Department of Botany and microbiology

Course specifications
Menoufia University Faculty of Science
Programme on which the course is given B.Sc. Botany, Botany & Chemistry
Major or minor element of programmes Major
Department offering the programme Botany
Department offering the course Botany

Academic year / Level First (1*)

Basic Information
Title: Plant anatomy and Morphology Code: B111
Credit Hours: 3 h Lecture: 2h Practical: 2h

Overall aims of course

After studying this course, the student should be able to define the plant
shapes and its external structure, the different plant seeds, and their
differences in both structure and function. The course aims also at
development the student’s knowledge about the differences of internal
structure between the dicotyledonous and monocotyledonous plants (roots,
stems and leaves).
The students should be also able to:
- Compare and contrast between the different plants and plant seeds based on
the macro morphology
- Summarize the functions of each tissue in the plant body
- Compare between the dicotyledonous and monocotyledonous plants from
their internal structure.
- Identify the different taxa of tracheophyta by using their external and
internal structure and use this as a tool in plant identification.

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Contents

No. Topics page

Introduction and characteristics of higher plants
1 3
Plant seed structure and seed types
2 5
Germination
3 8
Root system
4 26
Shoot system
5 34
Leaves
6 46
The plant cell
8 63
Formation of cell wall
9 66
Cytoplasm and membranous system in the cell
10 73
Tissues and tissue systems
11 80
Permanent tissue (epidermal tissue, Fundamental tissue)
12 85
Vascular or conducting tissue
13 102
Internal structure of stems
14 119
Internal structure of roots
15 129
Internal structure of leaves
16 139

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 Part I: Plant Morphology

Introduction

The branch of biology that deals with the structure of animals and
plants is Morphology

While, Plant morphology (phytomorphology) is the general term

for the study of physical form and external structure of plants. The
study of plant morphology is useful in the identification of plants.

Plant morphology represents a study of the development, form, and

structure of plants, and, by implication, an attempt to interpret
these on the basis of similarity of plan and origin.

There are major areas of investigation in plant morphology, and

each overlaps with another field of the biological sciences.

First, morphology is comparative, in which the morphologists

examine structures in many different plants of the same or different
species, then draws comparisons and formulates ideas about
similarities.

Secondly, plant morphology studies both the vegetative (somatic)

structures of plants, as well as the reproductive structures. The

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vegetative structures of vascular plants include the study of the
shoot system, ( stems and leaves), as well as the root system. The
reproductive organs are more varied, and are usually specific to a
particular group of plants, such as flowers and seeds.

The detailed study of reproductive structures in plants led to the

discovery of the alternation of generations found in all plants and
most algae. The study of plant morphology overlaps with the study
of biodiversity and plant systematic.

Thirdly, plant morphology studies plant structure at a range of

scales. At the largest scale is the study of plant growth habit, the
overall architecture of a plant. The pattern of branching in a tree
varies from species to species, as will the appearance of a plant as
a tree, herb, or grass.

Fourthly, plant morphology studies the pattern of development,

the process by which structures originate and mature as a plant
grows.

Plants constantly produce new tissues and structures throughout

their life. A living plant always has embryonic (meristematic)
tissues. The way in which new structures mature as they are

4
produced may be affected by the stage in the plants life when they
begin to develop, as well as by the environment to which the
structures are exposed.

The characters used in descriptions or for identification are called

diagnostic or key characters, which can be either qualitative or
quantitative.

1. Quantitative characters are morphological features that can be

counted or measured for example a plant species has flower petals
5-8 mm wide.
2. Qualitative characters are morphological features such as leaf
shape, flower color or hairiness. Both kinds of characters can be
very useful for the identification of plants.

A vascular plant begins from one celled zygote, formed after

fertilization of an egg cell by a sperm cell.

From that point, it begins to divide to form an embryo through the

process of embryogenesis. Through this happens, the resulting
cells will organize so that one end becomes the first root, while
the other end forms the tip of the shoot.

5
In seed plants, the embryo will develop one or more seed leaves
(cotyledons).

By the end of embryogenesis, the baby plant will have all the parts
necessary to begin in its life enclosed in solidified integuments
(seed coat) which is called in general the seed.

Seeds and seed functions

A seed is an embryonic plant enclosed in a protective outer

covering (solidified ovule integuments). The formation of the
seeds is part of the process of reproduction in seed plants, the
spermatophytes, including the gymnosperm and angiosperm plants.

Seeds are the product of the fertilized ripened ovule, after

fertilization by male gametes from pollen grain and some growth
within the mother plant. The embryo is developed from the zygote
and the seed coat from the integument (s) of the ovule.

Seeds with testa protect and nourish the embryo or baby plant.
Seeds usually nourish and give a seedling fast because of the larger
food reserves in the seed. Different plants have evolved many ways
to disperse their population through their seeds.

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Seed dispersal is often attributed mainly to fruits, however many
seeds aid in their own dispersal

An important function of most seeds is delaying germination

(dormancy) to allow time for dispersal and to prevent all seeds
from germinating at once when conditions are favorable.

Economic importance of seeds

Many seeds are edible (used in nutrition). Seeds also provide most
used oils. Many important nonfood oils are extracted from seeds,
many beverages, spices, and some important food additives.

Seeds are used to propagate many crops such as legumes, forest

trees, turf grasses ‫العشب‬, and pasture grasses ‫المرعى‬. Some seeds are
poisonous. The most important clothing fiber in the world grows
attached to cotton seed.

Seeds are the source of some medicines such that of castor oil, tea
tree oil.

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Germination

Germination is usually the emergence of radicle and plumule as a

result of the growth of a baby plant (Embryo) contained within a
seed; it results in the formation of the seedling.

Seeds remain dormant or inactive until conditions are favourable

for germination. All seeds need water, oxygen, and proper
temperature to germinate. Some seeds require proper light also.
Some seeds germinate better in full light while others require
darkness to germinate.

When a seed is exposed to the proper conditions, water and oxygen

are taken in through the micropyle in seed coat. The embryo's cells
start to divide and enlarge. Then the seed coat breaks open and a
radicle emerges first, followed by plumule that contains the leaves
and stem.

Many things can cause poor germination such as overwatering that

causes the plant to have not enough oxygen. Planting seeds too
deeply may cause them to use all stored energy before reaching the
soil surface. Dry conditions mean the plant does not have enough
moisture to start the germination process and keep it going.

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Some seed coats are so hard that water and oxygen cannot get
through until the coat breaks down. Other seeds need to be exposed
to proper temperatures.

Factors affecting seed germination

The factors affecting seed germination can be Abiotic

(environmental) or Biotic (internal).
A) Abiotic factors that control or affect seed germination can be
summarized as:
1) Temperature: every species of seed, have an optimal soil
temperature for germination, and at that temperature, the maximum
number of seeds will germinate in less time than at any other
temperature. Cold (low) temperature is not favorable for seed
germination. Seeds prefer higher temperature. The rate of seed
germination is directly proportional to the rise in temperature.
2) Moisture or water: Dry seeds do not germinate. They must
first imbibe water to start the process of germination.

For non-dormant seeds, germination starts when a seed is provided

with water as long as the temperature is appropriate. The uptake of
water by dry seed is called imbibition (imbibition means to drink:
seeds imbibe water). As seeds imbibe water, they expand and

9
enzymes and food supplies become hydrated. Hydrated enzymes
become active and the seed increase its metabolic activities to
produce energy for the growth process. In addition, the water
causes turgor pressure to increase in the cells and they are able to
enlarge.

The first part of the seedling to emerge from the seed coat is the

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In such soaked seeds it is an easy to remove the testa. On one
curve, the radicle fits into an inner pocket. The radicle is in a
somewhat unprotected position because of being so near the
micropyle, which is an area of weakness, and this pocket gives it
extra protection.

When the testa is removed, the naked embryo is fully exposed. It

consists of three parts: (1) Two seed- leaves, or cotyledons, which
are attached laterally to a very short primary axis. (2) The radicle,
or young root, which points directly downwards from the primary
axis. (3) The plumule, or rudimentary bud, which grows directly
upwards from the primary axis. It lies between the cotyledons at
right angles to the radicle.

The plumule develops into the plant’s whole system of leaf-

bearing, flower-bearing, and fruit-bearing parts.

In germination the pressure of the growing root tears the testa at

the point of weakness and its tip emerges in the region that was
once the micropyle .

When the root is well established in the ground, the plumule grows
up. Its tip is bent like a hook, so that the delicate leaves of this first

22
bud are not subjected to friction ‫اإلحتكاك‬. Once the tip has broken
through the surface of the soil it becomes erect by the straightening
of the stem.

The first leaves borne on the stem are simply stipules. These are
succeeded by compound leaves made up of two leaflets and a
rudimentary tendril. At the base of each compound leaf is a pair of
stipules. The later leaves have three pairs of leaflets and the
rudimentary tendril is still present.

The elongation of the stem of the plumule, the primary bud of the
plant, like the corresponding elongation of the axis of a resting-
bud, separates the nodes widely. The leaves, which were originally
crowded together, are now separated one from another by long
internodes, and the only crowding is that of the young leaves,
continually being formed from the growing point.

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Endospermic seed structure (Monocots): Poaceae (cereal grain,
caryopsis) – Zea mays and other cereals
Monocot species like corn and other cereal species have caryopses
(cereal grains) as propagation units. Caryopses are single-seeded
fruits in which the testa (seed coat) is fused with the thin pericarp
(fruit coat).
Cereal grains have highly developed embryos and the triploid
endosperm consists of the starchy endosperm (dead storage
tissue) and the aleurone layer (living cells).
Organs of the cereal embryo are the coleoptile (shoot sheath), the
scutellum, the radicle and the coleorrhiza (root sheath).
After germination, the scutellum functions in absorbing the
nutrients from the starchy endosperm and delivering them to the
growing seedling.
The coleoptile protects the shoot meristem and plumule when
growing through the soil.
The coleorhiza has a similar function for the radicle prior to
germination. The coleorhiza is the first structure that grows
through the pericarp, the radicle (the completion of germination)
then ruptures it.

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Germination and seedling establishment of cereal grains is
hypogenous. After the primary root emerges, the coleoptile is
pushed upward by elongation of the mesocotyl.
The coleoptile elongates and reaches the soil surface. Here it
ceases to elongate and the first leaves of the plumule emerge
through an opening at its tip.

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 Root System

In trcheophyta (vascular plants), the root is the organ of a plant

body that typically lies below the surface of the soil. This is not
always the case, however, since a root can also be aerial (growing
above the ground) or aerating (growing up above the ground or
especially above water). Therefore, it is better to define root as a
part of a plant body that lacks nodes and bears no leaves. There are
also important internal structural differences between stems and
roots.

