
16 1  Introduction to Plant Nutrition

1.5  Other Chemical Elements of Interest for Plant Nutrition

In addition to the elements considered essential to plant life, there are elements
considered beneficial, and also a group of toxic elements. The beneficial element is
defined as the one which stimulates plant growth, but is not essential or is essential
only for certain species or under certain conditions (Marschner 1986). Silicon,
cobalt, and selenium are considered beneficial to the growth of certain plants.
However, Malavolta (2006) considers only Si and Na to be beneficial. We note that
even a nutrient or beneficial element, when present in high concentrations in the soil
solution, can be toxic to plants. However, an element is considered toxic when it is
not qualified as nutrient or beneficial element. Potentially toxic elements present
high harm potential even at low concentrations in the environment as they accumu-
late in the trophic chain and slow growth, which may lead to plant death. We have
as examples Al, Cd, Pb, and Hg, among others, and their harmful potential depends
on the dose.
Aluminum has been extensively studied, considering that tropical soils have an
acid reaction with high concentrations of exchangeable Al+3, which is toxic.
However, it can even reduce the toxicity of other elements (Cu and Mn) at low con-
centration (0.2 mg L−1). Usually, excess Al in the soil is toxic for plants, constituting
the main limiting factor for food and biomass production in the world (Vitorello
et al. 2005). The presence of this element affects from germination (Marin et al.
2004) to root growth, interfering with nutrient uptake (such as P, Mg, Ca, and K)
(Freitas et  al. 2006). Aluminum stress increases the molecular mass of cell wall
hemicellulose, making it rigid and inhibiting root elongation (Zakir Hossain et al.
2006), besides increasing membrane leakage. However, the latter may be the conse-
quence of exposure to the element and not caused by damage of the root growth
(Yamamoto et al. 2002).
Al toxicity symptoms thicken roots, making them short and brittle, sometimes
developing a brown color (Furlani and Clark 1981). In the shoot, Al toxicity symp-
toms may not be clearly identifiable and may even be confused with nutritional
imbalance (P, Ca, Fe, or Mn). According to Malavolta (1980), it is similar to P and
K deficiency, that is, yellowing of the margin and drying of leaves. High Al content
in the plant can block plasmodemata (preventing the transport of solutes and water
via symplast), which is induced by callose production, a polysaccharide that plants
produce in the phloem when subjected to pathogenic or environmental stress (tem-
perature, Al or Ca in the cytosol).
Research on plant nutrition aims to select tolerant genotypes and the mecha-
nisms that plants use to mitigate the toxic effect of the element. Jo et  al. (1997)
indicate two types of mechanisms that Al-tolerant plants have, such as (1) external
mechanisms, in which tolerant plants release organic acids by the root, usually
citrate and malate, which bind to aluminum forming stable complexes that prevent
Al uptake by the plant; (2) internal mechanisms, in which aluminum is absorbed
into the plant and, consequently, into the cell, where it is inactivated by some
enzyme or isolated inside the vacuole. Menosso et al. (2001) observed that soybean
1.5  Other Chemical Elements of Interest for Plant Nutrition 17

cultivars considered Al-tolerant distinguish from sensitive cultivars by increased


