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Anatomy of Flowering Plants
Fourth Edition
Understanding plant anatomy is not only fundamental to
the study of plant systematics and palaeobotany but also an
essential part of evolutionary biology, physiology, ecology
and the rapidly expanding science of developmental
genetics. This modernized new edition covers all aspects of
comparative plant structure and development, arranged in
a series of chapters on the stem, root, leaf, flower, pollen,
seed and fruit. Internal structures are described using
magnification aids from the simple hand lens to the
electron microscope. Numerous references to recent
topical literature are included, and new illustrations reflect
a wide range of flowering plant species. The phylogenetic
context of plant names has been updated as a result of
improved understanding of the relationships among
flowering plants. This clearly written text is ideal for
students studying a wide range of courses in botany and
plant science, and is also an excellent resource for
professional and amateur horticulturists.

Paula J. Rudall is Research Professor at the Royal Botanic

Gardens, Kew, UK, and an international authority on the
evolution of plant form. Her research interests range from
the organization of flowers and the patterning of petal
surfaces to the intricate structure and development of the
stomatal pores on the surfaces of leaves. Her numerous
professional awards include the Dahlgren Prize, the
Linnean Society Gold Medal and election as Foreign
Member of both the Botanical Society of America and the
American Society of Plant Taxonomists. In addition to
several books, she has authored over 300 peer-reviewed
papers.
Anatomy of
Flowering Plants
An Introduction to Plant
Structure and Development
Fourth Edition

PAULA J. RUDALL
Royal Botanic Gardens, Kew
University Printing House, Cambridge CB2 8BS, United Kingdom

One Liberty Plaza, 20th Floor, New York, NY 10006, USA

477 Williamstown Road, Port Melbourne, VIC 3207, Australia

314–321, 3rd Floor, Plot 3, Splendor Forum, Jasola District Centre,

New Delhi – 110025, India

79 Anson Road, #06–04/06, Singapore 079906

Cambridge University Press is part of the University of Cambridge.

It furthers the University’s mission by disseminating knowledge in the pursuit of

education, learning, and research at the highest international levels of excellence.

www.cambridge.org
Information on this title: www.cambridge.org/9781108749121
DOI: 10.1017/9781108782104
© Paula J. Rudall 1987, 1992, 2007, 2020

This publication is in copyright. Subject to statutory exception

and to the provisions of relevant collective licensing agreements,
no reproduction of any part may take place without the written
permission of Cambridge University Press.

First edition published by Edward Arnold 1987

Second edition published by Cambridge University Press 1992
Third edition published by Cambridge University Press 2007
Fourth edition published by Cambridge University Press 2020

Printed in the United Kingdom by TJ International Ltd, Padstow Cornwall

A catalogue record for this publication is available from the British Library.
ISBN 978-1-108-74912-1 Paperback

Cambridge University Press has no responsibility for the persistence or accuracy of

URLs for external or third-party internet websites referred to in this publication
and does not guarantee that any content on such websites is, or will remain,
accurate or appropriate.
This book is dedicated to my friend and early mentor,
Dennis W. Stevenson
Contents

Preface page ix

1 Organs, cells and tissues 1

2 Stem 24

3 Root 41

4 Leaf 54

5 Flower 72

6 Seed and fruit 95

Glossary 107
Select bibliography 122
Index 135
Preface

Plant anatomy is concerned with the structural organization of plants

and the arrangements of their organs, cells and tissues. The study of
comparative plant anatomy remains highly relevant to the traditional
fields of systematics and palaeobotany and also to the relatively new
field of developmental genetics (including evolutionary develop-
mental genetics: evo-devo), which utilizes a combination of techni-
ques to examine gene expression in growing tissues23, 83. Modern
students can incorporate information from an increasingly wide
range of sources, most notably integrating morphological and mole-
cular data. The new edition of this book presents an introduction to
plant anatomy for students of botany and related disciplines.
Plant anatomy is today explored using a broad range of techni-
ques, from the compound microscope to high-resolution X-ray
computed tomography (HRCT). Although the simple optical lens
has been used for centuries to examine plant structure, detailed
studies of plant anatomy originated with the invention of the
compound microscope in the seventeenth century. Nehemiah
Grew (1641–1712) and Marcello Malpighi (1628–1694), physi-
cians working independently in England and Italy respectively,
were early pioneers of the microscopical examination of plant cells
and tissues. Their prescient work formed the foundation that
eventually led to the development of our understanding of cell
structure and cell division30. Other outstanding early figures
included Robert Brown (1773–1858), who discovered the
nucleus, and the plant embryologist Wilhelm Hofmeister
x Preface