The major functions of roots

1) Absorption and transportation of water and inorganic nutrients

2) Anchoring of the plant body to the ground

3) Roots often function in storage of food and nutrients.

Roots have four regions: a root cap; a zone of division (growing

zone); a zone of elongation; and a zone of maturation.

Root cap is interpreted as structure protecting the apical meristem

and assisting the growing root in penetrating the soil. It consists of

26
living parenchyma cells derived from the apical meristem by
divisions contributing cells a way from the apex. This part of the
apical meristem often appears as a distinct meristem called
calyptrogen. As new cells are produced, cells on the periphery of
the root cap are sloughed off ...‫تنسلخ‬. Root tips growing in the soil
are coated with more or less large amounts of mucilage.

The root cap appears to be the main source of this mucilage,

although the material occurs also on the surface of the young root
at levels free of the root cap and extending to the root hair zone.
The well-known effect of this coating mucilage is the adherence of
soil particles to the root tips and the root hairs.

The outer cells in the root cap secrete mucilage. The secretion
process is a function of dictyosomes, which produce large vesicles
containing the secretory product.

Above the root cap is the zone of division and above that is the
zone of elongation. The zone of division contains growing and
dividing primary meristematic cells. After each cell division, one
daughter cell retains the properties of the meristem cell, while the
other daughter cell (in the zone of elongation) elongates sometimes

27
up to as much as 150 times. As a result, the root tip is literally
pushed through the soil.

In the zone of maturation, cells differentiate and serve such

functions as protection, storage, and conductance. The zone of
maturation of many roots has an outer layer (the epidermis), a
deeper layers (the cortex), and a central region that includes the
conducting vascular tissue.

Early root growth is one of the functions of the apical meristem

located near the tip of the root. The meristem cells more or less
continuously divide, producing more meristem, root cap cells
(these are sacrificed to protect the meristem), and undifferentiated
root cells. The latter become the primary tissues of the root, first
undergoing elongation, a process that pushes the root tip forward
in the growing medium. Gradually these cells differentiate and
mature into specialized cells of the root tissues.

Roots will generally grow in any direction where the correct

environment of air, mineral nutrients and water exists to meet the
plant's needs. Roots will not grow in dry soil. Over time, given the
right conditions, roots can crack foundations, snap water lines, and
lift sidewalks. At germination, roots grow downward due to

28
geotropism, the growth mechanism of plants that also causes the
shoot to grow upward. In some plants (such as ivy), the "root"
actually clings to walls and structures.

Types of roots

A true root system consists of a primary root and secondary

(lateral) roots. The primary root originates in the radicle of the
seedling. It is the first part of the root to be originated. During its
growth, it re-branches to form the lateral roots. It usually grows
downwards. Generally, two categories are recognized:

The taproot system: the primary root is prominent and has a

single, dominant axis; there are fibrous secondary roots running
outward. Usually allows for deeper roots capable of reaching low
water tables. The taproot is most common in dicots. The main
function of the taproot is to store food.

The adventitious root system: the primary root is not dominant;

the whole root system is fibrous and branches in all directions. The
diffuse root is most common in monocots. The main function of
the fibrous root is to anchor the plant.

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The roots, or parts of roots, of many plant species have become
specialized to serve adaptive purposes besides the two primary
functions described above, these roots are modified for storage of
food or water

Storage roots: They include some taproots and tuberous roots.

Storage taproot

The primary root of taproot becomes fleshy and swollen due to

storage of food. These are of following types:

(a) Conical root is broad at the base and gradually tapers towards
apex e.g. Daucus carota (Carrot).

(b) Napiform. The food is accumulated only in upper parts to give

it a top-shaped appearance e.g. Beta vulgaris (beetroot) and
Brassica napus (turnip).

(c) Fusiform. The root is swollen in middle and tapers towards the
base and apex e.g. Raphanus sativus (radish).

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Adventitious roots

Adventitious root is any root arising, in an abnormal, unusual or

unexpected position or place.

Adventitious roots arise out-of-sequence from the more usual root

formation of branches of a primary root, and instead originate from
the stem, branches, leaves, or old woody roots. They commonly
occur in monocots and pteridophytes, but also in many dicots, such
as strawberry (Fragaria).

Most aerial roots are adventitious. In some conifers, adventitious

roots can form the largest part of the root system.

Respiratory or aerating roots (or pneumatophores): roots

rising above the ground, especially above water such as in
some mangrove genera (Avicennia, Sonneratia). In some
plants like Avicennia the erect roots have a large number of
breathing pores for exchange of gases.

Fibrous roots: A root system made up of many threadlike

members of more or less equal length, as in most grasses.

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Aerial roots: roots entirely above the ground, such as in ivy
(Hedera) or in epiphytic orchids. They function as prop roots,
as in maize or anchor roots or as the trunk in strangler fig.

Contractile roots: they pull bulbs or corms of monocots,

such as hyacinth and lily, and some taproots, such as
dandelion, deeper in the soil through expanding radially and
contracting longitudinally. They have a wrinkled surface.
Climbing root: any plant that in growing to its full height
requires some support. Climbing plants may clamber over a
support (climbing rose), twine up a slender support, or grasp
the support by special processes such as adventitious aerial
roots (English ivy,).
Haustorial roots: roots of parasitic plants that can absorb
water and nutrients from another plant, such as in mistletoe
(Viscum album) and dodder.
Propagative roots: roots that form adventitious buds that
develops into aboveground shoots, termed suckers, which
form new plants, as in cherry and many others.
Prop roots: these are adventitious support roots, that
develop on a trunk or lower branch that begin as aerial roots
but eventually grow into a substrate of some type, common

32
among mangroves. They grow down from lateral branches,
branching in the soil.
Tuberous roots: is a modified lateral root, enlarged to
function as a storage organ. It is thus different in origin but
similar in function and appearance to a tuber. Examples of
plants with notable root tubers include the sweet potato, and
Dahlia. It is a structure used to perennialize the plant for
survival from one year to the next.

33
 Shoot system
The shoot system refers to new fresh plant growth and does
include stems but to other structures like leaves or flowers.
The shoot originates in the embryo at the end opposite the root and
develops a complex shoot apex, different from that of the root.

Characteristics of Shoot Systems

The aboveground, conspicuous part of flowering plants constitutes

the shoot system, which is composed of erect stems on which are
attached leaves, flowers and buds. In most plants, stems are
located above the soil surface but some plants have underground
stems.

A stem is one of two main structural axes of a vascular plant. The

stem normally divided into nodes and internodes, the nodes hold
buds, which grow into one or more leaves, inflorescence
(flowers), or other stems etc. The internodes act as spaces that
distance one node from another. The upper angle between the stem
and the leaf at the node called the leaf axil.

Axillary (lateral) buds located in the leaf axils give rise to

vegetative branch stems or to flowers.

34
Terminal buds are present at the tips of the main stem and
branches and contain the apical meristem tissues.

Roots Shoots
Apical covered by a cup- exposed, no cellular cap
meristem shaped root cap
Apical absent produce primordia
appendages (leaves and buds)
Orientation of occurs in all planes oriented; two locations:
cell division inner ( corpus) and an
outer ( tunica)
Zones separate areas of no such recognizable
division, elongation, zones present
and differentiation

Stems have four main functions, which are

Support for and the elevation of leaves, flowers and fruits.

The stems keep the leaves in the light and provide a place for
the plant to keep its flowers and fruits.

35
 Transport of fluids between the roots and the shoots in the
xylem and phloem.
Storage of nutrients.
The production of new living tissue. The normal life span of
plant cells is one to three years. Stems have cells called
meristems that annually generate new living tissue.

Duration of plants: it is the length of time from establishment to

harvest
Annual: plant, which lives for one year or season, reproduces, and
then dies
Biennial: plant, which lives for two years or seasons, reproduces,
and then dies
Perennials are plants, which live for several to many years or
seasons. Perennials may be woody, with stems that persist
aboveground over the winter, or they may be herbaceous, with
stems that die back to the ground each year.
Evergreen: having leaves, which persist for two or more seasons.
Broadleaf evergreens usually have thick, leathery leaves.
Deciduous: having leaves which die and fall in the cold or the dry
season.

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Plant Habit: refers to the overall shape of a plant. It has a number
of components such as stem length and development, branching
pattern, and texture.

Herb: A plant in which all structures above the surface of the soil,
vegetative or reproductive, die back at the end of the annual
growing season, and never become woody. but may have
underground perennating structures; may be annual, biennial, or
perennial

Subshrub (=suffrutescent): lower stems woody but upper stems

herbaceous ("sub" means "almost")

Shrub: a woody low-stature ‫ القوام المنخفض‬perennial plant with one

to many slender trunks arising from near its base

Tree: a large woody perennial plant with one to several relatively

massive trunks and an elevated crown

Succulent: plant possesses thick, usually soft, watery leaves and/or

stems. Stem succulents & leaf succulents

Vine: a woody or herbaceous plant with a long, slender, more or

less flexible stem which cannot support itself

37
Stem may be: Erect having an essentially upright vertical habit or
position

Alternatively, prostrate laying flat on the ground. prostrate type of

stems may be: Creeping growing along the ground and not
producing roots at intervals along surface, or Runner an
elongated, slender branch that roots in the soil at the nodes or tip,

Metamorphosis of stems

Stems are often metamorphosed for storage, asexual reproduction,

protection or photosynthesis, including the following:

I: Underground stems

They are modified plant structures that derive from stem tissue but
exist under the soil surface. Plants use under ground stems to
multiply their numbers by asexual reproduction and to survive
from one year to the next, usually over a period of dormancy.
Plants produce these modified stems so they can survive a cold or
dry period, which normally is a period of inactive growth, and
when the cold or dry period is over the plants begin new growth
from the underground stems. Being underground protects the stems
from the elements during the dormancy period, such as freezing

38
and thawing in winter or extreme heat and drought in summer or
fire. They can also protect plants from heavy grazing pressure from
animals, the plant might be eaten to the ground but new growth can
occur from below ground that can not be reached by the
herbivores. A number of weedy species use underground stems to
spread and colonize large areas since the stems do not have to be
supported or strong, less energy and resources are needed to
produce these stems, often these weeds have more mass under
ground than above ground. Underground stem functions mainly in
reproduction but also in storage

1- Bulb: a short vertical underground stem with fleshy storage

leaves attached, e.g. onion, tulip.

Bulbs often function in reproduction by splitting to form new bulbs

or producing small new bulbs termed bulblets. Bulbs are a
combination of stem and leaves so may better be considered as
leaves because the leaves make up the greater part.