accumulation of organic acid (citric). Mendonça et al. (2005) observed in rice that
the Al-tolerant cultivar could adjust its proton balance more efficiently, in order to
absorb less Al and better tolerate the presence of this cation in the nutrient solution.
However, Braccini et al. (2000) found that change in rhizosphere pH is not related
to aluminum tolerance in coffee genotypes grown in soil.
Besides Al, other toxic elements are extensively studied to understand the mech-
anisms that tolerant plants use to minimize harmful effects. Research of this nature
allows introducing plants in areas with high load of heavy metals. As aforemen-
tioned, plants tend to accumulate metal in vacuoles and also form chelates with
these metals from two types of cysteine, phytochelatins or metallothioneins.
One of the beneficial elements most recently studied in Brazil is silicon. China,
USA, and Brazil are the countries that mostly publish scientific articles on silicon.
Plants normally absorb silicon in the form of acid (H4SiO4) with energy expenditure
(active) (Rains 1976), which then interacts with pectin and deposits on the cell wall,
remaining practically motionless in the plant. In the literature, Marschner (1995)
suggests a division of plants regarding their accumulation capacity for this element,
as follows: accumulating plants: contain 10–15% SiO2 contents, for example,
grasses, such as rice; intermediate plants: contain 1–5% SiO2 contents, with some
grasses and cereals; nonaccumulating plants: contain less than 0.5% SiO2 contents,
the majority of which are dicotyledons, such as legumes.
Ma and Takahashi (2002) propose criteria to differentiate nonaccumulating
plants from accumulating plants:
“Accumulators” have a Si concentration over 1% and a[Si]/[Ca] ratio >1.
“Excluders” have a Si concentration below 0.5% and a[Si]/[Ca] ratio <0.5.
Plants that do not meet these criteria are called “intermediates.”
We emphasize that all plants accumulate silicon, even those considered nonac-
cumulating. These plants accumulate silicon in the roots.
The evolution of the research allowed us to understand the absorption of Si by
plants. Si enters the plant from the external environment in the form of Si(OH)4
through specific influx channels (termed Lsi1), and efflux transporters (termed
Lsi2) mediate the loading of Si into the xylem and thus facilitate root-to-shoot trans-
location, which, in turn, moves Si to the aerial parts of the plant, where it deposits
as amorphous SiO2 (Ma and Yamaji 2015).
Miyake and Takahashi (1985) also add that there are other differences in plants
regarding Si uptake besides their content in the plant, referring to the transport rate.
Si-accumulating plants have high transport rate, intermediate plants have decreased
transport, and nonaccumulating plants concentrate absorbed Si in the roots.
Currently, Si is not considered a universal plant nutrient, although most authors
consider it beneficial, or, according to Epstein (2002), nearly essential.
The silica particles grow to a size of about 1 to 3 nm and are negatively charged
such that they can interact with the local environment of the cell walls of plants. It
has been suggested that the nucleation and growth of these structures are under the
control of specific proteins (Perry and Tucker 2000) and that a fraction of Si form
18 1  Introduction to Plant Nutrition

bonds with proteins, phenolic compounds (lignin, condensed polyphenols), lipids


and polysaccharides (cellulose) (Kolesnikov and Gins 2001). It is possible that sili-
con can replace structural compounds in the cell wall. This can decrease the energy
cost for the plant to form lignin or cellulose in the cell wall, favoring its growth.
In the literature, there are many reports on the benefits of Si for plants (Peixoto
et al. 2020). However, the most discussed benefits are its resistance against fungal
diseases in different crops, such as rice (Marschner 1986), and also decreased inci-
dence of pests (Goussain et  al. 2002) due to their increased difficulty to feed,
increased jaw wear, and increased mortality rate.
Si reduces diseases not only due to a physical factor (formation of compounds
that act as a physical barrier below the leaf cuticle) but also due to a chemical factor
(formation of compounds that are phytotoxic to pathogens). Recent studies indicate
that Si-rich plants may have holes in the epidermis without physical protection,
which hampers phytopathologists in the isolation of chemical compounds produced
by the plants.
The effects of Si on plants can be summarized as follows:
• Increases plant resistance to fungal diseases.
• Attenuates salinity (Calero Hurtado et al. 2019) and impacts the C:N:P stoichi-
ometry depending on the species (Calero Hurtado et  al. 2020) by increasing
enzyme activity (superoxide dismutase (SOD), catalase (CAT), and glutathione
reductase (GR)) and eliminating reactive oxygen species (Alves et al. 2020).
• Attenuates water deficit (Teixeira et al. 2020a), which can be seen visually in
sugarcane plants (Fig. 1.4) and forage (Rocha et al. 2021).

Fig. 1.4  Silicon attenuates water deficit in sugarcane plants


1.5  Other Chemical Elements of Interest for Plant Nutrition 19

• Increases cell stiffness through increased production of compounds such as lig-


nin in some species (Alvarez et al. 2018; Deus et al. 2019). Si can play a struc-
tural role similar to lignin, reducing its synthesis, which demands high
energy costs.
• Increased cell stiffness improves leaf architecture in the plant and favors photo-
synthesis (Flores et al. 2018).
• Reduces leaf senescence rates.
• Attenuates Fe, Mn, and Al toxicity. Si with salicylic acid attenuated B toxicity in
peas (Oliveira et al. 2020a).
• Attenuates the deficiency of K, Zn, Mn, Fe, and B (Prado et al. 2018); K, Ca, and
Mg in forages (Buchelt et  al. 2020); Mn (Oliveira et  al. 2019; Oliveira et  al.
2020b), B (Souza Júnior et al. 2019), Zn (Guedes et al. 2020), and Fe (Teixeira
et al. 2020b). The effect of Si can be seen visually in the attenuation of symptoms
of K deficit in quinoa plants (Fig.  1.5a), and Ca deficit in cabbage plants
(Fig. 1.5b) and also in tomato plants (Alonso et al. 2020) and cabbage (Silva et
al. 2021).
• Increased P use by plants (Silva and Prado 2021).

