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Anatomy and Morphology of Seed Plants

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Anatomy and Morphology
of Seed Plants
Gregor Fraser Barclay, Department of Life Sciences, The University of the West Indies,
St Augustine, Republic of Trinidad and Tobago
Seed plants, the most successful and diverse group of
higher plants, are distinguished from other plants by the
presence of a testa (seed coat) that protects the embryo.
Plant anatomy describes the structure and organisation
of the cells, tissues and organs of plants, whereas plant
morphology describes the external form and structure of
plants. The origin of these fundamental botanical sci-
ences is outlined, and illustrations and descriptions are
given of the main anatomical and morphological features
of plants, largely from a functional perspective. Meriste-
matic, dermal, support and vascular tissues are described.
Also, the fundamental organs, root, stem and leaf, as well
as some of their notable modifications are described.
Special attention is given to the flower and its modifica-
tions. Seed structure and dispersal mechanisms are
described, and a key to identifying the common types of
fruits is provided.
Introduction
The spermatophytes, or seed plants, form the most familiar
group of plants, which is comprised of the gymnosperms
(conifers and their relatives) and the angiosperms (flow-
ering plants). Seed plants range enormously in size, ranging
from species of the aquatic duckweed Wolffia in the family
Lemnaceae, with individual plants typically weighing just
150 mg, to the General Sherman Tree, a specimen of giant
sequoia (Sequoiadendron giganteum), which is the most
massive living thing on Earth. Its trunk alone is estimated
to weigh 1300 metric tons, 7 trillion times more than a
duckweed plant.
The angiosperm division is represented by at least
300 000 species, possibly 450 000 (Evert and Eichhorn,
2013), making it the largest group of photosynthetic
organisms. Yet, in spite of their diversity all seed plants are
constructed from just a few different types of cells, tissues
and organs. This article presents a general introduction to
eLS subject area: Plant Science
How to cite:
Barclay, Gregor Fraser (January 2015) Anatomy and Morphology of
Seed Plants. In: eLS. John Wiley & Sons, Ltd: Chichester.
DOI: 10.1002/9780470015902.a0002068.pub2
eLS & 2015, John Wiley & Sons, Ltd. www.els.net
Introductory article
. Introduction
Article Contents
. General Structure of Seed Plants
. The Organ System
. Acknowledgement
Online posting date: 27th January 2015
the anatomy (internal structures) and morphology (exter-
nal features) of seed plants. See also: Angiosperms;
Gymnosperms
The anatomy of plants was first described in the works of
Nehemiah Grew (1672, 1682) and Marcelli Malpighii
(1675). Among the terms that Grew introduced in 1672 are
parenchyma, radicle and plume (now termed plumule), and
he distinguished between cortical, lignous (lignified)
and pith regions of stems and roots. In 1682, Grew
described vessels which contained sap that was ‘sensibly
more sweet’ than that of the parenchyma, likely having
chanced upon sieve tubes of the phloem. He was the first to
describe the pollen grain in detail and to recognize its role in
reproduction of plants. See also: Malpighi, Marcello
The ancient Greek philosopher Theophrastus (c. 371–
287 BCE), regarded as the first to describe the morphology
of plants, was perplexed by the variations he observed
(Hort, 1896, p. 49):
In fact your plant is a thing various and manifold, and so
it is difficult to describe in general terms: in proof
whereof we have the fact that we cannot here seize on
any universal character which is common to all, as a
mouth and a stomach are common to all animals;
whereas in some plants some characters are the same in
all, merely in the sense that all have analogous char-
acters, while others correspond otherwise.
Despite his apparent confusion, Theophrastus did
recognize the notion of analogous structures in dissimilar
plants. For example, it is now known that succulent stems
can occur not only in plants of the Cactaceae but also in
plants of families as taxonomically distinct from each other
as the Eurphorbiaceae, Asteraceae and Asclepiadaceae
making these structures functionally analogous. Further-
more, cacti can bear spines that are modified leaves while
some species of the Eurphorbiaceae bear thorns that are
modified branches, which are analogous structures as well.
In contrast, petals and leaves have different functions but
the same origin, demonstrating homology.
The gifted literary figure Johann Wolfgang von Goethe
established plant morphology as a science in 1790 with the
publication of The Metamorphosis of Plants (Miller, 2009).
His concept of homology or shared ancestry of plant
structures became the central theme of plant morphology
in its classical sense. Goethe’s conjectures about homo-
logous relationships were rooted in phylogeny, the evolu-
tionary history of a species or other taxonomic group,
1
 Anatomy and Morphology of Seed Plants
especially with reference to ancestral lineage and relation-
ships among broader groups of organisms. The anatomy of
plants can be described in developmental, functional and
phylogenetic contexts, but plant morphology can embrace
all of these, even embodying anatomy itself. The philoso-
phical interpretation of plant morphology espoused by
Theophrastus persisted, notably in the works of Agnes
Arber (1950). See also: Goethe, Johann Wolfgang von
Although the morphology of some plant features can
result from their cellular structure (Dickison, 2000), plant
morphology is not necessarily related to plant anatomy.
The belief that they are intrinsically linked dates back to
Robert Hooke’s iconic depiction in Micrographia (Hooke,
1665) of a slice of bark. He commented on page 113 of this
work that the ‘‘_Cork is altogether fill’d with Air, and that
Air is perfectly enclosed in little Boxes or Cells distinct from
one another_’’ He was the first to recognize that plants are
comprised of cells, but he could not have known that the
dead, air-filled cork cells that form the outer bark are not
representative of most plant cells, which are interconnected
by pores termed plasmodesmata traversing their walls.
However, Grew (1682) thought that plant tissues resem-
bled needle work, for example, referring to ‘‘the Fibres of
the Pith running Horizontally, as do the Threds in a Piece
of Lace.’’ In fact, his drawings depict plant tissues with cell
walls consisting of interwoven, thread-like fibres, which he
considered to be the fundamental structures rather than the
cells (Arber, 1913). See also: Hooke, Robert
Grew’s curious misconception did not persist, but it was
nevertheless generally believed that tissues were con-
structed of freely permeable mesh networks of walls or
membranes through which food substances flowed. The
cells themselves were not regarded as discrete objects, and
more realistic observations of plant cells were not made
until the nineteenth century when microscope designs
improved significantly, leading to a difference of opinion
about the relationship between organisms and their cells.
The botanist Mattias Schleiden considered a plant to be
subordinate to its cells, consisting of ‘‘an aggregate of fully
individualized and isolated living beings’’, while the zool-
ogist Theodor Schwann declared ‘‘the reason for nutrition
and growth rests not with the organism as a whole but with
the individualized elementary particles, i.e., with the cells’’
(Schwann and Schleiden, 1847). Together they formalized
their interpretation of the structure of both plants and
animals in the cell theory, which Virchow (1859) popu-
larised with the epigram Omnis cellula e cellula (‘‘All cells
only arise from pre-existing cells’’). See also: Schleiden,
Matthias Jacob; Schwann, Theodor Ambrose Hubert
Wilhelm Hofmeister immediately challenged this
reductionist stance with his very first publication in 1847,
the same year that Schwann and Schleiden’s book
appeared. He demolished Schleiden’s fundamental claim
that the first cell in an embryonic plant arises from the tip of
the pollen tube by demonstrating that instead the first cell is
already present in the ovary of the flower before fertiliza-
tion occurs (Proskauer, 2008). Hofmeister argued that
plant development was organismal in nature, rather than
2 eLS & 2015, John Wiley & Sons, Ltd. www.els.net
cellular as Schleiden maintained: ‘‘The formation of new
cells in the vegetative point [apex] is accordingly a function
of the general growth, not its cause’’ (Hofmeister, 1867,
translated by Sharp, 1934). His interpretation was for-
malized in the organismal theory of plant multicellularity,
which considered plants to be living entities subdivided
secondarily into cells that remain incompletely separated
and function together.
It is notable that Hofmeister did not refer to plasmo-
desmata in any way although he must have seen these
traversing cell walls during his exhaustive studies. They
were undoubtedly observed before (Hartig discovered
sieve pores in sieve plates of the phloem in 1837), but
plasmodesmata were not discovered for what they were
until 1879 by Eduard Tangl (Köhler and Carr, 2006).
Hofmeister demonstrated the presence of connections
between cells of the endosperm of Phytelephas and Raphia
‘‘in the course of a lecture given during the winter of 1873–
4’’ (von Goebel, 1926 in Baker, 1952) but unfortunately
this event was not published. See also: History of Plant
Sciences
Hofmeister’s contributions, which include a logical,
space-filling explanation of how spiral phyllotaxy arises
(Hofmeister’s rule), and studies on tropic movements, did
not bring him the recognition he deserved. His pioneering
studies of embryogenesis directly motivated Gregor Men-
del to study hybridization in plants. His most significant
discovery was the alternation of haploid and diploid gen-
erations, which he showed was pervasive in non-seed plants
(which include the liverworts, mosses and ferns) and the
conifers. This amounted to a unifying theory of plant
evolution (Hofmeister, 1851) that prepared botanists to
accept Charles Darwin’s On the Origin of Species published
8 years later (Kaplan et al., 1996). Notably, Darwin had in
his library annotated copies of the English translation by
Currey (1862) of the second edition of Hofmeister’s (1851)
text and of his Die Lehre von der Pflanzenzelle published in
1867 (Biological Heritage Library, 2014). See also: Adap-
tation and Natural Selection: Overview; Darwin, Charles
Robert; Gametophyte and Sporophyte
The organismal theory has become largely the province
of plant morphologists, two of whom, Donald Kaplan and
Wolfgang Hagemann, presented substantial evidence to
support it, commenting that ‘‘application of cell theory to
plants is problematic at best’’ (Kaplan and Hagemann,
1991). They emphasized the independence of morphology
and anatomy, notably in the morphological similarity of
plants from very different groups ranging from simple
coenocytic (multinucleate but unicellular) green algae to
complex flowering plants. They pointed out that cells
divide differently in higher plants than in animals, which
undergo cleavage or pinching apart of the protoplast into
two cells. During mitosis of plant cells a separating layer of
Golgi vesicles arises, the cell plate, that radiates outwards
to fuse with the adjacent wall of the parent cell. The vesicles
fuse incompletely during cell wall maturation so that
plasmodesmata remain between adjacent cells, pre-
serving cytoplasmic continuity, analogous to coenocytic
 Anatomy and Morphology of Seed Plants