2- Corm: is a short, vertical, swollen underground plant stem that

serves as a storage organ used by some plants to survive winter or
other adverse conditions such as summer drought and heat.

39
A corm consists of one or more internodes with at least one
growing point, with protective leaves modified into skins or tunics.
The thin tunic leaves are dry papery, dead petiole sheaths, formed
from the leaves produced the year before which acts as a covering
that protects the corm from insects and water loss.

Internally a corm is mostly made of starch-containing parenchyma

cells above a circular basal node that grows roots.

Corms grow two different types of roots; from the bottom of the
corm normal fibrous roots are formed as the shoots grow, they are
produced from the basal area at the bottom of the corm, the other
type of roots are produced were the corm grows new corms. The
second type of roots is thicker layered roots that are able to pull the
corm deeper into the soil. These roots which are called contractile
roots are produced in response to a fluctuating soil temperature
and light levels e.g. Colocasia.

3- Rhizome: is a horizontal stem of a plant that is usually found

underground, often sending out roots and shoots from its nodes.
Rhizomes develop from axillary buds

40
In general, rhizomes have short internodes; they send out roots
from the bottom of the nodes and new upward-growing shoots
from the top of the nodes.

The plant uses the rhizome to store starches, proteins, and other
nutrients. These nutrients become useful for the plant when new
shoots must be formed or when the plant dies back for the winter.
This is a process known as vegetative reproduction and is used by
farmers and gardeners to propagate certain plants.

4- Tuber: is a thickened part of stolon that has been enlarged for

use as a storage organ.

In general, a tuber is high in starch, for example, the common

potato, which is a modified swollen, underground storage stem
adapted for storage and reproduction.

The tuber has all the parts of a normal stem, including nodes and
internodes, the nodes are the eyes and each has a leaf scar. The
nodes or eyes are arranged around the tuber in a spiral fashion
beginning on the end opposite the attachment point to the stolon.
Internally a tuber is filled with starch stored in enlarged

41
parenchyma like cells; also internally the tuber has the typical cell
structures of any stem, including pith, vascular zones and a cortex.

II: Metamorphosis for assimilation

Cladodes: A flattened stem that appears leaf like and is specialized

for photosynthesis with only one internodes (e.g. Asparagus) and

phylloclade: A flattened stem that appears leaf like and is

specialized for photosynthesis, with more than one internodes
(Cactus pads, Ruscus).

III: Climbing are stems that cling or wrap around other plants or
structures.

IV: Reduction of transpiration: The most Effective adaptation by

which desert plants reduce their water loss and adjust to variations
for water involves changes in the size of the transpiring body.
Seasonal reduction in the transpiring surface is an important factor
in maintaining the water balance of certain xerophytes
As a seedling, the plant has broad leaves, which wither over time.
When conditions are favorable in the wet season, the new growth
has broad leaves and soft spines. At the advent of the dry season,
however, adaptations that reduce water loss, leaves are shed, while

42
the spines are lignified and become stiff. The whole plant becomes
spiny and without leaves. Thorn is a reduced stem with a sharp
point and rounded shape. e.g. Alhagi, Zilla.

Buds

A bud is an undeveloped or embryonic shoot and normally occurs

in the axil of a leaf or at the tip of the stem. Once formed, a bud
may remain for some time in a dormant condition, or it may form a
shoot immediately.

The buds of many woody plants are protected by a covering of

modified leaves called scales, which tightly enclose the more
delicate parts of the bud, many bud scales are covered by a
gummy substance, which serves as added protection.

In many plants, scales are not formed over the bud, which is then
called a naked bud. Such naked buds are found in shrubs and in
herbaceous plants. In many of the latter, buds are even more
reduced, often consisting of undifferentiated masses of cells in the
axils of leaves.

43
A terminal bud occurs on the end of a stem and lateral buds are
found on the side. A head of cabbage is an exceptionally large
terminal bud.

Since buds are formed in the axils of leaves (axillary buds), their
distribution on the stem is the same as that of leaves. There are
alternate, opposite, and whorled buds, as well as the terminal bud
at the tip of the stem.

In many plants buds appear in unexpected places; these are known

as adventitious buds.

Types of buds

Buds are often useful in the identification of plants, especially for

woody plants in winter when leaves have fallen. Buds classified
and described according to different criteria: location, status,
morphology, function. Botanists commonly use the following
terms:

For location
o Terminal, when located at the tip of a stem (apical is equivalent
but rather reserved for the one at the top of the plant),

44
o Axillary, when located in the axil of a leaf (lateral is equivalent
but some adventitious buds may be lateral too),
o Adventitious, when occurring elsewhere, for example on trunk or
on roots (some adventitious buds may be former axillary ones
reduced and hidden under the bark, other adventitious buds are
completely new formed ones),

For morphology

Scaly or covered (winter), when scales (which are in fact

transformed and reduced leaves) cover and protect the embryonic
parts, Naked (summer), when not covered by scales, and Hairy,
when also protected by hairs (it may apply either to scaly or to
naked buds),

For function
o Vegetative, if only containing vegetative pieces (embryonic shoot
with leaves), Reproductive, if containing embryonic flower(s) (a
flower bud) and Mixed, if containing both embryonic leaves and
flowers.

45
 Leaves

A leaf is an aboveground plant organ, typically flat (laminar) and

thin, to expose the cells containing chloroplast to light over a broad
area, and to allow light to penetrate fully into the tissues
specialized for photosynthesis.

Leaves are also the sites in most plants where transpiration and
guttation take place. Leaves can store food and water, and are
modified in some plants for other purposes.

Parts of a Typical Leaf

The leaf consists of three parts namely, leaf base (usually
provided with a pair of stipules), petiole and leaf blade or
lamina.

(i) Leaf base (Hypopodium): Leaf base is the lower most part
of the leaf meant for attachment. It acts as a leaf cushion. In
most of the plants it is indistinct. Sometimes leaf base shows
different variations as follows:
(a) Pulvinate leaf base: In members of leguminosae the leaf is
swollen. Such swollen leaf bases are called pulvinate leaf
bases as seen in mango leaves.

46
(b) Sheathing leaf base: In grasses and many monocots, the
leaf base is broad and surrounds the stem as an envelope, such
a leaf base is called sheathing leaf base, e.g., Maize, Wheat and
Palms. In grasses (Maize, Wheat etc.) the sheathing leaf
base protects the intercalary meristem.
(c) Modified leaf base: The leaf bases in few plants perform
accessory functions and show modifications. In (Onion), the
leaf bases store food materials and become fleshy. They are
arranged concentrically to form a tunicated bulb.
In Platanus and Robenia, the leaf bases protect the axillary
buds and grow around them to form cup like structures.

The petiole attaches to the stem at a point called the "leaf axil". A
petiole may be absent, or the blade may not be laminar
(flattened).

After a period (i.e. seasonally, during the autumn), deciduous

trees shed their leaves. These leaves then decompose into the soil.

47
Arrangement on the stem

Different terms are usually used to describe leaf placement

(phyllotaxis): Alternate: leaf attachments are singular at nodes,
and leaves alternate direction, to a greater or lesser degree, along
the stem.

Opposite: The leaves on this plant are arranged in pairs opposite

one another, with successive pairs at right angles to each other
("decussate") along the stem or superposed if not rotated, but
two-ranked (in the same geometric flat-plane).

Whorled: three or more leaves attach at each point or node on the

stem. Opposite leaves may appear whorled near the tip of the stem.

A rosette: is a circular arrangement of leaves, with all the leaves at

a similar height. Though rosettes usually sit near the soil, their
structure is an example of a modified stem.

Divisions of the lamina (blade)

Two basic forms of leaves described considering the way the blade
is divided.

48
A simple leaf has an undivided blade. However, the leaf shape
may be formed of lobes, but the gaps between lobes do not reach to
the main vein.

A compound leaf has a fully subdivided blade, each leaflet of the

blade separated along a main or secondary vein. Because each
leaflet can appear to be a simple leaf, it is important to recognize
where the petiole occurs to identify a compound leaf.

Compound leaves are a characteristic of some families of higher

plants, such as the Fabaceae. The middle vein of a compound leaf,
when it is present, is called a rachis.

Palmately compound leaves have the leaflets radiating from the

end of the petiole, like fingers off the palm of a hand, e.g.
Cannabis (hemp).

Pinnately compound leaves have the leaflets arranged along the

main or mid-vein. This may be:

Odd-pinnate (imparipinnate): Leaflets are attached along an

extension of the petiole called a rachis; there is a terminal leaflet
and therefore an odd number of leaflets, e.g. Rosa.

49
Even-pinnate (paripinnate): Leaflets are attached along an
extension of the petiole called a rachis; there is an even number of
leaflets, e.g. Cassia.

Bipinnately compound leaves twice divided: The leaflets also

divided into leaflets, the leaflets arranged along a secondary vein
that is one of several branching off the rachis. Each leaflet called a
"pinnule". The pinnules on one secondary vein are called "pinna";
e.g. Albizia (silk tree).

Trifoliate: a pinnate leaves (also known as ternate leaves) are a

leaf shape characterized by a leaf divided into three leaflets, e.g.
Trifolium (clover).

Characteristics of the petiole

Petiolated leaves have a petiole. Sessile leaves do not. The blade
attaches directly to the stem. In clasping or decurrently leaves,
the blade partially or wholly surrounds the stem, often giving the
impression that the shoot grows through the leaf. In peltate leaves,
the petiole attaches to the blade inside from the blade margin.

In some Acacia species, the petioles are expanded or broadened

and function like leaf blades; these are called phyllodes.

50
Stipules

The stipules are the small lateral appendages present on either

side of the leaf base. They protect the young leaf or leaf
primordia. Leaves with stipules are called stipulate and those
without them are called exstipulate.

The stipules are commonly found in dicotyledons. In some

grasses (Monocots) an additional outgrowth is present between
leaf base and lamina. It is called ligule.

The leaves having ligules are called ligulate. Sometimes, small

stipules like outgrowths are found at the base of leaflets of a
compound leaf. They are called Stipules.

Types

According to their shape, size and location may be of the

following types: Adnate stipules: These are two lateral stipules
attached to the petiole up to some distance but the anterior part
remains free e.g., Rose and Lupin. Ochreate: When stipules of the
leaf make a hollow tube, which encircles (surrounds) the stem up
to the upper part of node, the structure is known as ochrea, e.g.,

51
Polygonum Rumex etc. Hairy stipules: These are hair like
stipules which are dry in nature, e.g.,Corchorus

Foliaceous stipules: These are two leafs by broad and green

stipules. Due to the absence of axillary buds in their axil these can
be distinguished from the leaves. In Lathyrus aphaca stipules are
modified into leaves and leaves into tendrils. In Pisum
sativum foliaceous stipules are much developed and upper leaflets
are modified into tendrils. Main function of these structures is to
synthesize food.

