Paula Rudall - Anatomy of Flowering Plants (2020)

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
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Cambridge University Press is part of the University of Cambridge.
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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
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
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
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
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
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
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
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
1.9 Stomata 17
Figure 1.10 Arabidopsis thaliana (eudicot: Brassicaceae), abaxial leaf surface,

showing a single stomatal pore (SEM). Scale = 10 µm
the guard cells. Specialized lateral subsidiary cells (e.g. in grasses

and palms) can supply the guard cells with solutes that help to
drive diurnal stomatal opening and closure90.
Classifications of stomatal types are based either on the presence
and arrangement of mature subsidiary cells or on their patterns of
development99, 116, 156. Types of mature stomata include anomocytic
(lacking subsidiary cells) and paracytic (possessing one or more pairs
of lateral subsidiary cells oriented parallel with the guard cells).
Lateral subsidiary cells of different species – though apparently
similar – can have different developmental origins; mesogene
lateral subsidiary cells are derived from the same cell lineage as
the guard-mother cell, whereas perigene lateral subsidiary cells are
derived from an adjacent cell lineage99, 116. During development
of the leaf epidermis in many eudicots (e.g. Arabidopsis), an iterative
Figure 1.11 Amborella trichopoda (ANA-grade: Amborellaceae), transverse

section of leaf epidermis through a stomatal pore. Scale = 10 µm
series of asymmetric divisions (amplifying divisions) results in

a spiral arrangement of mesogene epidermal cells with a stomatal
pore at the centre; the entire complex of cells is derived from the
same initial cell and hence is monoclonal92, 117.
1.10 Trichomes and papillae

Epidermal outgrowths include papillae, trichomes (hairs) and spe-
cialized domed cells that characterize many petal surfaces. Papillae
are small unicellular outgrowths from a single epidermal cell; some
leaf surfaces possess multiple papillae per cell. Trichomes can occur
on all parts of the plant surface; they vary widely in both form and
function, partly depending on their location in the plant. Trichomes
(Figure 1.12) can be unicellular or multicellular, branched or
unbranched; specialized forms include root hairs (Chapter 3),
glandular (secretory) hairs, scales, sensory hairs, stinging hairs
(Figure 1.13) and hooked hairs that facilitate seed dispersal.
Glandular trichomes usually possess a unicellular or multicellular
stalk and a secretory head consisting of one or more cells. Secreted
1.10 Trichomes and papillae 19
Figure 1.12 Plant hairs. (left) Quercus ilex (eudicot: Fagaceae), stellate non-
glandular trichome on abaxial leaf surface. Scale = 50 µm. (right) Salvia
involucrata (eudicot: Lamiaceae), trichomes on petal surface, including
glandular (secretory) hairs with short stalks and either four-celled or single-
celled glands. Scale = 20 µm.
Figure 1.13 Stinging hairs in Urtica dioica (eudicot: Urticaceae). (left) adaxial
leaf surface with many trichomes; the largest one is a stinging hair with
a glandular tip. Scale = 100 µm. (right) tips of stinging hairs, with apical
glands intact and broken (SEM). Scale = 10 µm.
substances such as volatile oils collect between the secretory cells and
a raised cuticle, which ultimately breaks to release the exudate. There
are many different types of glandular hair that secrete a variety of
substances, including salt, mucilage, essential oils and even
enzymes41, 154. Leaf glandular hairs of Cannabis sativa secrete
a resinous substance containing cannabinoids that act as plant defense
compounds. Glandular hairs of carnivorous plants such as Drosera
secrete both sticky mucilage and proteolytic enzymes that help to
capture and digest the prey. Salt-secreting glands help to modulate
ion concentration in leaves of salt-tolerant plants such as the man-
grove Avicennia70.
Some trichomes are specialized for water absorption rather than
secretion. Examples include the leaf scales that occur in many
epiphytic and rock-dwelling Bromeliaceae, which represent an
important source of water and mineral uptake to the plant. The
specialized hairs (hydropotes) that occur on the submerged sur-
faces of many water plants (e.g. waterlily, Nymphaea) initially
secrete mucilage, but following degeneration of the cap cells
they absorb water and minerals and accumulate heavy metals.
1.11 Vascular tissue

Vascular tissue consists of associated networks of cells that conduct
water (xylem) and nutrients (phloem). Primary vascular tissue is
derived from the procambium, which is itself produced by the
apical meristems. Secondary vascular tissue is derived from the
vascular cambium in eudicots, and from the secondary thickening
meristem in a few monocot species (Chapter 2.8). Both xylem and
phloem are complex tissues composed of many different cell types.
1.11.1 Xylem
The primary function of xylem is transport of water throughout the
plant, from roots via the stems to the leaves, where water is combined
with carbon dioxide to make carbohydrates and oxygen during
photosynthesis. Xylem is composed of several distinct cell types,
1.11 Vascular tissue 21
Figure 1.14 Vascular bundle. Lilium tigrinum (monocot: Liliaceae), transverse

section of stem vascular bundle with xylem (left) and phloem (right)
encircled by a ring of bundle sheath cells. bs = bundle sheath, c = companion
cell, mx = metaxylem vessel, px = protoxylem vessel, s = sieve tube element.
Scale = 50 µm.
often including parenchyma and fibres as well as vessels. Tracheids

and vessel elements (collectively termed tracheary elements) are the
water-conducting cells; they lack contents at maturity and are linked
into cell chains to form vessels. Tracheary elements are elongated cells
with thickened lignified walls. In a stem vascular bundle (Figure
1.14), the first-formed (protoxylem) elements often possess wall
thickenings that are either helical or arranged in rings (annular).
Later-formed primary tracheary elements (metaxylem) and second-
ary tracheary elements possess bordered pits in their lateral walls.
Bordered pits18 can be oval, polygonal or elongated (scalariform);
they can be organized in transverse rows (opposite pitting) or tightly
packed (alternate pitting).
The primary difference between tracheids and vessel elements is
that vessel elements possess large perforations in their adjoining end
walls, whereas perforations are absent from tracheids. Perforation
plates are generally either simple, with a single opening, or scalari-

form, with a ladder-like row of openings divided by a series of parallel
bars, or rarely a reticulate mesh. Vessel elements differ considerably in
diameter, not only between different species but also sometimes
across a single growth ring (e.g. in secondary xylem of oak).
1.11.2 Phloem
Phloem has complex roles in translocation of nutrients (sucrose and
electrolytes) and hormones throughout the plant39. Although com-
monly associated with xylem, phloem can develop precociously in
regions that require a plentiful supply of nutrients, such as devel-
oping sporogenous tissue. Phloem consists of conducting cells
(sieve elements) and associated specialized parenchyma cells (com-
panion cells) (Figure 1.14); these two closely interdependent cell
types are produced from a common parent cell (meristemoid) that
divides and develops asymmetrically to form a larger sieve element
and smaller companion cell. Most angiosperms possess sieve-tube
elements rather than the relatively unspecialized sieve cells.
At maturity, sieve elements lack nuclei and most organelles but
retain plastids and phloem-specific proteins (P-proteins).
Companion cells are densely cytoplasmic, retaining nuclei and
many active mitochondria. Sieve element plastids and P-proteins
occur in several morphological forms (amorphous, filamentous,
tubular and crystalline) that are often highly characteristic for parti-
cular plant families and are thus of systematic and evolutionary
value11,145. Sieve-element plastids are classified according to their
inclusions: starch (S-type plastids), protein (P-type plastids), or both.
Sieve elements are linked axially to form sieve tubes via sieve
plates. Slime plugs are formed when P-protein accumulates on
a sieve plate39. The walls of sieve elements are thin and possess
characteristic regions (sieve areas) that connect adjacent sieve ele-
ments; sieve areas consist of groups of pores and associated callose.
In sieve tube elements, the sieve areas are localized on the adjoining
end walls, forming sieve plates that are either simple or compound.
1.12 Secretory cells and laticifers 23
1.12 Secretory cells and laticifers

Secretory cells are specialized cells that release a broad range of
substances, often targeted at a particular function. Secretory cells
can occur throughout the plant body, either in the epidermis or in
internal tissues. Epidermal secretory cells include glandular tri-
chomes and nectaries, both extrafloral and floral. Internal secretory
structures include idioblasts and specialized ducts or canals that are
lined with cells that secrete substances such as resin or mucilage.
Characteristic resin canals occur in the secondary xylem of some
woody eudicots.
Laticifers are specialized latex-producing cells that permeate the
tissues of a few angiosperms (Figure 1.7). Latex is an aqueous,
often milky, solution that represents the source of commercial
rubber, which is traditionally harvested from the rubber tree,
Hevea brasiliensis. Laticifers of the opium poppy (Papaver somniferum)
produce latex that is the source of narcotic analgesics such as
morphine. Laticifers are traditionally classified into two cell
types: non-articulated and articulated, though both types can
occur within the same plant family (e.g. in Euphorbiaceae) and
overlap in some of their properties86,108,110. Non-articulated lati-
cifers (e.g. in Euphorbia, Figure 1.7) are highly branched cells that
are derived from a small group of initial cells in the cotyledonary
node of the embryo. These cells are coenocytes that undergo
repeated nuclear divisions without corresponding wall formation.
The resulting multinucleate cells grow intrusively between cells of
surrounding tissues, branching frequently and eventually ramify-
ing throughout the entire plant. In contrast, articulated laticifers
(e.g. in Hevea brasiliensis) form complex syncytia by cell fusion and
consist of linked chains of cells. Articulated laticifers can also grow
intrusively between adjacent cells, though not to the same extent as
the non-articulated cell type.
2
Stem
Stems are axes that are typically cylindrical, elongated and branch-
ing, though many modifications can occur in different species.
Shoot apical meristems are present at the tips of all the stem
branches; lateral branches are initiated from buds that are borne
in the axils of leaves. Stems are most commonly aerial, though
some stems occur below ground. Aerial stems are often green and
photosynthetic during early growth but subsequently turn brown
following radial stem thickening. Some underground stems are
modified into storage organs such as corms or rhizomes that allow
them to survive a harsh winter or dry season below ground.
2.1 Shoot apex

The shoot apex contributes to extension growth of the shoot and
initiates the leaf primordia at the nodes. Many shoot apices con-
tinue to grow indefinitely (i.e. they are indeterminate), though
some (e.g. those of shoot thorns and flowers) rapidly cease growth
and become determinate. In seed plants, lateral branches develop
from axillary buds located in the axils of leaves. Axillary buds
possess shoot apical meristems that remain in a dormant state
until they receive a growth stimulus.
The vegetative shoot apical meristem is typically dome-shaped
and partitioned by clear zones of activity (Figures 1.1, 2.1). The
outermost two (or more) cell layers are primarily maintained by
anticlinal cell divisions. In flowering plants, these outer layers are
termed L1 and L2 respectively, or collectively the tunica region.
2.1 Shoot apex 25
Figure 2.1 Diagram of angiosperm shoot apical organization
The inner corpus region (L3), in which cell divisions are more
randomly oriented, is the region proximal to the tunica. Thus, the
outer tunica region contributes to surface growth and the inner
corpus region to increase in volume, though there is often slight
intergradation between them39. The central region underlying the
corpus zone is a rib meristem, which gives rise to files of cells that
form the pith. It is surrounded by a peripheral flank meristem that
produces the leaf primordia cortex and procambium, which itself
gives rise to the primary vascular system.
Leaf primordia are initiated at or near the shoot apex in a regular
arrangement. In many species, a single leaf develops at each node,
though some species are characterized by a pair of leaves or
a whorl of leaves at each node. Extension growth of the stem
occurs at the internodes, either by diffuse cell divisions and cell
elongation throughout the internode, or sometimes in a more
restricted region, often localized at the base of the internode.
The reproductive shoot apex produces an inflorescence, and each
flower apex produces a single flower. At the transition to flowering,
there is an overall increase in mitotic activity in cells above the rib
meristem, resulting in a change in shape of the shoot apex39.
26 Stem
2.2 Primary stem structure

Primary vascular tissue in the stem typically consists of discrete
vascular bundles supported by parenchymatous ground tissue that
is present throughout the stem in the central pith and the outer
cortex. In older stems the pith sometimes breaks down to form
a central hollow cavity. Ground parenchyma cells frequently
become lignified as the plant ages. Ridged or angled stems often
possess strengthening collenchyma at the angles, located in the
outer cortex immediately inside the epidermis. Many young stems
are green, photosynthetic organs with a chlorenchymatous cortex.
In some stems, endogenous adventitious roots are initiated at the
junction between the cortex and the vascular region (the pericyclic
region).
Some plant stems possess secretory cells or ducts in the ground
tissue. For example, species of Euphorbia possess branched networks
of laticifers in the cortex, which extend throughout the ground
tissue of the stem and leaves. Plants with succulent stems (e.g.
many Cactaceae) possess large thin-walled cells that contain a high
proportion of water. Some stems that are specialized as storage or
perennating organs (e.g. corms of Crocus) store food reserves in the
form of starch granules, especially in the inner cortex. In some
species, the layer of cortical cells immediately adjacent to the
vascular tissue is packed with starch granules; this is termed
a starch sheath, or sometimes an endodermoid layer or endoder-
mis, although the component cells usually lack the suberinized
Casparian thickenings that are found in the root endodermis.
In eudicots and magnoliids, primary vascular tissue in the stem
typically consists of a ring of discrete collateral bundles, in which
the phloem is external to the xylem (Figure 2.2). Some stems also
possess medullary (pith) bundles or cortical bundles, which are
often associated with leaf vasculature. Within each primary vas-
cular bundle, discrete strands of xylem and phloem exhibit a range
of configurations: either adjacent to each other (in collateral vas-
cular bundles), with strands of phloem on each side of the xylem
2.2 Primary stem structure 27
Figure 2.2 Vicia faba (eudicot: Fabaceae), transverse section of stem.

Scale = 100 µm
(bicollateral vascular bundles), or with xylem surrounding the

phloem (amphivasal vascular bundles). Amphivasal bundles are
common in monocots but relatively unusual in eudicot stems. In
stems with bicollateral bundles (e.g. Cucurbita), both internal and
external phloem is present. Vascular cambium (Figure 2.3) is
located between the xylem and phloem in most woody eudicots
and magnoliids, but this meristem is absent from monocot vascu-
lar bundles (Figure 1.14).
In monocots, the stem vascular bundles are typically scattered
through the central ground tissue (Figures 1.14, 2.4), or sometimes
arranged in two or more distinct rings. Monocot vascular bundles
are collateral, bicollateral or amphivasal. Cortex and pith regions are
often indistinctly demarcated from each other, though the cortex
may be defined by an endodermoid layer or a cylinder of scler-
enchyma. The monocot vascular system can be highly complex163.
Each bundle, when traced on an upward course from any point in
the stem, often has several bridges or branches before passing into
a leaf; one of its major branches continues a similar upward course
28 Stem
Figure 2.3 Vascular cambium. Hyptis oblongifolia (eudicot: Lamiaceae),

transverse section of part of stem in region of vascular cambium.
Scale = 20 µm
towards the apex. In some palms, thousands of vascular bundles are

present in a single transverse section of the stem, though in most
monocots their number is much fewer.
2.3 Nodal anatomy

The vasculature of the leaf and stem is connected at the nodes.
Openings (lacunae or leaf gaps) occur in the stem vascular cylinder
beneath their point of contact (Figure 2.5). In eudicots and mag-
noliids, nodal anatomy is often characteristic of families,
particularly with respect to the number and arrangement of leaf
traces and leaf gaps. Nodes are unilacunar, trilacunar or
multilacunar, depending on the number of leaf gaps in the stem
vascular cylinder. This feature is most apparent in stems that
possess an otherwise continuous vascular cylinder, especially
where a limited amount of secondary thickening has taken place,
so that nodal anatomy is best understood in woody plants62. The
number of leaf traces departing from each gap is also generally
characteristic of a species, but can vary within a plant, especially in
2.3 Nodal anatomy 29
Figure 2.4 Monocot stem. Lilium tigrinum (monocot: Liliaceae), transverse

section of stem showing outer cortex surrounding central region containing
vascular bundles and demarcated by sclerenchymatous cylinder.
Scale = 100 µm
species with unilacunar and trilacunar nodes. For example, in

Clerodendrum (Lamiaceae) two traces depart from a single gap, and
in Prunus (Rosaceae) a single trace departs from each of three gaps
in the central vascular cylinder (Figure 2.5). In oaks (Quercus), up to
five traces depart from a trilacunar node. Leaf-trace bundles nor-
mally develop acropetally from the stem procambial system close
to the shoot apex and grow towards developing leaf primordia,
though in some species (e.g. Populus deltoides), subsidiary vascular
bundles are initiated at the base of each developing primordium
and grow basipetally to meet the stem procambial trace79.
Nodal vasculature is further complicated by the axillary bud
vascular traces, which are connected to the main stem vasculature
immediately above the leaf gaps. In most species, two traces
30 Stem
Figure 2.5 Nodal anatomy. Prunus lusitanica (eudicot: Rosaceae), transverse

section of part of stem at node, showing connection of petiole vasculature
with stem vascular cylinder. Scale = 100 µm
diverge to supply each bud or branch. In large woody eudicot

trees, the junction of the trunk and its branches is characterized by
a complex arrangement of secondary vascular tissue, which typi-
cally forms a collar around the base of the branch. This branch
collar is enveloped by a trunk collar that links the vascular tissue of
the trunk above and below the branch. There is no direct connec-
tion of xylem from the trunk above a branch into the branch
xylem, as the tissues are oriented perpendicular to each other. If
a branch dies, a protection zone forms around its base to prevent
a spread of infection into the trunk, and the branch is often shed.
2.4 Vascular cambium and secondary vascular tissue