Apical meristem

Protoderm
Procambium
Ground meristem
Vascular
cambium Phloem
Axillary bud
Epidermis Xylem

Transverse section
of stem
Vascular cambium
Epidermis

Phloem
Cork cambium
Xylem

Xylem
Epidermis
Phloem Transition zone
found where stem
Endodermis meets root
Phloem
Epidermis
Xylem
Cortex
Root hairs Xylem
Phloem
Endodermis

Transverse section
of root
Root cap
(a) (b)
Figure 1 Median longitudinal section of a typical eudicotyledonous plant and transverse sections of specific areas.

architecture. Furthermore, Cooke and Lu (1992) meticu-
lously tested Kaplan and Hagemann’s opinion of the
possible relationship between morphology and anatomy,
and showed that the shapes of cells do not determine the
form of plant organs. Unsurprisingly, the cell theory is
fundamentally more suited to animals whereas the orga-
nismal theory is more suited to plants.
Insights into the relationship between plant anatomy
and morphology and these seemingly conflicting theories
may be drawn from a scholarly review by Niklas (2014) of
the evolutionary developmental origins of multicellularity.
He concluded that evolutionary transformation to a mul-
ticellular plant body plan can follow two routes, from
unicellular to colonial to multicellular, or a more direct
siphonous to multicellular route.
General Structure of Seed Plants
Higher plants consist of roots which provide anchorage,
water and nutrients; stems, which give support; and leaves,
which produce food for growth and development. These
organs in turn consist of tissues which can be termed
meristematic, dermal, support or vascular. Organs can be
modified for different functions, most dramatically in the
form of the flower, which is really a group of leaves that are
specialized for reproduction.
Meristems
Because of their inherent immobility, plants must adapt to
changing environmental conditions in order to survive
where they are rooted. In addition, they can replace
essential parts that become damaged or lost, and so con-
tinue the cycle of flowering and reproduction as they grow.
Plants renew themselves continually through localized
growth and development using meristems, which are
sources of undifferentiated and genetically sound cells
(Figure 1). See also: Meristems
Meristems are either determinate, producing structures
of a certain size and shape such as leaves and flowers, or
indeterminate, producing continual growth in stems and
roots, thus allowing plants to increase in length (apices) or
girth (cambia). See also: Floral Meristems
Meristems appear first during the early development of
the embryo, and some plants have a distinct protoderm, the
primary meristem that forms the epidermis when their
embryos consist of only 16 cells. The ground meristem

eLS & 2015, John Wiley & Sons, Ltd. www.els.net 3

 Anatomy and Morphology of Seed Plants
Figure 2 Transverse section of a eudicotyledonous stem.
produces cortex and pith, and the procambium produces
the primary vascular tissues, while the primary shoot and
root apices appear later. Intercalary meristems are typical
of grasses where they occur at the nodes of stems and in the
basal regions of some leaves; they cause both of these
organs to elongate. Axillary buds arise in leaf axils and
normally give rise to branches but may form various
structures including tendrils, flowers, spines and thorns.
All of these are termed primary meristems because they
establish the pattern of primary growth in plants. See also:
Apical Meristems
Stems and roots add girth through the activity of vas-
cular cambium and cork cambium which are lateral mer-
istems that arise in secondary growth, a process common in
eudicotyledons (Figure 2). Many monocotyledons lack true
secondary growth. Cambium is in essence an intercalary
meristem because it lies between its derivatives. Vascular
cambium normally creates more xylem inwards and
phloem outwards by periclinal cell divisions. Eventually,
primary intercalary meristems stop producing new cells
and disappear, whereas vascular cambium is essentially
indeterminate and its activity is more complex. It produces
more xylem than phloem and thus it expands in cir-
cumference. Therefore, it must add new cells by anticlinal
divisions to maintain the integrity of the cambial cylinder.
See also: Lateral Meristems
Meristematic activity gives some plants the potential to
achieve states of near immortality and survive for
millennia.
Dermal layers
Epidermis is the outermost cell layer of the primary plant
body. It entirely covers their leaves, stems, flowers, fruits
and roots and creates an interface between the plant
and its external environment. Although it is usually just
one cell thick (uniseriate) some plants have a multiple
4 eLS & 2015, John Wiley & Sons, Ltd. www.els.net
Cuticle
Epidermis
Collenchyma
Stomatal complex
Cortex
Bundle cap of fibers
Phloem
Vascular cambium
Metaxylem
Protoxylem
Pith (parenchyma)
(multiseriate) epidermis consisting of several layers. Epi-
dermal cells may live for many years, and are modified
in various ways. See also: Epidermis: Outer Cell Layer of
the Plant
The outer wall of the epidermis is coated with cutin, a
complex fatty substance that forms the cuticle and is
indigestible by pathogens. The cuticle is impregnated with
long-chained waxes that render it very impermeable to
water. It is transparent like the epidermal cells it covers,
allowing light to reach the much less robust photosynthetic
tissues beneath. The cuticle selectively protects the plant
from potentially mutagenic ultraviolet sunlight. See also:
Cutin and Suberin; Plant Cuticle; Plant Waxes
Stomatal pores in the epidermis allow gas exchange
between the plant and the surrounding air. Each stoma
consists of a pair of bean-shaped guard cells that can bend
apart to create a stomatal pore. Guard cells have large
nuclei and numerous chloroplasts (usually they are the only
epidermal cells to have chloroplasts). The guard cells of
most monocotyledons are dumbbell shaped. See also:
Stomata
Trichomes arise from epidermal cells that extend out-
ward and typically divide repeatedly to form a single file of
cells. Their functions include shading the plant surface
from excessive light, reducing desiccation by slowing air
movement, and protecting the plant from insects and other
herbivores. Protection may be passive, through blocking
access to the plant surface or active, through secretion of
toxins. See also: Trichomes
Stinging nettle (Urtica dioica) is protected by trichomes
with brittle silica tips that readily break off to inject irri-
tating histamines into grazing animals or passing hikers.
The most elaborate trichomes are perhaps those which
occur on the leaves of the insectivorous sundew (Drosera
sp.). These excrete sticky nectar to attract and hold insects,
bend in concert with leaf-folding to trap them, excrete
digestive enzymes and finally absorb nutrients from their