Tendrillar stipules: are thin and modified into wire like structure.
Tendrillar stipules help in climbing of plants. These are usually
present in tendril climbing plants e.g., Smilax.

Spiny stipules: When stipules are converted into sharp pointed

structures, these are known as spiny stipules. These may be long
(as in Mimosa and Acacia) or curved (in Ziziphus). They protect
plants from wild animals.

Venation

Venation is the pattern of veins in the blade of a leaf. The veins

consist of vascular tissues, which are important for the transport

52
of food and water. Leaf veins connect the blade to the petiole, and
lead from the petiole to the stem. The venation pattern of a leaf is
classified as reticulated, parallel, or dichotomous.

In reticulated venation, the veins are arranged in a net-like

pattern, in that they are all interconnected like the strands of a net.

Reticulated venation is the most common venation pattern, and

occurs in the leaves of nearly all dicotyledonous Angiosperms,
whose embryos have two cotyledons (seed leaves).

In parallel venation, the veins are all smaller in size and parallel
or nearly parallel to one another, although a series of smaller veins
connects the large veins. Parallel venation occurs in the leaves of
nearly all monocotyledonous Angiosperms, whose embryos have
one cotyledon, as in grasses.

In dichotomous venation, the veins branch off from one another

like the branches of a tree. This is the rarest venation pattern, and
occurs in the leaves of some ferns and in the gymnosperms.

53
Leaf Blade Shape:

The overall shapes of leaves and leaflets are often characteristic of

individual species of plants and, together with reproductive
features, are used in plant identification. The terminology below is
used to describe the shape of laminae of simple and compound
leaves as well as the laminae of the leaflets of compound leaves.

Acicular: have a long and very narrow leaf shape, with sides that
are almost parallel to each other and are usually more than ten
times longer than broad. Acicular leaves are often borne on short
lateral branches called fascicles; Linear leaves: have a long and
very narrow leaf shape, with sides that are almost parallel with one
another and are usually more than four times longer than broad.

Oblong leaves: have a rectangular leaf blade two to four times

longer than it is wide, with sides that are almost parallel to each
other.

Lanceolate leaves: have a lance-shaped leaf, with the widest part

of the leaf near the base and the narrowest part near the apex.

Oblanceolate leaves: have a lance-shaped leaf, with the widest

part of the leaf near the apex and the narrowest part near the base.

54
Orbicular leaves: have a more or less circular leaf shape in which
the width and length of the lamina are equal, or nearly so.

Elliptical leaves: have a shape that looks like an ellipse, twice as

long as broad, with the widest part of the leaf near the middle.

Ovate leaves: are egg-shaped, with the widest part of the leaf
below the middle toward the base.

Obovate leaves: are egg-shaped, with the widest part of the leaf
above the middle toward the apex.

Reniform leaves: have a shape like a kidney.

Cordate leaves: are shaped like a heart, with the lobes of the heart
at the base of the leaf and the pointed portion at the apex.

Obcordate leaves: are also heart-shaped, but the lobes are at the
apex, and the basal lamina tapers into the petiole.

Sagittate leaves: have a shape like an arrowhead.

Hastate leaves: are also shaped like an arrowhead, but the basal
lobes diverge or extend away from the midrib, giving an outline
that resembles a halberd.

55
Spatulate leaves: have a spoon-shaped or spatula-shaped leaf
where the lamina is widest near the rounded apex.

Deltoid leaves: are delta-shaped, resembling an equiangular

triangle. Often the sides of the deltoid leaves are slightly curved
toward the apex.

Subulate leaves: are short, narrow, flat, stiff, awl-shaped leaves

that taper to a sharp point.

Scale leaves: are small, inconspicuous leaves that are typically

appressed tightly to the stem and have overlapping margins.

Lyrate: When the leaf blade is like a lyre ‫القيثارة‬, i.e., contains a big
terminal lobe and many smaller lateral lobes, e.g., Raphanus
sativus.

Leaf apices

The apex of a leaf lamina is opposite the petiole. Leaves apexes

vary greatly from plant to plant and can be useful in classification
and identification.

Acute: sharp, ending into a pointed apex with margins that form an
acute angle between 45 and 90 degrees. e.g., Mangifera indica.

56
Acuminate: a sharp-pointed apex with straight or convex margins
that form an angle less than 45 degrees.

Obtuse: a blunt apex, rounded end with margins that form an

angle greater than 90 degrees. e.g., Cassia obtusifolia.

Retuse: a rounded summit like obtuse but with a shallow

depression (shallow notch) at the apex, e.g., Pistia stratiotes.

Emarginate: with a shallow depression at the apex, not exceeding

1/15 of the distance to the centre of the leaf blade. e.g., Bauhinia

Mucronate: Rounded apex with a small extension of the midrib

barely extending beyond the blade apex e.g., Calotropis gigantea.

Leaf margins

The margin of a leaf is another name for the structure of the leaf's
edge. There are many different variations, and they can be:

Entire: A leaf margin that has a continuous, unbroken and smooth

edge, without teeth, lobes or indentations

57
Crenate: the margins of a leaf (or leaflet or other organ ) have
relatively large rounded, blunt projections, but not so large as to be
considered lobes.

Serrate: A leaf margin forming a row of small sharp outward

projections pointing toward the apex of the leaf resembling the
teeth of a saw.

Dentate: A leaf margin with sharp tooth-like projections pointing

outward

Sinuate: Strongly wavy margins with shallowly rounded divisions

within the same plane of the blade.

Spinose: Has spines along leaf margins. Often these spines occur
on the margin where the leaf veins terminate at serrations at the
leaf margin.

58
Terminology in plant morphology

The seeds are the dispersal and propagation units of the

Spermatophytes (seed plants): Gymnosperms (conifers and related
groups) and angiosperms (flowering plants).

Seeds are mature, fertilized ovules. Ovules are structures of seed

plants containing the female gametophyte with the egg cell, all
being surrounded by the nucellus and 1-2 integuments.

Double fertilization results in formation of the diploid embryo

and the triploid endosperm.

Testa (seed coat): Outer protective layer of the seed developed

from the integuments of the ovule, diploid maternal tissue, Seed
coats help protect the embryo from injury and also from drying
out. Seed coats can be thin and soft as in beans or thick and hard as
in locust or coconut seeds.

Hilum and funiculus: Funicular scar on seed coat that marks the
point at which the seed was attached via the funiculus to the ovary
tissue.

59
The Micropyle is a canal or hole in the coverings (seed coat) of
the nucellus through which the pollen tube usually passes during
fertilization. Later, when the seed matures and starts to germinate,
the micropyle serves as a minute pore through which water enters.
The micropylar seed end has been demonstrated to be the major
entry point for water during seed imbibitions and germination.
During germination the testa ruptures at the micropylar end and the
radicle protrudes through the micropylar endosperm.

Chalaza: Non-micropylar end of the seed. At the base of an ovule,

bearing an embryo sac and surrounded by integuments.

Fruits are mature, ripened ovaries containing seeds. The pericarp

("fruit coat") is diploid maternal tissue

Embryo: Young saprophyte, diploid (2n), results of fertilization.

The mature embryo consists of cotyledons (seed leaves),
hypocotyls (stem-like embryonic axis below the cotyledons), and
radicle (embryonic root).

Endosperm: Food storage tissue, triploid (3n), results of double

fertilization. A temporary food supply is packed around the
embryo in the form of special leaves called cotyledons or seed

60
leaves. These generally are the first parts visible when the seed
germinates.

Endospermic seeds: The endosperm is present in the mature seed

and serves as food storage organ.

Non-endospermic seeds: The cotyledons serve as sole food

storage organs as in the case of pea (Pisum sativum). During
embryo development the cotyledons absorb the food reserves from
the endosperm. The endosperm is almost degraded in the mature
seed and the embryo is enclosed by the testa.

61
 Part II: plant anatomy

Introduction

The plant anatomy is the study of the internal structure of the

various parts of the plant, and began a little over 300 years ago
with the work of Grew and Malpighi. Their work involved careful
well-illustrated descriptions of plant material.

The highly organized plant body of a seed plant begins its

existence usually with the fertilized egg, the zygote, which
develops into the embryo by characteristic steps resulting in the
adult organization. The root represented by its formative tissue of
root, the radical. Similarly, the apical formative tissue of the shoot
may or may not initiate the development of a shoot above the
cotyledons. If a primordial shoot is present, it is called epicotyls,
plumule.
Cell divisions in the embryo initiate the organization of tissue
systems. The component tissues are still formative (meristematic)
but their position and cytological characteristics indicate a relation
to mature tissues appearing in the subsequently developing
seedling.

62
The plant cell

Cells make up the smallest level of a living organism. The cell is

called the fundamental unit of life because the cellular level of an
organism is where the metabolic processes occur that keep the
organism alive.

Each cell is capable of not only make up living things but also it is
living things. The cell is the unit of construction of plant, just as
atoms are the units of molecules.

Plants, like animals, are composed of cells. Some consists of only

one cell, but the flowering plants, with which our study is
concerned, are made up of many cells, which at maturity differ
greatly in structure.

Recognition of the cellular construction of plants goes back to the

seventeenth century and knowledge of the cell and its contents has
progressed hand-in hand with the development of the microscopes
employed for its observation.
By the middle of the nineteenth century, it had been realized that
all organisms consist of cells, and further, that all such units are
derived from the division of existing cells. It was shown that

63
chromosomes were present in the nucleus, and that these divided
during nuclear divisions.
The most important characteristic of a cell is that it can reproduce
by dividing. Cell division is the process by which cells duplicate
and replace themselves. Nuclear divisions were of two kinds:
1- Mitosis division; those that gave rise to the somatic cells of the
plant, in which, the chromosomes duplicated and the daughter cells
had the same number of chromosomes as the original cells.
2- Meiosis division; those that gave rise to the reproductive cells of
the plant, in which, the daughter cells had only half the original
number of chromosomes.
The cell was known to consist of a cell wall and its inner contents,
the protoplast.
The protoplast comprised a more or less spherical body, the
nucleus, containing chromosomes, which were the bearers of the
hereditary units or genes, embedded in the granular matrix, the
cytoplasm. Various other inclusions were also observed with the
microscope, cell organelles.
The various components of the plant cell that are known at the
present time will now be discussed.

64
As summarized below the main components of the plant cell are
the cell wall, cytoplasm, and nucleus. The cytoplasm includes the
different cell organelles. In the following, we will discuss briefly
each of the cell components.