(1824–1877), who first described the alternation of generations

in the life cycle of land plants. During the nineteenth and twentieth
centuries, plant anatomy became an important element of studies
of both physiology and systematic biology, and an integral aspect
of research in the developing field of anatomical palaeobotany, led
by such luminaries as Dukinfield Henry Scott (1854–1934). The
physiologist Gottlieb Haberlandt (1854–1945) utilized anatomi-
cal observations in his groundbreaking work on photosynthetic
carbon metabolism. Notable plant anatomists of the twentieth
century included Agnes Arber (1879–1960), whose books
included works on monocotyledons4, and Katherine Esau
(1898–1997), recognized particularly for her work on the struc-
ture and development of phloem and her influential textbooks on
plant anatomy37. Other important botany textbooks include works
on anatomy28, 40, 41, embryology26, 82, morphology44 and
palaeobotany44, 84, 134.
The invention of the transmission electron microscope (TEM)
in the mid- twentieth century allowed greater magnification than
any optical microscope, and hence revitalized studies in cell
ultrastructure39 and pollen morphology36, 125. The subsequent
development of the scanning electron microscope (SEM) pro-
vided greater image clarity and much greater depth of focus than
light microscopes when examining surface structure, and thus
further increased accessibility of minute structures, including
seeds, pollen grains and organ primordia. More recent innova-
tions, including fluorescence microscopy, differential interfer-
ence contrast (DIC) microscopy and confocal imaging, have
allowed enhanced visualization of tissue structure. Others,
including HRCT and nuclear magnetic resonance (NMR) ima-
ging, facilitate enhanced visualization of three-dimensional
objects. The most effective studies employ more than one of
the techniques available today.
To study plant evolution using comparative data, an under-
standing of taxonomy is essential. Throughout this book, species
Preface xi

are assigned to families according to a modern understanding of

their classification. In textbooks published before 1990, extant
angiosperms were consistently subdivided into two major
groups – dicotyledons (dicots) and monocotyledons (monocots),
based partly on the number of cotyledons in the seedling. This
dichotomy was long considered to represent a fundamental diver-
gence at the base of the angiosperm evolutionary tree. However,
the expansion of molecular phylogenetics through the 1990s
demonstrated that some species that were formerly classified as
primitive dicots do not belong to either category, though the
monophyly of monocots was confirmed. Thus, although the
dicot/monocot distinction remains useful for generalized descrip-
tions of angiosperm groups, current evidence suggests that it does
not represent an entirely natural classification. It is now widely
accepted that several relatively species-poor angiosperm lineages
(early divergent angiosperms) evolved before the divergence of
the three major lineages that led to the magnoliids, monocots and
the remaining dicots (now termed eudicots, or sometimes
tricolpates).
Early divergent angiosperms are a small but highly diverse
assemblage of taxonomically isolated lineages that probably
represent the surviving extant members of their respective
clades, accounting for only about one per cent of extant
species139. They include the New Caledonian shrub Amborella
and the water lilies (Nymphaeaceae). The magnoliids include
woody families such as Magnoliaceae and Lauraceae and
herbaceous or climbing families such as Piperaceae and
Aristolochiaceae. Monocots account for approximately one
quarter of all flowering plant species. They dominate signifi-
cant parts of world ecosystems and are of immense economic
importance, including the staple grass food crops (wheat,
barley, rice and maize) and other important food plants
such as onions, palms, yams, bananas and gingers. Eudicots
represent about 75 per cent of extant angiosperm species and
xii Preface

encompass a wide range of morphological diversity, espe-

cially in the two largest eudicot subclades, Rosidae (rosid
eudicots) and Asteridae (asterid eudicots). Thus, understand-
ing of this revised and updated phylogenetic context is essen-
tial for credible interpretation of phenotypic patterns.
 1

Organs, cells and tissues

1.1 Organs
Plants are essentially modular organisms; each individual plant
consists of distinct but connected organs. In their turn, the organs
are composed of cells, which are mostly grouped into tissues.
Vegetative organs support photosynthesis and plant growth, and
reproductive organs enable sexual reproduction. In seed plants, the
primary vegetative organs are the root, stem and leaf (Figure 1.1).
Roots and stems have well-defined growing points at their apices,
but the leaves are determinate lateral organs that stop growing when
they reach a particular size and shape. When a seed germinates, the
seed coat (testa) is ruptured and the embryonic structures emerge
from opposite poles of the embryo: a seedling root (radicle) grows
downwards from the root apex and a seedling axis (hypocotyl)
bears the first leaves (cotyledons) and the shoot apex, which ulti-
mately develops new foliage leaves.