prey. See also: Coevolution: Plant–Insect; Plant Defences
against Herbivore Attack
The shoot epidermis protects the plant from desiccation,
whereas the root epidermis allows the plant to extract water
and ions from the soil aided by their root hairs, which are
delicate trichomes. The root epidermis has a thin cuticle
but its waxes have shorter chains than that of the shoot
epidermis, and therefore it is much more permeable to
water.
Plants in secondary growth can replace their epidermis
with periderm, which is a more substantial and complex
tissue. Periderm usually arises just beneath the epidermis
and consists of three layers: a middle layer of phellogen
(cork cambium, a lateral meristem) that produces an outer
layer of phellem (cork cells) and an inner layer of phello-
derm (which contributes to the cortex). Cork cells are dead
at maturity and their walls are layered with suberin and
sometimes with lignin, conferring resilience to desiccation
and insect and pathogen attack. Periderm also develops in
roots where it usually arises from the pericycle, which is just
outside the phloem. See also: Cork
Lenticels are gaps in the phellem that allow exchange of
gases between the atmosphere and the tissues that the
periderm protects so well. They arise beneath stomata in
the epidermis and persist as blisters in tree bark. Unfor-
tunately, they provide a constantly open route for infection
and herbivory.
‘Bark’ generally means all of the tissues outside the
vascular cambium (secondary phloem, phloem fibres,
cortex and the periderm). It forms many different patterns
because periderm forms at different rates at different places
on the stem. Also, the cork breaks apart as the stem
expands in circumference. New cambia arise under outer
ones, pushing the old periderms outwards. The fibrous
bark of the hydrostat-like trunk of the baobab tree
(Adansonia sp.) is unusual because it provides significant
support (Chapotin et al., 2006). See also: Bark
Protective cell layers occur in roots, stems and some-
times leaves. The most prominent such layer is the endo-
dermis that is commonly found in roots, where it forms the
innermost cell layer of the cortex and separates it from the
stele (vascular cylinder). The endodermal cell wall contains
suberin which is normally restricted to the Casparian band,
a narrow, thin strip running around the middle of each cell
in its radial and tangential walls. Mature endodermal cells
are often extensively lignified and form characteristic U-
shaped (‘phi’) thickenings that arch around each cell to
meet the Casparian bands. See also: Ecology of Water
Relations in Plants; Plant–Water Relations
The Casparian band effectively prevents uncontrolled
entry of water and ions that are absorbed from the soil and
into the stele through the apoplast (the cell walls and
intercellular spaces). It directs transport through the sym-
plast of the endodermal cells instead, which subject it to
physiological influence, allowing selective uptake by the
plant. The symplast is the cytoplasmic pathway, the net-
work of interconnected protoplasts and plasmodesmata
that traverse living cells and tissues of plants.
eLS & 2015, John Wiley & Sons, Ltd. www.els.net
Anatomy and Morphology of Seed Plants
An endodermis may occur in the stem where it usually
lacks a Casparian band or phi thickening. Stem endo-
dermis is sometimes referred to as a starch sheath, but since
the starch content which characterizes it may itself be
absent, the term is not always useful. Some aquatic plants
do have an obvious endodermis in their stems and in their
leaves as well. See also: Endodermis and Exodermis in
Roots
Exodermis (a type of hypodermis) forms the outermost
layer of the cortex of some roots and lies directly beneath
the epidermis. It can be lignified and can have a Casparian
band, and is often more obvious in monocotyledons than in
eudicotyledons. Exodermis may function to reduce water
loss from roots. Normally a feature of stems and roots, the
needle-like xerophytic leaves of conifers have a prominent
exodermis and endodermis.
Support tissues
Several tissues provide support in plants including par-
enchyma, which is composed of undifferentiated, isodia-
metric cells with thin cell walls, large vacuoles, and parietal
cytoplasm; it occupies much of the cortex and pith of stems
and roots. While parenchyma in roots often has a storage
function, in stems it can contribute support if it is turgid.
The swollen protoplast of each cell presses outwards
against its cell wall and all of the parenchyma cells press out
against their neighbours. Indeed, pressure from the par-
enchyma of the cortex and pith may contribute to stem
growth (Kutschera and Niklas, 2007). See also:
Parenchyma
The cells of collenchyma tissue have primary cell walls
that are unevenly thickened to provide support for seed-
lings and growing stems. It occurs in the cortex just beneath
the epidermis of stems in primary growth and in the mid-
ribs of eudicotyledon leaves; sometimes collenchyma is
photosynthetic. It seldom occurs in eudicotyledon roots
and is comparatively uncommon in monocotyledons.
Collenchyma provides plastic support by growing with
plants and allowing them to deform without breaking. This
plasticity results from the relatively high hemicellulose
content of its cell walls. Because collenchyma tissue rarely
contains lignin it contributes support only if the plant is not
water stressed; thus it does not prevent wilting. See also:
Collenchyma
Epidermis provides support to stems and petioles (Nik-
las, 1992), largely as a result of its thick outer wall. The
epidermis together with peripheral collenchyma is a sig-
nificant strengthening component of the shoot system.
Sclerenchyma tissue provides the most robust support.
Its cells have thick lignified secondary walls, which make it
both strong and waterproof. This tissue helps prevent
wilting, but it is metabolically expensive for the plant to
make. It is a more permanent tissue than collenchyma and
provides elastic support to maintain the established shape
of the plant. See also: Sclerenchyma
Sclerenchyma is widely distributed, occurring as a bun-
dle cap outside the phloem in vascular bundles and as a
5
 Anatomy and Morphology of Seed Plants
bundle sheath in monocotyledons. The bundle cap physi-
cally protects the inner tissues of the stem. The bundle
sheaths in grass leaves may extend to the epidermis where
they form bundle sheath extensions. Sclerenchyma cells
only mature when the cells surrounding them stop growing.
They are usually dead at maturity, although the lumens of
the cells remain connected by pits.
Sclerenchyma cells occur in two forms: fibres, which are
long (up to 55 cm in the case of fibres of hemp, Cannabis
sp.) with tapered ends, and sclereids, which are more or less
isodiametric. Brachysclereids (‘stone cells’) form clumps in
the flesh (mesocarp) of the Bartlett pear (Pyrus communis)
giving it a characteristic grittiness. They occur in a dense
layer forming the endocarp (‘shell’) of the coconut (Cocos
nucifera). Many seed coats (testas) are made of a double
layer of sclereids, especially those of legumes.
Branched astrosclereids occur in the petioles and blades
of water lily leaves (Nymphea sp.) making them leathery
and resistant to the tearing forces of waves and currents.
Some botanists distinguish between support scler-
enchyma (fibres and sclereids) and conducting scler-
enchyma (vessels and tracheids). Xylem vessels provide
both functions, but one comes at the expense of the other
because thicker walls reduce internal vessel diameter.
Vascular tissues
Higher plants have two transport systems, the xylem and
the phloem, which comprise their vascular tissue. They
have somewhat different functions but generally arise
Plastid
Parietal membranes
Filamentous strands
Sieve plate
Sieve pore lined
with callose
(a)
Figure 3 (a) Sieve element and companion cell. (b) Xylem vessels with different patterns of lignin deposition.
6 eLS & 2015, John Wiley & Sons, Ltd. www.els.net
together and are nearly always found running side by side
within all organs of the plant.
Xylem transports water and various dissolved ions from
the roots upward through the plant. Phloem transports a
solution of metabolites (mainly sugars, amino acids and
some ions) from ‘sources’ of production such as fully
expanded leaves to ‘sinks’, such as developing leaves, fruits
and roots. Both tissues contain long tubular cells (and some
other associated cells) joined end-to-end that are respon-
sible for transport. See also: Xylem: Differentiation, Water
Transport and Ecology; Xylem Structure and Function
Phloem is predominantly a living tissue consisting of
sieve tubes, companion cells (Figure 3a), phloem par-
enchyma, and phloem fibres. Each sieve tube consists of a
file of sieve elements, called sieve tube members in
angiosperms. Sieve element is the collective term for the
sieve tube members, in most angiosperms and the sieve cells
in nonangiosperms. But this discussion will not deal with
the somewhat different and much less understood sieve cell,
and the term sieve element is used here rather than sieve
tube member. See also: Phloem Structure and Function
Sieve elements are joined by thick end walls, termed sieve
plates, which are pierced by large, modified plasmo-
desmata (cytoplasmic bridges between cells) called sieve
pores. Sieve pores are lined with the complex carbohydrate
callose, which rapidly proliferates to seal the pores in
response to damage to the phloem such as that caused by
grazing animals. Sieve elements contain only a little cyto-
plasm dominated by plastids that contain starch grains or
protein and filamentous proteins. When mature, they lack
a nucleus and tonoplast but retain a plasmalemma
(b)