The plant cell

The cell wall The protoplast

Protoplasm Ergastic substances

The cytoplasm The nucleus

Hyaloplasm Cell organelles

65
The cell wall is not a living system, but in absence of the protoplast
that formed it, is merely a non- living shell. The wall is formed
during the growth of the cell. Plant cell walls are of considerable
importance to man. Cell walls themselves constitute timbers and
also used directly as fibers, cotton, etc.; materials extracted from
them serve as glue, food and food additives.

Formation of cell wall

The cell wall consists of a crystalline poly-saccharide, which in
higher plants is cellulose. The cellulose occurs in bundles of chains
which comprise fiber like structures, the microfibrils.
The cell wall is formed during the process of cell division. There is
evidence that the presence of the nucleus is necessary for wall
formation.
The cell division takes place through two steps namely
karyokinesis (nuclear division) and cytokinesis (division of the
cytoplasm).
During nuclear division, a plate is gradually produced at the
position of the equator of the spindle. Vesicles formed by
dictyosomes apparently fuse to form the cell plate, and this process
continues until the cell plate reaches the exciting cell walls.

66
The cell plate gives rise to the middle lamella, which, is composed
of pectic substances. The pectic substances react with calcium and
magnesium ions already present in the cell, to form the middle
lamella which become a rigid substance (cementing material). The
middle lamella holds together the primary walls of adjacent cells.

The primary cell wall

The primary wall is the first wall to be formed by the cell, and is
deposited on either side of the middle lamella by the contiguous
cells.
Chemically it consists mainly of cellulose, hemicellulose and other
polysaccharides. Layers of pectic substances may separate the
cellulose lamellae.
All meristematic cells and many mature cells still have living
contents, have primary walls. Since the wall is formed when the
cell is young, it must undergo considerable growth. The wall is
characterized by plasticity.

67
Secondary walls
Secondary walls are usually formed after a cell has completed its
elongation; therefore do not normally extend to any considerable
degree.
Where a secondary wall is formed it is deposited on the inner side
of the existing wall, next to the cell lumen. It consists mainly of
cellulose and other polysaccharides.
Various other substances, notably lignin may be deposited in the
wall.
Among the organic substances, cutin, subrin, and waxes are found
most commonly in the protective surface tissues of the plant which
imparts rigidity to the cell wall
The secondary wall often consists of three layers, so that a cell wall
may consist altogether of five layers

Pits:
Pits are relatively thinner portions of the cell wall that
adjacent cells can communicate or exchange fluid through. Pits are
characteristic of cell walls with secondary layers.

Generally each pit has a complementary pit opposite of it in the

neighboring cell. These complementary pits are called "pit pairs".

68
Pits are composed of three parts: the pit chamber, the pit aperture,
and the pit membrane. The pit chamber is the hollow area where
the secondary layers of the cell wall are absent. The pit aperture is
the opening at either end of the pit chamber. The pit membrane is
the primary cell wall and middle lamella, or the membrane
between adjacent cell walls, at the middle of the pit chamber.

The primary cell wall at the pit membrane may also have
depressions similar to the pit depressions of the secondary layers.
These depressions are primary pit-fields, or primary pits.

In the primary pit, the primordial pit provides an interruption in the

primary cell wall that the plasmodesmata can cross. The primordial
pit is the only aperture in the otherwise continuous primary cell
wall.

Plasmodesmata are thin sections of the endoplasmic

reticulum that traverse pits and connect adjacent cells. These
sections provide an avenue of transport through the pits and
facilitate communication

Types of pits

Though pits are usually simple and complementary, a few more

pit variations can be formed:

69
 Simple pits: A pit pair in which the diameter of the pit chamber
and the diameter of the pit aperture are equal.

Bordered pits: A pit pair, in which the pit chamber is over-arched

by the cell wall, creating a larger pit chamber and smaller pit
aperture.

Half bordered pits: A pit pair in which a bordered pit has a

complementary simple pit. Such a pit pair is called half
bordered pit pair.

Blind pits: A pit pair in which a simple pit has no complementary

pit.

Compound pits: A pit pair in which one cell wall has a large pit
and the adjacent cell wall has numerous, small pits.

70
The architecture of the cell wall
Cellulose forms the framework interpenetrated by the matrix
represented by the non-cellulosic carbohydrates.
The cellulose framework is a system of fibrils composed of
cellulose molecules.
The fibrils are of different classes of magnitude. The largest are
visible with a light microscope and are called macrofibrils.
With an electron microscope these fibrils are resolvable into
microfibrils. Subunits of the microfibrils are referred to as
micelles.
Cellulose has crystalline properties as a result of orderly
arrangement of cellulose molecules. Less regularly arranged
glucose chains occur between and around the micelles and
constitute the paracrystalline regions of the microfibril.

Methods of wall building

The mechanisms of wall building, to increase the thickness, occur
by two methods of deposition of wall material: Apposition and
intussusceptions.

71
In apposition, the building units are placed one on top of another
and this is usually centripetal, i.e. it occurs from outside towards
the center of the cell and so the lumen of the cell becomes narrow
In intussusception, the units of new material are inserted into the
existing structure. Intussusception is probably the rule when lignin
or cutin is incorporated into the wall.

Intercellular spaces
A large volume of the plant body is occupied by an intercellular
space system, which is most characteristic of mature tissues.
The most common intercellular spaces develop by separation of
contiguous primary walls through the middle lamella. Two types of
intercellular space develop in plant tissues.
1- Schizogenous in which spaces result from separation of cell
walls from each other along more or less extended areas of their
contact.
In such cases, the intercellular substance dissolves partly and an
intercellular space develops and becomes quite big in size.
Schizogenous cavities form an intercommunicating system of long
intercellular canals, which facilitate diffusion of gases and liquids
from one part of the plant body to the other. The resin ducts in the

72
Coniferales, and the secretory ducts in the Compositae are the
typical examples.

2- Lysigenous intercellular spaces (lysis, loosening, Greek). These

spaces arise through dissolution of entire cells. These cavities of
intercellular spaces store up water, gases and essential oils in them.
The examples are commonly found in water plants and many
monocotyledonous plants. The secretory cavities in Citrus and
Gossypium are good examples.

The cytoplasm
The cytoplasm, physically, is a viscous substance, forming a
colloidal system, which is more or less transparent in visible light.
Chemically the structure of cytoplasm is very complex; the major
component is water (forms 85-90% in active cells and about 10-
12% in cells of dormant seeds) with soluble and non-soluble
components (hyaloplasm).

Membranous system in the cell

The cell contains a number of membranes forming system
particularly in the well - organized cells. All the membranes have a

73
more or less the same chemical structure namely lipoprotein
referred to as a unit membrane. Each of these membranous
systems functions according to their location.

The nucleus
The nucleus is present in all different types of cells except the
sieve tube cell of higher plants and the red blood cells. It is
considered the most important component of the cell. It contains
the hereditary material, which control the activity of the cell.
Eukaryotes possess a definite nucleus.
Prokaryotes such as bacteria and others do not have easily
recognizable nuclei, but they contain certain nuclear material. The
eukaryotic nucleus consists of nuclear membrane, chromatin
material, nucleolus and nuclear sap.

Endoplasmic reticulum
It is a membranous system, which branches inside the cytoplasm to
form a network-like structure. The endoplasmic reticulum is
present in all adult plant. Under the electron microscope, it appears
as double unit membranes, which, chemically made of lipoprotein.
It forms channels that branch and penetrate throughout the

74
cytoplasm. Endoplasmic reticulum serves as a channel for
transportation to various parts of the cell of different nutrients,
protein including enzymes.
Two types of endoplasmic reticulum are found:
1) Granular due to the presence of small dense granules, which are
the ribosomes. The granular ER, is related to protein synthesis due
to the presence of the ribosomes,
2) A granular endoplasmic reticulum, similar to the rough E. R.,
but lacks the ribosomes. The smooth ER is normally related to
steroid, lipid and glycogen metabolism. The manufactured protein
can penetrate into cavities of the endoplasmic reticulum.

Plastids
They are only present in plant cells. They appear under the light
microscope as a small, rounded, oval or disc-shaped bodies
embedded in the cytoplasm. The plastids are considered as part of
the membranous system. The principal types of plastids are
chloroplasts, chromoplasts and leucoplasts.
Chromoplasts are usually yellow, orange, or red because of the
carotenoid pigments. They occur in petals, in some ripe fruits, and
some roots (e.g. carrot roots). Leucoplasts are non-pigmented

75
plastids, usually located in tissues not exposed to light; they store
the plant products such as starch (amyloplasts), proteins
(proteinoplasts), and fats (elaioplasts). Leucoplasts of tissues
which become exposed to light may develop into chloroplasts (e.g.
in potato tuber).
Chloroplasts are green because of pigment chlorophyll, which
predominates in them. The chloroplastids are bounded by an
envelope consisting of two unit membranes. Internally the plastids
contain a membrane system embedded in a proteinaceous matrix
or stroma.
The membrane system is in the form of flattened sacs called
thylakoids. The thylakoid system consists of grana and frets
(stroma thylakoids). The grana are interconnected by the frets that
transverse the stroma. In addition to the light harvesting system,
the chloroplasts contain the enzymes responsible for the fixation of
carbon dioxide into sugar. Each granum is composed of a series of
disc-like thylakoids stalked one upon the other like a pile of coins.

76
The Golgi apparatus
The Golgi apparatus appears, under EM, as membranes in the form
of
a) System of flattened sacs (Cisternae)
b) Vesicles and
c) Large vacuoles surrounded by membranes.
In many cells of higher plants and in some animal cells, the Golgi
apparatus appear to consist of many separate units called
dictyosomes.
Golgi apparatus performs a number of important functions such as
Secretion of large molecules in plant cells e.g. the growing region
in root tips secretes pectic material which take part in cell plate
formation and collection and accumulation of certain substances
such as lipids, proteins and enzymes.

Mitochondria
They are found lying free in the cytoplasm of nearly all plant and
animal cells. They appear, under the light microscope, as rods or
spherical granules freely distributed in the cytoplasm. The
mitochondria are surrounded by a double envelope consisting of an
outer and an inner membrane.

77
The outer membrane is smooth and stretched around, while the
inner one is greatly folded. These folds are known as
mitochondrial cristae. The cristae produce a large surface to
volume ratio, which facilitates the function of the mitochondrion.
The interior of the mitochondria is filled with a certain liquid
known as the matrix.
The matrix includes variety of soluble substances such as sugars,
organic acids, mineral salts, vitamins as well as many respiratory
enzymes. The membranes of mitochondria are made of proteins
and lipids.
The main function of mitochondria is considered as the site where
the respiratory reactions take place. The mitochondria contains the
cytochromes (electron transfer agents), the dehydrogenises
enzymes (oxidoreductases) associated with them, respiratory
pigments and some enzymes involved in the krebs cycle.