1.2 Cells and cell walls

Plant cells consist of a living protoplast contained within a protein-
rich plasma membrane, which is itself enclosed by a cell wall. During
the cell-division cycle, cells undergo a series of phases that are broadly
grouped as interphase, nuclear division (mitosis or meiosis) and
cytokinesis (cleavage of the cytoplasm and cell-wall formation).
During mitosis, nuclear division occurs first, followed by progressive
deposition of membranes in the cytoplasm into a cell plate that is
2 Organs, cells and tissues

Figure 1.1 Ligustrum vulgare (eudicot: Oleaceae), longitudinal section of

vegetative shoot apex. Scale = 100 µm

located in the equatorial zone between the two daughter nuclei138.

The cell plate extends to join the cell wall, thus depositing a new wall.
The primary cell wall consists mostly of carbohydrates: microfi-
brils of cellulose and hemicelluloses embedded in a matrix of pectins
(Figure 1.2). Cell walls of adjacent cells are linked together by
a pectin-rich middle lamella37, 39. Following cell enlargement and
elongation, a secondary cell wall is deposited on the inside surface of
the primary wall. Secondary cell walls often appear layered and can
contain deposits of complex organic polymers such as cutin, lignin
and suberin. Cutin is the primary component of the plant cuticle,
which covers the aerial epidermis (Section 1.8). Lignin provides
strengthening and rigidity to sclerenchyma cells, especially in the
secondary xylem, but also in fibres close to the vascular bundles in the
stem and leaf. Suberin is a complex hydrophobic lipid that provides
a protective water-resistant lipophilic barrier in periderm cells.
1.3 Cytoplasm, plastids and photosynthesis 3

Figure 1.2 Diagram of a generalized plant cell

Primary pit fields are thin regions of the primary cell wall that
correspond with similar regions in the walls of neighbouring cells.
Pits have protoplasmic strands (plasmodesmata) passing through
them, connecting the protoplasts of adjacent cells. The connected
living protoplasts are collectively termed the symplast. Primary pit
fields often persist as thin areas of the wall even after a secondary
wall has been deposited. In simple pits, which occur on relatively
non-specialized cells such as parenchyma, the pit cavity is of more
or less uniform width. In bordered pits, which are present in
tracheary elements, the secondary wall arches over the pit cavity
so that the opening to the cavity is narrow and the outer rim of the
primary pit field appears as a border around the pit opening when
viewed through a light microscope18.

1.3 Cytoplasm, plastids and photosynthesis

The living cell protoplast consists of cytoplasm that encloses
a complex range of membrane-bound internal structures termed
4 Organs, cells and tissues

Figure 1.3 Lapageria rosea (monocot: Philesiaceae), stomatal pore and guard
cells (TEM). Scale = 10 µm

organelles; they include mitochondria, the nucleus, plastids and

vacuoles, as well as small particles termed ribosomes and sometimes
inorganic contents such as oil, starch grains or crystals (Figure 1.3).
Most plant cells possess a single nucleus that is bounded by
a pair of membranes; the outer membrane is continuous with the
endoplasmic reticulum. At interphase, one or more nucleoli can
sometimes be distinguished, together with the uncondensed chro-
matin within the nuclear sap. During nuclear division, the chro-
matin becomes condensed into chromosomes that bear the
hereditary information. Vacuoles are membrane-bound structures
that contain a watery sap; they can vary considerably in size and
shape during the life history of a cell. They can accumulate storage
products and soluble pigments such as anthocyanins.
Plastids are large cell organelles that develop from proplastids. They
each contain their own genome, which is much smaller than the
nuclear genome and normally heritable via the maternal parent. Each
plastid is bounded by a pair of membranes, and many contain a system
1.4 Inorganic cell inclusions 5