indicating that they are living cells. Sieve elements have
primary cell walls through which pass abundant plasmo-
desmata connecting them to small companion cells
clustered around each sieve element. Companion cells
contain large nuclei, dense cytoplasm with abundant
organelles (especially mitochondria) and many small
vacuoles. These features are related to the functional
association of companion cells and sieve elements. See also:
Companion Cells
Xylem consists of vessels, tracheids, xylem parenchyma
and fibres. The conducting cells of the xylem are the vessels
and tracheids, which are collectively called the tracheary
elements. These have secondary walls cross-linked with the
complex polymer lignin. The deposited lignin creates dif-
ferent patterns with increasing age, the youngest being
annular and the oldest being pitted (Figure 3b). As the
degree of lignification of the tracheary elements increases,
the resistance to water stress increases. Vessels consist of
cylindrical vessel elements (also termed vessel members)
joined together by large openings in their end walls forming
simple perforation plates, with ridges representing the
remnants of degraded end walls. See also: Plant Cell Wall
Biosynthesis
Although transport can occur freely through the per-
foration plates of vessels, movement also takes place lat-
erally among adjacent vessels and with adjoining xylem
parenchyma cells. This happens across areas of primary
wall where no secondary wall has been deposited. Vessels
with annular thickenings provide the greatest area for lat-
eral transport with less area becoming available as the
degree of lignification increases.
Lateral transport in pitted vessels is restricted to pits.
These occur as two main types, simple pits and bordered
pits. Simple pits are areas of bare primary wall in vessels
otherwise covered with lignin, whereas bordered pits con-
tain pressure-sensitive valves that prevent air embolisms
from interrupting flow between cells.
The other type of tracheary element is the tracheid,
which has a thin secondary wall, tapered ends, and no
perforation plates. It is connected to adjoining cells only by
pits. While tracheids are found in both gymnosperms and
angiosperms, only the most advanced gymnosperms con-
tain vessels.
The Organ System
Root
Roots anchor plants in the soil (rather than support them),
absorb water and ions from it, produce plant growth reg-
ulators, and store sugars and starch. Structurally, roots
have four external zones, beginning at the tip with the root
cap, which protects the root apical meristem and secretes
mucigel. This absorbs water from the soil to facilitate ion
diffusion into the root, fosters growth of bacteria that
release more nutrients, and lubricates the root to ease
passage through the soil. Behind the root cap is the
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Anatomy and Morphology of Seed Plants
elongation zone a few millimetres long where cells elongate
to add length to the root; it is the only part of a root system
that grows longer. This is followed by the root hair zone.
Root hairs are fragile outgrowths of epidermal cells that
live just a day or two. They greatly increase the volume of
soil that can be mined by a plant for nutrients. Last is the
lateral root zone or maturation zone characterized
internally by the presence of an endodermis enclosing the
stele (pericycle and vascular tissue) and giving rise to sec-
ondary or lateral roots (Figure 4a and b). See also: Roots
and Root Systems
Root growth is indeterminate and governed by the
moisture, fertility, composition, and homogeneity of
the soil such that growth occurs in patches making root
architecture variable. In general, a seedling produces
a primary root that grows straight down and gives rise
to secondary lateral roots. These may produce tertiary
roots, which may branch in turn, with the process con-
tinuing almost indefinitely; new roots arise endogenously
from the pericycle of the main root. Lateral roots may
exist as primordia termed seminal roots in the embryo
before germination. See also: Lateral/Secondary Roots;
Primary Root
Root modifications
Tap roots like those of carrot (Daucus carota) are modified
for storage. Another storage root is the root tuber, which is
really a swollen adventitious root (sweet potato, Ipomea
batatas).
Aerial roots occur on epiphytes including many orchids
which have a multiseriate epidermis termed velamen made
of dead cells with suberized walls. Velamen apparently
absorbs water and nutrients during wet conditions and
retains water during dry conditions. Some aerial roots are
photosynthetic.
Pneumatophores (breathing roots) occur on species of
mangrove in the genera Avicennia and Sonneratia. They
grow upwards through the substrate, usually anaerobic
mud, and possess lenticels that allow gas exchange when
exposed at low tide.
Prop roots are a type of adventitious root that provides
more transport capacity as well as extra support for plants.
In monocotyledons they arise from the lowest nodes of
stems and grow downwards through the air to the soil,
where they may contract slightly to help anchor the plant
(maize, Zea mays (Figure 5a); screw pine, Pandanus veichii).
See also: Adventitious Roots; Roots: Contribution to the
Rhizosphere
Contractile roots shrink to pull growing corms, bulbs
and rhizomes down in the stable soil environment, hiding
them from herbivores (ginger, Zingiber officinale (Figure
5b); dandelion, Taraxacum officinale).
Root nodules occur mostly on legumes and harbour
Rhizobium bacteria, which receive nutrients from the host
plant and in return fix nitrogen gas to nitrates for the host
to use. See also: Root Nodules (Legume–Rhizobium
Symbiosis)
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 Anatomy and Morphology of Seed Plants

Epidermis

Exodermis

Cortex

Endodermis

Pericycle

Xylem
Phloem

Pith

(a)

Cortex
Endodermis
Pericycle
Phloem
Metaxylem
Protoxylem

(b)
Figure 4 (a) Transverse section of a monocotyledonous root. (b) Transverse section of eudicotyledonous root.

Haustorial roots of parasitic plants are functional ana-
logues of true roots, differing from them anatomically and
morphologically, and invade the cortex and vascular tissue
of host plants, instead of the soil. Birdvine (Pthyrusa stelis)
does not invade the phloem and so only partly depends on
its host for food, while dodder (Cuscuta sp.) does invade it,
depending completely on its host for food.
Stem
The stem and its branches distribute leaves to maximize
exposure to sunlight and minimize self-shading, and flow-
ers to best attract pollinators. Branching arises from the
activity of apical and axillary buds. While branching is a
complex topic, four basic patterns can be identified.
See also: Shoots and Buds in Arabidopsis
Monopodial plants have a rhythmically active shoot
apical meristem with axillary shoots that remain secondary
and regulated by the main shoot apex; most conifers exhibit
monopodial branching (Figure 6a). See also: Phyllotaxy
In plants with sympodial branching, the shoot apex
becomes reproductive or aborts. One axillary shoot grows
upward and becomes the main stem. Its shoot apex
becomes reproductive or aborts, and so on. The char-
acteristic pagoda shape of the seaside almond (Terminalia
catappa, Figure 6b) results in part from this growth habit.
In dichasial (or dichotomous) branching, the terminal bud
splits into two buds on opposite sides of the primary or
stem. These grow at a similar rate and then branch again,
resulting in a repeatedly forked appearance. A dichot-
omously branching stem may appear to diverge equally in
two, but rarely does a stem apex split seamlessly and this is
often simply a form of sympodial growth. Many plants dis-
play such superficial dichasial branching, notably the fran-
gipani tree (Plumeria sp. Figure 6c); relatively few, including
some palms and cacti, exhibit true dichasial branching.
Although shoots usually arise from apical or axillary buds
they may appear endogenously from any organ to create
adventitious branches. Sweet potato (I. batatas, Figure 6d)
owes its scrambling habit to stems that arise from the roots
and give the plant a disorganized appearance.
Stem modifications
Stems provide other functions besides support. Some are
modified for storage. Bulbs consist of flattened, short stems