Fig. 1.5  Silicon attenuates symptoms of K deficit in quinoa plants (a) and Ca deficit in brachiaria
grass (b) and cabbage plant (c)
20 1  Introduction to Plant Nutrition

• Attenuates ammonium toxicity (Barreto et al. 2016, 2017; Viciedo et al. 2019b;
Silva Júnior et al. 2019; Viciedo et al. 2020a).
• Increases nitrogen nutrition efficiency, as it increases nitrate reductase activity
(Silva et al. 2020).
• Reduces excessive transpiration.
• Increases reproductive growth (Miyake and Takahashi 1978) and pollen grain
production and viability.
• Increases C utilization efficiency (Frazão et al. 2020; Lata-Tenesaca et al. 2021).
Si-accumulating plants such as rice can accumulate 250  kg ha−1 of Si, being
more absorbed than N or K (Körndorfer et al. 2002). Thus, there is a relationship
between Si in the plant and rice production, reaching 95% of the maximum with leaf
content equal to 34 g kg−1 in US organic soil (Korndörfer et al. 2001).
Beneficial effects of Si may occur for sugarcane with steel slag application as
corrective material and Si source, which promoted a linear effect on stem produc-
tion. Leaf spraying is an alternative for supplying Si with soluble sources and
increasing yield (Flores et al. 2018; Deus et al. 2019; Felisberto et al. 2020) and
growth of seedlings of sugarcane (Santos et al. 2020), soybeans, and common beans
(Souza Júnior et al. 2020), besides biofortified vegetables (Souza et al. 2018).
Plant response to silicon is more significant in production systems with some
type of stress, whether biotic (diseases/pests; poorly erect cultivar, etc.) or abiotic
(water deficit; excess metals, such as Al; low pH, etc.).
Si toxicity in field conditions is not known. There are reports on orchids with leaf
applications of high Si concentration for 18 months (Mantovani et al. 2018, 2020).
This effect may have occurred due to decrease in leaf gas exchange.
Sodium is also considered a beneficial element, as it was a nutrient for a salt-­
tolerant halophyte (Atriplex vesicaria), according to Brownell and Wood (1957). It
also showed beneficial effect for other plants, such as asparagus, barley, broccoli,
carrots, cotton, tomatoes, wheat, peas, oats, and lettuce (Subbarao et al. 2003). One
important aspect of sodium is its ability to replace part of K in nonspecific func-
tions, such as vacuolar K when K supply is limited.
Thus, Na would replace K in its contribution to the solute potential and, conse-
quently, in the generation of cellular turgor. This occurs significantly only in a spe-
cific group of plants, such as beets, spinach, savoy, coconut, cotton, cabbage, lupine,
and oats (Lehr 1953). In these crops, it is possible to use potassium fertilizers with
increased proportion of sodium (lower cost) in nonsodic soils. In addition, Na can
affect photosynthesis, especially in C4 plants, although this role is not fully under-
stood. Na would increase CO2 concentration and chloroplast integrity in leaf sheath
mesophilic cells (Brownell and Bielig 1996), as well as regenerate phosphoenol-
pyruvate (PEP) in the chloroplast and participate in chlorophyll synthesis. However,
Na can impair the enzymatic action of K at high concentrations, dislodging it from
enzyme action sites.
There are indications in the literature that Se is essential in organic compounds
such as amino acids, proteins, volatile compounds, ferrodoxins, and hydrogenases
1.5  Other Chemical Elements of Interest for Plant Nutrition 21

Fig. 1.6  Lettuce control plants (a) with symptoms of selenium toxicity (128 μM of Se in the form
of selenite) (b)

(Malavolta 2006), transporting RNA (Wen et al. 1988) and activating some enzymes,
such as superoxide dismutase, catalase, glutathione reductase, and guaiacol/ascor-
bate peroxidase (Djanaguiraman et al. 2005). Se decreases the leaf senescence rate
by decreasing peroxidase activity, which can increase N utilization efficiency, ben-
efiting crop production. However, it promotes oxidative stress/nutritional distur-
bance at high concentrations, damaging photosynthesis (Ferreira et  al. 2020),
inducing toxicity symptoms in the plant, reducing growth, and causing leaf chloro-
sis and small and brittle roots (Fig. 1.6).
Se biofortification through fertilization is feasible by increasing Se content in
food. This is important because Se is a nutrient for humans and animals, although
being highly toxic if ingested in excess, and this fact is rarely addressed in bioforti-
fication studies (Prado et al. 2017)

1.6  H
 ydroponic Cultivation: Preparation and Use
of Nutritional Solutions

The term hydroponics (from Greek hydro = water and ponos = work) is relatively
new, designated as soilless cultivation.
The alternative cultivation system of hydroponics can optimize production with
an increased number of crops per year (Jensen and Collins 1985) and increased
production compared to the conventional system (Table 1.5). However, hydroponic
cultivation is restricted to crops with fast cycle and small size, such as vegetables
and flowers, among others.
22 1  Introduction to Plant Nutrition