of membranes termed thylakoids. Plastids such as amyloplasts, chlor-

oplasts and chromoplasts play specialized roles within the cell.
Amyloplasts are the source of starch grain production. Chromoplasts
contain carotenoid pigments that produce some of the colours found
in some plant organs, such as flower petals. Mitochondria – the sites of
respiration within the cell – are smaller than plastids; they also contain
their own heritable genetic material and are enclosed within a double-
membrane system. Both plastids and mitochondria originated via
endosymbiotic events at an early unicellular stage in plant evolution50.
Chloroplasts are highly specialized plastids containing green
chlorophyll proteins that absorb energy from sunlight. During
photosynthesis, plants convert light energy into chemical energy
in energy-storage molecules such as adenosine triphosphate
(ATP), which they use to make carbohydrates from carbon dioxide
(CO2) and water, releasing oxygen as a by-product. Chloroplasts
occur in all green cells but are most abundant in the leaf meso-
phyll. In most plants, photosynthetic carbon reduction is achieved
via a three-carbon compound (C3 cycle), but some plants capture
inorganic carbon more effectively using CO2-concentrating
mechanisms via a C4 cycle or Crassulacean acid metabolism
(CAM)31. Some C4 plants possess a distinctive leaf anatomy,
termed Kranz anatomy (Section 4.9).

1.4 Inorganic cell inclusions

Many cells possess non-protoplasmic contents such as mucilage,
oils, tannins, starch granules, calcium oxalate crystals and silica
bodies. Both oil and mucilage are produced in isolated specialized
secretory cells (idioblasts). Tannins are phenol derivatives that are
widely distributed in plant cells; they are amorphous and appear
yellow, red or brown in cells of sectioned material due to oxidation.
Starch is also widespread in plant tissues but especially common
in storage tissues such as endosperm or in parenchyma adjacent to
a nectary. Starch granules are formed in specialized plastids
6 Organs, cells and tissues

Figure 1.4 Idioblastic cells containing crystals of calcium oxalate. (a) Dioscorea
sosa (monocot: Dioscoreaceae), raphide crystals in a mucilage sheath. (b)
Cissus rhombifolia (eudicot: Vitaceae), druse (cluster crystal). (c) Atriplex
hymenelytra (eudicot: Amaranthaceae), druse. Scale = 10 µm

(amyloplasts). They often appear layered due to the successive

deposition of concentric rings and possess characteristic shapes.
The unusual and specialized starch grains present in laticifers in
some species of Euphorbia possess highly characteristic elongated
rod- or bone-like shapes compared with the more rounded starch
grains of neighbouring parenchyma cells85.
Calcium oxalate crystals (Figure 1.4) occur within crystal idio-
blasts; they can be distributed in almost every part of both vegetative
and reproductive organs and are often located near veins, possibly
reflecting transport of calcium through the xylem. They form within
the vacuoles of actively growing cells and are usually associated with
1.4 Inorganic cell inclusions 7

membrane chambers, lamellae, mucilage and fibrillar material.

Contrasting crystal shapes can be highly characteristic of different
plant families; for example, styloid crystals characterize the Iris family
(Iridaceae). Common crystal types include solitary needle-like or
rhomboidal crystals (styloids), bundles of needle-like crystals borne
together in the same cell (raphides), aggregate crystalline structures
that have precipitated around a nucleation site (druses) and numerous
fine particles of crystal sand. In some woody eudicot species, crystals
occur in the secondary phloem or secondary xylem; for example,
crystal cells are common in the ray parenchyma of some woods
(Chapter 2). Cystoliths are calcareous bodies that are mostly located
in leaf epidermal cells (Figure 4.4).
Most plants deposit silica in at least some of their tissues101. Some
species accumulate silica in large quantities and deposit it as discrete
bodies of solid silicon dioxide in the lumina of specific plant cells.
Opaline silica bodies, commonly termed ‘phytoliths’, are

Figure 1.5 Oryza sativa, rice (monocot: Poaceae), H-shaped silica bodies
(phytoliths) in a leaf epidermis (SEM). Scale = 10 µm
8 Organs, cells and tissues

a characteristic feature of some flowering plant groups. These

groups include several commelinid monocots, notably the grass
family Poaceae118, in which silica bodies are almost exclusively
restricted to the epidermis, and the palm family Arecaeae, in
which they are primarily restricted to the vascular bundle sheath
cells. Grass phytoliths occur in various shapes that can characterize
different species, so they can have immense significance as diag-
nostic markers in studies of grassland palaeoecology (Figure 1.5).
Silica bodies are often associated with sclerenchyma. In some
woody eudicot species they can occur in the secondary xylem.