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(a)
Figure 5 (a) Prop roots on a corn plant (Zea mays). (b) Ginger plant with a rhizomatous storage stem and bulbous contractile roots.
with thick, fleshy non-photosynthetic storage leaves (onion,
Allium cepa). Corms are vertical, enlarged fleshy stem bases
with scale leaves called cataphylls (dasheen, Colocasia
esculenta, Figure 7a; Gladiolus sp.). Rhizomes are fleshy
stems, also with scale leaves (arrowroot, Maranta arundi-
nacea; ginger, Z. officinale, Figure 5b) that normally grow
horizontally but some plants, including some orchids, have
rhizomes that cling vertically to tree trunks. Stem tubers are
thick storage stems, usually horizontal (yam, Dioscorea
alata; Irish potato, Solanum tuberosum). See also: Plant
Storage Products (Carbohydrates, Oils and Proteins)
All of these stems serve as both storage organs and
organs of perennation. They enable plants to survive
periods of drought or cold by storing food reserves away
from most herbivores in a relatively stable environment.
The stem of sugar cane (Saccharum officinale, Figure 7c)
is an aerial stem tuber. Storage stems are propagative,
and may produce new shoots and roots. See also: Plant
Reproduction
Stolons (which bear photosynthetic foliage leaves) and
runners (which have only protective scale leaves) allow
plants to spread vegetatively above ground. These are
indeterminate propagative stems with long, thin internodes
that readily root at the tip to form new plants if growing
conditions are favourable (Paspalum sp., Saxifraga sp.,
Chlorophytum comosum). They have no storage or support
function.
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Anatomy and Morphology of Seed Plants
(b)
Although many stems photosynthesize to augment
their leaves, some stems are more specifically designed
to replace them. The cladode is flattened to look vaguely
leaf-like (prickly pear cactus, Opuntia sp.), but it is swollen
for water storage (succulence), too. The conifer-like nee-
dles found on the she oak (Casaurina sp.) are better
examples of photosynthetic stems. They are not enlarged
for storing or supporting anything, and they bear tiny scale
leaves.
Some stems provide support by wrapping around
another plant or other support. Many members of the
Convolvulaceae (the morning glory family) are vines, with
twining stems that wrap especially well round chain link
fences.
Leaf
Leaves are designed to provide plants with photosynthates,
but they are modified to provide protection (spines), sup-
port (tendrils), storage (onion bulb), nitrogen acquisition
(insect traps), and pollinators (flower petals). Typical
eudicotyledon leaves have a large surface area to maximize
photosynthesis and a thin profile to expose lots of chlor-
ophyll to sunlight. At the same time they must dissipate
wind forces and heat, retain water, allow gas exchange for
photosynthesis and respiration, and exclude pathogens.
See also: Evolution of Photosynthesis; Leaf and Internode;
9
 Anatomy and Morphology of Seed Plants

(a) Monopodial branching (b) Sympodial branching (Seaside almond,

(Bunya pine, Arucaria bidwillii) Terminalia catappa)

(c) Dichasial branching (Frangipani, Plumeria sp.)

(d) Adventitious branching in

sweet potato (Ipomea batatas)
Figure 6 Branching patterns. (a) Monopodial branching (Arucaria bidwilli). (b) Sympodial branching (Seaside almond, Terminalia catappa). (c) Dichasial
branching (Plumeria sp.), details. (d) Adventitious branching in sweet potato (Ipomea batatas).

(a)
Figure 7 (a) Dasheen (Colocasia esculenta) plant with a basal corm bearing cormels. (b) Stem of a noble cane variety of sugar cane, comprised of
internodes forming an aerial stem tuber.
Photosynthesis; Photosynthesis: Ecology; Photosynthesis:
Light Reactions; Photosynthesis: The Calvin Cycle
By extending the leaf blade (lamina) away from the stem
by a flexible leaf stalk (petiole) shading from the stem is
reduced and the leaf can flutter. Fluttering promotes
cooling, decreases insect and pathogen attack, dissipates
wind forces to reduce damage, and increases carbon
dioxide absorption. Monocotyledon leaves are usually
narrow and belt-shaped, with a sheathing base, and in
addition they move easily in the wind. See also: Energy,
Radiation and Temperature Regulation in Plants
The adaxial epidermis of most eudicotyledon leaves has
a thicker cuticle and fewer stomata than the abaxial epi-
dermis. Palisade mesophyll cells, the major sites of pho-
tosynthesis, are arranged within the leaf above spongy
mesophyll cells, where gas exchange occurs (Figure 8a). The
adaxial and abaxial surfaces of many monocotyledon
leaves are more or less the same and their internal tissues
are more homogeneous than stratified (Figure 8b).
Leaf modifications
Mature spines consist entirely of compact bundles of
sclerenchyma fibres. They can, if densely packed, shield the
plant against intense sunlight and the drying effect of wind.
Spines are usually but not always classed as modified leaves
rather than stems or other structures, but the reason is not
always obvious. For example, the fearsome pointed
structures on the rough lemon tree (Citrus jambhiri) are
subtended by leaves (Figure 9a), suggesting they are stem
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Anatomy and Morphology of Seed Plants
Adventitious
roots
Axillary bud
Leaf scar
(b)
spines, or simply thorns in common usage. But, in this
plant they arise from prophylls (the first leaves on axillary
shoots) so they are leaf spines after all; either way they both
provide protection from herbivores.
Emergences, which are neither modified roots or stems,
nor leaves, often form pointed sclerophyllous structures
called prickles; these can be ephemeral and are analogous
to spines and thorns. Emergences appear in unexpected
places including leaf midribs or laminas, fruits and stems,
and arise mainly from epidermal tissue (Solanum sp.,
Figure 9b; Soursop, Annona muricata, Figure 9c). Rose bush
thorns are really prickles, in spite of their robust appear-
ance (Figure 9d).
Stipules are typically inconspicuous, paired structures
arising from the base of a petiole, but are modified to
form spines, which become hollow to allow habitation by
ants on some species of Acacia in return for added pro-
tection. Bud scales are sessile (lacking a petiole), non-
photosynthetic leaves (sometimes modified bracts) that
protect dormant apical or axillary buds from herbivores
and drying winds (Figure 9e). They are common on tem-
perate trees and in the tropics they occur on the para rubber
tree (Hevea brasiliensis), magnolia (Magnolia sp.) and
mahogany (Swietenia macrophylla). Bud scales may pro-
duce cork cambium that makes a thin bark, the only type of
leaf to do so (Mauseth 1988). They are often called cata-
phylls, which are typically protective but deciduous, and in
some plants have a storage function; cotyledons (the
embryonic leaves in eudicotyledon seeds) are sometimes
referred to as a type of cataphyll.
11
 Anatomy and Morphology of Seed Plants

Cuticle
Upper epidermis

Collenchyma Palisade mesophyll

Spongy mesophyll

Xylem

Stomatal complex

Cambium

Phloem
Collenchyma

Lower epidermis

(a)

Upper epidermis Guard cells

Cuticle
Stomatal pore

Xylem
Phloem

Vascular bundle
(b) Bundle sheath extension
Figure 8 (a) Transverse section of the midrib of a eudicotyledonous leaf. (b) Transverse section of a monocotyledonous leaf.

Tendrils lack a lamina (leaf blade) like spines, but
photosynthesize, never stop growing, and help to support
plants (Figure 9f). They are touch-sensitive not light-
sensitive and coil around things especially stems of other
plants. Coiling results from growth occurring faster on
the outer side of the tendril away from the inner side in
contact with the support. Tendrils occur on many plants
including members of the Cucurbitaceae (cucumber,
melon), Passifloraceae (passion fruit, Passiflora edulis),
and the Convolvulaceae (morning glory family). Both
spines and tendrils can be modified stems rather than
modified leaves. Indeed, spines arise from adventitious
roots on a species of mangrove and some tendrils even arise
from inflorescences.
Sclerophyllous leaves are xerophytic and designed to
withstand a desert-like environment. Like spines they have
abundant fibres, but unlike them sclerophyllous leaves
retain photosynthetic ability. Because of their significant
lignin content, sclerophyllous leaves are expensive for the
plant to make and so are retained for a long time. Ordinary
leaves by contrast are cheap and mass-produced by many
plants and are expendable. Sclerophyllous leaves are
especially common among the monocotyledons, for
example Agave sisalana (Figure 9g). The fibres that make
leaves of this plant so durable are used to make sisal rope.
See also: Hot Deserts
Succulent leaves are also xerophytic and conse-
quently have few air spaces and thick cuticles. Typically
their mesophyll is isolated from the leaf surface by
cells with large vacuoles, which filter heat from intense
sunlight. Succulent leaves often contain mucilage that
binds water so it can be stored in the leaf. Their vascular
tissue is said to be conserved. There is not much of it
because little transport occurs in these leaves. Aloe vera is a

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 Anatomy and Morphology of Seed Plants

(a) (b) (c)

(d) (e) (f)

(g) (h) (i)

Figure 9 (a) Leaf spines on a rough lemon tree (Citrus jambhiri). (b) Emergences on leaf of Solanum sp. (c) Emergences on fruit of soursop (Annona
muricata). (d) Prickles on stem of a rose bush (Rosa sp.). (e) Bud scales on a chestnut branch (Castanea sp.). (f) Leaf tendrils on Passion fruit plant (Passiflora
edulis). (g) Sclerophyllous leaves of Agave sisalana. (h) Succulent leaves of Aloe vera. (i) Pitcher of a Nepenthes villosa plant.