Table 1.5  Production of some vegetables grown in a greenhouse with hydroponic system and in
the field
Hydroponic system Field
Crop t/ha Number of crops t/ha/year t/ha/year
Broccoli 32.5 3 97.5 10.5
Green beans 11.5 4 46.0 6.0
Savoy 57.5 3 172.5 30.0
Chinese cabbage 50.0 4 200.0 –
Cucumber 250.0 3 750.0 30.0
Aubergine 28.0 2 56.0 20.0
Lettuce 31.3 10 313.0 52.0
Bell pepper 32.0 3 96.0 16.0

In commercial cultivation, hydroponics can be used in the production of valu-


able, high-quality (free of microorganisms and pesticides) crops with increased
added value to the product, such as various vegetables.
Thus, the hydroponic system has some advantages and disadvantages, as follows:
• Advantages
Requires less operational work.
Elimination of soil preparation (fuel costs and machinery purchases).
Elimination of crop rotation.
Reutilization of the culture medium.
Increased production without competition for nutrients and water.
Uniform plants regarding development.
Better root development and products with higher quality and longer shelf life.
Low loss of water and nutrients.
Reduction of pests and diseases (decreased spraying).
Better use of the agricultural area.
Implementation without restriction to the type of area (shallow soils, low drain-
age, or high slope).
Immune to climatic adversities (frost/hail).
• Disadvantages
Increased costs and initial work.
Higher risk of losses due to lack of electricity in automatic systems.
Requires technical skill and knowledge of plant physiology.
Inadequate balance of the nutrient solution can cause serious problems for plants.
Need to support roots and shoots.
Only inert materials should come into contact with the plants.
Regular routines.
Good drainage to prevent root death.
Water contamination affects the entire system.
In research, hydroponics can be used in several studies, such as:
1.6  Hydroponic Cultivation: Preparation and Use of Nutritional Solutions 23

Demonstration of nutrient essentiality.


Definition of symptoms of nutritional disorder, whether due to deficit or excess
(toxicity).
Knowing plant nutritional requirement.
Selection of plants tolerant to nutritional stress.
Ionic uptake, transport, and redistribution mechanisms.
Disease control.
Quality of products in hydroponics (e.g., nitrate accumulation).
For hydroponic cultivation, appropriate use of the nutrient solution is fundamen-
tal for its success, either for experimental or for commercial purposes. The nutrient
solution is a homogeneous system with nutrients and oxygen dispersed in appropri-
ate proportions and quantities, in ionic or molecular form. The homogeneity of the
nutrient solution is altered when it comes into contact with the plant roots, as there
are organic compounds from microbial activity, especially from decomposition of
root fragments or other impurities (coming with the plant or the hydroponic sys-
tem), or even exudates of organic acids from the roots.
One of the classic studies in the field of nutrition employing hydroponics is the
use of induced deficit or missing element technique. This technique was widely
used to investigate nutrient functions through the effects of the lack of nutrient on
the biochemical activity of the plant, besides being used for teaching in the disci-
pline of plant nutrition. In this technique, a complete solution is used minus the
nutrient under study, cultivating the plant until the characteristic symptoms of the
missing element appear, identifying the deficit (Fig. 1.7). When the plant is sub-
jected to stress, such as nutritional stress, it attempts to acclimatize with a series of
changes in the hormonal system (Morgan 1990). If the stress continues, a series of
events occur before the injury is visible, and if it occurs, this is the last biological
event, that is, at tissue level (Fig. 1.8). At this stage, half the production should be
compromised depending on the species. In species with long cycle, production
losses can be less than 50%. The form of symptoms depends on the role that the
respective nutrient plays in the plant and the place of occurrence (old or new leaf)
depends on its mobility in the plant phloem.
Besides the description of symptoms, we monitor plant responses throughout the
crop through growing plants with the complete nutrient solution and deficient solu-
tion (usually with total omission or with 10% of the appropriate concentration)
based on growth evaluations (height, stem diameter, leaf area, and dry matter).
Some plants are sensitive to nutrient deficit and excess. There are plants that indi-
cate nutrient deficiency, such as N (maize, apple), P (lettuce, barley), K (potato), Ca
(alfalfa, peanuts), Mg (cauliflower, broccoli), S (cotton, alfalfa), Zn (citrus, peach),
B (beetroot, turnip, celery, cauliflower), Mn (apple, cherry, citrus), Cu (citrus,
plum), Fe (cauliflower, broccoli), and Mo (tomato, lettuce, spinach) (Malavolta
et al. 1997). There are several studies in the literature with the omission of nutrients
and the features caused by nutritional disorder in crops such as cotton (Rosolem and
Leite 2007), brachiaria (Monteiro et  al. 1995), sorghum (Santi et  al. 2006), rice
(Alves et al. 2002), maize (Coelho et al. 2002), bean plant (Cobra Netto et al. 1971;
24 1  Introduction to Plant Nutrition