1.5 Meristems
Meristems are the growing points of the plant. They represent
localized regions of thin-walled, tightly packed living cells that
undergo frequent mitoses and often continue to divide indefinitely.
Most of the plant body is differentiated at the meristems, though cells
in other regions may also occasionally divide.
Apical meristems are located at the shoot apex (Figures 1.1, 2.1),
where the primary stem, leaves and flowers differentiate, and at the
root apex (Figures 3.1, 3.2), where primary root tissue is produced. In
flowering plants, the shoot and root apical meristems are highly
organized but differ from each other in many respects. Both shoot
and root apical meristems contribute to extension growth and are self-
renewing. The shoot apical meristem also initiates lateral organs
(leaves) at its flanks in a regular nodal arrangement, each node bearing
a single leaf, a pair of leaves or a whorl of leaves. Subsequent elonga-
tion of the shoot axis occurs at the stem internodes, either by diffuse
cell divisions and growth throughout the youngest internodes (unin-
terrupted meristem) or in a restricted region, often at the base of the
internode (intercalary meristem). Both intercalary and uninterrupted
meristems represent growth in regions of differentiated tissues.
Lateral meristems are important for stem thickening growth;
they include vascular cambium and the primary and secondary
1.5 Meristems 9

thickening meristems (Chapter 2). Lateral meristems occur in

localized regions parallel to the long axis of a shoot or root, most
commonly in the pericyclic zone, at the junction between vascular
tissue and cortex. The phellogen (cork cambium) is a lateral mer-
istem that occurs in the stem or root cortex, where it forms
a protective corky layer (Figure 2.10); a phellogen can also
develop in the region of a wound or at the point of leaf abscission.
Meristemoids are isolated and densely protoplasmic cells that
reactivate embryonic activity to allow tissue differentiation.
Typically, a meristemoid either itself undergoes unequal (asym-
metric) cell division or is the smaller daughter cell that results from
an asymmetric division16 (Figure 1.6). Asymmetric divisions are
caused by cell polarization resulting from organized arrangements
of actin filaments in the dense cytoplasm during cell plate
alignment49. Examples of asymmetric cell divisions include

Figure 1.6 Amborella trichopoda (ANA-grade: Amborellaceae), developing leaf

epidermis, highlighting a pair of cells that have recently undergone
asymmetric mitosis to form a dense meristemoid and its larger sister cell
(TEM). The meristemoid will form a guard-mother cell and will ultimately
divide symmetrically to form a pair of guard cells. Scale = 2 µm
10 Organs, cells and tissues

formation of a root hair initial (trichoblast), microspore division into

a larger vegetative cell and smaller generative cell, a protophloem
mitosis to form a larger sieve tube element and smaller companion
cell, and division of a protodermal cell into two cells of unequal sizes,
the smaller of which is the guard-mother cell, a meristemoid that will
divide symmetrically to form the paired guard cells of a stoma.
Other types of localized meristems occur in some species, in
which differentiated plant cells can become de-differentiated and
meristematic. One example of such cell-fate plasticity includes
localized meristems that occur on the leaf margins of some succu-
lent Crassulaceae; these meristematic cells can give rise to entire
plantlets131. Plants can also regenerate tissues and organs at the site
of a wound by cellular proliferation on callus tissue, a process that
is regulated by the plant hormones auxin and cytokinin. Callus
cells formed from roots and from some aerial organs resemble the
apices of lateral roots derived from the pericycle81, 137. Isolated
callus tissue can be used in laboratory conditions to artificially
grow a new plant using tissue culture methods.

1.6 Cell growth and expansion

Cells develop and expand in different ways depending partly on
their location and surrounding tissue and partly on their cell-wall
properties. Those in close contact with each other are initially glued
together by a pectin-rich middle lamella and hence have a mutual
influence on shape during early expansion. Parenchyma cells,
which are largely thin walled and isodiametric, typically expand
relatively evenly. Different rates of expansion of adjacent cells can
result in the formation of lobes and intracellular air spaces, as in the
spongy mesophyll of the leaf 5. Adjacent tissues can also expand at
different rates; for example, leaf epidermal cells continue to enlarge
after the subepidermal mesophyll cells have ceased growth, influ-
encing development of a substomatal cavity and anticlinal cell-wall
undulations in abaxial epidermal cells. Epidermal cells, which
1.7 Tissues 11