Ribosomes
Ribosomes appear under the EM as small granules. They are
found either attached to the endoplasmic reticulum or free in the
cytoplasm. The ribosomes are being formed within the nucleolus.

78
They are chemically composed of RNA known as ribosomal RNA,
and protein.

Lysosomes
Lysosomes are vacuoles surrounded by a unit membrane formed
by the Golgi apparatus. There are primary and secondary
lysosomes.
The primary is formed on the rough ER (endoplasmic reticulum).
The secondary lysosomes are formed on the smooth ER.
Lysosomes are containing high concentration of number of
digestive enzymes.
These enzymes are capable of hydrolyzing different classes of
macromolecules.
The lysosomes control the release of these digestive enzymes to
the cell. In case of abnormal situation, lysosomal enzymes may
escape into the cytoplasm and either kill the cell (autolysis), or
bring about dramatic metabolic changes

79
Tissues and tissue systems
The morphologic units of multicellular organisms, the cells, are of
many different kinds and acquire diversity in structure and
function. Similar cells are associated in various ways with each
other forming coherent masses or tissues. Besides being similar in
form and function, the cells of a tissue have a common origin.

A group of vegetative cells is a vegetative tissue, and group of

reproductive cells is a reproductive tissue.

The tissues of more highly organized plants are distinguished as

regards their stage of development into:

1- Meristematic or formative in which growth is taking place.

2- Permanent tissues, those which lost, at least temporarily, their

power of division.

Meristematic tissues

Meristems are specialized tissues responsible for the formation of

new cells and localized in certain organs in plants. The cells of
meristems differ from mature tissues and characterized by:

1- They have abundant cytoplasm.

2- Vacuoles small or lacking,

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3- Large nuclei compared with cell size.

4- Thin cellulosic walls.

5- No intercellular spaces.

6- The ability of repeating division.

According to the mode of their origin, meristematic tissues are

classified as primary and secondary.

1- Primary meristems arise by the division of the germ cell and at

the very early stage of plant life comprise the whole embryo. Later,
they become localized at the apices of stems and roots and the
primordial of leaves and similar appendages. In primary
meristems, promeristem do always the earliest stage and transition
stages to mature tissues constitute the remainder of the meristem.

2- Secondary meristem, on the other hand, are derived either from

the inactive remains of the primary meristem or are newly
originated from cells of the permanent tissues by cell division.
Such secondary meristems, which get the name cambium, give
rise to the secondary growth increasing thickness of woody plants,
as represented by:

1- New tissues in medullary rays (interfascicular cambium) or

2- Periderm or cork tissue (cork cambium)

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On the other hand, as regard their position in plant body meristems
are classified into apical, intercalary and lateral.

+ Apical meristems: are those which lie at the apices of the axis
and of the appendages and are commonly called growing points
e.g. tips of roots, stems and often leaves of vascular plants. The
activity of apical meristems brings increase in the length of these
organs, building the primary body of the plant.

+ Intercalary meristems are portions of apical meristems that

have become separated from the apex during development by
layers of permanent tissues. The best known intercalary meristems
are those which lie at the basal regions of internodes and leaves
of many monocotyledons, e.g. grasses. Intercalary meristematic
regions ultimately disappear, as being wholly transformed into
permanent tissues.

+ Lateral meristems are situated laterally in an organ such as the

cambium and the cork cambium. The cells of these latter
meristems divide chiefly in one plane (periclinally), thus
increasing the diameter of the organ in which they occur.

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Structural development and differentiation of plant tissues

In higher plants, the apical meristem consists of a small apical

portion, the promeristem. The remainder of the apical meristem
represents the early stages of the tissues formed by the
promeristem. These tissues gradually pass over into the permanent
stage, while the promeristem retains its power of division.

In the stem apex, the region below the promeristem is

distinguished into:

1- An outermost layer, protoderm which, in the mature region,

can be recognized as the epidermis. This layer which is uniseriate
external layer sometimes termed the dermatogen.

2- The inner cell become distinguished into procambium which

appears as strands (procambial strands) very close to the apex of
the stem and consists of elongated, slender cells. This procambium
is a central core which forms the pith and the primary vascular
tissue, this central core sometimes termed the plerome.

3- The remaining undifferentiated meristematic tissue is composed

of layer of cells with intercellular spaces and is known as the
ground meristem. From the ground meristem the primary cortex,

83
medullary rays and photosynthetic elements are all formed. This
meristematic tissue is sometimes termed the periblem.

Along the sides of the meristem are outgrowths of meristemaic

cells which have arisen by the accelerated divisions of some of the
outermost cells of the meristems. Such outgrowths are the
beginnings of young leaves and thus referred to as leaf primordia.

Branch primordial cells arise in the same way in the axils of

immature leaves. The cells of the leaf, after a time, lose their
meristematic properties, while the axillary branch retains its
growing point.

Unlike the growing point of the shoot, that of the root lacks lateral
appendages and segregation into nodal and internodal regions.
The thin-walled meristematic cells of the growing point are
protected by a special organ composed of permanent tissue, and
called the root cap, in which the outer cell walls become
mucilaginous and this makes the forward passage of the root
easier. The root cap has an independent origin, from initial cells
called calyptrogen in monocotyledons, while in dicotyledons the
formation of the root cap results from the periclinal division of the
dermatogen itself.

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Tissue systems

In specialization of cells from meristem changes in shape, structure

and nature of the cell wall and of the protoplast produce various
cell types; accordingly, there are two tissue types:

1- Simple, when cells of a single type occur together in a uniform

or homogenous mass, e.g. parenchyma, collenchyma and
sclerenchyma.
2- Complex or Heterogenous, when cells of more than one type
are associated in a tissue, e.g. xylem and phloem.

Permanent tissues

Permanent tissues are those in which growth has ceased at least

temporarily. Permanent tissues may again become meristematic,
also sometimes when permanent tissues mature their cells become
completely dead. Permanent tissues are recognized into three tissue
systems.

1) Epidermal tissue system (dermal – boundary tissue).

2) Ground tissue system (fundamental tissue).
3) Vascular tissue system (conducting tissue).

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Epidermal or Boundary Tissue

The epidermal tissue system consists of three elements; epidermal

cells, stomata, and hairs or trichomes.

The epidermis is the outermost layer or layers of cells on all plant

parts during primary growth. It is thus in direct contact with the
environment and is subject to structural modification by various
environmental factors.

The epidermis of the stem, leaves and floral parts originates from
the surface layer of the shoot apical meristem. That of the root
originates from a layer of cells in the root apical meristem that is
covered by the root cap.

Usually the epidermis consists of only one layer of cells, but in a

few species the cells of this layer may divide periclinally to give
rise to a several layered or multiple epidermis.

The epidermis of both root and shoot may be differentiated into

various kinds of cells. In both instances, epidermal cells may
elongate at an angle to the surface of the organ to give rise to
hairs.

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In the epidermis of the leaf, and often also of the stem, stomata
may be present; in some species cork cells containing silica are
also differentiated.

Epidermal cells are living cells tubular in shape, barrel-shaped or

lens-shaped in cross section and sometimes elongated or nearly
isodiametric.

Each epidermal cell has a large central vacuole and thin

peripheral cytoplasm. Minute leucoplasts are present and
chloroplasts are absent except in hydrophytes and those of deeply
shaded habitats.

The epidermal cells in many leaves and petals have wavy lateral
walls, which increase the firmness of the union of the cells.

Epidermal cell walls vary in thickness in different plants and in

different parts of the same plant.

Epidermal cell wall may become silicified as in grasses. Primary

pits and plasmodesmata generally occur in the anticlinal and inner
periclinal walls of the epidermis.

Very often, a layer of fatty material, or cutin, is deposited on the

surface of the epidermal cell wall, forming the cuticle. This

87
substance is for the most part impervious to water and may have a
protective function.

The cuticle is extremely resistant to microorganisms, thus in the

living plant it affords some protection, perhaps largely mechanical,
against infection by pathogens, and in fossilized plant remains it
may by extremely well preserved, being resistant to decay.

In many plants, conspicuous deposits of wax are formed on the

surface of the cuticle. It is this wax, which gives the “bloom” to
some fruits, e.g. grapes. Waxes may be deposited in large or small
flakes ‫ رقائق‬or granules, rods, tubes or sheets, which may then be
sculptured into ridges.

Cutin and waxes are synthesized in the living protoplasm and

migrate to the surface through the cell wall. The cuticle, the
cutinized cell wall beneath it and the surface wax serve to reduce
loss of water.

Stomata (sing. Stoma) occur on most of the aerial parts of plants,

though predominantly on leaves and young stems. They are the
ports for exchange of oxygen and carbon dioxide gas for
photosynthesis, but also release excess water into the air.

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This process of water loss maintains a steady flow of water and
minerals from the roots to the leaves. To minimize the water loss,
many plants regulate the duration and time of day when stomata
are open. A stoma consists of a pore surrounded by two guard
cells.

The stoma arises by the division of one of the young epidermal

cells (protodermal cell); one of the products of this division forms
the guard mother cell. The guard mother cell divides by a vertical
wall to form the two guard cells.

The epidermal cells adjoining the guard cells often differ in size or
arrangement from the rest of the epidermal cells; such cells called
subsidiary cells. The stoma, together with the subsidiary cells, is
sometimes termed the stomata complex.

Stomata are clearly of great importance to the physiology of the

plant, being sites of gaseous exchange during transpiration,
photosynthesis and respiration.

The guard cells differ from other cells of the epidermis in the more
richly cytoplasm, the prominent nucleus and existence of
chloroplasts and starch grains.

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The guard cells of dicotyledons and monocotyledons except
Gramineae (Poaceae) and Cyperaceae are commonly crescent-
shaped (kidney-shaped), with rounded ends, as seen from the
surface, and have ledges ‫ حواف‬of wall material on the upper or both
upper and lower sides. The cells are covered with cuticle that
extends over the surfaces facing the stomatal pore and the
substomatal chamber.

Stomata may be completely covered with wax. Among the guard

cells of monocotyledons, those of Poaceae have a rather uniform,
specific structure. As seen from surface, the cells are narrow in
the middle and enlarged at both ends, i.e. dumb-bell-shaped.
The middle portion is thick walled and rigid whereas the enlarged
or bulbous ends are thin walled. Although the guard cells of the
major taxa have their distinguishing characteristic, they chair the
feature that the anticlinal wall away from the pore (dorsal wall) is
thinner, and then more flexible than the other walls. This feature
is commonly cited as having a causal relation to the ability of the
guard cells to change their shape in response to turgor changes and
thus to control the size of the stomatal opening.