Fig. 1.7  Plants grown in


nutrient solution with all
nutrients (complete – with
phosphorus) (+P) and
minus phosphorus (−P)
(beans) (a), demonstrating
visual deficiency
symptoms

Dantas et al. 1979), green beans (Osório et al. 2020), sunflower (Prado and Leal
2006 ), sugarcane (Mccray et al. 2006), mallow (Fasabi 1996), Cyclanthera pedata
(Fernandes et al. 2005), coffee (Haag et al. 1969), acacia (Dias et al. 1994; Sarcinelli
et  al. 2004), açaí palm (Viegas et  al. 2004a, b), cupuassu (Salvador et  al. 1994),
Myrciaria dubia (H.B.K.) McVaugh (Viegas et al. 2004a, b), stevia (Lima Filho and
Malavolta 1997), eucalyptus (Rocha Filho et al. 1978), rubber tree (Amaral 1983),
teak (Barroso et al. 2005), guava tree (Salvador et al. 1999), soursop (Avilán 1975;
Batista et al. 2003), passion fruit tree (Avilán 1974), Spondias tuberosa (Gonçalves
et al. 2006), Cinchona officinalis (Viegas et al. 1998), castor bean (Lavres Júnior
et al. 2005), black pepper (Veloso and Muraoka 1993; Veloso et al. 1998), peach
palm (Silva and Falcão 2002), beetroot (Alves et  al. 2008), basil (Borges et  al.
2016), aubergine (Flores et al. 2014), soybean (Malavolta et al. 1980), watermelon
(Cavalcante et al. 2019), and orchid (David et al. 2019).
Soilless cultivation techniques can be divided into several categories due to use
of different substrates (materials other than soil) (Castellane and Araújo 1995):
Water culture or hydroponics: plant roots are immersed in a solution formed by
water and nutrients called NFT (Nutrient Film Technique) nutrient solution.
Sand culture: plants are supported by a solid substrate, with particles from 0.6 to
3.0 mm diameter.
Gravel culture: the substrate is solid with particles larger than 3 mm in diameter.
1.6  Hydroponic Cultivation: Preparation and Use of Nutritional Solutions 25

Decreased speed of metabolic processes

Interruption and compromise of biological processes


Molecular level

Not visible
Alteration of membranes, cell wall, organelles
Subcellular level

Cell deformation
Cell level

Alteration of tissues Visible


Tissue level = symptom (chlorosis / necrosis)

Fig. 1.8  Sequence of biological events in nutrient-deficient plants

Vermiculite culture: when the substrate is vermiculite or its mixture with other
materials.
Rockwool culture: uses rockwool, glasswool, or similar material as substrate (high
porosity and water uptake, with nutrients and air close to the roots). Rockwool
has an inconvenience regarding the environmental aspect at disposal, since its
decomposition is very slow.
We highlight that cultivation in a nutrient solution without solid components is
the most used technique in plant nutrition studies.
It is difficult to consider that a nutrient solution is ideal for all crops, with a for-
mula that ensures maximum development, and that all nutrients are supplied exactly
in the proportion in which they should be absorbed. Thus, nutrient solution’s com-
position is influenced by a number of factors, namely: plant species (nutritional
requirements are genetically controlled); plant age and growth stage; time of year
(length of light period); environmental factors (temperature, humidity, light); part of
the plant harvested, etc. In order to calculate the chemical composition of a nutrient
solution, besides crop requirements during cultivation, the environment should be
considered, as it affects the luminosity and temperature, directly affecting the tran-
spiration rate. Transpiration rate is important as conditions favoring high transpira-
tion would increase the loss of water solution at a faster rate than the nutrient uptake,
which may cause a salt effect. Thus, the higher the transpiration rate predicted for
the crop, the lower nutrient concentration in the nutrient solution. For example, if a
26 1  Introduction to Plant Nutrition