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 Anatomy and Morphology of Seed Plants
good example (Figure 9h). Both succulent and scler-
ophyllous leaves have a small surface area to volume ratio
to reduce transpiration losses.
Trap leaves occur on insectivorous plants, which to-
lerate nitrogen poor soil by trapping and digesting
insects. The highly modified prey-trapping leaves of pitcher
plants (species of Nepenthes, Darlingtonia, Cephalotus
and Sarracenia), are typically red brown like carrion
and are lined with glands that release digestive enzymes.
They may have lids to exclude rain, downward-pointing
trichomes, and abundant flaky wax to prevent the escape
of insects. In species of Nepenthes (Figure 9i) the petiole
of the trap is elongate, wide and flat, like a normal
lamina, forming a phyllode. This structure compensates
for the reduced photosynthetic role of the pitcher.
Other kinds of trap leaves occur on the Venus flytrap
(Dionaea sp.), sundew (Drosera sp.), and bladderwort
(Utricularia sp.).
Trees and wood
A stump of a tree trunk usually displays a distinct outer
ring of lighter sapwood containing functional secondary
xylem, surrounding a large area of darker heartwood, the
nonfunctional secondary xylem. The dark colour results
from accumulated tannins and phenolic compounds. A
thin band of secondary phloem produced by an imper-
ceptible layer of vascular cambium circles the sapwood,
and this in turn is surrounded by a narrow layer of cortex
and then the periderm.
Concentric rings appear in the wood of the stump if there
is annual variation in the growth rate caused by seasonal
differences in climate large enough to cause the vascular
cambium to produce secondary xylem at different rates.
Growth rings can even be apparent in the secondary phloem
and in the periderm. With some exceptions growth rings
provide an accurate way to age trees while trees without
growth rings, which can occur in tropical rainforests close to
the Equator, are essentially impossible to age.
In general, ‘hardwood trees’ (angiosperms such as oak
(Quercus sp.), ash (Fraxinus sp.), teak (Tectona grandis),
and mahogany (Sweitenia sp.) are rich in fibres, making
them denser, heavier and stronger than ‘softwood trees’
(gymnosperms such as Caribbean pine, Pinus caribaea and
white spruce, Picea glauca), which lack fibres and are rich
in tracheids.
Reproductive structures
In the gymnosperms (meaning ‘naked seeds’ in Greek)
pollen is produced by male cones and ovules are produced
by female cones. In the conifers, the major group of gym-
nosperms, pollen is usually carried passively by the wind
from pollen cones to ovulate cones to allow pollination.
The seeds of angiosperms (derived from ‘vessel’ and ‘seed’)
are protected by the wall of the ovary in which they form.
Angiosperm flowers are designed to attract pollinators
14 eLS & 2015, John Wiley & Sons, Ltd. www.els.net
(mostly insects) with which they have coevolved for many
millions of years.
Flower
A flower is a shoot system consisting of a determinate axis
with laterally borne, concentric rings of four kinds of leaves
designed for sexual reproduction: sepals, petals, stamens
and carpels (Figure 10). Green sepals closely resemble leaves
while petals and coloured (petaloid) sepals contain chro-
moplasts instead of chloroplasts and have a poorly devel-
oped vascular system, no palisade mesophyll and little
sclerenchyma. Most stamens and carpels do not resemble
leaves but in the more primitive flowers (species of Drimys,
Magnolia and Michelia) they may be wide and flattened
which points to their leafy origin. See also: Arabidopsis:
Flower Development and Patterning; Flowers
Petals and sepals are not necessary for reproduction and
are termed accessory parts; hence stamens and carpels are
essential parts. The sepals together form the calyx of the
flower and the petals together make up the corolla, with both
whorls together comprising the perianth. The stamens
(comprised of filament and anther) are the male parts
(androecium) and the carpels (comprised of ovary, style and
stigma) are the female parts (gynoecium). See also: Carpels;
Petals; Sepals; Stamen and Pollen Development; Stamens
Flowers arise in the axils of leaves which are usually
small and are known as bracts. Some flowers have con-
spicuously coloured bracts that supplement the petals or
even replace them (Poinsettia (Euphorbia pulcherrima),
Bougainvillea sp.). A stem bearing a single flower is a
pedicel and the point of attachment of the flower to the
pedicel is the receptacle, while the stem of an inflorescence,
a branched system of flowers, is a peduncle.
Floral modifications
Loss of parts
A complete flower possesses all four whorls of floral leaves;
an incomplete flower lacks one or more whorls. Perfect or
hermaphrodite flowers have both carpels and stamens but
may lack sepals, petals, or both. An imperfect flower lacks
either carpels or stamens. A staminate (male) flower has
only stamens, and no carpels; a carpellate (female) flower
has only carpels.
The ear of a maize plant (Zea mays) is a carpellate
imperfect inflorescence. Its silks are greatly elongated
styles, with sticky stigmatic surfaces for catching airborne
pollen. The tassel is a staminate imperfect inflorescence.
Maize is monoecious because it bears both male and female
flowers on the same plant; many cucurbits are monoecious,
too. Papaya (Carica papaya) is dioecious, with male and
female flowers borne on different plants. See also: Zea
Mays (Maize, Corn)
Fusion of parts
Within whorls, if the sepals are fused to form a tube, the
flower is gamosepalous (Hibiscus sp., Figure 10). If the

Staminal tube
Style
Staminal tube
Bracteole
Nectary
(a)
Figure 10 Half flower drawing and floral diagram of Hibiscus (Hibiscus rosa-sinensis)
petals are fused it is gamopetalous (yellow bell, Allamanda
cathartica) but if the petals are not fused it is polypetalous.
The usual fusion between whorls is between stamens and
sepals; such stamens are adnate to the petals, producing an
epipetalous flower.
If the filaments of the stamens are fused into a tube they
are connate and the androecium is aldelphous. If the car-
pels are fused the gynoecium is syncarpous but if they are
free it is apocarpous; fusion of gynoecium and androecium
produces a gynostegium. The flowers of many legumes (of
the Papilionoideae) and all Compositae have their stamens
fused to form a tube.
Ovary structure
Ovules arise from placentas that appear as swellings inside
the ovary (Figure 11). A simple ovary (with one carpel) has
marginal placentation with ovules arising along the junc-
tion of the two margins of the carpel. If there is more than
one carpel the placentation becomes parietal but if the
ovules are attached to the central axis formed by the carpels
it is axile. In central placentation there is only one locule
and the ovules are borne on a central axis; in free central
placentation the axis is incomplete. In basal placentation
there is also one locule but the ovules are attached to the
base of the ovary. See also: Gametogenesis
Ovary position
A flower that is hypogynous (meaning that the other flower
parts are ‘below the gynoecium’) has a convex receptacle
(Figure 11c) and its ovary is superior to the rest of the flower.
A perigynous (‘around the gynoecium’) flower has a
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Anatomy and Morphology of Seed Plants
(b)
concave receptacle and the ovary is below the other floral
parts making it half superior. In epigynous (‘above the
gynoecium’) flowers the ovary is inferior and embedded in
the receptacle (Figure 11d; Table 1, part C).
Aestivation
Flowers show variation in the horizontal arrangement of
their calyx and corolla (aestivation). In a valvate flower the
petals or sepals meet without overlapping. In a contorted
or regular flower the petals or sepals all overlap in the same
direction. In an imbricate or irregular flower the petals or
sepals overlap in both directions so that one sepal or petal is
wholly inside the ring and at least one other is wholly
outside the ring.
Floral shape
Actinomorphic flowers exhibit radial (multilateral) sym-
metry. All the floral parts in each whorl are alike in size and
shape, and any longitudinal cut across the centre creates
two mirror image halves (Hibiscus sp., Cucurbita sp.).
Zygomorphic flowers have bilateral symmetry and only
one specific longitudinal cut across them will produce
mirror image halves. This symmetry results from differ-
ences in the size, shape, or loss of some of the parts
(Orchidaceae, Leguminoseae). In asymmetric flowers there
is no symmetry to the arrangement of the parts (Canna sp.).
Fruits
Fruits differ botanically from what may be construed from
the popular notion of ‘fruits and vegetables’. The green
bean (Phaseolus vulgaris) is a fruit not a ‘vegetable’, the
15
 Anatomy and Morphology of Seed Plants