Increase in stem length, achieved by extension growth at the shoot
apical meristem, is followed by radial growth that results in increase
2.4 Vascular cambium and secondary vascular tissue 31
Figure 2.6 Secondary thickening in eudicots. Cissus alata (eudicot: Vitaceae),

transverse section of stem, showing early growth in thickness. Scale = 200 µm
in stem thickness129 (Figure 2.6). In most woody angiosperms,

secondary vascular tissue (both xylem and phloem) is generated by
the vascular cambium, which becomes active a short distance behind
the stem apex, initially as a fascicular meristem formed within the
primary vascular bundles. Subsequently, an interfascicular vascular
cambium is formed in the parenchymatous region between the
primary vascular bundles, resulting in a vascular cylinder.
The vascular cambium is a bifacial meristem (Figure 2.3) that
generates secondary xylem (wood) at its inner edge (centripetally)
and secondary phloem at its outer edge (centrifugally). It consists of
both fusiform initials and ray initials, which form the axial and radial
systems respectively. Ray initials are isodiametric cells that divide
periclinally to form ray parenchyma cells in both xylem and phloem.
Fusiform initials are axially elongated cells with tapering ends. These
undergo periclinal divisions to form the axial elements of the
32 Stem
Figure 2.7 Wood. Quercus robur (eudicot: Fagaceae), wood block at edge of
transverse and tangential longitudinal surfaces, showing large early wood
vessels (SEM). Scale = 100 µm
secondary tissues, including tracheary elements, fibres and axial

parenchyma in secondary xylem, and sieve elements, companion
cells and fibres in secondary phloem. In addition, fusiform initials
sometimes give rise to new ray initials as the stem increases in
circumference and new rays are formed. Occasionally, transitional
cell types can occur. For example, perforated ray cells, an unusual
feature of some woods, can link two vessel elements and themselves
resemble and function as vessel elements; their perforation plates
correspond with those of the adjacent vessel elements, though they
are formed from ray initials.
The degree of secondary growth depends on the growth habit
of the plant. In some species, including all monocots, some mag-
noliids and some herbaceous eudicots (e.g. Ranunculus), a vascular
cambium is entirely absent. In many woody temperate plants,
cambial activity is seasonal (usually annual), resulting in the
2.4 Vascular cambium and secondary vascular tissue 33
Figure 2.8 Wood. Alnus glutinosa (eudicot: Betulaceae), transverse section of

secondary xylem, showing differences in fibre thickness on either side of
growth-ring boundary. Scale = 100 µm
formation of growth rings. Xylem of early wood formed in the

spring is generally less dense and consists of thinner-walled cells
than that of late wood (summer wood). In ring-porous woods
such as oak, the early wood vessels are considerably larger than
those of late wood (Figure 2.7). In diffuse-porous woods, the
main distinction between early and late wood is in relative fibre
size and wall thickness (Figure 2.8).
Atypical activity of the vascular cambium characterizes some
woody angiosperms with anomalous secondary growth, such as
many climbing plants (lianes)129. For example, in some species
(e.g. Avicennia) isolated regions of phloem (included or interxylary
phloem) are formed embedded in the xylem, either in islands or in
alternating concentric bands. Other species have irregularly
34 Stem
Figure 2.9 Anomalous secondary growth. Tynanthus elegans (eudicot:

Bignoniaceae), transverse section of woody stem, showing four regions of
phloem embedded in secondary xylem. Scale = 1 mm
divided or deeply fissured areas of xylem and phloem caused by

differential cambial activity in different radial sectors (Figure 2.9).
2.5 Secondary xylem

Secondary xylem (wood) is produced by the vascular cambium. It is
composed of a matrix of cells, some arranged axially (fibres, vessels
and axial parenchyma) and others radially. The ray parenchyma
forms the wood rays that radiate from the vascular cambium towards
the pith. As woody plants age and their trunks increase in girth, the
central area becomes non-functional with respect to water transport
or food storage; the vessels frequently become blocked by tyloses that
are formed when adjacent parenchyma cells grow into them through
common pit fields. The central non-functional area of the trunk, the
2.6 Secondary phloem 35
heartwood, is generally darker than the outer living sapwood; it

provides a supporting skeleton for the living parts of the tree.
The precise cellular arrangement of wood is often highly character-
istic of the species (Figures 2.7, 2.8). To observe their structure,
woods are sectioned transversely and in two longitudinal planes:
along the rays (radial or median longitudinal section: RLS or MLS)
and across the rays (tangential longitudinal section: TLS). When
viewed in transverse section, vessels can appear solitary (e.g. the
early vessels of oak, Quercus) or arranged in clusters or radial chains.
Axial parenchyma cells can be independent of the vessels (apotracheal
parenchyma) or associated with them (paratracheal parenchyma), or
they can sometimes occur in regular tangential bands. The relative
abundance of axial parenchyma differs between species, ranging from
sparse or absent to rare cases such as balsa wood (Ochroma pyramidale), in
which axial parenchyma cells are often more abundant than fibres.
Rays are termed uniseriate if they are one cell wide tangentially,
and multiseriate if they are more than one cell wide. Both uniseriate
and multiseriate rays can co-occur in the same wood (e.g. in oak,
Quercus: Figure 2.7). In radial longitudinal section, ray cells can be
homogeneous (homocellular) or variously shaped (heterocellular).
Other aspects of variation in the structure of hardwoods include the
presence of either axial or radial secretory canals in some woods,
storied (stratified) appearance of the rays and differences in the fine
structure of the vessel walls (intervascular pitting, perforation plates
and wall thickenings). For example, in lime wood (Tilia cordata), the
vessel element walls are helically thickened, and in many legumes
(Fabaceae) the pit apertures are surrounded by characteristic warty
protuberances, termed vesturing.
2.6 Secondary phloem

Secondary phloem is also produced by the vascular cambium. It
consists of both axial and radial systems that are formed from the
fusiform initials and ray initials, respectively. In transverse section,
36 Stem
phloem rays are radially continuous with xylem rays. Close to the
vascular cambium, they can be either uniseriate or multiseriate, but
they become dilated towards the cortex as a result of cell divisions that
accommodate increase in stem thickness (Figure 2.6), and older ray
cells often become lignified. At their outer periphery, parenchyma-
tous ray cells are indistinguishable from cortical cells and form part of
the bark. The axial system of the secondary phloem not only consists
of sieve elements and companion cells, as in primary phloem, but also
includes fibres, sclereids and axial parenchyma cells. In some species,
the phloem fibres are formed in groups at regular intervals, resulting
in characteristic tangential bands of fibres alternating with groups of
sieve elements and parenchyma cells.
2.7 Periderm and bark

Periderm is a specialized protective tissue of corky (suberinized)
cells that is produced in the outer layers of the cortex of a stem or
root that has increased in thickness. Bark is a more generalized
term that includes all tissues located outside the vascular cambium;
it is commonly applied to the outermost layers of stems of woody
plants, especially trees3.
Periderm forms an impervious layer to prevent water loss and
protect against injury. It is commonly formed in the cortex of secon-
darily thickened stems, replacing the epidermis, which splits and peels
away (Figure 2.10). It can also form in response to a wound, for
example forming a protective layer over a site of leaf abscission. In
stems and roots, periderm consists of up to three distinct layers,
termed phellogen, phellem and phelloderm39. The phellogen layer is
a uniseriate lateral meristem composed of thin-walled cells that divide
to produce phellem to the outside and (in some cases) phelloderm to
the inside. Phellem represents the most prominent region of periderm;
it is arranged in tightly packed rows of cells. Phellem cells lack contents
at maturity and possess cell-wall deposits of suberin and sometimes
lignin, especially on their inner faces. In contrast, the phelloderm cells
2.7 Periderm and bark 37
Figure 2.10 Periderm formation. Sambucus nigra (eudicot: Adoxaceae),

transverse section of slightly thickened stem, showing periderm forming in
the subepidermal outer cortex. Scale = 100 µm
located inside the phellogen layer are non-suberinized and can con-
tribute to the secondary cortex of a stem. The initial phellogen
originates either in the subepidermal (hypodermal) layer or deeper
in the cortex, more rarely within the epidermis itself.
Bark tissues include secondary phloem, fibres, cortex and
periderm3. In some woody trees, the bark can become very thick
and two distinct regions are distinguishable: inner bark, consisting of
secondary phloem tissues, and outer bark (rhytidome), consisting of
cortex and periderm tissues. In some species, several phellogens can
form sequentially or almost simultaneously; they frequently overlap
each other within the bark. The pattern of periderm formation largely
dictates the appearance of the bark of a woody plant. For example, the
smooth papery bark of a birch (Betula) tree results from regular
formation of abscission bands causing thin sheets to flake off. In cork
oak (Quercus suber), the initial phellogen can maintain activity indefi-
nitely and produces seasonal growth rings. In trees grown for the
38 Stem
production of commercial cork, the initial phellogen is removed after

about 20 years to make way for a second, more vigorous phellogen.
Many species of woody eudicots and magnoliids produce lenti-
cels, which are restricted regions of loose cells formed within the
periderm that protrude outwards from the surface of the bark.
Lenticels also occur on the surface of tubers such as potato. They
are initially formed in cells underlying stomata in the epidermis
and have a similar role in allowing passage of gases and water
vapour across the stem, at least during the summer months,
though their role may change with the seasons52. In a few species
(e.g. several Bombacoideae–Malvaceae), localized regions of the
outermost phellogen formed proliferated cell layers that result in
pronounced rigid outgrowths of the bark (prickles).
2.8 Primary and secondary thickening meristems

in monocots
In monocots, which lack a vascular cambium, a limited degree of
radial growth is achieved by a primary thickening meristem (PTM)
Figure 2.11 Diagram of monocot shoot apical organization showing PTM

near shoot apex
2.8 Primary & secondary thickening meristems in monocots 39
near the vegetative shoot apex (Figure 2.11). The PTM is especially
extensive in species with short internodes and crowded leaves, such
as bulbous taxa27, 109. In monocots, the PTM originates in ground
tissue in the pericyclic region of the stem. It is a tiered meristem,
consisting of a zone of meristematic cells that produces radial deri-
vatives. The majority of its radial derivatives are centripetal, consisting
of both parenchyma and discrete vascular bundles, though it also
produces some parenchyma towards the outside. In addition to
primary stem thickening, the PTM forms linkages between root,
stem and leaf vasculature, especially in relatively condensed bulbous
plants. It typically ceases activity at a short distance behind the apex,
and subsequent stem thickening is limited. However, in some spe-
cies, this region retains some meristematic potential and resumes
meristematic activity further down the stem; it represents the site of
adventitious root production in some monocots109.
Although monocots are predominantly herbaceous, some species
achieve considerable height and girth. Tree-forming palms possess
an extensive PTM that results in a large sunken shoot apex; in these
species, further stem thickening occurs by subsequent division and
enlargement of ground parenchyma cells, termed diffuse secondary
growth. A PTM is not confined to monocots; similar pericyclic
meristems also occur in some eudicots with thick stems (e.g.
Cactaceae14). In some woody monocots of the order Asparagales
(e.g. Agave, Aloe, Cordyline, Yucca), considerable increase in stem thick-
ness is achieved by means of a secondary thickening meristem
(STM) (Figure 2.12). The STM resembles the PTM in that it is
radially located in the pericyclic region of the stem within the cortex
and it produces radial derivatives, but it occurs further from the
shoot apex; the two meristems are best regarded as developmental
phases of the same meristem. The STM produces secondary vascular
bundles that are mostly amphivasal and radially elongated. In some
woody monocots (e.g. Beaucarnea recurvata, Cordyline terminalis), the PTM
and STM are axially discontinuous, whereas in others (e.g. Yucca
whipplei) they are axially continuous29, 132, 133.
40 Stem
Figure 2.12 Monocot growth in thickness. Cordyline indivisa (monocot:

Asparagaceae), transverse section of stem, showing outer cortex and inner
region containing scattered vascular bundles. The STM has produced
secondary vascular bundles that are radially aligned. Scale = 100 µm
The PTM and STM, although sometimes termed the monocot

cambium, are not homologous with the vascular cambium. The
PTM and STM originate from the pericyclic region, whereas the
vascular cambium originates inside stem vascular bundles (fasci-
cular cambium) and subsequently spreads to interfascicular
regions. The PTM and STM produce discrete vascular bundles
centrifugally, whereas the vascular cambium is a bifacial meristem
that produces xylem centrifugally and phloem centripetally.
3
Root
Roots are typically branching cylindrical structures that develop

underground to facilitate extraction of moisture and nutrients
from the soil, often in association with hyphal networks of soil-
dwelling fungi. In contrast, epiphytes and epiliths, which grow
entirely above the ground, often develop aerial roots that absorb
moisture from their environment. In vascular plants, the root
apex is a growing tip where both the root cap and the primary
root tissues are produced. Lateral roots are initiated some dis-
tance from the root apex, by cell divisions in the pericycle,
among differentiated cells that have retained meristematic
capacity.
3.1 Root apex and root cap

The root apex contributes to extension growth of the root and also
differentiates the terminal protective root cap (Figure 3.1). In
contrast with the shoot apex, the root apex does not produce
lateral appendages and grows relatively uniformly, lacking nodes
and internodes7, 42. The junction between the root cap and the
root apical meristem is either clearly defined by a distinct cell
boundary (closed organization, as in Arabidopsis thaliana and Zea
mays) or ill-defined (open structure, as in Vicia faba), though inter-
mediates exist (e.g. in carrot, Daucus carota) and the boundary
between the cap and the rest of the root is sometimes unstable55
(Figure 3.2).
42 Root
Figure 3.1 Diagram of root apical organization in Zea mays (monocot:

Poaceae)
The root apex is structured into zones of cells of contrasting size

and differential activity. The quiescent centre is a group of relatively
inactive cells that divide infrequently; their role is to maintain initial
cells in an undifferentiated state. Probably the region of greatest
mitotic activity is the promeristem surrounding the quiescent centre,
though actively dividing cells extend from the apex into older parts of
the root. In some species (e.g. Vicia faba) there is an undifferentiated
common initiating region for all root tissues131, but in others (e.g.
Arabidopsis thaliana), the initial cells lie in clearly defined regions. In root
apices with closed organization, the root epidermis is associated
either with cortical cells (in most monocots) or with cap initials (in
most eudicots); in root apices with open organization the precise
origin of the root epidermis is relatively difficult to determine19.
3.2 Primary root structure 43
Figure 3.2 Longitudinal sections of root apices. Left: Vicia faba (eudicot:
Fabaceae), open organization. Right: Zea mays (monocot: Poaceae), closed
organization. Scales = 100 µm
The root cap consists of several layers of parenchymatous cells.

In maize (Zea mays), a species with “closed” root apical structure
(Figures 3.1, 3.2), the cap initials become established and inde-
pendent from the apical meristem at an early stage in seedling
development7. The cap meristematic cells, located adjacent to the
quiescent centre, produce derivatives that are eventually displaced
towards the outside of the root cap and are subsequently worn
away, in the process contributing to the external slime that helps
the root push between soil particles. Cells are generated in the root
cap at approximately the same rate as they are discarded.
3.2 Primary root structure

Primary structure retains a characteristic ground plan that differs
from that of the stem. Each root possesses clearly defined radial
organization with concentric tissue regions. The outermost dermal
tissue (epidermis, rhizodermis or velamen) develops specialized
44 Root
Figure 3.3 Root anatomy. Ranunculus acris (eudicot: Ranunculaceae),

transverse section of root, with detail of central vascular region (inset).
Scales = 100 µm and (inset) 20 µm
water-absorbing root hairs in well-defined regions. Inside the der-

mal layer lies the cortex, which includes a specialized innermost
cortical layer, the endodermis. Finally, the central cylinder consists
of a pericycle surrounding the central vascular tissue (Figure 3.3).
3.3 Root epidermis and root hairs

The root epidermis (rhizodermis) is typically uniseriate, though in
some species, especially in aerial roots of aroids and orchids, it
becomes a multiseriate tissue, termed a velamen, which has a role
in water storage. Velamen cells lack a protoplast and can become
saturated with water; their cell walls are often partly thickened and
sometimes lignified.
3.4 Root cortex and endodermis 45
In most angiosperm species, root hairs are located about

a centimetre from the apex, in a region where cells have ceased
division but are still enlarging. This region of the root is the most
active in absorption of water and solutes; the root hairs provide
a much greater surface area for this purpose. Root hairs are out-
growths of epidermal cells that form by apical intrusive growth
between adjacent soil particles. The root hairs persist for a limited
period before they wither, and new hairs are formed closer to the root
apex. Root hairs are mostly unicellular and unbranched, but they can
sometimes branch. Long root hairs characterize species that do not
form ectomycorrhizal associations (e.g. sedges). In some species (e.g.
rice, Oryza), any rhizodermal cell is capable of root hair formation, but
in others (e.g. lilioid monocots and waterlilies), root hairs are pro-
duced only by specialized root epidermal cells (trichoblasts) in an
alternating pattern within each cell file. In other species (e.g. Arabidopsis
thaliana), root hairs are formed only in alternate axial cell files25.
3.4 Root cortex and endodermis

The cortex, located between the pericycle and the epidermis, con-
sists primarily of ground tissue formed by sequential periclinal cell
divisions. The number of cell layers in the cortex ranges from one or
two in some species (e.g. Arabidopsis) to ten or more in others. The
final, innermost cortical layer becomes specialized as the endoder-
mis. Some plants possess a dimorphic hypodermal layer adjacent to
the epidermis, consisting of alternating long and short cells73, 127.
In older roots, the epidermis often becomes worn away and its
protective role is taken over by the outermost cortical cells, which
can become suberinized or lignified to form an exodermis, espe-
cially in monocots. In woody eudicot species with lateral thickening
growth, a periderm often forms in the outer cortex.
Apart from these specialized layers, most cortical cells are par-
enchymatous. Cortical cells located in the root hair region facilitate
radial passage of solutes from the epidermis to the vascular tissue;
46 Root
Figure 3.4 Lateral roots. Vicia faba (eudicot: Fabaceae), transverse section
of root with three developing lateral roots. Scale = 100 µm
in other parts of the root they often have a storage role. In some
plants (e.g. root crops such as carrot), the tap root is a modified
swollen storage organ with a broad cortex. Contractile roots occur
periodically in many plants with modified underground stems such
as corms, bulbs or rhizomes (e.g. Crocus); these specialized roots
grow downwards, then contract axially and expand radially to draw
the stem deeper into the soil67. Contractile roots have a wrinkled
surface; they possess two or three clearly distinct concentric regions
of cortical parenchyma, distinguishable by cell size, including
a region of collapsed outer cortical cells interspersed with occa-
sional thicker-walled cells (Figure 3.4). In some species, centripetal
collapse of outer cortical cells results from loss in turgidity follow-
ing transpiration, which produces a differential between atmo-
spheric pressure and relatively low xylem pressure.
The root endodermis is a uniseriate cylinder of innermost cortical
cells surrounding the central vascular region, adjacent to the pericycle.
3.5 Pericycle and vascular cylinder 47
Endodermal cells are characterized by deposition of a band of suberin

or lignin in their primary walls, termed a Casparian strip, which
represents a barrier against non-selective passage of water through
the endodermis. Occasional endodermal cells (passage cells) remain
thin-walled, allowing selective passage of water between the cortex
and vascular region. In older endodermal cells, the inner periclinal
waIl becomes thickened and lamellated so that the Casparian strip is
obscured. The secondary wall is often lignified and/or suberized, and
acts as an effective barrier to water loss.
3.5 Pericycle and vascular cylinder