Figure 1.7 Euphorbia eyassiana (eudicot: Euphorbiaceae), longitudinal section

of stem showing branched non-articulated laticifer. Scale = 50 µm

require additional strength in their protective role, display differ-

ential growth between their anticlinal and periclinal walls64, 78, 126.
Some specialized cells are capable of apical intrusive growth (tip
growth) that allows them to separate the middle lamella between
adjacent cell walls and intrude between them, prior to the onset of
cell-wall thickening66, 94. Fibres, which collectively confer tensile
strength to tissues such as secondary xylem, extend axially at both
ends by apical intrusive growth. Similarly, non-articulated laticifers
(e.g. in Euphorbia) branch and ramify throughout the plant by apical
growth (Figure 1.7). Other specialized cells capable of apical intrusive
growth include root hairs, pollen tubes and star-shaped astrosclereids.

1.7 Tissues
Simple tissues, such as parenchyma, collenchyma and sclerenchyma,
consist of regions of similar individual cells, often interspersed with
isolated specialized cells (idioblasts43) and secretory cells or canals.
The bulk of the primary plant body (the ground tissue) consists of
12 Organs, cells and tissues

simple tissues. In contrast, complex tissues incorporate elements of

several different simple cell and tissue types. Complex vascular tissues
(phloem and xylem) can incorporate parenchyma, sclerenchyma and
vascular cells. The epidermis is a dermal tissue that encloses the entire
plant and is continuous between the various organs. Primary tissues
are produced by an apical meristem, and secondary tissues are pro-
duced by lateral meristems such as the vascular cambium.

1.7.1 Parenchyma and other thin-walled cell types

The term parenchyma incorporates the relatively unspecialized
cells that occur in both primary and secondary tissues of the
plant body; it includes many cells with living contents that do
not fit readily into other categories. Parenchyma cells are thin
walled and often loosely packed together, leaving small air spaces
between them where their walls are not in contact with each other.
Individual parenchyma cells are polyhedral or amorphous, some-
times axially elongated or even lobed.
Chlorenchyma cells are green thin-walled cells that contain chlor-
oplasts. In leaves (Chapter 4), chlorenchyma tissue is termed meso-
phyll. Mesophyll is often differentiated into two distinct zones:
palisade mesophyll on the adaxial side of the lamina and spongy
mesophyll on the abaxial side. Palisade mesophyll consists of rows
of cells that are tightly packed together and anticlinally elongated;
spongy mesophyll consists of cells of various shapes that are rela-
tively loosely packed, with many intercellular air spaces.
Aerenchyma is a specialized parenchymatous tissue that char-
acterizes many aquatic plants. It represents a regular, well-
developed system of cells interspersed with considerably enlarged
intercellular air spaces that facilitate internal diffusion of gases
(Figure 1.8). Contrasting developmental patterns of aerenchyma
exist in different species38. Lysigenous aerenchyma involves pro-
grammed cell death during development, leaving regular spaces
(e.g. in the cortex of a maize root). In contrast, in schizogenous
aerenchyma, which is relatively complex and more structured, the
1.7 Tissues 13

Figure 1.8 Nymphaea miniata (ANA-grade: Nymphaeaceae), transverse section

of petiole showing aerenchyma. Scale = 100 µm

large air spaces are formed by differential growth of surrounding

tissues. For example, in leaves, stems and roots of some aquatic
plants (e.g. Hydrocharis), schizogenous aerenchyma is associated
with a system of transverse septa or diaphragms that provide
mechanical resistance to compression.
Transfer cells are specialized plant cells that facilitate transfer of
soluble substances across tissue boundaries. They typically possess
cell-wall ingrowths protruding into their protoplasts, thus increas-
ing their surface area. They occur in companion cells in phloem
(especially at the node of a stem), in root nodules, in the haustoria
of parasitic plants and in the epidermis of water plants98.