crop requires an appropriate N content equal to 50 g kg−1 dry matter, associated with
a transpiration rate of 300 L kg−1 dry matter, it would have 50 g of K in 300 L of
water or 166 mg L−1 of K. If the transpiration rate was 400 L kg−1 dry matter, the
solution would need dilution, that is, 50 g per 400 L or 125 mg L−1 of K.
Therefore, the chemical composition or ideal formula of the nutrient solution is
the one that meets the nutritional requirements of the species at all stages of the
production cycle. The nutrient solution may contain other elements besides nutri-
ents, reaching up to 20 elements (Jones 1998).
Santos (2000) tested four nutrient solutions using the NFT hydroponic system.
From the results, the solution proposed by Castellane and Araújo (1995) had the
best performance with increased yield, followed by the one proposed by Furlani
(1995). Thus, the author recommends the two solutions for lettuce cultivation in a
hydroponic system in the State of Rio Grande do Sul, Brazil. Although this study
indicated differences between solutions, this fact may not occur frequently, as the
solutions studied and more than a hundred solutions indicated in the literature derive
from the Hoagland and Arnon (1950) solution (Table 1.6), one of the solutions with
the highest concentration of salts. However, it has low Fe and Mn concentration,

Table 1.6  Chemical composition of some nutrient solutions, namely: Castellane and Araújo
(1995), Furlani (1995), and Hoagland and Arnon (1950), water solubility (cold and hot), and
salt index
Castellane and Furlani Hoagland and Solubillityc (g L−1) Salt
Components Araújo (1995) (1995) Arnon (1950)b (0, 5, and 100 °C) index
g/1000 L of solution
Calcium nitrate 950 1000 1200 1212 and 6598 53
Monoammonium – 150 150 224 and 1730 30
phosphate
Potassium phosphate 272 –
monobasic
Potassium chloride – 150 250 277 and 561 116
Potassium nitrate 900 600 260 134 and 2471 74
Magnesium sulfate 246 250 500 700 and 906 2
Manganese chloride – 1.17
Manganese sulfate 1.70 –
Zinc sulfate 1.15 0.44
Copper sulfate 0.19 0.10
Boric acid 2.85 1.02 19.5 and b389
Sodium molybdate 0.12 0.13
Fe-EDTAa 1 (L) 0.5 (L)
a
Fe-EDTA was used as iron source in both solutions, obtained by dissolving 24.1 g of iron sulfate
in 400 mL of water and 25.1 g of sodium-EDTA in 400 mL of hot water (80 °C), mixing the two
solutions when cold and completing the volume to 1 L
b
Solution of micronutrients (L/1000 L of solution) and Fe-EDTA (L/1000 L of solution)
c
Solubility of other salts (in g L−1) (0.5 and 100 °C): ammonium nitrate (1183 and 8711); diam-
monium phosphate (426 and 1063); ammonium sulfate (704 and 1033); magnesium sulfate (700
and 906); and potassium sulfate (67 and 239)
1.6  Hydroponic Cultivation: Preparation and Use of Nutritional Solutions 27

which can affect demanding plants, such as grasses. It should be noted that all nutri-
tional solutions provide the essential elements for plants. Franco and Prado (2006)
observed similarity in four nutritional solutions tested (Hoagland and Arnon;
Sarruge; Castellane and Araújo; Furlani) in the growth of guava seedlings. In a simi-
lar study with star fruit seedlings, it was observed that the use of these nutrient solu-
tions affected plant nutritional efficiencies (Rozane et al. 2007).
In the literature, the nutrient concentration indicated to formulate a nutrient solu-
tion is highly variable, regardless of nutrient, such as (in mg L−1) N-NO3: 70 to 250;
N-NH4+: 0 to 33; P: 15 to 80; K: 150 to 400; Ca: 70 to 200; Mg: 15 to 80; S: 20 to
200; B: 0.1 to 0.6; Cu: 0.05 to 0.3; Fe: 0.8 to 6.0; Mn: 0.5 to 2.0; Mo: 0.01 to 0.15;
Zn: 0.05 to 0.5; and Cl: 1 to 188 (Cometti et al. 2006).
The nutrient solution shown (Table 1.6) at full concentration (100%) is used in
plants with some development or in very cold periods as these solutions are very
concentrated, with the risk of physiological damage (salt effect) for young plants in
early growth stages. We use diluted solutions (25–75%) at the beginning of growth
and less diluted solutions are used as the plant develops, until reaching full concen-
tration (100%).
Besides nutrients, the beneficial element can also be used in the nutrient solution,
such as Si. Thus, the Si concentration used is 0.5 mmol L−1 in Na2SiO3 9H2O form,
which must be added first, maintaining a low pH in solution at this moment (Epstein
1995). The maximum Si concentration without risk of polymerization is 2.0
mmol L−1.
In basic plant nutrition studies using nutrient solution, the concentration of an
element in the solution does not always explain plant growth. This is because the ion
availability for plant uptake, that is, its activity in the solution can be influenced by
several factors, such as the solution ionic strength, the pH value, and chelate types
(Cometti et al. 2006). Ionic strength is more important when working with heavy
metals, especially Al, where the ion activity is reduced by increasing the ionic
strength. In a nutrient solution, unlike the soil solution, there is high pH variation
throughout cultivation, which can change free and complex forms of the element.
High pH (>6.0) decreases the availability of macronutrients Ca and P and micronu-
trients Mn, Cu, Zn, and B, due to the formation of precipitates, besides reducing
nutrient transport into the cells. The use of Fe chelates in the solution can chelate
Cu, Zn, and Mn. Thus, use of the Fe-EDDHA (ethylenediamine-N,N′-bis(2-
hydroxyphenylacetic acid) chelating agent will only partially chelate Cu, while use
of diethylenetriamine pentaacetic acid (DTPA) or ethylenediaminetetraacetic acid
(EDTA) chelates also forms complexes with Zn and Mn, especially at pH >5.5 for
Zn and pH >7.0 for Mn. This is important because micronutrients Cu, Mn, and Zn
are absorbed in the free form, and the quality of the Fe chelate can induce deficiency
of Fe and other micronutrients (Zn, Mn, and Cu). Fe-EDDHA (very stable) prevents
chemical reactions with other elements in the solution, although possibly releasing
less Fe+2 in the cell cytosol, depending on the species, causing deficiency.
Thus, in order to choose the appropriate solution, it is necessary to consider the
management factors of the nutrient solution, so as to increase the productive effi-
ciency of hydroponic systems, such as:
28 1  Introduction to Plant Nutrition