The central vascular cylinder is surrounded by a layer of thin-walled
cells, the pericycle, which is mostly uniseriate, especially opposite
the phloem poles (Figure 3.3). Both the pericycle and vascular
tissue are derived from cells located on the proximal (shoot) side
of the quiescent centre at the root apex. The pericycle is the site of
lateral root initiation; it retains its meristematic potential for some
distance away from the apex.
Primary vascular tissue in the root consists of several strands of
phloem alternating with radially arranged rays (archs) of xylem. The
xylem region often appears star-shaped in transverse section; it
extends to the centre of smaller roots, which lack a central parench-
ymatous region. Roots possess two, three, four or more protoxylem
poles, in which case they are diarch, triarch, tetrarch or polyarch
respectively. There is often variation in the number of xylem poles,
sometimes even within the same plant, depending on the diameter of
the root. Eudicot roots possess a relatively small number of xylem
poles (most commonly two, three or four) compared with many
monocots. In monocots with polyarch roots (e.g. Iris), the centre of
the root is parenchymatous, often becoming lignified in older roots.
During root development, protoxylem elements are the first to be
formed and the narrowest in diameter; they are located at the tips of
the archs, nearest to the pericycle. The larger metaxylem elements
48 Root
are located along the archs, towards the root centre. Both xylem and
phloem are exarch in the root (i.e. they mature centripetally).
3.6 Lateral and adventitious roots

Lateral roots are branches of the primary root, whereas adventitious
roots are initiated in other parts of the plant, most commonly in
stem tissue (Figures 3.4, 3.5). Both lateral roots and adventitious
roots are common in eudicots, but monocots and waterlilies rely
mostly on adventitious roots, and the primary root is ephemeral.
Lateral roots are initiated some distance from the root apex, often
in acropetal sequence, so that the most recently formed lateral roots
are those closest to the root apex. They have a deep-seated (endo-
genous) origin in the parent root (Figure 3.4). They are initiated in
groups of founder cells in the pericycle, following an auxin-drive
stimulus100. In many eudicots, lateral root initiation in the pericycle
occurs at a point adjacent to a protoxylem pole, though in diarch
Figure 3.5 Adventitious roots. Ligustrum vulgare (eudicot: Oleaceae), transverse

section of stem with two developing adventitious roots. Scale = 100 µm
3.7 Secondary growth in roots 49
roots (as in Arabidopsis) lateral roots are initiated opposite the xylem
poles. In polyarch roots of monocots, the site of lateral root initia-
tion is difficult to determine; it can be opposite either protoxylem or
phloem poles88. The founder cells undergo a series of periclinal and
anticlinal divisions to form a lateral root primordium. The first
anticlinal divisions are asymmetric, resulting in daughter cells of
unequal sizes. In many species, some subsequent cell divisions
occur in the endodermis, so that ultimately both the pericycle and
the endodermis contribute to the tissues of the lateral root. The
emerging lateral root pushes its way through the cortex and epi-
dermis of the parent root by mechanical and/or enzymatic action.
Adventitious roots have various sites of origin within a stem; they
are typically endogenous (Figure 3.5), but more rarely are exogen-
ous, arising from superficial tissues such as the epidermis (e.g. in
surface-rooting Begonia leaves) or from callus tissue at the site of
a wound. In woody species, adventitious roots are often formed at
nodes on the stem, which is why horticultural cuttings are commonly
taken from just below a node. In most monocots, adventitious roots
arise from cell divisions in the pericyclic region of the stem.
3.7 Secondary growth in roots

In woody eudicots and magnoliids, thickening and strengthening of
the root system below ground is important in supporting the
enlarging stem above ground. Most eudicot roots possess at least
minimal secondary thickening (Figure 3.6), with the exception of
some herbaceous species such as Ranunculus (Figure 3.3). In contrast,
secondary (lateral) growth in roots is extremely rare in monocots,
even among arborescent or woody species that possess a secondary
thickening meristem; the only reported exception is Dracaena, in
which a limited region of secondary tissue is formed in the root141.
In the roots of most woody eudicots and magnoliids, secondary
vascular tissue (both xylem and phloem) is produced by
a bidirectional vascular cambium, as in the stem. This tissue is
50 Root
Figure 3.6 Secondary thickening in roots. Ulmus sp. (eudicot: Ulmaceae),

transverse sections of roots with (top) early secondary growth and (bottom)
more extensive secondary thickening. Scales = 200 µm
initiated in the region located between the primary xylem and

phloem of younger roots, and subsequently in derivatives of cell
divisions within the pericycle, close to the xylem poles. As a result
of this sequence of cambial activity, the xylem cylinder soon
appears circular in transverse section and the outer layers of cortex
and epidermis begin to split apart. Further pericyclic cell divisions
result in the formation of a secondary cortex. In many cases
a periderm forms in the outermost layers, particularly where
secondary growth is extensive (Figure 3.6). Root secondary
xylem can differ in some respects from stem secondary xylem in
3.8 Roots associated with microorganisms 51
the same plant. For example, in Quercus robur (oak), stem wood is
ring porous, with earlywood vessels markedly larger than late-
wood vessels, but root wood is diffuse porous, with vessels that
maintain a relatively even size across each growth ring.
3.8 Roots associated with microorganisms

Many vascular plants form symbiotic relationships with soil
microorganisms that invade the cells of the host root and promote
the uptake of mineral nutrients to the plant, especially phosphate
and nitrogen, in exchange for carbon. Fungal symbioses occur in
a high proportion of land plants89. In some temperate woody
forest species, especially in the families Fagaceae and Betulaceae,
ectomycorrhizal fungi form a mantle that entirely envelopes the
lateral roots. Ectomycorrhizal fungal hyphae do not enter the cell
lumen but grow between the host epidermal and cortical cells,
forming a mantle termed a Hartig net. In contrast, arbuscular
Figure 3.7 Mycorrhizal roots. Neottia nidus-avis (monocot: Orchidaceae),

transverse section of root with fungal hyphae in outer cortex. Scale = 100 µm
52 Root
mycorrhizae invade the host cortical cells and form intracellular

structures (arbuscules) (Figure 3.7).
Some soil microorganisms can induce the formation of modified
lateral roots. In some eudicots, including many legume species,
symbiotic nitrogen-fixing bacteria (rhizobia) enter the root cortex
through root hairs and stimulate meristematic activity and cell pro-
liferation in the cortex and pericycle to form a specialized structure
termed a root nodule22, 87. Within the nodule, the rhizobia invade
host cells, where they are surrounded by a plasma membrane and
resemble organelles that reduce nitrogen to ammonia. Legume root
nodules can be either determinate and spherical (e.g. in soybean
plants) or indeterminate and more elongated (e.g. in pea plants).
Indeterminate nodules develop an apical meristem and peripheral
vascular bundles connected to the central root vasculature.
3.9 Root haustoria of parasitic angiosperms

A few angiosperms are parasitic on the roots, stems and leaves of
other species, including members of the families Convolvulaceae
(dodders, e.g. Cuscuta), Loranthaceae (mistletoes, e.g. Viscum),
Orobanchaceae (e.g. broomrapes, Orobanche, and toothworts,
Lathraea) and Santalaceae (sandalwoods, Santalum). Parasitic plants
produce specialized root haustoria that penetrate the host tissue to
transfer nutrients from the host to the parasite. Some hemiparasites
such as Rhinanthus and Bartsia are similarly haustorial. Haustoria are
formed either directly from the root apex of the parasite or from
a modified lateral or adventitious root or stem63.
In many parasitic species, the root haustorium penetrates the
host tissue to the xylem and phloem and forms a continuous
vascular bridge (Figure 3.8). Haustorial epidermal cells in contact
with the host become elongated and secretory, and the centre of
the haustorium develops an intrusive process that grows into the
host tissue by enzymatic and mechanical action. The developing
parasitic haustorium often forms a mantle of parenchymatous
3.9 Root haustoria of parasitic angiosperms 53
Figure 3.8 Parasite haustoria. Cuscuta epithymum, dodder (eudicot:

Convolulaceae) parasitic on host stem of Trifolium, clover (eudicot:
Fabaceae), with several haustoria penetrating host vascular tissue.
Scale = 200 µm
tissue around the host organ. In some parasitic plants of mistletoe

family (Loranthaceae), the haustorial mantle around the host
tissues influences the host to develop a woodrose, which is
a woody outgrowth of vascular tissue that forms a conduit for
nutrients to the parasite.
4
Leaf
Leaves are determinate lateral organs that are usually dorsi-

ventrally flattened and lack a growing tip. Foliage leaves are
green; they are typically borne on stems above ground level
because they require sunlight for photosynthesis.
Angiosperm leaves consist of a sheathing leaf base that clasps
the stem at the node and a distal zone that extends away
from the stem to capture light effectively20, 54, 72, 114, 120. In
eudicots and magnoliids, the sheathing lower zone is often
reduced or sometimes absent and the distal zone forms the
bulk of the leaf, consisting of a petiole and an elliptical blade
(lamina) with net-like (reticulate) venation (Figure 4.1). The
margins of the lamina can be smooth, lobed or toothed.
Some species possess compound leaves with individual leaf-
lets either borne on a central stem-like axis (pinnate leaves;
e.g. tomato: Solanum lycopersicum) or radiating from a single
point at the distal end of the petiole (palmate leaves; e.g.
aroids such as Arisaema). In monocot leaves, a sheathing leaf
base is typically well developed and the upper zone can be
thick or radial in outline, or even reduced to a short, thick-
ened apex. Some underground stems bear reduced scale-like
leaves. Underground bulbs possess swollen leaves or leaf
bases that contain storage products such as starch to allow
the plant to survive below ground during harsh conditions.
4.2 Leaf initiation and development 55
Figure 4.1 Leaf venation. Hypenia pauliana (eudicot: Lamiaceae), cleared leaf
with reticulate venation. Scale = 2 mm.
4.1 Leaf anatomy

The mature lamina consists of an adaxial and abaxial epidermis enclos-
ing several layers of mesophyll cells interspersed with a network of
vascular bundles (Figure 4.2). The majority of species have bifacial
(dorsiventral) leaves, in which the adaxial and abaxial surfaces differ
and the mesophyll is differentiated into palisade and spongy regions.
In unifacial (isobilateral) leaves, the epidermis is similar on both
surfaces and the mesophyll is relatively undifferentiated. In monocots
with unifacial leaves, the leaf base is bifacial and the lamina is unifacial
and either flattened (e.g. in Acorus and most Iridaceae) or rounded in
transverse section (e.g. terete leaves of some Allium species).
4.2 Leaf initiation and development

Leaves are initiated from groups of founder cells located close to
the shoot apex, either in the outermost cell layers (L1 and L2) or in
56 Leaf
Figure 4.2 Leaf anatomy. Ficus cordata (eudicot: Moraceae), transverse section
of leaf blade. Scale = 50 µm
the layers immediately below them (L3) (Section 2.1). Founder

cells undergo periclinal divisions to form small conical projections
that represent the leaf primordia. In monocots, a leaf primordium
rapidly develops into a bifacial hood-like structure, its base par-
tially or wholly encircling the stem to form a leaf sheath71, 114, 160.
In a simple leaf, subsequent meristematic activity occurs in an
adaxial transition zone between the precursor tip and sheath. In
some monocots such as grasses, epidermal outgrowths (ligules)
develop from an abaxial cross zone in this region.
The adaxial marginal cells divide rapidly to form a flattened leaf
blade, though marginal growth is suppressed in the region that
later becomes the petiole. Marginal growth can occur at the same
time as apical growth and is subsequently replaced by cell divisions
across the leaf blade; by this developmental stage, the number of
cell layers is established and the entire lamina functions as a plate
4.3 Leaf epidermis 57
meristem. Subsequent cell divisions are mainly anticlinal, resulting

in regular layers of cells that are disrupted only by the differentia-
tion and maturation of the vascular bundles.
Different rates of growth and cell division can occur in different
parts of the leaf, resulting in variations in leaf shape. Individual
leaflets of a compound leaf can be produced either acropetally or
basipetally. For example, in tomato (Solanum lycopersicum), the
uppermost (distal) leaflets are initiated first, followed by middle
and lower leaflets in a basipetal sequence65. Smaller intermediate
leaflets are formed later, in a more chaotic sequence.
In some monocots, meristematic activity governing leaf elonga-
tion is restricted to a basal rib meristem, which results in axial files
of cells of increasing maturity towards the distal end of the leaf.
The unifacial leaves of some monocots (e.g. Acorus), which possess
a bifacial sheathing leaf base and a unifacial upper blade, result
from suppressed marginal growth and the presence of an adaxial
(ventral) meristem in the transition zone72.
4.3 Leaf epidermis

The leaf epidermis is generally uniseriate, though in a few eudicot
and magnoliid species (e.g. Ficus and Peperomia) it can proliferate
into several cell layers to form a multiple epidermis, which resem-
ble a hypodermis. Specialized elements of the leaf epidermis
include stomata, trichomes, papillae, surface sculpturing, epicuti-
cular wax and variously differentiated pavement epidermal cells.
In surface view, pavement epidermal cells range in shape from
elongated to isodiametric. Cells over veins are often elongated,
especially in eudicots with reticulate venation, in which the costal
regions (over veins) develop before the intervening (intercostal)
regions. In the linear leaves of many monocots, the epidermal cells
are elongated along the long axis of the leaf. Anticlinal cell walls can
be straight or undulating; they are often more sinuous on the abaxial
than the adaxial surface of the same leaf (Figure 4.3). Epidermal cells
can also differ in size and wall thickness in different parts of the same
58 Leaf
Figure 4.3 Leaf surface. Vicia faba (eudicot: Fabaceae), abaxial leaf surface
with undulating anticlinal walls. Scale = 20 µm
leaf. In some grasses, restricted regions of the leaf epidermis contain

enlarged bulliform cells that play a role in the unrolling of the leaf in
response to turgor pressure and water availability.
Specialized epidermal cells can contain crystals or silica bodies
(Section 1.4). For example, cystoliths are calcareous bodies that
occur either solely in epidermal cells (e.g. in the family Opiliaceae)
or across both the epidermis and underlying mesophyll (e.g. in
Ficus: Figure 4.4), occasionally protruding above the leaf surface.
Discrete bodies of silicon dioxide (silica phytoliths) occur in
specialized epidermal cells in some monocot families (e.g. sedges
and grasses). The grass epidermis is characterized by dimorphic
long and short cells resulting from asymmetric divisions (Figure
4.5); each short cell often divides symmetrically to form a short-
cell pair that includes a silica cell (Figure 1.5) and a cork cell118.
Stomata (Section 1.9) occur on almost all angiosperm leaves,
except the submerged leaves of some aquatic plants, which lack
stomata entirely (e.g. Ceratophyllum and Podostemaceae). In species
with amphistomatic leaves, stomata are present on both leaf sur-
faces, whereas in species with hypostomatic leaves they are
4.4 Cuticle and wax 59
Figure 4.4 Ficus elastica (eudicot: Moraceae), transverse section of leaf blade
showing adaxial epidermis and cystolith. Scale = 50 µm
restricted to the abaxial surface. The floating leaves of many aquatic
plants are epistomatic, stomata being restricted to the adaxial surface.
4.4 Cuticle and wax

In mesomorphic leaves, the cuticle is thin and almost transparent,
but many xeromorphic plants possess a thick leaf cuticle that often
appears lamellated in transverse section. In some species, the outer
surface of the leaf cuticle possesses characteristic patterns of ridges,
folds or striations. These striations can be short or long and
oriented randomly or in a regular pattern, sometimes radiating
around stomata or trichomes (Figure 4.6). Surface patterning has
biological significance in relation to mechanical and optical prop-
erties and wettability of the surface.
Plant leaves have a covering of epicuticular wax over the surface
of the cuticle, ranging from a thin, almost imperceptible film to
crystalloid particles or a dense surface crust that gives the leaf
surface a whitish appearance9 (Figure 4.7). Some species of
Euphorbia have wax chimneys surrounding the stomatal pore,
60 Leaf
Figure 4.5 Grass epidermis. Coix lacryma-jobi (monocot: Poaceae), abaxial

leaf surface showing short-cell pairs (scp), stomata with guard cells (gc)
and adjacent lateral subsidiary cells (lsc). Scale = 20 µm
derived from an extended crust. Individual epicuticular wax par-
ticles display considerable morphological diversity that can be
characteristic of the species, including irregular granules, straight
or coiled rodlets, hollow tubules and flat platelets. These charac-
teristic structures are determined by their chemistry; for example,
coiled rodlets are predominantly composed of β-diketones.
Epicuticular wax particles can be variously arranged in aggregates,
rows or rosettes and are often oriented at a specific angle to the
surface. Like the cuticle, epicuticular wax is hydrophobic and
protects the plant surface from water loss, especially in species
that occupy dry habitats.
4.5 Trichomes and papillae 61
Figure 4.6 Surface patterning. Schisandra rubriflora (ANA-grade:

Schisandraceae), abaxial leaf surface showing cuticular striations radiating
around stomata (SEM). Scale = 30 µm
4.5 Trichomes and papillae