Midrib vascular
bundle of carpel

Septum

Placental vascular bundle

(b)
(a)

(c) Hypogynous Perigynous Epigynous (d) Unilocular tricarpellary ovary

Petals

Anthers
Stigma
Style

Ovule Ovary
Sepals
(e)

Calyx lobe Petal bundle

Sepal bundle
Dorsal bundle
Stamens and styles of carpel
Endocarp

Outer limit
of carpel
(core line)
(f)
Seed
Floral tube
Figure 11 Ovary structure and position in flowers. (a) Transverse section of a trilocular tricarpellary ovary. (b) Unilocular ovary. (c) Ovary position. (d)
Uniocular tricarpellary ovary. (e) Apple flower and (f) fruit structure.
 Anatomy and Morphology of Seed Plants

Table 1 A classification of fruits

A. Simple fruits: Fruit derived from a solitary carpel in a single flower.
Dry fruits: Having a mesocarp that is definitely dry at maturity
(i) Indehiscent fruits
(a) Arising from one carpel
Achene – small, with seed single attached at one point to a thin pericarp (Buttercup, Ranunculus sp.).
Cypsela – similar to achene, but arises from an inferior ovary (characteristic of the Compositae),
making it a false fruit.
Caryopsis or grain – similar to an achene, but pericarp and testa of single seed are fused (all grasses).
Samara – Testa modified into a wing-like structure, may be one-seeded (elm, Ulmus sp.) or two
(maple, Acer sp.).
(b) Arising from a compound gynoecium with several carpels
Nut – Ovary has several carpels, all but one atrophy. One-seeded, with woody pericarp of sclereids
(hazelnut, Corylus sp.).
(ii) Dehiscent fruits
(a) Arising from one carpel
Follicle – pod-like fruit, splits open on one side (milkweed, Asclepias sp.; cotton, Gossipium sp.).
Legume – breaks open on both sides (all beans and peas of the Leguminosae).
(b) Arising from a compound gynoecium with many carpels
Silicule (or silicula) – two sides split, seeds remain attached to a false central partition (replum). It is
as broad as, or broader than, it is long. The silique (siliqua) is a silicule type of fruit that is longer than
broad. Both are characteristic of the Brassicaceae, the mustard family.
Capsule – carpels split apart in different ways (mahogany, Swietinia sp., iris (Iris sp.), lily
(Liliaceace), common poppy (Papaver rhoeas); akee (Blighia sapida) may be considered a fleshy
capsule).
(iii) Schizocarpic fruits: Arising from a compound ovary; the locules split into separate (usually dry) fruits
Schizocarpic mericarp – twin mericarpic fruits joined at one point (Apiaceae, parsley family).
Fleshy fruits: Having a mesocarp that is soft at maturity
Berry – (true berry or bacca) – endocarp, mesocarp, and exocarp all soft and easily distinguishable at
maturity (blueberry, Vacinnium vacillans; tomato, Solanum lycopersicum).
Drupe – similar to a berry, but endocarp is woody (mango, Mangifera indica; peach, Prunus persica).
Pome – (also classed as a false fruit) – similar to drupe, except with a papery endocarp (apple, Pyrus
malus).
Pepo – similar to berry, but with a thick, tough exocarp. May also be classed as false fruits
(watermelon (Citrullus lanatus; pumpkin, squash, Cucurbita sp., many other Cucurbitaceae).
Hesperidium – leathery exocarp with oil glands (most citrus).
B. Compound fruits (Syncarps): Fleshy fruits composed either of the fruits of several flowers, or of several carpels of a single flower.
Aggregate fruits. Formed from one flower with many ovaries maturing together, some types fusing with receptacle.
Etaerio – consists of an aggregate of achenes, berries, or drupes (Red raspberry, Rubus idaeus;
strawberry (Fragaria sp.) is also classed as a pseudocarp, a kind of false fruit).
Multiple fruits. Developing from the ovaries of several flowers of an inflorescence which mature together.
(a) Fleshy multiple fruits (may also be classed as accessory fruits)
Sorosis – fruits arise on a common axis and derived from the ovaries of several flowers that are
usually coalesced (mulberry, Morus sp.; pineapple, Ananas comosus; breadnut, Artocarpus atilis).
Syconium – a syncarp with achenes attached to the inside of an infolded receptacle (Ficus sp.).
(b) Dry multiple fruits
Strobilus – multiple fruit of achenes including bracts (hop, Humulus lupulus).
C. Accessory/false fruits: The ovary wall can be augmented by various tissues that protect the seeds, including the receptacle of
soursop, the perianth of breadnut and the scaly bracts of pineapple. Fruits that develop from inferior
ovaries are false fruits. In an apple the mesocarp and receptacle merge together with no exocarp
discernible, because the carpels become fused to the accessory tissue during fruit ontogeny. In the flesh
of the apple two sets of vascular bundles, one set previously leading to sepals, the other set to petals
before the apple formed, remain visible. The true fruits of a strawberry are the tiny ‘seeds’, which are
really achenes, on the surface of the swollen red receptacle.

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 Anatomy and Morphology of Seed Plants
coconut is not a nut (although both coconuts and nuts are
fruits) and the banana (Musa acuminata) is an example of a
berry. See also: Fruit Structure and Diversity; Seeds
A true fruit is the mature ovary of an angiosperm. The
maturation of the ovule(s) to form the seed(s) within the
ovary is normally accompanied by a thickening of the
ovary wall to form the three layers comprising the pericarp.
The outer layer (exocarp) may simply be a layer of epi-
dermis as in grape (Vitis vinifera) and the middle layer
(mesocarp) is commonly soft like the flesh of the mango
(Mangifera indica). The inner layer (endocarp) can vary
from gelatinous (tomato, Solanum lycopersicum) to stony
(the ‘pit’ or ‘stone’ of the peach (Prunus persica)). See also:
Angiosperms
In general, fruits may be classified according to four
criteria:
. The number of flowers involved in their formation.
. The number of ovaries.
. The degree of hardness of the mesocarp – dry and hard
or soft and fleshy.
. The ability of the fruit to dehisce (split open when
mature) or not.
See Table 1 for classification of fruits.
Seed
The seed is the mature, fertilized ovule of a flower. It con-
tains an embryonic plant together with food reserves in a
resting state before initiation of germination. An envel-
oping seed coat (testa) derived from the integument cell
layer(s) of the ovary within the nucellus (megasporangium)
provides protection. A pore remains at the apex (the
micropyle) through which the pollen tube reaches the
ovule. Beside the micropyle is the hilum which is a scar
created when the seed breaks free of the seed stalk (funi-
culus). Seeds that store food are albuminous seeds; those
that do not are exalbuminous seeds. Food stores (carbo-
hydrates, proteins and lipids) can be endospermous
(derived from the large central cell of the nucellus), peri-
spermous (proliferated nucellus tissue), or both. See also:
Seed Germination and Reserve Mobilization; Seeds
Embryos of many eudicotyledon seeds absorb their food
stores well before germination but with notable exceptions
including castor bean (Ricinus communis) which has oily
seeds. However, many monocotyledon seeds retain a pro-
minent endosperm like the grasses wheat, oats, barley, rice
and corn. The grass embryo has a large scutellum that
excretes enzymes to digest the endosperm to provide the
embryo with food.
Seed dispersal
Reproductive success is very much enhanced if seeds are
distributed to new habitats. To this end seeds and their
associated fruits show a variety of adaptations to take
advantage of various vectors of dispersal (Figure 12).
See also: Dispersal: Biogeography
18 eLS & 2015, John Wiley & Sons, Ltd. www.els.net
Animals
Many berries have attractive and edible pericarps and
small, slippery seeds protected by hard woody testas
well suited to being consumed and surviving digestion.
Testas and pericarps may have spines, burrs, or other
emergences designed to engage animal fur or hikers’ socks
as vectors of dispersal (railway daisy, Bidens pilosa, Figure
12a).While the testa of a drupe is thin its woody endocarp
protects the seed making it suitable for ingestion. The
pericarps of true nuts are entirely woody and indehiscent,
making them favourites of rodents for their food stores
(hazelnut, Corylus sp., Figure 12b). Arils are normally fle-
shy, edible accessory outgrowths, typically of the funicu-
lus, and aid dispersal by animals of seeds of the akee tree
(Blighia sapida), and the monkey pot tree (Lecythis ollaria).
They also occur on seeds of better-known fruits like
tomatoes.
Water
The coconut fruit is buoyant and seaworthy because of the
air held in its hollow seed protected by a tough endocarp
made of sclereids and by the air in its fibrous mesocarp,
which in turn is protected by a tough, waxy exocarp.
Growth of the beached, germinating seed is aided by
rainwater retained by the mesocarp. Seeds of the red
mangrove (Rhizophorus mangle) are viviparous (with no
dormant period). They produce leaves and a prominent
torpedo-shaped root while still attached to the parent tree;
seedlings eventually break free to be carried away by the
tide.
Wind
Seeds dispersed by the wind must be small and light and
produced in great numbers to reach suitable habitats
(mustard seed, Figure 12c). To this end some orchids pro-
duce dust-like seeds by the million; their mass is measured
in nanograms. Seeds of milkweed (Asclepias sp.) have
plumes that are really extended dead trichomes which
equip them for dispersal. The dandelion seed is really a
cypsela fruit and its plume is a pappus, a modified calyx.
The fruit of mahogany (Swietenia sp.) is a type of capsule
that splits open when mature to release seeds which bear
winged testas so they can rotate and float away from the
parent tree (Figure 12d).
Self-dispersal
The pericarps of some legumes split open explosively to
release their seeds (purple vetch, Vicia benghalensis; Scotch
broom, Cytisus scoparius). This phenomenon results from
the sudden release of tension created in the maturing
pericarp by layers of fibres in the pericarp shrinking in
opposite directions when the carpels are broken apart.
Hydrostatic pressure built up in the fruits of dwarf mis-
tletoe (Arceuthobium pusillum) can expel seeds as far as
15 m from the parent plant.
 Anatomy and Morphology of Seed Plants