Guard cells may occur at the same level as the adjacent epidermal
cells. These are usually present in mesophytes, while raised types

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of stomata, which protrude above epidermal cells levels, are
confined to some water plants (hydrophytes) and sunken stomata
(below the surface of epidermis) are confined to desert plants
(xerophytes).

In leaves with parallel veins such as monocotyledons and some

dicotyledons and in the needles of conifers, the stomata are
arranged in parallel rows and attain the stripped pattern. While in
leaves with reticulate venation (dicots) the stomata are distributed
in no particular order and the stomata are scattered.

The mechanism of stomata movement

During stomata opening, starch disappears from chloroplasts, at the

same time as K+ ions enter the guard cells, and during stomata
closure, the reappearance of starch parallels the loss of K+ ions.

The early theory that the breakdown of starch contributes to the

increase of osmotic pressure in the guard cells, because of
formation of sugar, has been replaced by the concept that starch
hydrolysis may provide the organic anions with which potassium
uptake is associated.

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Types of stomata according to subsidiary cells

In most species, the number and arrangement of the subsidiary

cells around the stomata are relatively constant. Various
classifications of stomata according to these arrangements have
been drown up, and are sometimes useful to taxonomists. Four
types of stomata complex have been described; these types have
given descriptive names as follows.

1- Anomocytic or irregular-celled (Ranunculaceous): the

surrounding cells are indefinite in number and do not differ
from the other epidermal cells.
2- Anisocytic or unequal-celled (Cruciferous): usually three
subsidiary cells surround the stoma, one cell being
considerably smaller or larger than the other two.
3- Diacytic or cross-celled (Caryophyllaceous): two subsidiary
cells surround the stoma with their common wall at right
angles to the guard cells.
4- Paracytic or parallel-celled (Rubiaceous): one or more (often
two) subsidiary cells are present, with their long axes parallel
to the guard cells.

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Trichomes or hairs

Trichomes are highly variable appendages formed by the

outgrowth of epidermal cells. They are formed in all parts of the
plant, including stamens (e.g. Tradescantia) and seeds (e.g.
cotton). Root hairs are morphologically typical hairs

Hairs may be unicellular or cell division may take place, so that

the hair becomes multicellular.

The unicellular hair, which may have the form of papillae,

represents the simplest form and they are common upon petals and
many leaves. The unicellular hairs may be branched or
unbranched.

The more advanced types or forms of hairs are the multicellular,

which range from simple linear hairs of few cells to complex
branched or massive structures involving considerable areas of
the epidermis.

Stinging hairs (peltate hairs) and many others are complex

multicellular structures. Unicellular, multicellular and peltate
hairs may be glandular, producing secretions which are often of
the nature of ethereal oils. The glandular head may be unicellular
or multicellular, the head constitutes the secretory part of the

93
hair. The glandular hair formed of living cells and those of the
head have dense protoplasmic contents and large nuclei. The cells
of hairs may be dead in other types.

The cell walls of trichomes are commonly of cellulose and are

covered with a cuticle. They may be lignified. Plant hairs often
produce thick secondary walls, e.g. the cottonseed hairs are
cellulose, and the walls of hairs may be impregnated with silica or
calcium, e.g. stinging hairs.

Cystolith and other crystals may develop in hairs. Among the

diverse types of hairs are those associated with absorption of water
from the atmosphere, e.g. the unicellular absorbing hair of
Diplotaxis harra.

Hairs perform very different functions.

1- Dense coverings of hairs bring about a decrease in the rate of

transpiration.

2- Root hairs serve for the absorption of water and other

substances.

3- Vesiculated hairs on leaves of Atriplex halimus (salty plant)

serve to remove salts from the leaf tissue and thus prevent an
accumulation of toxic salts in the plant.

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5- Trichomes may provide a defense against insects.

Ground tissue or Fundamental tissue

Fundamental or ground tissue is that in which the vascular bundles

are embedded or is the tissue, which occupies all the spaces in the
plant organs, which are not occupied by the vascular or
conducting tissues. The ground tissue originates from the periblem
and plerome i.e. it constitutes the cortex and pith.

The ground tissue in the different plant organs is differentiated

into cortex, pith, and medullary rays and in leaf is represented by
the mesophyll, which is differentiated in dicotyledonous leaves
into palisade and spongy tissues. The ground tissue is therefore
responsible of different activities such as photosynthesis, storage,
assimilation, and respiration. It is considered a heterogenous tissue
consisting of parenchyma, collenchyma, sclerenchyma and
secretory elements.

Parenchyma

Parenchyma is the main representative of the ground tissue system

and is found in plant organs forming a continuous tissue, as in the

95
cortex and pith of stems, cortex of roots, and ground tissue of
petioles, and mesophyll of leaves; or individual or groups of cells
in the complex tissue systems, xylem, and phloem.

Parenchyma origin may thus be diverse: from the apical

meristems of stem or root, the marginal meristems of leaves or
from the vascular cambium or even the phellogen in more
mature organs with secondary growth.

Parenchyma cells are living cells variable in their morphology and

physiology, but generally having a polyhedral shape. They may be
isodiametric or elongated or even folded, lobed or armed as the
parenchyma of the mesophyll.

The cell walls in active vegetative ground parenchyma, including

the mesophyll of leaves are relatively thin and are classified as
primary. Sometimes the primary wall becomes thick, a feature
prominently displayed by the storage parenchyma of some seeds,
as those of Coffea arabica and Phoenix dactylifera. The
carbohydrates of such walls are regarded as reserve material
utilized by the embryo during germination. Lignified cell walls
also occur in parenchyma cells, e.g. lignified secondary wall of
lignified xylem parenchyma.

96
The variation in activities of parenchyma cells are usually
reflected in variations in the balance of protoplasmic components.

Chlorenchyma contains numerous chloroplasts are intensively

engaged in photosynthesis. Chlorenchyma finds its highest
expression in the leaf mesophyll, but chloroplasts may be
abundant also in the cortex of stem and may occur in deeper
tissues, even in the pith.

The storage parenchyma is rich in organic contents such as

sugars, starch, fatty oils, or proteins. Many succulent plants as
Aloe and Mesembryanthemum contain water storage parenchyma.

Aerenchyma is tissues, notably in many aquatic plants, with

intercellular spaces exceptionally well developed and form a
connected system throughout the entire plant.

Collenchyma

Collenchyma cells have living protoplasts and thickened

cellulosic walls. They are extensible ‫ قابل لالمتداد‬cells with a
considerable degree of plasticity, and function as supporting
tissue in growing organs.

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They may contain chloroplasts and carry out photosynthesis. They
thus differ from parenchymatous cells chiefly in their thick-walled
nature, and in being usually somewhat elongated in a plane
parallel to the long axis of the organ in which they occur.

Collenchyma cells differ from sclerenchymatous fibers both in

their possession of living contents at maturity and in cellulosic
nature of their walls.

Collenchyma usually occupies a peripheral position in the organs

in which it occurs. In stems, it may lie immediately beneath the
epidermis, or below a few outer parenchymas. The
collenchymatous cells may form a complete cylinder near the
periphery of the stem or they may occur in the form of discrete
strands, especially in ridged structures such as the petiole of celery
(Apium graveolens) or many stems as in Calendula, Senecio.

Collenchyma cell wall thickening begins in the corners of the

cells, but it may spread from there in various ways in different
species.

The main types of collenchyma are recognized, according to the

deposition of the wall thickening, these are:

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a) Angular collenchyma, in which the wall thickening is
deposited predominantly at the corners, or angles of the cells, e.g.
Celery petiole, stems of Datura and Dahlia.

b) Lamellar collenchyma, in which the thickening is deposited

more heavily on the tangential than on the radial walls of cells,
e.g. stem of Sambucus.

c) Lacunar collenchyma, in which the thickening is deposited

primarily around the intercellular spaces between the cells and
hollow rod-like structures are formed.

Sclerenchyma

This tissue consists of thick-walled elements, which are normally

hard and lignified. The cell walls are thickened secondary walls
and the cells usually have no living protoplasts when mature.

Sclerenchyma is distinguishable from collenchyma both in this

lake of living contents and in being lignified; it has a similar
function in the plant, however, namely that of support.

Sclerenchyma may be subdivided into fibers and sclereids

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A) Fibers vary considerably in length; they are typically many
times longer than broad. Most fibers are elongated elements with
pointed tips, a narrow lumen, and thick secondary walls.

Fibers may occur in roots, stems, leaves, and fruits, in association

with a number of different tissues. They may be present in the
xylem or phloem, as a sheath or bundle cap associated with
vascular bundles, especially in leaves, or in the parenchymatous
tissues of the pith or cortex.

Fibers are sometimes classified into two groups: xylem fibers, and
extraxylary fibers, the latter including all those fibers which
occur in tissues other than the xylem, i.e. phloem, cortical and
perivascular fibers.

In monocotyledonous leaves fibers may be present not only as a

sheath around the vascular bundle but also extending between the
bundles and the upper and lower epidermis.

Fibers are polygonal, in transverse section, the walls are usually

lignified. The lumen of fiber is small. Sometimes it is a very
narrow channel or at certain levels, no lumen exists.

100
B) Sclereids are widely distributed in seed coats and fruits. They
are dead cells with strongly thickened walls which are often
lignified and accordingly the lumen is narrow.

Sclereids vary greatly in type, thickness of wall and number of

pits. Various terms have been applied to the different types

1- Macrosclereids (rod cells), more or less columnar in shape,

lumen narrow, constituting the palisade layer of many seeds
and fruits and occurring in some xerophytic leaves and stem
cortices.
2- Brachysclereids (stone cells), short roughly, isodiametric
sclereids, resembling parenchyma cells in shape. Widely
distributed in cortex, phloem, and pith of stems and in fleshy
fruits. Stone cells may be solitary or may lie loosely together
closely packed.
3- Osteosclereids (bone cells), of bone like or barrel-shape or
columnar cells enlarged at ends, constituting hypodermal layers
in many seeds and fruits and frequent in xerophytic leaves.
4- Astrosclereids (stellate cells), they are found in stems and
leaves of xerophytic organs, having extreme lobes or arms.

101
5- Trichosclereids (internal hairs), branched, thin walled
sclereids with lobes projecting like hairs into cellular spaces, in
leaves and stems of hydrophytes.

Vascular or conducting tissue

The vascular or conducting systems are special tracts, which are

constructed to suit the function of translocation of water, and food
from one organ to another in the plant body.