Sources of Fertilizers

Nutrient sources depend on the nature of cultivation, with commercial fertilizers


being used for commercial cultivation and pro analyze sources, with increased
degree of purity, intended especially for basic research in plant nutrition. We empha-
size that pro-analysis sources must be purchased from reputable companies, with
special attention to choose products free of inert materials or other elements that
may compromise the results and scientific rigor of the study.
We indicate ammonium molybdate or molybdic acid as Mo source, as sodium
molybdate is very alkaline, which may cause precipitation reactions with other
micronutrients.

Water

In commercial hydroponics, drinking water can be used, while distilled water is


generally used in scientific hydroponics. However, in some cases, deionized (up to
twice) water is better, as in assays aimed to induce nutritional deficiency, especially
of micronutrients.
If the water comes from the urban network, we recommend let it rest for about
24 h to eliminate the chlorine used in its treatment (Martinez and Silva Filho 2004).

Order of Nutrient Addition

In order to avoid precipitation reactions of fertilizers (poorly soluble), which would


become unavailable to plants, it is necessary to follow a certain order of nutrient
addition. Before adding fertilizers (nutrients), the water pH value must be adjusted
to appropriate standards (discussed in the subsequent item).
1. Add calcium-free macronutrient fertilizers, such as potassium phosphate, potas-
sium nitrate, and magnesium sulfate. This procedure is necessary due to incom-
patibility between calcium nitrate and salts containing phosphorus and sulfur,
forming precipitated compounds with low solubility.
2. Calcium salts (calcium nitrate).
3. Iron-free micronutrients, such as fertilizer sources based on Mn, Zn, Cu,

B, and Mo.
4. Iron source (Fe-EDTA, Fe-EDDHA, or other).
Then, immediately adjust the pH value and determine the EC (electrical conduc-
tivity) (discussed in the subsequent item).
1.6  Hydroponic Cultivation: Preparation and Use of Nutritional Solutions 29

Thus, if a stock solution is formulated, it is necessary to have a solution A based


on calcium nitrate, another solution B with other macronutrients, and a solution C
for micronutrients.

Maintenance of Adequate pH in the Nutrient Solution

As for pH, there are the following concerns:


Calibrate the instrument before each measurement.
The pH value should be adjusted daily to keep the solution pH value in the
5.5–6.5 range. In case of a pH value above this range, add drops of an acid at 0.1 M
(HCl; HNO3, H2SO4; H3PO4) or, otherwise, add a 0.1  M base (NaOH; KOH).
However, the best way to adjust the pH in the solution is the management of nitro-
gen sources (NO3- or NH4+) by using part of N in the NH4+ form, in the proportion
of 10–20% of the total N.
The use of solution with N in ammonium and nitrate form may initially decrease
the pH value until almost all ammonia in the solution was absorbed by the plant,
with subsequent increased nitrate uptake and increased pH value. Maintenance of
the solution pH is important as low values (~ 3–4) affect membrane integrity (H+
affects the permeability of cell membranes), which may cause loss of nutrients
already absorbed, besides affecting cation availability and uptake. In a solution with
high pH (>7), there may be problems with the electrochemical gradient and proton-­
anion cotransport across the membranes, besides the loss of phenolic compounds
and electrolytes from cells and anion uptake reduction. In addition, high pH may
cause undesired chemical reactions in the solution, leading to nutritional deficien-
cies in plants (Fe, Mn, B, and P).
We observed that lettuce cultivation in acid pH caused increased damage to plant
growth in relation to alkaline pH, compared to the appropriate range. In the pH
range from 5 to 7, plant dry matter production (relative) was 100.0%, while in other
pH ranges, 4–5, 3–4, 2–3, and 7–9, the production was 94.5, 1.2, 0.6, and 60.0%,
respectively.