(a) (b)

(c) (d)

Figure 12 (a) Railway daisy (Bidens pilosa). (b) True nut (Hazelnut, Corylus sp.). (c) Silique (Mustard family). (d) Capsule with winged seeds (Mahogany,
Swietenia sp.). Illustrations are not drawn to scale.

Acknowledgement Baker JR (1952) The cell-theory: a restatement, history, and cri-

I would like to thank Lisa Teelucksingh for supplying the
original artwork for this article.
References
Arber A (1913) Nehemiah Grew 1641–1712. In: FW Oliver (ed.)
Makers of British Botany, pp. 44–66. Cambridge: Cambridge
University Press.
Arber A (1950) The Natural Philosophy of Plant Form. Cam-
bridge: Cambridge University Press.
tique. Part III. The cell as a morphological unit. Quarterly
Journal of Microscopical Science 93(2): 157–190.
Biological Heritage Library (2014) Charles Darwin’s Library.
http://www.biodiversitylibrary.org/collection/darwinlibrary
Chapotin SM, Razanameharizaka JH and Holbrook NM (2006)
A biomechanical perspective on the role of large stem volume
and high water content in baobab trees (Adansonia spp; Bom-
bacaceae). American Journal of Botany 93(9): 1251–1264.
Cooke TJ and Lu B (1992) The independence of cell shape and
overall form in multicellular algae and land plants: cells do not
act as building blocks for constructing plant organs. Interna-
tional Journal of Plant Sciences 153: S7–S27.

eLS & 2015, John Wiley & Sons, Ltd. www.els.net 19

 Anatomy and Morphology of Seed Plants
Currey (1862) Germination, Development, and Fructification of the
Higher Cryptogramia, and on the Fructification of the Coniferae.
London: Robert Hardwicke for the Ray Society. (Original
edition published by Hofmeister (1851).
Dickison WC (2000) Integrative Plant Anatomy. New York, NY:
Academic Press.
Evert RF and Eichhorn SE (2013) Raven Biology of Plants, 8th
edn. New York, NY: WH Freeman.
von Goebel K (1926) Wilhelm Hofmeister: The Work and Life of a
Nineteenth-Century Botanist. London: The Ray Society.
Grew N (1672) The Anatomy of Vegetables Begun. London:
Spencer Hickman.
Grew N (1682) The Anatomy of Plants. London: W Rawlins.
Hartig T (1837) Vergleichende Untersuchungen über die Orga-
nisation des Stammes der Einheimischen Waldbäume. Jahres-
berichte Über die Fortschritte der Forstwissenschaft und
Forstlichen Naturkunde 1: 125–168.
Hofmeister WFB (1847) Untersuchungen des Vorganges bei der
Befruchtung der Oenothereen. Botanische Zeitung 5: 785–792.
Hofmeister WFB (1851) Vergleichende Untersuchungen der Kei-
mung, Entfaltung und Fruchtbildung höherer Kryptogamen
(Moose, Farn, Equisetaceen, Rhizocarpeen und Lycopodiaceen)
und der Samenbildung der Coniferen. Leipzig: W Engelmann.
Hofmeister WFB (1867) Die Lehre von der Pflanzenzelle. In:
WFB Hofmeister (ed.) Handbuch der Physiologischen Botanik
I-1, pp. 1–664. Leipzig: W Engelmann.
Hooke R (1665) Micrographia. London: John Martyn and James
Allestrey.
Hort A (1896) Theophrastus, Enquiry into Plants. London: William
Heinemann.
Kaplan DR, Cooke TJ and Todd J (1996) The genius of Wilhelm
Hofmeister: the origin of causal-analytical research in plant
development. American Journal of Botany 83(12): 1647–1660.
Kaplan DR and Hagemann W (1991) The relationship of cell and
organism in vascular plants. BioScience 41: 693–703.
Köhler P and Carr DJ (2006) A somewhat obscure discoverer of
plasmodesmata: Eduard Tangl (1848–1905). In: M. Kokowski
(ed.) Proceedings of the 2nd ICESHS, Cracow, Poland.
Kutschera U and Niklas KJ (2007) The epidermal-growth-control
theory of stem elongation: an old and a new perspective. Journal
of Plant Physiology 164(11): 1395–1409.
20 eLS & 2015, John Wiley & Sons, Ltd. www.els.net
View publication stats
Malpighii M (1675) Anatome Plantarum. Londini: Johannis
Martyn.
Mauseth JD (1988) Plant Anatomy. Menlo Park: Benjamin/
Cummings.
Miller G (2009). The Metamorphosis of Plants. Cambridge: MIT
Press.
Niklas KJ (1992) Plant Biomechanics: An Engineering Approach to
Plant Form and Function. Chicago: The University of Chicago
Press.
Niklas KJ (2014) The evolutionary-developmental origins of
multicellularity. American Journal of Botany 101(1): 6–25.
Proskauer J (2008) Hofmeister, Wilhelm Friedrich Benedikt.
Complete Dictionary of Scientific Biography. http://www.ency-
clopedia.com/topic/Wilhelm_Hofmeister.aspx
Schwann T and Schleiden MJ (1847) Microscopical Researches
into the Accordance in the Structure. Growth of Animals and
Plants. (Trans H. Smith). London: The Sydenham Society.
(Original works published separately 1839,1838)
Sharp LW (1934) Introduction to Cytology, 3rd edn. London:
McGraw-Hill.
Virchow R (1859) Vorlesungen über Cellularpathologie in ihrer
Begründung auf physiologischer und pathologischer Gewebe-
lehre. Berlin: Verlag von August Hirschwald.
Further Reading
Beck CB (2010) An Introduction to Plant Structure and Develop-
ment: Plant Anatomy for the Twenty-First Century, 2nd edn.
Cambridge: Cambridge University Press.
Bell AD (2008) Plant Form: An Illustrated Guide to Flowering
Plant Morphology. London: Timber Press.
Evert RF (2006) Esau’s Plant Anatomy. Meristems, Cells, and
Tissues of the Plant Body: Their Structure, Function, and
Development, 3rd edn. London: John Wiley & Sons.
Heywood VH, Brummitt RK, Culham A and Seberg O (2007)
Flowering Plant Families of the World. Kew: Royal Botanic
Gardens.
Niklas KJ (1997) The Evolutionary Biology of Plants. Chicago:
University of Chicago Press.