The presence, distribution and detailed structure of the leaf trichome
complement (indumentum) can be highly characteristic of a species
(Figures 1.12, 1.13). Trichomes are sparse or absent in glabrous
leaves. Trichomes can occur over the entire leaf surface, or they can
be restricted to particular regions, such as abaxial grooves (Figure
4.8) or leaf margins. In many species, several different types of
trichome occur on the same leaf. For example, mints (Lamiaceae)
characteristically possess two or more sizes of glandular trichome and
non-glandular trichomes that are either branched or unbranched.
Specialized leaf hair types include stinging hairs, sensory hairs,
water-absorptive leaf scales and salt-secreting glands that accumu-
late excess sodium chloride, allowing plants to colonize highly
saline soils (e.g. in Avicennia, Limonium and Tamarix). Stinging hairs
characterize five families of eudicots91 and are epitomized by the
common stinging nettle (Urtica dioica). They are hollow structures
that are rigid (silicified or calcified) and taper to a spherical tip
(Figure 1.13). The tip is readily broken off in contact with an
62 Leaf
Figure 4.7 Surface waxes. Yucca queretaroensis (monocot: Asparagaceae), leaf

surface showing epicuticular waxes around stomata (SEM). Scale = 50 µm
intruder, enabling the residual sharp point to penetrate the skin

and release a toxic fluid, thus forming an effective deterrent.
Sensory trigger hairs occur on leaves of some carnivorous plants,
such as the Venus flytrap (Dionaea muscipula), which traps insects in
a highly specialized bilobed leaf structure. A mechanical stimulus
such as the touch of an unwary insect on two or more of the six long
trichomes generates an electrical signal; a second stimulus rapidly
activates the trap to capture the insect150. Trichomes of other
carnivorous plants (e.g. Drosera) secrete both sticky mucilage and
proteolytic enzymes that help to capture and digest the prey.
4.6 Extrafloral nectaries, domatia and hydathodes

Localized regions of the leaf epidermis can be specialized for
particular roles, including extrafloral nectaries and water
pores156. Some plants exude water from their leaves during
humid conditions when there is sufficient soil moisture; this
guttation process accelerates passage of nutrients through the
plant when transpiration is low (e.g. at night). Guttation is achieved
4.6 Extrafloral nectaries, domatia and hydathodes 63
Figure 4.8 Erica carnea (eudicot: Ericaceae), transverse section of leaf showing
abaxial groove containing trichomes and stomata. Scale = 50 µm
via specialized leaf epidermal regions termed hydathodes, which are

water glands located most commonly at the leaf margins, especially
at the leaf teeth. Hydathodes typically consist of a region of modified
stomata located over a region of loosely arranged mesophyll cells
(epithem) associated with a vein ending. They can also be involved
in absorption of water from rain and dew.
Some plants provide food rewards that attract insects, especially
ants, which can protect the plant against potential herbivores.
Food bodies or pearl glands are scattered isolated globular tri-
chomes that occur on leaves of some tropical eudicots and mag-
noliids. They are rich in carbohydrates, lipids and proteins and are
harvested by ants95. Extrafloral nectaries are specialized epidermal
regions that secrete sugary nectar, which is attractive to small
invertebrates such as ants and mites (Figure 4.9). These localized
structures consist either of groups of glandular trichomes (e.g. in
some Hibiscus species) or raised regions of anticlinally elongated
secretory epidermal cells, often associated with underlying vascu-
lar tissue, in which case they resemble hydathodes. Extrafloral
nectaries can be located over veins, in the angles between primary
and secondary veins, on inrolled leaf margins or at the proximal or
distal ends of the petiole, sometimes situated in specialized pits or
64 Leaf
Figure 4.9 Omphalea triandra (eudicot: Euphorbiaceae), extrafloral nectary on

abaxial leaf surface. Scale = 20 µm
hair-lined pockets (domatia) that provide accommodation for

invertebrates. Increased herbivory can stimulate the plant to pro-
duce more extrafloral nectaries, and in some cases, the insects
themselves can stimulate production.
4.7 Mesophyll
Chlorophyll is contained in chloroplasts in the mesophyll, which is
the primary photosynthetic tissue of the leaf. In many plant species the
mesophyll is demarcated into distinct regions, termed palisade and
spongy tissues (Figures 4.2, 4.4), though in some species the meso-
phyll is relatively undifferentiated and homogeneous throughout the
leaf. Palisade mesophyll is located on the adaxial side of the leaf and
spongy mesophyll on the abaxial side. Palisade cells are anticlinally
elongated and possess relatively few intercellular air spaces; they often
occur in a single layer, but can be arranged in two or more layers. In
contrast, spongy mesophyll consists of several layers of cells of various
shapes interspersed with numerous air spaces.
In thick leaves, the central mesophyll cells are often large,
undifferentiated and non-photosynthetic. In the thick, fleshy
4.7 Mesophyll 65
Figure 4.10 Succulent leaves. Aloe somaliensis (monocot: Asphodelaceae),

transverse section of leaf margin showing outer chlorenchyma, inner
hydrenchyma, and two rows of small vascular bundles that lack
sclerenchyma. Scale = 20 µm
leaves of succulent plants such as many Aloe species (Figure 4.10),

the inner leaf mesophyll cells have relatively elastic wall properties,
enabling water storage in a specialized hydrenchyma tissue that can
be utilized during periods of drought1. In the thick “keel” or
“midrib” of Crocus leaves there is a region of large parenchymatous
cells with thin walls, which often break down to form a cavity
(Figure 4.11); this effect causes the characteristic white stripe visible
in the midrib region of Crocus leaves.
In some plants (e.g. Ficus, Figure 4.4) subdermal mesophyll layers
located immediately within the adaxial (or more rarely the abaxial)
epidermis are modified into a hypodermis. Hypodermal tissue con-
sists of one or more layers of non-photosynthetic cells that are usually
slightly larger and thicker walled than adjacent mesophyll cells and
frequently resemble epidermal pavement cells in transverse section.
Mesophyll is often interspersed with tissues that provide mechan-
ical strengthening; for example, sclerenchyma is frequently present
66 Leaf
Figure 4.11 Crocus cancellatus (monocot: Iridaceae), transverse section of leaf.

Scale = 100 µm
at leaf margins and extending as girders from the vascular bundles to

the epidermis. Fibres are often present as vascular bundle caps,
especially associated with the phloem poles.
Various types of idioblast are often present in mesophyll, includ-
ing crystal idioblasts and non-articulated laticifers, which permeate
all parts of the plant in some eudicots such as Euphorbia and Ficus
(chapter 1.12). Sclereids occur in the leaf mesophyll in many species;
for example, star-shaped astrosclereids occur in the lamina of
Nymphaea and the petiole of Camellia (Figure 1.9) and bone-shaped
osteosclereids are present in the palisade tissue of some species with
centric leaves. In some eudicot species (e.g. some Melastomataceae),
sclereids are associated with veinlet endings; some terminal sclereids
(e.g. in Mouriria) resemble swollen tracheids with spirally thickened or
pitted walls and are widely interpreted as storage tracheids43.
4.8 Leaf vasculature

Many eudicots and magnoliids are characterized by densely reti-
culate leaf vein architecture (venation) that enables greater tran-
spiration efficiency15, 20, 60, 123. Leaves with reticulate, net-like
4.8 Leaf vasculature 67
venation possess a primary vein (midrib) extending from the base to

the tip (Figure 4.1). The midrib is linked to smaller secondary veins
that radiate outward. They either extend to the leaf margins, where
they can terminate at a leaf tooth, or loop round to join the super-
adjacent secondary vein. In their turn, smaller veins branch from
the second-order and subsequent-order venation, thereby forming
a complex network. The regions circumscribed by the smallest veins
in the leaf are termed areoles. In many species, small veins branch
into the areoles to form veinlet endings; the relative number of
veinlet endings per areole represents a variable aspect of leaf venation.
In contrast, venation is parallel in many monocots with narrow,
linear leaves. In these species, the primary veins run parallel to each
other for most of their length and merge or fuse at the leaf tip.
There are typically numerous small veins interconnecting the
larger veins, but relatively few vein endings in the mesophyll.
Petioles also possess characteristic venation patterns. The simplest
form of petiole vasculature appears in transverse section as a vascular
crescent, with xylem on the adaxial side and phloem on the abaxial
side (Figure 4.12). In some species, the vascular crescent is inrolled
at the ends; in others it forms a ring or is divided into separate
bundles. Some species possess additional bundles outside the main
vascular crescent. In transverse sections of the lamina, vascular
bundles are typically arranged in a single row (Figure 4.2), though
two or more rows of leaf vascular bundles are present in species
with very thick, succulent leaves, such as Agave and Aloe (Figure
4.10). Lamina bundles are most commonly collateral, with adaxial
xylem and abaxial phloem, but orientation can vary, so that some
bundles are bicollateral or even amphivasal (with xylem surrounded
by phloem). The isobilateral leaves of some monocots possess two
rows of vascular bundles with opposite orientation to each other;
the xylem poles towards the leaf centre. Centric leaves possess a ring
of vascular bundles.
Leaf vasculature develops acropetally from a primordial procam-
bial strand20, 79. The central trace develops first and ultimately
68 Leaf
Figure 4.12 Petiole anatomy. Prunus lusitanica (eudicot: Rosaceae), transverse

section of petiole. Scale = 100 µm
becomes the primary vein (midrib). The timing of initiation of

lower-order venation differs among different species, depending
on ultimate leaf shape and venation pattern. Both xylem and phloem
tissues are present in the leaf blade, even in minor veins, though the
smallest vascular bundles often have only one or two rows of xylem
tracheids and a few files of phloem sieve-tube elements. The xylem
conducting system can consist entirely of tracheids with helical or
annular thickenings, though in some leaves both vessel elements
and xylem parenchyma are also present.
4.9 Bundle sheath and Kranz anatomy

Most minor vascular bundles in angiosperm leaves are surrounded
by a bundle sheath, which extends even to the very smallest veins.
The bundle sheath usually consists of thin-walled parenchymatous
4.10 Ecological adaptations in leaves 69
cells, often arranged in a single layer. Some monocots possess distinct

inner and outer bundle sheaths, of which the outer sheath is parench-
ymatous and the (partial) inner sheath is sclerenchymatous, forming
a sclerenchyma cap that is usually located at the phloem pole. Grasses
possess either a single sheath consisting of an outer layer of thin-
walled cells containing chloroplasts or a double sheath consisting of
an outer layer of thin-walled cells and an inner layer of thicker-walled
cells. This distinction can represent an important distinguishing fea-
ture between different grass groups; for example, double sheaths
often occur in festucoid grasses and single sheaths in panicoid grasses.
Leaves of many plants possess regions of sclerenchyma or parenchyma
that extend from the vascular bundle sheaths towards either or both
epidermises. These bundle sheath extensions, termed girders if they
reach the epidermis, can afford mechanical support to the leaf.
Some plants that undergo the C4 pathway of photosynthesis,
including both monocots (e.g. some grasses) and eudicots (e.g.
some Chenopodiaceae), display modified leaf anatomy termed
Kranz anatomy (Section 1.3)31, 124. Kranz anatomy is character-
ized by elongated mesophyll cells radiating from an outer bundle
sheath consisting of a single layer of large parenchymatous cells
that contain starch and enlarged chloroplasts; this specialized
sheath itself surrounds an inner sheath (Figure 4.13). Carbon
assimilation occurs in the radiating mesophyll cells, where the
enzyme phosphoenolpyruvate carboxylase (PEP carboxylase) is
localized, followed by carbon reduction in the outer bundle
sheath, where the carboxylating enzyme Rubisco is localized.
4.10 Ecological adaptations in leaves

Features that are often associated with water plants (hydrophytes)
include the absence of sclerenchyma and the presence of large regular
air spaces in the ground tissue (aerenchyma). Submerged leaves
typically lack stomata entirely. In floating leaves, stomata are restricted
to the upper (adaxial) surface and water-absorptive hairs (hydro-
potes) are often present on the submerged surface (Section 1.10).
70 Leaf
Figure 4.13 Kranz anatomy. Eleusine coracana (monocot: Poaceae), transverse

section of leaf. Scale = 50 µm
In contrast, plants that grow in dry, seasonally dry or otherwise

nutrient-deficient habitats (collectively termed xerophytes) can
possess a range of specialized features that minimize collapse
during drought periods; contrasting xeromorphic traits character-
ize different xerophytes. In some species, well-developed scler-
enchyma provides mechanical support. Some xeromorphic species
possess thick or succulent leaves (e.g. Aloe: Figure 4.10). Others
have terete (centric or cylindrical) leaves, or hairy leaves, or even
rolled or folded (plicate) leaves (Figure 4.14). Thick and terete
leaves possess a reduced surface/volume ratio to reduce water loss.
Leaf folding within a bud helps to minimize water loss in unex-
panded developing leaves; on subsequent expansion, the leaf can
achieve a large surface area. Succulent plants contain a high pro-
portion of large thin-walled cells that play a role in water storage,
often containing a high proportion of mucilage.
In many xeromorphic species, protected stomata restrict water
loss by transpiration. For example, individual stomata can be raised
4.10 Ecological adaptations in leaves 71
Figure 4.14 Ammophila arenaria (monocot: Poaceae), transverse section of

rolled leaf. Scale = 200 µm
or sunken with respect to the surface, or groups of stomata can be

confined to hair-lined grooves or depressions on the abaxial leaf
surface, thus trapping a pocket of water vapour in an air chamber
either externally or internally (Figure 4.8). A hypodermis or thick
epidermis and cuticle can reduce the intensity of light that reaches
the photosynthetic tissue. Well-developed palisade tissue in multi-
ple layers is sometimes correlated with high light intensity.
Most leaf stomata are associated with a substomatal cavity in the
underlying mesophyll; these cavities are also relatively high in water
vapour and represent important sites for exchange of solutes and
gases. During early development, the stomata are initiated over the
junction between underlying cortical or mesophyll cells. Epidermal
cells continue to enlarge after the mesophyll cells have ceased
expansion. Differential expansion between the guard-mother cell
and developing mesophyll cells influences several aspects of leaf
anatomy, including the substomatal cavity5, 49, 78.
5
Flower
Flowers are borne on reproductive axes, either as solitary struc-

tures or on inflorescences, which can be unbranched or variously
branched (Figure 5.1). In a determinate inflorescence, the inflor-
escence apex is terminated by a flower, whereas in an indetermi-
nate inflorescence it maintains growth until the apical meristem
becomes exhausted32, 152. Each flower is often subtended by one
or two modified leaf-like sterile bracts borne on the inflorescence
axis, though bracts are entirely absent from some species. Some
species also possess one or more leaf-like bracteoles on the flower
axis. At the onset of flowering, the shoot apical meristem under-
goes structural modification that transforms it from a vegetative
apex to a reproductive apex.
5.1 Flower structure

The floral axis (receptacle) is unbranched, determinate and highly
condensed. It bears several distinct organ regions in a consistent
radial sequence10, 32, 107, 152. The densely crowded floral organs
are arranged either in distinct whorls or in a spiral pattern on the
axis. Each organ series consists of one or more whorls. In a typical
flower bud, an outer region of sterile organs (either tepals, or
sepals and petals) encloses the pollen-bearing organs (stamens),
which in turn are outside the ovule-enclosing organs (carpels).
The outer sterile organs represent the perianth, and the stamens
and carpels form the androecium and gynoecium respectively. In
5.1 Flower structure 73
Figure 5.1 Inflorescence of Arabidopsis thaliana (eudicot: Brassicaceae) (SEM).

Scale = 500 µm
species with a syncarpous ovary, the gynoecium terminates the

short floral axis, though a residual floral axis exists in some species
with free carpels (Figure 5.2).
Different organ arrangements characterize different angiosperm
lineages. For example, some species possess unisexual flowers, in
which one of the reproductive organ types is either absent or
sterile: female flowers lack functional stamens and male flowers
lack carpels. In the grass family Poaceae, the flowers are highly
reduced structures (florets) that are grouped together in spikelets.
Each floret is subtended by a sterile leaf-like structure, the palea,
which surrounds two (occasionally three) reduced structures
termed lodicules; these are widely interpreted as representing
a single modified perianth whorl2, 75, 119.
The degree of fusion of individual floral organs within each
flower differs between species. Individual organs can be fused
together within the same whorl (connate) or fused with other organs
in adjacent whorls (adnate). A typical example of adnation occurs in
74 Flower
Figure 5.2 Nigella bucharica (eudicot: Ranunculaceae), scanning electron

micrograph of developing flower bud with sepals removed, showing
primordia of eight petals and multiple stamens in spiral arrangement and
a single whorl of five free carpels surrounding a residual apex.
Scale = 500 µm
flowers where the stamens are borne on the perianth. An example of

connate organs includes species in which the petals are fused
together (at least at their bases) to form a corolla tube or perianth
tube, as in many asterid eudicots. Similarly, in many angiosperms,
the carpels are fused into a single unified gynoecium (Figure 5.3).
5.2 Floral ontogeny

During early flower development, the lateral organ primordia are
generally initiated in centripetal (acropetal) sequence, so that the
latest primordia to be initiated are located close to the floral apex.
For example, in the magnoliid Drimys (Winteraceae), the
5.2 Floral ontogeny 75
Figure 5.3 Flagellaria indica (monocot: Flagellariaceae), longitudinal section

of flower. Scale = 500 µm
innermost stamens are the last to be initiated in the developing

flower bud, even though they ultimately grow larger than the
other stamens and are the first to dehisce in the open flower142.
Some species deviate from a strict centripetal progression, so that
some floral organs are initiated in a distal-to-proximal (centrifugal
or basipetal) sequence, either within or between organ zones113.
For example, in Lythrum salicaria, the outer sepal whorl is initiated
after the inner sepal whorl, the inner stamen whorl is initiated after
the gynoecium, and the petals are initiated after both the
76 Flower
gynoecium and the androecium. In some palm flowers that con-

tain large numbers of stamens, the inner stamens are initiated
before the outer ones144. These examples suggest that the floral
apex is pre-patterned into zones even before the organ primordia
become visible. Changes in auxin levels across the apex can result
in different sequences of organ initiation. In some species, groups
of floral organs are initiated in clusters (fascicles) on shared
primordia, which are themselves arranged in whorls; for example,
stamen fascicles are common in the eudicot families Malvaceae and
Hypericaceae (Figure 5.4).
Following initiation, floral organs expand both axially and later-
ally to assume their ultimate shape. In many flowers, organ expan-
sion continues after bud opening; for example, many petals
continue to expand in size and increase in colour after the flower
bud is open. Organ fusion is important in determining flower shape
and presentation to potential pollinators. In some species, floral
organs are initiated separately and become fused during develop-
ment, a process termed ‘postgenital fusion’. Postgenital carpel
fusion characterizes many monocots, in which the contacting
Figure 5.4 Hypericum empetrifolium (eudicot: Hypericaceae), SEM of

developing flower bud with outer organs removed, showing primordia of
petals, stamen fascicles (sf) and syncarpous gynoecium. Scale = 20 µm
5.3 Floral vasculature 77
epidermal cells of adjacent developing carpels become pressed

together and eventually lose their epidermal identity106. In contrast,
many floral organs are profoundly united from their inception,
a mode of development that is termed congenital fusion or synor-
ganization. An extreme example of congenital fusion occurs in the
reproductive organs of most orchids, which form an integrated
structure termed a gynostemium. Another example is the perianth
tube of many asterid eudicots, in which the petals are congenitally
fused at their bases.
5.3 Floral vasculature