1.7.2 Thick-walled tissues: collenchyma

and sclerenchyma
Collenchyma is a strengthening tissue that consists of groups of
axially elongated, tightly packed cells with unevenly thickened
walls. Collenchyma tissue typically occurs as primary ground
14 Organs, cells and tissues

tissue, often located in the angles of young stems, or in the leaf

midrib or petiole. Collenchyma cells differ from fibres in that they
often retain their living protoplasts and their cell walls are rela-
tively unlignified, though they may ultimately become lignified as
the tissue ages.
Sclerenchyma cells lack contents at maturity and possess evenly
thickened walls that are usually lignified, with simple pits that are
often slit-like. Sclerenchyma occurs in both primary and secondary
tissues, either in groups or individually as idioblasts interspersed in
other tissue types. Fibres are long, narrow sclerenchyma cells that are
elongated along the long axis of the organ and possess tapering ends.
Fibres generally occur in groups, often located at the phloem poles of
vascular bundles (Figures 2.2, 2.3, 2.6). The bulk of the secondary
xylem (wood) in woody eudicots consists of fibres that confer
considerable tensile strength. Bast fibres are formed in the secondary
phloem and cortex in some species; they are used to make textiles or
rope in species such as flax (Linum) and hemp (Cannabis). Gelatinous
fibres are formed in the secondary xylem or sometimes in the bark of
some woody plants; they are typically formed in response to stress
caused by asymmetric branch thickening (compression wood or
tension wood). In gelatinous fibres, the inner secondary wall is non-
lignified and rich in polysaccharides, making it more flexible39.
Sclereids (stone cells) are variously shaped cells that develop
thick secondary walls as the plant matures; they can occur
throughout the plant, either as isolated cells (idioblasts) or in
groups (e.g. groups of stone cells in the fruit endocarp of
peach). Brachysclereids are isolated isodiametric cells dispersed
among parenchyma. Astrosclereids develop projections that
grow intrusively into adjacent intercellular air spaces or along
middle lamellae during the growth phase of the organ, prior to
cell-wall thickening. The shapes of astrosclereids are dictated by
the cellular arrangement of surrounding tissues; they are often
star-shaped (Figure 1.9), though sclereids located in palisade
mesophyll can be bone-shaped (osteosclereids).
1.8 Epidermis 15

Figure 1.9 Camellia japonica (eudicot: Theaceae), transverse section of leaf

midrib with branched sclereid. Scale = 20 µm

1.8 Epidermis
The outermost (dermal) cell layer, the epidermis, covers the entire
plant surface; in the root it is sometimes termed rhizodermis. The
aerial epidermis is covered with a hydrophobic cuticle layer that
consists primarily of a complex polymer, cutin, which forms
a protective barrier to water and CO2 and also prevents adhesion
between cells of adjacent organs in tightly packed structures such
as the bud. The cuticle permeates the outermost cell wall and also
forms an outer skin of varying thickness, often associated with
epicuticular waxes70 (Figure 4.7). The cuticle can be striated or
variously ornamented.
The epidermis is a primary tissue that is initially derived from
the protodermal cells of the apical meristem. In developing organs,
both anticlinal divisions and cell elongation extend the epidermis
to accommodate organ elongation and even some organ
16 Organs, cells and tissues

thickening. The epidermis is usually uniseriate, with rare excep-

tions: a few species (e.g. Ficus elastica) develop a multiseriate
epidermis in their leaves, and the aerial roots of many orchids
possess a specialized multiseriate epidermis termed velamen. The
epidermis generally persists throughout the life of the plant
except in regions of lateral thickening growth, where it is often
replaced by a periderm. In older roots, it is sometimes worn away
by friction with the soil, and the root is subsequently protected
by an exodermis formed by cell-wall thickening in the outer
cortical layers.
The epidermis can develop various specialized cell types. For
example, in leaves, relatively undifferentiated pavement epidermal
cells are interspersed with several specialized cell types, including
stomata and trichomes.

1.9 Stomata
Stomata are specialized pores in the epidermis through which
gaseous exchange (water release and CO2 uptake) takes place.
They occur not only on most aerial plant surfaces, especially on
green photosynthetic stems and leaves, but also on some floral
organs. Each stoma consists of two guard cells surrounding
a central pore (Figures 1.10, 1.11). Guard cells are either kidney-
shaped (in most plants) or dumbbell-shaped (in grasses, sedges and
allied families); their inflation by increased osmotic potential results
in opening of the pore, and their deflation causes its closure.
Cuticular ridges extending over or under the pore from the outer
or inner edges of the guard-cell walls help to seal the closed pore. In
contrast with most epidermal cells, guard cells typically contain
plastids that function as both chloroplasts and amyloplasts; these
specialized plastids are smaller than those of mesophyll cells and
contain photoreceptors that help to control stomatal opening80, 122.
Epidermal cells adjacent to the guard cells are termed sub-
sidiary cells if their shape differs from that of other pavement
epidermal cells; they can be either lateral or polar with respect to
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