 aintenance of Adequate Osmotic Pressure


M
in the Nutrient Solution

The osmotic pressure in the nutrient solution must be in the range of 0.5–1.0 atm, as
high values may indicate excess salts in the solution, with serious damage to
the roots.
30 1  Introduction to Plant Nutrition

Maintenance of Adequate Temperature in the Nutrient Solution

In order to avoid heating of the nutrient solution (>25–30  °C), it is important to


avoid light incidence, storing the solution in a shaded and ventilated location. We
also note that light incidence in the solution allows algae to proliferate.

Maintenance of Oxygenation in the Nutrient Solution

It is a consensus that oxygenation of the nutrient solution is mandatory to maintain


adequate O2 contents for roots (~3 ppm O2). The adequate oxygenation level ranges,
with low demand (rice = 3mg L−1 O2) and high demand (tomato = 16mg L−1 O2) of
oxygen in the nutrient solution according to the crop. As increasing temperatures in
the nutrient solution decrease the dissolved O2 content, it is important to avoid high
temperatures. Aeration is applied according to each hydroponic system, since only
in the fall when the nutrient solution returns to the reservoir or, in the case of iso-
lated pots, the injection of compressed air can supply the necessary oxygen to
the plants.

Maintenance of Adequate EC in the Nutrient Solution

During cultivation of plants in nutrient solution, the elements of the solution are
depleted, ranging according to the nutrient. This is due to differences in the uptake
rate of nutrients, which can be fast (N, P, K, and Mn), intermediate (Mg, S, Fe, Zn,
Cu, and Mo), or slow (Ca and B) (Bugbee 1995). Thus, nutrients need to be replaced
in the solution, which is performed using electrical conductivity data.
The conductivity of an electrolyte solution is the quantitative expression of its
ability to transport electrical current. It is defined as the inverse of the electrical
resistance of 1 cubic cm of liquid at a temperature of 25 °C. Electrical conductivity
is the unit equivalent to 1 mhos = 1 Siemens = 103 mS = 106 μS (mS = milliSiemens;
μS = microSiemens).
Normally, the nutrient solution’s EC ranges from 1.5 to 4.0 mS/cm according to
the solution chosen for the respective cultures. The EC obtained in a nutrient solu-
tion is the sum of the EC of all fertilizers used in the formula of that solution.
Castellane and Araújo (1995) obtained an EC of 2.6 to 2.8 mS/cm. As 1 mS/cm
corresponds to 640 ppm of nutrients, we note that the use of this variable during
cultivation would prevent the nutrient solution from having a low nutrient concen-
tration, which could lead to a nutritional deficiency. Normally, when the electrical
conductivity reduces to a certain level of the initial solution (approximately
30–50%), it is recommended to replace it. Backes et al. (2004) suggested replacing
the solution when it decreased to 50% of the initial EC. In commercial cultivation,
1.6  Hydroponic Cultivation: Preparation and Use of Nutritional Solutions 31

it is possible to use EC to manage the nutrient solution as follows (Carmello and


Rossi 1997):
(a) Add daily an amount of new solution equivalent to the amount of solution that
reduced in the recipient. After 21 days, renew the nutrient solution if the con-
ductivity reaches 4 mS/cm.
(b) Add only water to replace the amount of evaporated solution and monitor con-
ductivity; when it reaches less than 1 mS/cm, add the salts to recompose it or
change it. Renew the nutrient solution in 21 days.

Disease Prevention Measures

Diseases may occur in hydroponic cultivation, especially fungal, requiring preven-


tive measures. Thus, it is important to disinfect materials for each crop. We can use
sodium hypochlorite or calcium hypochlorite, based on 1.000 mg/L or 10.000 mg/L
of active chlorine, respectively. It is necessary to rigorously wash the materials, as
Cl residues (>0.5 mg/L) can injure plants, especially with the use of ammoniacal
salts in the nutrient solution (Martinez and Silva Filho 2004). The nutrient solution
can also be disinfected through pasteurization. Pasteurization occurs as follows:
heat the nutrient solution from 95 to 105 °C for a period of 30 s and then cool it
quickly until reaching room temperature (in 30 s) (Martinez and Silva Filho 2004).

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