In the majority of flowers, each organ is supplied by one or more
vascular strands that diverge from the central vascular cylinder of
the floral axis (Figure 5.5). Vascular traces often branch above
their level of divergence from the stem vasculature; perianth traces
are often highly branched, forming a vascular network that
Figure 5.5 Flagellaria indica (monocot: Flagellariaceae), transverse section of

central region of flower, showing a stamen with a single central vascular
bundle and the style with three vascular bundles surrounding a central
transmitting tissue. Scale = 50 µm
78 Flower
Figure 5.6 Transverse sections of styles. (left) Salvia pratensis (eudicot:

Lamiaceae), with two vascular bundles and a central pollen-tube-
transmitting tissue (pttt). (right) Cordyline cannifolia (monocot: Asparagaceae),
with three vascular bundles and three mucilage-lined channels.
Scales = 50 µm
develops as the organ expands laterally. Most stamens possess

a cylindrical filament that bears a single central vascular strand,
though some families are characterized by three or four stamen
traces and a few species possess diverse and often branching sta-
men vasculature. Many carpels are supplied by three vascular
bundles, of which the dorsal carpellary trace, which is typically
unbranched, passes along the style and supplies the stigma, and
two ventral carpellary traces, which are often highly branched,
supply the placenta and ovules6, 30, 143 (Figure 5.10). In many
species, the two ventral carpellary bundles are fused together
above the level of carpel closure. The number of dorsal carpellary
bundles in the style of a syncarpous gynoecium is often an indi-
cator of the constituent number of fused carpels; many tricarpel-
late monocots possess three stylar bundles whereas many
bicarpellate eudicots possess only two (Figure 5.6).
5.4 Perianth
Perianth organs are dorsiventrally flattened. They consist of an
abaxial and adaxial epidermis enclosing three or four (or more)
layers of undifferentiated isodiametric or elongated mesophyll
5.5 Androecium 79
cells, interspersed with a row of vascular bundles. In most eudi-

cots, the perianth is differentiated into two distinct series or
whorls: an outer sepal whorl (calyx) and an inner petal whorl
(corolla). In contrast, there is little clear morphological distinction
between the inner and outer perianth organs of many early diver-
gent angiosperms, monocots and some basal eudicots; such undif-
ferentiated perianth organs are termed tepals rather than petals and
sepals. Sepals, which envelop the unopened flower bud, are typi-
cally green and photosynthetic, with stomata and trichomes com-
monly present on their surfaces. Petals are typically large and
colourful in insect-pollinated plants but reduced or absent in
many wind-pollinated plants. Tepals often display a gradual tran-
sition from outer, green (i.e. sepaloid) organs to inner, coloured
(i.e. petaloid) organs33, 151.
Flower colour is primarily controlled by pigment chemistry.
Anthocyanins, betalains and ultraviolet-absorbing flavonoids are
largely confined to the epidermal cells of the petal, whereas
other pigments, such as carotenoids, mostly occur in the
mesophyll74. Petal surfaces frequently lack stomata and the
petal epidermal cells are often domed or papillate (Figure 5.7).
The effect of the domed cell surface is to guide incident light
into the petal, where it is reflected outwards from the inside
walls of the epidermal cells or from the multifaceted walls of
mesophyll cells, thus passing through the pigments inside the
cell. Many petal surfaces possess a strongly striated cuticle, which
further scatters incident light. In contrast, some petal surfaces are
smooth and mirror-like, strongly reflecting incident light and
resulting in a glossy effect (e.g. in buttercup and mirror
orchids74, 146, 147).
5.5 Androecium
Stamens typically consist of a stalk-like filament that supports
the pollen-producing anther24. Most stamen filaments are
80 Flower
Figure 5.7 Arabidopsis thaliana (eudicot: Brassicaceae), scanning electron

micrograph showing domed cells on petal surface. Scale = 5 µm
slender and cylindrical, but in some species they are flattened

and leaf-like (e.g. in Nymphaea odorata) or even branched (e.g. in
Ricinus communis). In some polyandrous angiosperms (bearing
multiple stamens), the stamens are borne in clusters, termed
fascicles, each fascicle developing from a single primordium on
the early floral apex (Figure 5.4). The filament surface can bear
trichomes and stomata and sometimes displays cuticular pattern-
ing. In transverse section, the filament contains parenchymatous
ground tissue surrounding a single vascular bundle that extends
into the connective tissue between the anther locules.
The anther bears four sporangia separated into two pairs
(thecae), linked together by a connective. Each theca possesses
two sporangia or anther locules divided by a septum24. The
anther wall consists of several cell layers: the outer epidermis
encloses the endothecium, which in turn encloses one or more
middle layers; the tapetum is the innermost layer, which sur-
rounds the anther locule (Figures 5.8 and 5.9). The endothecium
and tapetum are the most specialized layers; their enlarged cells
5.5 Androecium 81
Figure 5.8 Flagellaria indica (monocot: Flagellariaceae), transverse section of

anther containing mature pollen; by this stage the tapetum has largely
degenerated. Scale = 50 µm
often crush the intervening middle layers in older anthers.
Endothecial cells develop fibrous wall thickenings that contribute
to the anther dehiscence mechanism.
In some species, a layer of tapetal cells persists around the anther
locule; this type of tapetum is termed the secretory type. In contrast,
in species with a plasmodial tapetum, the tapetal cells undergo
programmed degeneration and their protoplasts fuse to form
a multinucleate tapetal plasmodium (a periplasmodium) inside
the anther locule. Transitional types of tapetum occur in some
species, especially among early divergent angiosperms45. Tapetal
cells are often multinucleate and contain dense cytoplasm. They can
produce a range of substances, including sporopollenin, which
forms a major component of the pollen wall (exine), and also
proteins and lipids that are retained in chambers in the exine and
allow sporophytic recognition when the pollen arrives on a stigma.
82 Flower
Figure 5.9 Lilium martagon (monocot: Liliaceae), transverse section of

anther wall containing developing microspores and tapetum. Scale =
50 µm
5.6 Gynoecium
Carpels are the ovule-bearing floral organs. The number of carpels
per flower ranges from a single carpel to multiple carpels in
different species. Monocots typically possess three carpels
(Figure 5.10), whereas the most common carpel numbers in
eudicots are two and five34. In the majority of flowering plants,
the carpels are fused together to form a syncarpous gynoecium,
which represents the central region of the flower. In contrast, the
carpels remain unfused (free) in apocarpous species, especially
among early divergent angiosperms (e.g. Amborella, Illicium), but
also in many eudicots, magnoliids and monocots34.
A syncarpous gynoecium consists of an ovary, style and one or
more stigmas (depending on the degree of fusion of the carpels).
The stigma provides a receptive surface for pollen grains. Stigmatic
epidermal cells are usually secretory; they are often domed or
possess variously elongated papillae and a specialized cuticle58, 59
(Figure 5.11). Stigmas of some species possess little or no surface
secretions (termed dry stigmas), whereas wet stigmas exude
copious surface secretions that play a role in pollen recognition
Figure 5.10 Asparagus officinale (monocot: Asparagaceae), transverse section of
central region of ovary, which consists of three postgenitally fused carpels.
Scale = 100 µm
Figure 5.11 Arabidopsis thaliana (eudicot: Brassicaceae), scanning electron

micrograph of stigma. Scale = 50 µm
84 Flower
and germination. The stigmatic cuticle is often layered, with

a lamellated outer layer and reticulate inner layers. Some Crocus species
possesses a chambered cuticle, and in some Euphorbia species the cuticle
is fenestrated.
Style anatomy can differ between species. Many syncarpous eudi-
cots possess solid styles, in which there is a central specialized
secretory tissue, termed transmitting tissue or compitum, which
serves as a nutrient-rich tract for pollen-tube growth from the stigma
to the ovary140 (Figure 5.6). Transmitting tissue is derived from the
superficial tissue of the fused carpel margins. In contrast, most
syncarpous monocots possess open styles with a central stylar canal
that is typically filled with mucilage.
The ovary is the ovule-bearing part of a syncarpous gynoecium. In
many syncarpous species, the transmitting tract (compitum) allows
the pollen tubes to reach any of the ovules located in the ovary locules.
Many species possess a proliferation of secretory tissue around the base
of the funicle of each ovule; this papillate tissue (obturator) provides
nutrients for the developing pollen tubes and helps to guide them into
the micropyles140. Ovules are borne on placentas, which are meriste-
matic regions located at the carpel margins within the ovary locules128.
Ovule primordia are initiated as small swellings on the placenta in the
developing ovary. Placental arrangement can differ between species. In
species with two or more ovary locules and axile placentation, the
ovules are borne on placentas located on the central axis, but in species
with unilocular ovaries, the placentas can be located on the ovary wall
(parietal placentation), at the base of the ovary (basal placentation) or
on a central column of tissue that is linked to the ovary wall only at the
top and bottom (free-central placentation).
5.7 Floral secretory structures

Many flowers bear specialized secretory structures, such as nec-
taries, elaiophores and osmophores, which secrete nectar, oil and
scent respectively. Their secretion products help to attract potential
5.7 Floral secretory structures 85
pollinators. These specialized secretory structures are often

formed after the primary floral organs have already been
initiated.
Nectaries are localized areas of tissue that regularly secrete
a sugary nectar. They consist of secretory epidermal cells with
dense cytoplasm, sometimes modified into trichomes.
Adjacent subepidermal cells can also be secretory; in some
species, nectar passes to the surface through modified stoma-
tal pores. The nectary is associated with vascular tissue that
consists predominantly of phloem, which transports sugars to
the secretory region. Nectaries can occur on any floral organ,
or they can form part of an entire modified organ, or even
a novel structure, such as a nectar spur35. Most species of the
mint family (Lamiaceae) possess an enlarged nectariferous
disc surrounding the base of the ovary, which is derived
from proliferated ovary tissue. In most Brassicaceae (e.g.
Arabidopsis thaliana) the nectary is located at the base of the
stamen filament. Septal nectaries are characteristic of many
monocots; these secretory regions occur at the unfused carpel
margins in monocots with postgenitally fused ovaries106
(Figure 5.10). Nectar produced from septal nectaries is
exuded from secretary epidermal cells and emerges from slit-
like openings on the surface of the gynoecium112.
Flowers of some insect-pollinated species lack nectaries but
possess other secretory structures that attract potential pollinators.
For example, some flowers bear oil-secreting glands, termed
elaiophores (Figure 5.12), which resemble nectaries in their mor-
phology. Osmophores are modified floral structures that produce
volatile scents; the scents released by some orchid flowers can
resemble insect pheromones149. They typically possess
a relatively thick, domed or papillate epidermis with densely
cytoplasmic contents. In Platanthera bifolia (the Lesser Butterfly
orchid) the epidermis of the labellum secretes a nocturnal scent,
and in several Ophrys species osmophores on the labellum consist of
86 Flower
Figure 5.12 Tigridia meleagris (monocot: Iridaceae), elaiophores on tepal

surface. (left) SEM, (right) transverse section. Scales = 10 µm
regions of dome-shaped, papillate, dark-staining epidermal cells.

Flowers of Narcissus emit pollinator-specific volatiles that are
derived from the colourful corona.
5.8 Pollen development

During development inside the anther, primary sporogenous cells are
derived from the same initials as the primary parietal cells of the
anther wall. The sporogenous cells give rise – either directly or by
successive mitotic divisions – to microsporocytes, which are the
parent cells of the microspores. Each microsporocyte undergoes two
meiotic divisions, either successively or almost simultaneously, to
form a tetrad of four haploid microspores, a process termed micro-
sporogenesis. Pollen tetrads can occur either in tetrahedral or tetra-
gonal arrangement, depending on whether microsporogenesis is
simultaneous or successive. A wall of callose (a β-1,3-linked glucan)
develops during microsporogenesis, forming a transient but highly
significant and effective barrier between the diploid somatic tissue
and the haploid developing spores. In the successive microsporogen-
esis type, callose walls are formed after both meiosis I and meiosis II,
5.9 Pollen grains at anthesis 87
forming four microspores embedded in callose. In the simultaneous

type, cytokinesis and callose deposition does not occur until both
meiotic nuclear divisions are complete. Microsporogenesis is simul-
taneous in most eudicots, whereas both successive and simultaneous
types occur in monocots and early divergent angiosperms12, 48.
In most species, the tetrad separates into individual microspores,
though in some species, microspores are released as permanent
tetrads or even remain clumped into larger clusters (e.g. orchid
pollinia can each contain many thousands of pollen grains68).
Before anthesis, each microspore undergoes an unequal division
to form a larger vegetative cell and a smaller generative cell. The
generative cell migrates into the cytoplasm of the vegetative cell and
subsequently divides to produce two sperm, a mitosis that occurs
either inside the pollen grain or in the pollen tube. The mature
pollen grain is therefore usually bicellular, or rarely tricellular.
Development of the pollen-grain wall is also highly regulated12, 103.
Initially, a primexine layer is formed around the microspore within the
callose wall. The primexine is a cellulosic matrix traversed by a radial
system of rod-like probacula56, 77. It accumulates pollen-wall material
that causes it to become undulating within the callose shell, forming
the basis for the pollen exine sculpturing of the dispersed pollen grain.
The innermost layer of the exine and the intine are the final wall layers
to be formed. The transient callose wall is digested by enzymes; in
many species, further sporopollenin secreted by the tapetum or the
tapetal plasmodium is deposited on the pollen surface. In many plants,
pollen-wall apertures, which represent regions that lack sporopollenin
and have reduced primexine deposition, are located in sites that are
predetermined by the contact points of the microspores during the
tetrad phase12, 47, 77.
5.9 Pollen grains at anthesis

At anthesis, pollen grains are radially or bilaterally symmetrical
bodies that represent units of dispersal from the anther to the
88 Flower
stigma (Figure 5.13). They differ considerably in size and shape in

different species, though they are predominantly spherical or
ellipsoidal. Each individual grain consists of a layered wall enclos-
ing a bicellular (or tricellular) microspore, which represents the
male gametophyte12, 77. The pollen-grain wall consists of two
distinct domains: an exceptionally hard and durable outer exine,
which is composed mainly of sporopollenin (a carotenoid poly-
mer), and a relatively soft inner intine, which consists of poly-
saccharides. The exine is itself often differentiated into an outer
sculptured ectexine (sexine) and an inner non-sculptured endex-
ine (nexine). In some monocots, such as Heliconia, the exine is
reduced to isolated spinules of sporopollenin and the intine is
correspondingly thick and channelled77, 135.
Typically, openings (apertures) are present in the exine layer that
reveal the underlying intine. Apertures range in shape from elon-
gated, slit-like furrows to circular pores. Some apertures are partly
covered by an operculum, which is an island of exine that resembles
Figure 5.13 Eschscholzia californica (eudicot: Papaveraceae), scanning electron

micrograph of pollen grain with six apertures (colpi). Scale = 5 µm
5.9 Pollen-tube growth and fertilization 89
a lid46. In spiraperturate pollen grains (e.g. in some Crocus species),

the aperture spirals around the grain. Apertures that lie along the
distal face of the pollen grain (normally the face that was directed
outwards in the tetrad) are termed sulci, whereas apertures located
along the equatorial face of the pollen grain, as defined during the
tetrad phase of development, are termed colpi (Figure 5.13). Sulcate
pollen grains characterize monocots and basal angiosperms. Pollen
with three equatorial apertures (tricolpate pollen) is highly char-
acteristic of eudicots. In most eudicots, microspore tetrads are
tetrahedral with three apertures (colpi) arranged equidistantly
around the equator of the microspore, though the number of
colpi is increased to four or more in many species47.
There are many different patterns of pollen exine sculpturing.
Wind-borne pollen grains are generally small and light, with
relatively little surface sculpturing. Water-dispersed pollen often
possesses adaptations such as a slime coating (e.g. in
Hydrocharitaceae77). In other species, the exine can be reticulate
or areolate, or it can possess surface holes (puncta), granules, warts
or spines102. The surface patterning ensures elasticity of the wall
and resistance to collapse. In species with animal-dispersed pollen,
substances derived from the tapetum – including lipids, proteins
and carbohydrates – are retained in the intercolumellar spaces of
a deeply chambered exine. These substances can confer odour or
cause grains to aggregate into sticky masses, which aids animal
dispersal. Following anther dehiscence, pollen grains dehydrate
after contact with the air, and the exine contracts. Subsequently,
rehydration and exine expansion occur on the stigmatic surface. In
some species (e.g. in Brassicaceae), exine-borne substances derived
from the tapetum are released onto the stigma after rehydration,
allowing species recognition on a compatible host stigma96.
5.9 Pollen-tube growth and fertilization

Pollen tubes emerge from the apertures of pollen grains that have
landed on a receptive stigma and carry the male gametes to the
90 Flower
ovules, a process termed siphonogamy. Typically, each pollen

grain produces a single unbranched pollen tube, but some species
produce multiple or branched pollen tubes69.
A pollen tube is an extension of the vegetative cell of the male
gametophyte. The generative cell is contained within the cyto-
plasm of the vegetative cell; it divides into two sperm cells, either
inside the pollen grain or in the pollen tube itself. The two sperm
cells are often fusiform in shape, allowing them to move with the
cytoplasmic streaming of the vegetative cell, close to the tip of the
pollen tube. Some angiosperms produce callose plugs that block
the pollen tube as it extends, sealing off the growing tip from the
older part of the tube158. The pollen-tube wall represents an
extension of the intine; it consists of an outer pectic layer and an
inner callosic layer57, 97, 130.
Germinating pollen tubes (Figure 5.14) grow rapidly by intru-
sive apical growth between the stigmatic papillae. In species with
a dry stigma, the pollen tube tip uses enzymes to pass through the
stigmatic cuticle. The pollen tubes extend through the style. In
syncarpous eudicots with closed styles, pollen tubes grow along
the middle lamellae of the loosely packed cells of the stylar trans-
mitting tissue. In monocots with open styles, pollen tubes grow
along the mucilagenous surface of the hollow stylar canal. In both
cases, they obtain nourishment (polysaccharides and proteins)
from these richly cytoplasmic stylar tissues. When pollen tubes
reach the ovary, they are nourished by specialized secretory tissues
such as the obturator or the outer integument of the ovule.
A pollen tube typically enters an ovule through its micropyle,
though in a few species it enters via the chalaza69. After discharging
the male gametes into the embryo sac, the pollen tube begins to
degenerate. One of the discharged male gametes enters the haploid
egg cell and fuses with its nucleus to form a diploid zygote. Nuclear
fusion between the second male gamete and the diploid central cell
of the embryo sac results in a triploid primary endosperm nucleus.
This process, termed double fertilization, represents a key feature
5.10 Ovule and embryo sac 91
Figure 5.14 Gymnadenia frivaldii (monocot: Orchidaceae), scanning electron

micrograph of pollen mass (pollinium) on stigmatic surface, with many
germinating pollen grains. Scale = 20 µm
of angiosperms. Following double fertilization, the zygote and

primary endosperm nucleus undergo mitoses to form the embryo
and endosperm respectively (Chapter 6).
5.10 Ovule and embryo sac

Each carpel typically contains one or more ovules, though in
some syncarpous ovaries only one carpel is fertile. Each ovule
is attached to the placenta by a stalk (funicle or raphe), which
contains a vascular strand that usually terminates in the cha-
lazal region, where the funicle, integuments and nucellus meet
(Figure 5.15). The chalaza is often differentiated into specia-
lized structures. In some species, the nucellus proliferates in
this central chalazal zone to form a hypostase, which consists
of suberized or lignified tissue (e.g. in many monocots, such
as Acorus and Crocus).
92 Flower
Figure 5.15 Lomandra longifolia (monocot: Asparagaceae), longitudinal section

of ovule. Scale = 50 µm
During early ovule development, a nucellus is formed from the

apex and body of the ovule primordium. Subsequently, one or two
integuments are initiated around the primordium base and ulti-
mately grow to encircle its apex, forming the micropyle. The inner
integument is usually initiated before the outer integument. The
micropyle is a narrow opening in the ovule formed by one or both
integuments, located at the opposite end of the embryo sac to the
chalaza. The possession of two integuments is the most common
condition in angiosperms, but a single integument characterizes
some eudicots; a few species even lack integuments entirely. In
some species, the dermal nucellar cells surrounding the embryo
sac degenerate and the innermost epidermal layer of the inner
integument becomes differentiated to form an endothelium (inte-
gumentary tapetum) that envelopes the embryo sac and transfers
nutrients from the integument. The endothelial cells are enlarged
and densely cytoplasmic, sometimes becoming polyploid.
5.10 Ovule and embryo sac 93
Within the nucellus, a single hypodermal cell becomes

a primary sporogenous cell (archesporial cell, or archespore).
Occasional species (e.g. Brassica campestris105) possess
a multicellular archespore, of which one cell produces the mega-
gametophyte. The archesporial cell either gives rise directly to the
megaspore mother cell (megasporocyte) or undergoes mitosis to
form a primary parietal cell and a megasporocyte. The megaspor-
ocyte then undergoes two meiotic divisions to form a tetrad of
four megaspores, which are usually either in a linear or T-shaped
arrangement, a process termed megasporogenesis (Figure 5.16).
In the majority of angiosperm flowers, only one of the four
megaspores is functional, most commonly the chalazal one. The
functional megaspore gives rise to the mature embryo sac by
further mitotic divisions, and the other three megaspores
degenerate84, 157, 159. This type of development is termed mono-
sporic or Polygonum-type. In relatively few angiosperms, two or
four megaspores play a role in embryo sac formation; these types
are termed bisporic or tetrasporic respectively.
Figure 5.16 Diagram showing stages of development of the

megagametophyte and megasporophyte, resulting in an eight-nucleate
embryo sac
94 Flower
In most angiosperms, the mature embryo sac (the megagame-

tophyte) possesses eight nuclei arranged in seven cells, though
types with four and sixteen or more nuclei have also been
recorded. At the binucleate stage, the two nuclei migrate to the
micropylar and chalazal poles respectively and subsequently divide
further to form four nuclei. One of the two micropylar nuclei
divides to form the synergids, and the other divides to form the
egg cell and one of the polar nuclei. The two chalazal nuclei each
divide so that one forms two antipodal cells and the other forms an
antipodal and a polar nucleus. The two polar nuclei migrate to the
centre and fuse to form a diploid fusion nucleus. Subsequent
cellularization results in a mature megagametophyte with seven
cells and nuclei: three antipodal cells at the chalazal pole, a central
cell with a fusion nucleus and two synergids plus an egg cell at the
micropylar pole. The synergids and the egg cell are tightly com-
pressed together and are collectively termed the egg apparatus. The
synergids help to direct the pollen tube through the micropyle into
the embryo sac; they are calcium-rich and normally possess a series
of wall thickenings, the filiform apparatus, which extends into the
micropyle. In many species, the antipodal cells degenerate at an
early stage, but in others they persist, and sometimes undergo
endoreplication or cell division, as in many grasses.
6
Seed and fruit
The seed represents the dispersal unit of a plant. Seeds are dis-
persed from the flower either as separate units or enclosed inside
a fruit. Each seed develops from a fertilized ovule. Fruits can
develop from various structures, including a single ovary (simple
fruits, found in the majority of angiosperms), a flower with multi-
ple free carpels (aggregate fruits, e.g. in Ranunculus), a single carpel
(e.g. the monocarpellate pod of the legume family Fabaceae) or
even from an entire inflorescence (e.g. in pineapple, Ananas
comosus).
Seeds and fruits often possess specialized structures that are
related to dispersal13, 76, 136. For example, some wind-dispersed
seeds possess outgrowths that function as wings. Some animal-
dispersed seeds are fleshy and often edible, whereas others possess
structures that are attractive to animals, such as arils and elaio-
somes. Some plants, especially parasitic or mycoheterotrophic
plants such as Monotropa or orchids, produce large numbers of
highly reduced dust seeds from each ovary; these minute seeds
can be blown over long distances.
6.1 Seed coat

The seed coat is formed from ovule tissues, including the chalaza,
raphe (funicle) and integuments. Its primary role changes over
time. During early development, the seed coat can provide nutri-
ents for the developing embryo and endosperm; during seed
96 Seed and fruit
dispersal it protects these soft tissues from dehydration or preda-

tion. Many angiosperms have bitegmic seeds with two distinct
layers, testa and tegmen, which are derived from the inner and
outer integuments respectively13, 21, 153 (Figure 6.1). In contrast,
unitegmic seeds possess a single integument and lack a tegmen.
Seed-coat vasculature consists of a single vascular bundle that
passes through the stalk (raphe) to the chalaza. The vascular bundle
often remains unbranched and terminates in the chalaza, but in
some species it can extend further and even ramify, especially in
larger seeds, though the extent and degree of branching can vary.
The seed coat of many orchids lacks vasculature (Figure 6.2).
At dispersal, the seed coat typically includes a hard, protective
mechanical region. In exotestal seed coats, this mechanical layer
forms from the outer epidermis of the outer integument. In
endotegmic seed coats, it forms from the inner epidermis of the
inner integument. In some species, the mechanical layer consists
Figure 6.1 Diagram of a generalized campylotropous bitegmic

dicotyledonous seed with perisperm
6.1 Seed coat 97
Figure 6.2 Cypripedium calceolus (monocot: Orchidaceae), minute “dust”

seed, with thin testa surrounding a globular embryo. Scale = 100 µm
of one or more rows of elongated, palisade-like cells, such as the

macrosclereids in the exotesta of many legumes (Fabaceae). In
fleshy seeds, the fleshy part of the seed coat, termed the sarcotesta,
is most commonly formed from part of the outer integument. Arils
are fleshy outgrowths of the funicle.
Seed surfaces exhibit a variety of cellular patterns8 and some-
times display characteristic surface sculpturing that is papillate or
striate (Figures 6.3 and 6.4). Some seeds possess epidermal
trichomes; the long, twisted seed-coat hairs of Gossypium (cotton)
represent an important source of textile fibres. Many indehiscent
seeds and fruits produce mucilage when they become wet,
providing a sticky adhesive glue for animal-mediated dispersal,
a phenomenon termed myxospermy (in seeds) or myxocarpy
(in fruits)121, 161. Mucilage is produced in specialized cells in the
seed coats of flax (Linum usitatissimum), plantain (Plantago) and
dodder (Cuscuta)153. In the fruit (nutlet) walls of Coleus and
some other Lamiaceae, the epidermal cells absorb water and
rupture, producing large amounts of slime interspersed with
coiled thread-like protuberances.
98 Seed and fruit
Figure 6.3 Silene nutans (eudicot: Caryophyllaceae), entire seed (left) and
detail of seed surface (right) showing papillate epidermal cells with sinuous
anticlinal walls. Scales: left = 100 µm, right = 20 µm
6.2 Fruit wall (pericarp)

In simple fruits, the pericarp is derived from the ovary wall.
The pericarp displays a range of variation depending on
whether the fruit is dry or fleshy and dehiscent or indehiscent.
It is typically divided into three layers: the outer exocarp,
central mesocarp and inner endocarp, though in some fruits
the three layers are not readily distinguishable. At least one
layer of the fruit wall usually consists of thick-walled lignified
cells (Figure 6.5), though in some fleshy fruits (berries), such
as those of Vitis vinifera (grape), the entire endocarp consists of
thin-walled succulent cells. In other fleshy fruits (drupes),
such as those of Prunus persica (peach), the endocarp cells are
thick walled, and only the mesocarp is fleshy, the exocarp
being a narrow epidermal layer. In many fruits (e.g. Olea
europaea, olive) the fleshy mesocarp is textured with thick-
walled sclereids.
6.3 Grass caryopsis 99
Figure 6.4 Papaver rhoeas (eudicot: Papaveraceae), SEM of entire seed.

Scales = 100 µm
6.3 Grass caryopsis

The grass dispersal unit, often called a seed, is actually an indehis-
cent one-seeded fruit (a caryopsis) in which the testa and pericarp
are fused together (Figure 6.6). The grass caryopsis is an indehis-
cent fruit (an achene) in which the seed coat has undergone
further reduction104, 119. After fertilization, the pericarp consists
of a few cell layers and the integuments largely disintegrate,
leaving only a hyaline membrane derived from the outer layer of
the inner integument. Grass seeds possess highly differentiated
embryos with a unique highly characteristic prominent outgrowth
of the embryo, termed the scutellum, which represents a modified
cotyledon. Grass embryos are well differentiated within the seed, prior
to germination. They characteristically possess a sheath (coleoptile)
surrounding the epicotyl and plumule, and a well-developed radicle
also surrounded by a sheath (the coleorhiza). Some grass species
possess an outgrowth opposite the scutellum, termed the epiblast,
which has been variously interpreted as a second cotyledon, an
100 Seed and fruit
Figure 6.5 Conium maculatum, hemlock (eudicot: Apiaceae), transverse section

of one of a pair of endospermous seeds. Scale = 100 µm
outgrowth of the first cotyledon or the coleorhiza. In some grasses, the

outermost layer of the endosperm (the aleurone layer) is a specialized
tissue of enlarged cells containing protein bodies and large nuclei.
6.4 Endosperm
Endosperm forms a food-storage tissue contained within the seed,
facilitating dispersal and promoting longevity and growth. Mature
endosperm generally consists of tightly packed cells that contain
food reserve materials such as starch grains or protein bodies. In
contrast, the liquid milk of the coconut palm (Cocos nucifera) is
a syncytium that contains many free endosperm nuclei in addition
to oil droplets and protein granules. In the majority of angiosperms,
endosperm is a triploid tissue formed by fusion of a single male
nucleus with two female polar nuclei. Endosperm is present in most
angiosperm seeds but in greatly varying amounts; for example,
endosperm formation is negligible in orchid seeds but extensive
in grass seeds, in which it forms the basis for several crucially
important food sources (e.g. barley, maize, oats, rye).
6.4 Endosperm 101
Figure 6.6 Triticum vulgare, wheat (monocot: Poaceae), longitudinal section

of caryopsis, with detail of aleurone layer (inset). Scale = 500 µm
Early endosperm development is traditionally classified into three

types, termed nuclear, cellular and helobial, based on the timing
and degree of cell-wall formation148. Nuclear endosperm, which
characterizes many eudicots (e.g. Arabidopsis thaliana) possesses both
a syncytial (free-nucleate) phase and a cellular phase. In this type,
early cell divisions are not followed by cell-wall formation, and the
nuclei are initially free in the cytoplasm of the embryo sac, usually
102 Seed and fruit
surrounding a central vacuole. Cell walls eventually form, but some-

times the nuclei at the chalazal end remain free. In the cellular type
of endosperm formation, which occurs in some eudicots (e.g.
Acanthaceae), even the earliest nuclear divisions are followed by
cell-wall formation. In the helobial endosperm type, which is
restricted to monocots, the primary endosperm nucleus undergoes
division to form two unequal chambers, normally a small chalazal
chamber and a large micropylar chamber. The nucleus of the
micropylar chamber migrates to the top of the embryo sac, and
its initial divisions are not accompanied by cell-wall formation,
though cell walls are formed with later mitoses. The chalazal
chamber has far fewer nuclear divisions, and its nuclei remain
free in the cytoplasm; it often has a haustorial role.
Endosperm haustoria can develop in all three types of endosperm.
Haustoria assist nutrient absorption and sometimes invade adjacent
tissues. For example, most species of the mint family (Lamiaceae)
possess both chalazal and micropylar haustoria (Figure 6.7), which
can be either free-nucleate or cellular, sometimes even amoeboid115.
In these species, the first division of the primary endosperm nucleus
is longitudinal, followed by formation of a transverse wall. The
chalazal nucleus forms a small chalazal haustorium close to the
antipodals, and the micropylar nucleus divides further to form
a micropylar haustorium and a central cellular endosperm. The
micropylar haustorium transfers nutrients from the integument to
the embryo and cellular endosperm. The chalazal haustorium trans-
fers nutrients from the vascular bundle to the endosperm.
6.5 Perisperm
In some plants, especially monocots, parts of the nucellus enlarge or
proliferate after fertilization and have a role as a regulating or storage
tissue for the developing embryo. Seed storage tissues derived from
the nucellus are termed perisperm111. In some monocots, endosperm
is entirely absent from the mature seed, and perisperm represents the
primary storage tissue (e.g. in some Poales119). Seeds of Yucca possess
6.6 Embryo 103
Figure 6.7 Prunella grandiflora (eudicot: Lamiaceae), longitudinal section of

young developing seed, showing globular proembryo and two endosperm
haustoria. Scale = 50 µm
perisperm that contains membrane-bound protein and oil bodies
within the cells, together with reserve carbohydrates in the thick cell
walls61. Some members of the ginger order (Zingiberales) possess
perisperm, but this is often entirely compressed in the mature seed,
only the cell walls remaining (e.g. in Musa51). In other Zingiberales
(e.g. Canna53), mitotic activity during ovule development causes the
chalazal region of the nucellus to become massive; this region is
sometimes termed a pachychalaza. In Acorus the perisperm is dermal
in origin, formed from nucellar epidermal cells that elongate and
become filled with clear, transparent, proteinaceous cell contents.
6.6 Embryo
In a typical angiosperm reproductive system, the embryo develops
from the diploid fertilized egg cell (zygote). Following fertilization,
104 Seed and fruit
the zygote often undergoes a change in volume, either shrinkage or

enlargement, before cell division commences. The initial cell divi-
sion is usually transverse and sometimes asymmetric, to form
a small apical and larger basal cell155. The pattern of subsequent
cell division differs between species and has been classified into
several types93. Most embryos eventually differentiate into an undif-
ferentiated globular mass of cells (the proembryo) attached to the
embryo sac wall by a stalk (the suspensor) (Figure 6.8). In Arabidopsis,
the apical cell gives rise to the proembryo, which ultimately forms
the bulk of the embryo, and the basal cell produces the suspensor
and the hypophysis, which is the precursor of the root cortex initials
and the central region of the root cap17. The proembryo can be
massive (e.g. in Degeneria), or small, as in Capsella, in which it consists
of only eight cells131.
Figure 6.8 Diagram showing stages of embryo development in a eudicot

(e.g. Arabidopsis), from a zygote to a two-celled stage resulting from an
unequal cell division, followed by growth of the suspensor, globular
proembryo stages, to a heart-shaped embryo
6.6 Embryo 105
The angiosperm suspensor exhibits considerable diversity in

angiosperms; it can be uniseriate or multiseriate, and filamentous,
spherical or irregular93, 162. Cells of large suspensors, such as those
of Phaseolus, are often endopolyploid. Suspensors of some species
are secretory, and those of others (e.g. Sedum and Tropaeolum) pro-
duce haustoria that invade surrounding endosperm tissue.
The suspensor ultimately degenerates, and the globular proem-
bryo undergoes a process of irregular meristematic activity that
results in a shift from radial to bilateral symmetry. The proembryo
eventually becomes organized into a structure with root and shoot
apices at opposite ends of an embryonic axis (the hypocotyl).
Many dicotyledonous embryos become bilobed or heart-shaped
as the two cotyledons differentiate (Figure 6.9). Monocot embryos
develop a single, often elongated cotyledon. The degree of differ-
entiation of mature embryos varies considerably; for example, in
orchids the embryo remains a simple undifferentiated mass of cells
(Figure 6.2). Some highly differentiated embryos possess, in addi-
tion to the hypocotyl and cotyledons, a short primordial root
(radicle), often with a root cap, and a shoot bud or short shoot
(the epicotyl) developed beyond the cotyledons.
Figure 6.9 Linum usitatissimum (Linaceae), transverse section of seed.

Scale = 100 µm
106 Seed and fruit
6.7 Seedling
At germination, the testa is ruptured and the seedling radicle
emerges through the micropyle and pushes through the substrate.
Seedlings possess a root (radicle) and a hypocotyl, which bears the
cotyledons (seed leaves) and shoot apex. The hypocotyl varies in
size and form, from a swollen food-storage organ to a very short
structure, in which case the radicle extends almost to the cotyle-
donary node. Following the emergence of the radicle, either the
hypocotyl elongates and the cotyledons and shoot apex emerge
(epigeal germination), or the cotyledons remain enclosed in the
testa and the internode above them (the epicotyl) elongates,
pushing the shoot apex upwards (hypogeal germination).
Epigeal germination is the most common type in angiosperms;
the cotyledons are borne above ground and are usually photosyn-
thetic. In contrast, some larger-seeded eudicots such as the legume
pea or bean (e.g. Vicia faba) are hypogeal and possess fleshy,
swollen cotyledons.
In monocots, the seedling radicle withers at an early stage, and
subsequent roots are shoot-borne (adventitious); they are each
initially surrounded by a sheath (coleorhiza), which develops
from the outer cortical tissue by cell elongation. The cotyledons
are usually morphologically different from the first foliage leaves,
often possessing simpler vasculature with a single vascular bundle
(Figure 6.7). In monocot seedlings, the cotyledon typically con-
sists of three parts: a basal sheath, a ligule or ligular sheath, and
a limb, though the relative differentiation of each region varies; for
example, in Tigridia seedlings both the hypocotyl and the basal
sheath are extremely reduced.
Glossary
abaxial: away from the axis. cf. adaxial

abscission layer: well-defined region of tissue separation, e.g. for
abscission
acropetal: towards the apex. cf. basipetal
adaxial: towards the axis. cf. abaxial
adventitious roots: stem- or leaf-borne roots
aerenchyma: specialized parenchymatous tissue associated with
aquatic plants, characterized by a regular, well-developed
system of intercellular air spaces
aleurone layer: specialized outermost cell layer of endosperm in
some grasses
amphistomatic [leaf]: with stomata present on both surfaces
amphivasal [vascular bundle]: with xylem surrounding phloem
amyloplast: plastid containing starch
anisocytic [stomata]: surrounded by three unequal subsidiary
cells
anomalous secondary growth (e.g. xylem with included
phloem): growth in thickness that is secondary (i.e., not
produced by an apical meristem) but does not fit the typical
pattern of xylem and phloem production
anomocytic [stomata]: lacking subsidiary cells
anticlinal: perpendicular to the surface. cf. periclinal
antipodal cell: part of megagametophyte. One of a group of
(most commonly three) cells at the chalazal end of the mature
embryo sac
108 Glossary
aperture [of pollen grain]: thin or modified region of pollen

exine, through which the pollen tube grows at germination
apocarpous gynoecium: one in which carpels are not fused
apoplast: cell areas not bounded by plasmalemma (i.e. cell wall,
middle lamella, intercellular spaces)
apotracheal: parenchyma (in secondary xylem): axial
parenchyma not associated with vessels
archesporial cell: primary sporogenous cell
areole: region of mesophyll between smallest veins in leaf
aril [in seeds]: fleshy outgrowth of the funicle
articulated laticifer: one composed of several cells, their
adjoining walls often partly broken down
astrosclereid: star-shaped or highly branched sclereid
bark: region of woody stem located outside the secondary
xylem; i.e. including vascular cambium, phloem, cortex and
periderm
basipetal: towards the base; i.e. away from the apex. cf. acropetal
bast fibre: extraxylary fibre in stem; i.e. cortical or phloem fibre
bicollateral vascular bundle: one with phloem on both sides of
the xylem
bifacial (dorsiventral) leaf: one with both adaxial and abaxial
surfaces, usually differing morphologically from each other
bitegmic seed: one with two seed-coat layers, derived from two
integuments
brachysclereid (stone cell): more or less isodiametric sclereid
bulliform cells (usually in grass leaves): groups of epidermal
cells that are markedly larger than neighbouring epidermal
cells
bundle sheath: layer of cells surrounding leaf vascular bundles
callose (β–1,3–Glucan) product of plasma membrane that is
primarily a component of the cell wall; acts as a permeability
barrier or sealant in developing tissues such as microspore
tetrads or pollen tubes, or in response to wounding or
pathogens
Glossary 109
callus tissue: undifferentiated mass of thin-walled cells; usually

wound tissue
cambium: meristematic band of cells; e.g. cork cambium or
vascular cambium
campylotropous [ovule]: bent through 90º during development
and fused to the funicle
Casparian strip, or thickening: band of suberin in primary walls
of cells of root endodermis (sometimes also found elsewhere,
e.g. in root exodermis, or stalks of glandular hairs)
cellulose: a carbohydrate; a crystalline compound that is a major
component of plant cell walls
centric (terete) leaf: one that is cylindrical or circular in transverse
section
centrifugal: outwards (from the inside)
centripetal: inwards (from the outside)
chalaza: region of the ovule or seed where nucellus and
integuments merge, opposite the micropyle
chlorenchyma: photosynthetic tissue; specialized parenchyma
containing chloroplasts
chlorophyll: green photosynthetic pigment contained within
chloroplasts
chloroplast: plastid containing chlorophyll, the site of
photosynthesis
coenocyte: multinucleate cell; i.e. one in which cell division has
occurred without cell-wall formation, e.g. non-articulated
laticifer
coleoptile (in seedlings): parenchymatous sheath enclosing
plumule
coleorhiza (in monocot seedlings, especially grasses):
parenchymatous sheath covering primary root
collateral: vascular bundle: one with xylem and phloem adjacent
to each other
collenchyma: strengthening tissue, consisting of groups of
axially elongated, tightly packed cells with unevenly thickened
walls
110 Glossary
colpus (pl. colpi): aperture in pollen grain wall, aligned

equatorially during the tetrad stage, and usually elongated (slit-
like)
companion cell: parenchymatous cell associated with sieve-tube
element in phloem.
compitum: opening in transmitting tissue of ovary, near
micropyle
cork: suberized tissue (periderm)
cortex: region in stems and roots between epidermis and central
vascular region
cotyledon: first leaf of the embryo
cuticle: non-cellular layer of a fatty substance (cutin), covering
surface of epidermis
cystolith: calcareous body found in epidermal cell, or in leaf
mesophyll
cytokinesis: cytoplasmic cleavage following nuclear division
diacytic [stomata]: with one or more pairs of subsidiary
cells with their common walls at right angles to the guard
cells
diarch [root]: with two protoxylem poles
dictyosome or Golgi body: cell organelle associated with
secretory activity
distal: located away from the centre of a body or its point of
attachment; sometimes terminal on axis (cf. proximal)
domatia: specialized pockets or tufts of hairs on some leaf
surfaces, providing shelter for small insects; sometimes
associated with extrafloral nectaries
dorsiventral or bifacial [leaf]: with the two surfaces
morphologically different from each other
druse: cluster crystal, or compound crystal
ectomycorrhizal [fungal mycelium on roots]: associated with the
surface (cf. endomycorrhizal)
ectexine (ektexine or sexine): outer, sculptured part of exine in
pollen grain wall
Glossary 111
egg apparatus (in mature embryo sac): egg cell and two synergids
egg cell: part of megagametophyte; haploid cell at micropylar end
of mature embryo sac that will fuse with male haploid nucleus
to form a zygote
elaiophore: oil-secreting trichome or tissue in flower
embryogenesis: embryo development
endexine (nexine) inner layer of exine in pollen grain wall
endocarp: inner layer of fruit wall (pericarp)
endodermis: innermost cell layer of cortex (mainly in roots)
endogenous: of deep-seated (internal) origin (cf. exogenous)
endomycorrhizal [fungal mycelium on roots]: invading tissues
and cells (cf. ectomycorrhizal)
endoplasmic reticulum (ER): series of membranes permeating
the cytoplasm
endosperm: seed storage tissue, formed by fusion of one male
nucleus with two female polar nuclei (i.e. usually triploid)
endotegmic seed coat: one with thickened, mechanical layer
derived from the inner epidermis of the inner integument
endothecium: anther wall layer immediately within the
epidermis; often possessing characteristic thickenings
epiblast: in grass embryos, outgrowth opposite scutellum
epicotyl: seedling axis above cotyledons
epidermis: outermost layer of cells, covering entire primary plant
surface
epigeal germination: seedling germination type in which
cotyledons are green and borne above ground
epigynous flower: one with inferior ovary (i.e. the ovary is
attached to the receptacle above the level of insertion of the
stamens and perianth parts)
epistomatic leaves: with stomata found on adaxial surface only
epithem: tissue (often loosely packed parenchyma) in hydathode
between epidermis and vascular tissue
equatorial (pollen grain aperture): located at or crossing a line
midway between the two poles of a microspore or pollen grain
112 Glossary
exarch root: one that matures centripetally

exine: outer coat of pollen grain, often differentiated into outer
ectexine and inner endexine
exocarp: outermost layer of pericarp
exodermis: outer few cell layers of root cortex that have become
thicker walled and lignified
exogenous: of superficial (external) origin (cf. endogenous)
exotestal: seed coat: one with mechanical layer formed from the
outer epidermis of the outer integument
fibre: axially elongated, thick-walled cell, usually occurring as part
of a group, lacking contents at maturity, and with simple pits
fibre-tracheid: cell type that is transitional between a fibre and a
tracheid, possessing bordered pits
filament: (in flower) stalk of stamen
filiform apparatus: wall thickenings in synergid cells of mature
embryo sac
funicle (funiculus): stalk attaching ovule to placenta in ovary
fusiform: elongated with pointed ends
generative cell: part of male gametophyte; divides (usually
within pollen tube) to form two sperm cells
girder (or bundle sheath extension): in leaves, usually a group of
cells (parenchymatous or sclerenchymatous) linking a vascular
bundle sheath with either or both epidermises
graft: union (by cell differentiation) of tissues of two different
individuals so that one (the scion) can survive through
dependency on the other (the stock)
ground tissue: tissue (usually parenchyma) surrounding vascular
tissue
growth ring: (in secondary xylem) a distinct growth increment
caused by differential rates of growth during a growing season
guard cell: one of a pair of cells of a stoma, together surrounding
a pore
guttation: secretion of water droplets, often occurring at
hydathodes
Glossary 113
gynoecium (in flower): collective term for carpels; including

ovary, style(s) and stigma(s)
hair: epidermal appendage, or trichome
haustorium (pl. haustoria): cellular process that penetrates
adjacent tissues and has a role in nutrient transport. For
example, haustoria of parasitic plants are modified roots;
endosperm haustoria are specialized absorptive cells
hemicellulose: one of the carbohydrate constituents of plant cell
walls
heterocellular ray: one composed of cells of different shapes and
sizes
hilum: scar on seed indicating point of attachment of funicle to
ovary wall
histogenesis: tissue differentiation
homocellular ray: one composed of cells which are the same
shape and size
hydathode: region of secretion of water droplets (usually on leaf
margin)
hydrophyte: water plant, sometimes displaying hydromorphic
features (e.g. aerenchyma)
hypocotyl: seedling axis bearing cotyledons and shoot apex
hypodermis: (often applied to leaves) distinct cell layer(s)
immediately within epidermis
hypogeal germination: type in which cotyledons remain enclosed
in seed coat after radicle has emerged
hypogynous flower: one with superior ovary; i.e. ovary attached
to the receptacle above the level of insertion of the stamens and
perianth parts
hypophysis: uppermost cell of the suspensor
hypostase: proliferation of nucellus at chalazal end of embryo sac,
often bearing antipodals
hypostomatic leaves: with stomata found on abaxial surface only
idioblast: isolated specialized cell that differs from cells of
adjacent tissue in size, shape and function
114 Glossary
included phloem: areas of phloem embedded in secondary

xylem
inaperturate (microspore or pollen grain): one lacking a clearly
defined aperture on the surface, so that the pollen tube can
potentially emerge at any point
integument: structures enclosing the embryo sac and nucellus
intercalary growth: cell divisions in region apart from apical
meristem or other well-defined meristems
internode: region separating two nodes on stem
intine: inner layer of pollen grain wall
isobilateral leaf: one with both surfaces similar, or with palisade
tissue on both sides (abaxially and adaxially)
Kranz anatomy: (in some plants with C4 photosynthesis)
distinctive leaf anatomy with mesophyll cells radiating from
the vascular bundle sheaths
lateral roots: branches of the tap root; they have an endogenous
origin
latex: fluid contained within laticifers, consisting of a suspension
of fine particles
laticifer: latex-secreting cell
lenticel: region of loose cells in the periderm (bark)
lignin: substance often deposited in cell walls of strengthening
tissues (e.g. fibres), giving rigidity
ligule: outgrowth of the abaxial epidermis of a leaf in the region
between the sheath and petiole (or between sheath and lamina
if the petiole is absent)
macrosclereid: elongated sclereid often found in seed coat
megagametophyte: mature embryo sac, most commonly
consisting of eight nuclei
megaspore: female haploid cell resulting from meiosis;
usually one of two or four, of which only one is
functional
megasporogenesis: process of megaspore formation from a
megaspore mother cell
Glossary 115
megasporocyte: a diploid cell that will give rise to (usually) four

haploid megaspores following meiosis
meiosis: two successive divisions of a diploid nucleus to form a
haploid gamete
meristem: region of cell division and tissue differentiation (e.g.
apical meristem, intercalary meristem, lateral meristem,
vascular cambium, primary and secondary thickening
meristems)
meristemoid: isolated meristematic cell, usually the smaller cell
resulting from an asymmetric division (e.g. guard-mother
cell)
mesocarp: middle layer of pericarp
mesogene cell: stomatal subsidiary cell derived from
meristemoid
mesomorphic: displaying no xeromorphic or hydromorphic
characteristics
mesophyll: ground tissue of leaf; mainly consisting of
parenchyma or chlorenchyma; often differentiated into
palisade and spongy mesophyll
metaxylem: primary xylem formed after protoxylem
microfibril: thread-like component of cell wall, usually of
cellulose
micropyle: opening at one end of the ovule, usually formed by
the integuments
microsporangium: pollen sac, contained within anther
microspore (male spore): individual haploid cell that will give
rise to male gametophyte (pollen grain)
microsporocyte: a diploid cell that will give rise to four haploid
microspores following meiosis
microsporogenesis: developmental process leading to the
production of four haploid microspores from a diploid
microsporocyte by meiosis and cytokinesis
middle lamella: layer between the walls of neighbouring
cells
116 Glossary
mitochondrion (pl. mitochondria): respiratory cytoplasmic

organelle
mitosis: cell division to form two cells of equivalent chromosome
composition to parent cell
mucilage (slime): compound that swells in water
multiseriate: consisting of more than one layer or row
of cells
nectary: (floral or extrafloral) localized cell or cells that secrete a
sugary liquid (nectar)
nexine: inner, non-sculptured part of exine
node: part of stem where leaves are attached
non-articulated laticifer: one composed of a single
multinucleate coenocytic cell
nucellus: ovule cell layer(s) immediately surrounding
megagametophyte
obturator: proliferation of (usually) ovary tissue near micropyle,
with secretory function, to guide growing pollen tubes into
micropyle
ontogeny: development; differentiation and growth
osmophore: scent-producing gland
osteosclereid: bone-shaped sclereid
papilla (pl. papillae): epidermal appendage; small unicellular
trichome
paracytic [stomata]: with one or more subsidiary cells at either
side of the guard cells
paratracheal parenchyma: (in secondary xylem) associated with
the vessels
parenchyma: tissue composed of thin-walled cells with living
contents
passage cell: an endodermal cell that remains thinner walled than
neighbouring endodermal cells
pearl glands (pearl bodies): secretory, often globular, leaf
emergences or trichomes that provide food (carbohydrates,
lipids and proteins) for ants
Glossary 117
perforated ray cell: (in secondary xylem) ray cell linking two
vessel elements, and itself resembling and functioning as a
vessel element
perforation plate: opening in end wall of vessel element
perianth: outer sterile part of flower
pericarp: fruit wall
periclinal: parallel with the surface. cf. anticlinal
pericycle: in roots, a distinct layer of thin-walled cells located
within the endodermis
periderm: cork tissue
perigene cell: in stomatal complex, neighbour cell that is
not derived from the same cell lineage as the guard
cells
periplasmodium: coalescent mass in anther locule, formed from
protoplasts of tapetal cells
perisperm: food-storage tissue in the seed, derived from part of
the nucellus
phellem: external derivatives of phellogen
phelloderm: internal derivatives of phellogen
phellogen: cork cambium, or cork meristem
phloem: tissue that transports food in the form of assimilates;
either primary (produced by the apical meristem) or secondary
(produced by the vascular cambium)
phyllotaxis: the pattern of arrangement of organs on an axis, e.g.
leaves on a stem, flowers on an inflorescence
pith: central parenchymatous region of stem
pits: thin areas of the primary and secondary cell wall, often
corresponding with pits in adjacent cells (pit-pairs)
placenta: region of attachment of ovules on ovary wall
placentation: arrangement of placentae and locules in ovary (e.g.
axile, basal, free central, parietal)
plasma membrane (plasmalemma): cell membrane (within cell
wall) that encloses protoplast
118 Glossary
plasmodesmata: protoplasmic strands passing through primary

pit fields between adjacent cells and connecting their
protoplasts
plastid: cell organelle contained within cytoplasm, often with
specialized function (e.g. chloroplast, amyloplast)
polar nucleus: one of a pair of nuclei of the mature
megagametophyte, often in a central position
pollen grain: male gametophyte; bicellular (or rarely tricellular)
at maturity
pollen tube: tube emerging from germinating pollen grain on
stigma, which will transport male nuclei to megagametophyte
polyarch root: one with several (more than four) protoxylem
poles
primordium: newly differentiating organ
procambium: primary tissue near (shoot or root) apex that gives
rise to primary vascular tissue
proembryo: young globular embryo prior to differentiation of
cotyledons and hypocotyl
promeristem: (in root apices) region of greatest mitotic activity
protoplast: living part of cell, surrounded by a plasma membrane
protoxylem: first-formed primary xylem
proximal: situated closer to the centre of a body or its point of
attachment (antonym: distal)
quiescent centre: region of cells at root apex, with little or no cell
division activity
radicle: first-formed root of seedling
raphe: stalk attaching seed to ovary (directly derived from the
funicle)
raphide: fine, needle-like crystal, one of a group of several
raphides formed within a single cell
ray: (in secondary xylem) tissue of radially oriented cells, usually
parenchymatous
root cap: protective covering of cells over root apex
root hair: water-absorbing hair on root epidermis
Glossary 119
sarcotesta: fleshy part of seed coat

scale (peltate hair): a modified trichome, consisting of a fused
disc of cells attached to the epidermis by a stalk
sclereid: thick-walled sclerenchymatous cell, usually lacking
contents at maturity
sclerenchyma: strengthening tissue, consisting of cells with
thickened lignified walls, usually lacking contents at maturity
scutellum: specialized structure in grass embryos, often
interpreted as a modified cotyledon
sieve tube element: conducting cell in phloem; possessing sieve
areas and sieve plates in walls. Groups of sieve tube elements
are linked axially to form sieve tubes
sporopollenin: a highly resistant complex polymer that forms the
outer wall of a pollen grain
stipules: appendages at base of leaf sheath, often paired and
sometimes leafy
stoma (pl. stomata): pore in aerial epidermis
stone cell: isodiametric sclereid
styloid: elongated prismatic crystal
suberin: fatty compound sometimes deposited in cell walls, e.g.
in cork cells
subsidiary cells: epidermal cells adjacent to stomata or other
subsidiary cells, which differ from surrounding epidermal
cells
sulcus (pl. sulci): aperture in pollen grain wall on its distal
face (i.e. the face that is normally directed outwards in the
tetrad)
suspensor: row of cells attaching globular proembryo to wall of
embryo sac
symplast: connected living protoplasts of adjacent cells
syncarpous gynoecium: one in which carpels are more or less
fused, though stigmas and styles may remain separate
syncytium: cytoplasmic region enclosed by a single plasma
membrane and bearing several nuclei
120 Glossary
synergid: part of megagametophyte; one of a pair of cells at

micropylar end of mature megagametophyte
tannins: a broad group of phenol derivatives
tap root: main central root, formed directly from seedling radicle
tapetum: layer of nutritive tissue between the microsporocytes or
developing pollen grains and the wall of the anther locule
tegmen: the inner layer of the seed coat, formed from the inner
integument
testa: seed coat; or in bitegmic seeds, the outer layer of the seed
coat, formed from the outer integument
tetrad: group of four microspores or megaspores, the daughter
cells of a single microsporocyte or megasporocyte
tetrarch (root): one with four protoxylem poles
tracheid: xylem water-conducting cell, usually possessing
bordered pits but lacking perforation plates
transfer cell: specialized plant cell that facilitates transport of
soluble substances across tissue boundaries
transmitting tissue (stigmatoid tissue): secretory tissue through
which pollen tubes grow, linking stigma with centre of ovary
triarch (root): one with three protoxylem poles
trichoblast: root epidermal cell with dense cytoplasm, which will
give rise to a root hair
trichome: epidermal outgrowth (hair)
tricolpate (microspore or pollen grain): one with three colpi
tunica-corpus: regions of central shoot apical organization
tylose: (in secondary xylem) outgrowth of the wall of an axial
parenchyma cell into a vessel element through a pit; eventually
often completely blocking vessel
unifacial leaf: one with both surfaces similar, sometimes derived
from a single (usually abaxial) surface
uninterrupted meristem: region of diffuse cell divisions that is
continuous with the apical meristem; producing extension
growth of the axis
uniseriate: consisting of a single layer or row of cells
Glossary 121
vacuole: cavity
vascular cambium: bifacial meristem that produces secondary
vascular tissue (phloem and xylem)
vascular tissue: conducting tissue (phloem and xylem)
vegetative cell: one of two cells of the male gametophyte
velamen: outer dermal layer on the aerial roots of some tropical
epiphytes
venation: arrangement of vascular bundles in leaf (e.g. parallel or
reticulate venation)
vessel element: water-conducting cell of xylem, possessing
bordered pits on lateral walls and perforation plates on end
walls; groups of axially linked vessel elements form a vessel
vestured pitting (in secondary xylem): bordered pits surrounded
by numerous warty protuberances
wax: fatty substance often deposited on the surface of the cuticle
whorl: ring of organs; sometimes applied to a region of a similar
organ type, e.g. the stamen whorl
wood: secondary xylem
xeromorphic: showing characteristics that are often associated
with dry environments
xerophyte: plant that grows in a dry (xeric) environment
xylem: water-transporting tissue, consisting of several different
cell types
zygote: cell formed by fusion of megasporocyte and
microsporocyte; will eventually divide to form the proembryo
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Index
Acanthaceae, 102 Bromeliaceae, 20

Acorus, 55, 57, 91, 103 bulliform cell, 58
Adoxaceae, 37 bundle sheath, 69
adventitious root, 48, 49
aerenchyma, 12, 13, 69 Cactaceae, 26, 39
Agave, 39, 67 calcium oxalate crystal, 5
aleurone layer, 100 callose, 86, 90
Allium, 55 callus, 10, 49, 109
Alnus, 33 Camellia, 15, 66
Aloe, 39, 65, 67, 70 Canna, 103
Amborella, 9, 18, 82 Cannabis, 14, 18
Ammophila, 71 Capsella, 104
amyloplast, 5, 6, 16 carnivorous plant, 20, 62
Ananas, 95 carotenoid, 79
anther locule, 80 carpellary bundle, 78
anthocyanin, 79 Caryophyllaceae, 98
Apiaceae, 100 caryopsis, 99, 101
Arabidopsis, 17, 41, 42, 45, 48, 73, 80, 83, Casparian strip, 46
85, 101, 104 cellulose, 2, 109
archesporial cell, 93 Ceratophyllum, 58
areole, 67 chalaza, 90, 91, 96
aril, 97 Chenopodiaceae, 69
Arisaema, 54 chlorenchyma, 12, 26, 65
Asparagaceae, 40, 62, 78, 83, 92 chloroplast, 5
astrosclereid, 11, 14, 66 chromoplast, 5
Atriplex, 6 Cissus, 6, 31
Avicennia, 20, 33, 61 Clerodendrum, 29
axial parenchyma, 32, 34, 35 Coix, 60
coleoptile, 99
Bartsia, 52 coleorhiza, 99, 106
Beaucarnea, 39 Coleus, 97
Begonia, 49 collenchyma, 11, 13, 26
Betula, 37 colpus, 89
Betulaceae, 33, 51 companion cell, 21, 22, 32, 36
bordered pits, 3, 21 compitum, 84
Brassica, 93 contractile root, 46
Brassicaceae, 17, 73, 80, 83, 85, 89 Convolvulaceae, 52
136 Index
Cordyline, 39, 40, 78 exine, 81, 87

cork, 37 exocarp, 98
cortex, 26, 29, 37, 45 exodermis, 45
cotyledon, 1, 99, 105, 106 extrafloral nectary, 63
Crassulaceae, 10
Crassulacean acid metabolism (CAM), Fabaceae, 27, 35, 43, 46, 53, 58, 95,
5 97
Crocus, 26, 46, 65, 66, 84, 89, 91 Fagaceae, 19, 32, 51
crystal sand, 7 fibre, 14, 34
Cuscuta, 52, 53, 97 Ficus, 16, 56, 57, 58, 59, 65,
cuticle, 2, 15, 18, 59, 79, 90 66
cuticular striations, 59 filiform apparatus, 94
cystolith, 7, 58, 59 Flagellariaceae, 75, 77, 81
cytokinesis, 1, 132 floral apex, 74, 76
funicle, 91, 97
Daucus, 41 fusiform initial, 31
Degeneria, 104
Dionaea, 62 glandular trichome, 18
Dioscorea, 6 Gossypium, 97
domatia, 64 guard cell, 16
double fertilization, 90 Gymnadenia, 91
Dracaena, 49 gynostemium, 77
Drimys, 74
Drosera, 20, 62 haustorium, 53, 102
druse, 7 Heliconia, 88
druses, 6, 110 helobial endosperm, 102
Hevea, 23
ectomycorrhizal fungi, 51 Hibiscus, 63
egg cell, 94 hydathode, 63
elaiophore, 84, 85, 86 Hydrocharitaceae, 89
Eleusine, 70 hydrophyte, 69
embryo, 95, 97, 99, 103, 104 hydropote, 20, 69
embryo sac, 93, 94 Hypenia, 55
endocarp, 98 Hypericaceae, 76
endodermis, 46 Hypericum, 76
endoplasmic reticulum, 4 hypocotyl, 1, 105, 106
endosperm, 5, 90, 95, 100, 102, 103, 105, hypodermis, 57, 65, 71
107 hypogeal germination, 106
endothecium, 80 hypophysis, 104
endothelium, 92 hypostase, 91
epiblast, 99 Hyptis, 28
epicotyl, 99, 105
epicuticular wax, 59, 60 idioblast, 5, 6, 11, 14, 23, 66
epidermis, 7, 15 Illicium, 82
epigeal germination, 106 included phloem, 33
epithem, 63 integument, 92
Erica, 63 intine, 87, 88
Ericaceae, 63 Iridaceae, 7, 55, 66, 86
Eschscholzia, 88 Iris, 7, 47
Euphorbia, 6, 11, 23, 26, 59, 66, 84
Euphorbiaceae, 11, 23, 64 Kranz anatomy, 5, 69, 70
Index 137
Lamiaceae, 19, 28, 29, 55, 61, 78, 85, 97, Neottia, 51
102, 103 Nigella, 74
Lapageria, 4 nitrogen-fixing bacteria, 52
lateral meristem, 8 nodal anatomy, 28
lateral root, 47, 48 nucellus, 91
Lathraea, 52 nuclear endosperm, 101
laticifer, 6, 11, 23, 26, 66 nucleus, 4, 90
leaf gap, 28 Nymphaea, 13, 20, 66, 80
leaf primordium, 56
lenticel, 38 obturator, 84
lignin, 2, 36 Ochroma, 35
Ligustrum, 2, 48 Olea, 98
Liliaceae, 21, 29, 82 Oleaceae, 2, 48
Lilium, 21, 29, 82 Omphalea, 64
Limonium, 61 Ophrys, 85
Linum, 14, 97, 105 Opiliaceae, 58
Lomandra, 92 Orchidaceae, 16, 44, 51, 77, 79, 85, 87,
Loranthaceae, 52, 53 91, 95, 96, 97, 100, 105
Lythrum, 75 Orobanchaceae, 52
Orobanche, 52
Malvaceae, 38, 76 Oryza, 7, 45
marginal growth, 56 osmophore, 84, 85
megagametophyte, 93, 94
megaspore, 93 pachychalaza, 103
megasporocyte, 93 palisade mesophyll, 12, 64, 71
megasporophyte, 93 Papaver, 23, 99
meristem, 8 Papaveraceae, 88, 99
meristemoid, 9, 22 papillae, 18
mesocarp, 98 parallel venation, 67
mesophyll, 55, 64, 78 parenchyma, 11, 12
metaxylem, 21, 47 pearl gland, 63
micropyle, 92, 106 Peperomia, 57
microspore, 82, 86 perforated ray cell, 32
microsporocyte, 86 perforation plate, 21
microsporogenesis, 86 pericarp, 98
middle lamella, 2, 10, 11 pericycle, 44, 45, 46, 47
mitochondria, 4, 5, 22 periderm, 36, 37, 45
Monotropa, 95 periplasmodium, 81
Moraceae, 56, 59 perisperm, 102
mucilage, 5, 6, 18, 20, 23, 62, 70, Phaseolus, 105
97 phellogen, 9, 36, 37, 38
multiple epidermis, 57 phloem, 12, 20, 22
multiseriate epidermis, 16 pigment, 79
Musa, 103 pith, 26
mycorrhizae, 51, 52 placenta, 84
myxocarpy, 97 Plantago, 97
myxospermy, 97 plasma membrane, 1
plasmodesmata, 3
Narcissus, 86 plastid, 4, 5, 16
nectar, 63 Platanthera, 85
nectary, 5, 64, 84, 85 plumule, 99
138 Index
Poaceae, 7, 42, 43, 60, 70, 71, 73, 100, Schisandraceae, 61

101 sclereid, 14, 15, 66
Podostemaceae, 58 sclerenchyma, 11, 12, 14, 70
polar nucleus, 94 scutellum, 99
pollen aperture, 88 secondary phloem, 31, 35
pollen grain, 87, 88 secondary thickening meristem (STM), 39,
pollen tetrad, 86 40
pollen tube, 89, 90 secondary xylem, 14, 31
pollen-tube transmitting tissue, 84 Sedum, 105
pollen-tube-transmitting tissue (pttt), seed coat, 95
78 septal nectary, 85
Populus, 29 shoot apex, 1, 2, 8, 24, 25, 29, 38, 39,
primary thickening meristem (PTM), 106
38 sieve element, 22, 32, 36
proembryo, 103, 104, 105 sieve element plastid, 22
promeristem, 42 sieve plate, 22
protoplast, 1, 3 Silene, 98
protoxylem, 21, 47 silica, 16
Prunella, 103 silica body (phytolith), 7, 58
Prunus, 29, 30, 68, 98 Solanum, 54, 57
sperm cell, 90
Quercus, 19, 29, 32, 35, 37, 50 spongy mesophyll, 12, 64
quiescent centre, 42 sporopollenin, 81, 87, 88
stamen fascicle, 80
radicle, 1, 99, 105, 106 starch, 5
Ranunculaceae, 44, 74 stigma, 82, 83
Ranunculus, 32, 44, 49, 95 stinging hair, 61
raphides, 6, 7 stomata, 4, 16, 17, 58, 60, 61, 63,
ray, 35 70, 79
ray initial, 31 stylar bundle, 78
ray parenchyma, 34 style anatomy, 84
receptacle, 72 styloid, 7
reticulate venation, 67 suberin, 2, 36, 46
Rhinanthus, 52 sulcus, 89
rhizodermis, 15, 43, 44 suspensor, 104, 105
ribosome, 4 symplast, 3
Ricinus, 80 syncarpous gynoecium, 78, 82
root apex, 1, 8, 41, 45 synergid, 94
root cap, 41, 105
root hair, 44 Tamarix, 61
root haustoria, 52 tannin, 5
root nodule, 52 tapetum, 80, 81, 87
Rosaceae, 29, 30, 68 tegmen, 96
testa, 1, 96, 97, 99, 106
salt-secreting gland, 61 thecae, 80
Salvia, 19, 78 Tigridia, 86, 106
Sambucus, 37 Tilia, 35
Santalaceae, 52 tracheary element, 3, 20, 21, 32
Santalum, 52 transfer cell, 13
sarcotesta, 97 transmitting tissue, 90
Schisandra, 61 trichoblast, 9, 45
Index 139
trichome, 18, 19, 61, 63, 79, Vicia, 27, 41, 42, 43, 46, 58,
85, 97 106
Tropaeolum, 105 Viscum, 52
tunica-corpus, 24 Vitaceae, 6, 31
tylose, 34 Vitis, 98
Ulmaceae, 50 Winteraceae, 74
Ulmus, 50
Urtica, 19, 61 xerophyte, 70
xylem, 12, 20
vacuole, 4
vascular cambium, 27, 28, 31, 34, Yucca, 39, 40, 62, 102
35
veinlet ending, 67 Zea, 41, 42, 43
velamen, 16, 43, 44 zygote, 90, 103

Paula Rudall - Anatomy of Flowering Plants (2020)

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Anatomy of Flowering Plants

Paula J. Rudall is Research Professor at the Royal Botanic

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

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

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

This publication is in copyright. Subject to statutory exception

First edition published by Edward Arnold 1987

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

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

1 Organs, cells and tissues 1

6 Seed and fruit 95

Plant anatomy is concerned with the structural organization of plants

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

are assigned to families according to a modern understanding of

encompass a wide range of morphological diversity, espe-

Organs, cells and tissues

1.2 Cells and cell walls

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

located in the equatorial zone between the two daughter nuclei138.

Figure 1.2 Diagram of a generalized plant cell

1.3 Cytoplasm, plastids and photosynthesis

organelles; they include mitochondria, the nucleus, plastids and

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

1.4 Inorganic cell inclusions

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

membrane chambers, lamellae, mucilage and fibrillar material.

a characteristic feature of some flowering plant groups. These

thickening meristems (Chapter 2). Lateral meristems occur in

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

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

1.6 Cell growth and expansion

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

require additional strength in their protective role, display differ-

simple tissues. In contrast, complex tissues incorporate elements of

1.7.1 Parenchyma and other thin-walled cell types

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

large air spaces are formed by differential growth of surrounding

1.7.2 Thick-walled tissues: collenchyma

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

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

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

Figure 1.10 Arabidopsis thaliana (eudicot: Brassicaceae), abaxial leaf surface,

the guard cells. Specialized lateral subsidiary cells (e.g. in grasses

Figure 1.11 Amborella trichopoda (ANA-grade: Amborellaceae), transverse

series of asymmetric divisions (amplifying divisions) results in

1.10 Trichomes and papillae

1.11 Vascular tissue

Figure 1.14 Vascular bundle. Lilium tigrinum (monocot: Liliaceae), transverse

often including parenchyma and fibres as well as vessels. Tracheids

plates are generally either simple, with a single opening, or scalari-

1.12 Secretory cells and laticifers

2.1 Shoot apex

Figure 2.1 Diagram of angiosperm shoot apical organization

2.2 Primary stem structure

Figure 2.2 Vicia faba (eudicot: Fabaceae), transverse section of stem.

(bicollateral vascular bundles), or with xylem surrounding the

Figure 2.3 Vascular cambium. Hyptis oblongifolia (eudicot: Lamiaceae),

towards the apex. In some palms, thousands of vascular bundles are