Since an adequate supply of water and food is obviously essential

for growth, it is at once evident that the vascular system is
functionally very important within the plant. The possession of
vascular tissues separates the higher plants from some of the more
primitive groups of plants, which are devoid of comparable
conducting tissues. The vascular system of the plant is made up of
xylem and phloem.

The arrangement of xylem and phloem to each other differs

according to the organ in which they detected. They are arranged
either in radial arrangement or in vascular bundles.

102
a) Radial arrangement is characteristic for roots in which xylem
and phloem are not on the same radius, but phloem alternate
with the xylem ridges.
b) Vascular bundles: the primary vascular system of seed plants
consists of strands variable in size and degree of distinctness.
Discrete individual strands are commonly referred as vascular
bundles. This type of arrangement is characteristic for stems.
The phloem and xylem show variations in their relative position in
vascular bundles as shown below:

1) Collateral is the prevalent ‫ السائد‬arrangement, in which the

phloem occurs on one side (abaxial) of the xylem.

In a typical dicotyledonous stem, a band of very narrow cells, the

cambium exists between xylem and phloem and the vascular
bundle is thus called opened vascular bundles. In
monocotyledonous stems, the bundles contain no cambium
between xylem and phloem, so called closed vascular bundles
and they have not the possibility of formation of secondary
elements.

2) Bicollateral in which one part of the phloem occurs on the outer

side and another on the inner side of the xylem and the two
complements of phloem are referred to as the external (abaxial)

103
and the internal (adaxial) phloem. This is represented in certain
ferns and in some dicotyledons (Apocynaceae, Asclepiadaceae,
Convolvulaceae, Cucurbitaceae and Solanaceae).

3) The concentric vascular bundle in which either the phloem

surrounds the xylem (amphicribral, amphiphloic bundle) or the
xylem surrounds the phloem (amphixylic, amphivasal bundle).

The amphivasal bundles are found in certain positions in stems of

some dicotyledons (medullary bundles in Rumex,
Mesembryanthemum and Begonia) and in monocotyledons
(Araceae, Liliaceae, Juncaceae and Cyperaceae).

Amphicribiral bundles are most common in ferns but are found

also in angiosperms. Small bundles in flowers, fruits, and ovules
may be amphicribral.

Xylem

It is the principal water conducting tissue in a vascular tissue.

Developmentally, it is convenient to distinguish between primary
and secondary vascular tissues.

104
The primary tissues differentiate during the formation of the
primary plant body, and the meristem directly concerned with the
formation of the primary vascular tissue is the procambium.

The secondary vascular tissue are produced during the second

major stage of plant development in which an increase in thickness
results from lateral additions of new tissues to the axial parts of the
plant (i.e. stem and root) and their larger branches. It results from
the activity of the vascular cambium. Both the primary and
secondary xylem is complex tissues containing at least water
conducting elements and parenchyma cells and usually other
types of cells, especially supporting cells.

In stems, leaves and floral parts, the primary xylem and the
associated primary phloem commonly occur in strands, the
vascular bundles. Panels of parenchyma, the interfascicular
regions, occur between the vascular bundles in stems; these panels
are often called medullary rays and are considered part of the
ground tissue.

In the root, the primary xylem forms a core with or without

parenchyma in the center or is arranged in strands.

105
Developmentally, the primary xylem usually consists of an ealier
part, the protoxylem and a later part, the metaxylem. Although the
two parts have some distinguishing characteristics, they integrate
so that the delimitation of the two can be made only
approximately.

The protoxylem differentiates in the parts of the primary plant

body that have not completed their growth and differentiation.

The protoxylem usually contains only tracheary elements

embedded in parenchyma that is considered to be part of the
protoxylem.

The metaxylem is somewhat more complex than the protoxylem

and may contain fibers in addition to tracheary elements and
parenchyma cells. The tracheary elements of the metaxylem are
retained after the primary growth is completed but become non-
functioning after some secondary xylem is produced.

In plants lacking secondary growth, the metaxylem remains

functional in mature organs.

Xylem, which is a complex tissue, consists of several types of

cells, living and non-living. These are tracheary elements

106
(tracheids and vessel members), xylem fibers (wood fibers) and
xylem parenchyma (wood parenchyma).

1) Tracheids
It is the fundamental cell type in xylem. The tracheid is “an
elongate cell with tapering ends which when mature is non-
living, i.e. without protoplast”.
The walls of tracheids are hard and usually lignified. They are
angular in cross section. The lignified secondary walls are usually
pitted and the pits are of the bordered type.
The tracheid is well adapted to its functions, which are primarily
water conduction and secondarily support. It is a long, empty,
firm-walled tube, extending parallel with the long axis of the
organ. It is in communication with contiguous tracheids as well as
other types of cells, living, and non-living by means of pits.
In gymnosperms, xylem is formed mainly of tracheids, and it is
considered as a primitive element than the vessel elements. In
addition, it is formed in monocots between the metaxylem vessels.
Tracheids differ from vessels in that they are not perforated.

107
2) Vessels
Vessels or tracheae constitute the fundamental conducting
elements of the xylem of the angiosperms. Vessels formed of a
series of vessel members whose end walls are dissolved or
perforated.
The length of xylem vessels differ greatly depending on the kind of
plant, the location in the organ and apparently with the rate of
growth of the organ. Vessels are formed from series of xylem
mother cells, procambial cells, in formation of primary xylem, and
from cambial derivatives in formation of secondary xylem. From
meristematic stage, the vessel elements enlarge rapidly, increase
greatly in diameter. At the last stages of development, the cells
fuse together end to end. This fusion involves the loss of the end
walls or perforation of these walls so that the lumina of the series
of cells are freely open into one another and with the walls forms a
long tube.
The wall area bearing the perforation is called the perforation
plate. The perforation may be simple with one pore or multiple
with more than one. Simple perforation plates are thought to have
been derived from the multiple types, by loss of the bars of

108
thickening. Typically, the protoplasts die after the perforation is
formed.

Structure of secondary wall in a vessel

Generally in the first formed part of the primary xylem a more

limited area of the primary wall is covered by secondary wall
layers than in the later formed primary xylem and in the secondary
xylem.
Beginning with the earliest primary xylem, the secondary
thickening are deposited in the successive elements as rings
(annular), continuous helices (helical or spiral), then as
networks (reticulate). When the meshes of the net are rather
distinctly elongated transversely, the thickening is called
scalariform thickening. Much advanced elements have extensive
secondary walls, which are interrupted only in the pit areas and
thus called pitted.

3) Xylem fibers (wood fibers)

Xylary fibers are elongated elements with pointed ends. They
usually have thicker walls and the pits may have smaller borders
than the tracheids of the same species.

109
Wood fibers may retain ‫ تحتفظ بــ‬living protoplasts for as long as
twenty years, although they are usually defined as non-living
elements. There may be transitional types between fibers and
tracheids.
Two types of xylem fibers are recognized: the fiber tracheids and
the libriform fibers.
The libriform fibers are longer and have thicker walls than the
fiber-tracheids. The pits in libriform fibers are simple and have no
borders. Fibers of both categories may be septate.
These fibers are widely distributed in dicotyledons and usually
retain their protoplasts in mature sapwood, being concerned with
storage of reserve materials.

4) Xylem parenchyma
Parenchyma cells are a common constitute of the xylem of most
plants. They may be thin-walled or thick-walled. If a secondary
wall is present, the pit pairs between the parenchyma cells and the
tracheary elements may be bordered, half-bordered, or simple.
Xylem parenchyma cells have a variety of contents, e.g. stored
food materials such as starch and oils especially at the end of a
growth season; tannins, crystals and various other substances.

110
Parenchyma cells are present in two forms:
a) Axial or vertical series of more or less elongated cells placed end
to end and are known as wood or xylem parenchyma.
b) Radial transverse series and are known as xylem ray
parenchyma.

Phloem

The food conducting tissue of seed plant, the phloem, is

associated with the xylem in the vascular system.
The phloem consists of several types of cells and may be classified
into a primary and a secondary tissue according to the stage of
development.
The primary phloem is derived from procambium, while the
secondary phloem originates in the vascular cambium.
Phloem is a complex tissue; it made up of
a- Sieve cells and phloem parenchyma, only as in simple and in
more primitive lower vascular plants as Ptridophytes.
b- Sieve cells, parenchyma and phloem fibers as in some
gymnosperms.

111
c- Sieve tubes, companion cells, phloem parenchyma and one or
more kinds of phloem fibers, sclereids and secretory cells as in
angiosperms.

I) Sieve elements
Parallel with the classification of the tracheary elements into
primitive tracheids and more advanced vessel members, the
conducting elements of the phloem segregated into the less
specialized sieve cells and the more specialized sieve tube
elements. Sieve cells and sieve tube elements are morphologically
equivalent and are alike in fundamental structure and function.
1) Sieve cells
The sieve cells are elongate, living cells with thin peripheral layer
of cytoplasm, having no nucleus when the cell is mature. Sieve
cells taper at their ends or have steeply inclined end walls.
Sieve cells have no sieve plates and the conduction of food takes
place through small openings found on their lateral walls.
Sieve cells have no companion cells. The sieves cells originate
from a normal meristematic cell filled with cytoplasm and have a
nucleus. The meristematic cell enlarges and a big vacuole is
formed. The nucleus is then lost and the sieve areas are developed.

112
2) Sieve tube elements or members
They are composed of two constituents; these are sieve tubes and
companion cells.
a) Sieve tubes
Sieve tubes are longitudinal series or files of thin walled cells
connected with each other through the cross walls which are
perforated to form sieve plates.
They resemble the sieve cells in that, they elongate, are living cells
with a thin layer of peripheral cytoplasm and no nucleus is present
when the cells mature. They also perform the function of
conduction in absence of the nucleus.
The sieve tubes originate from the mother cells of sieve elements,
which are either short cylindrical, elongate, or tapering. They
elongate and the cytoplasm becomes highly vacuolated, the wall
thickens; the cytoplasm strands become prominent and increase in
size. Callus develops around the connecting strands.
As the sieve tube elements reaches maturity in size, the walls
become thinner, and the nucleus disintegrates, the connecting
strands increase in diameter, peripheral cytoplasm becomes

113
extremely thin. At this stage, the function of conduction apparently
begins.
The vacuole of sieve tube contains a strand of slime, which often
appears to be accumulated at the sieve plate in the form of slime
plugs, which is regarded as an indication that the cell has been
injured. Slime in normal cases consists of a relatively viscous
substances dispersed in the vacuolar sap and consists mainly of
proteins.

Sieve areas and sieve plates

The sieve areas, which resemble a sieve, are wall areas with
clusters of pores, through which the neighboring sieve elements are
interconnected by cytoplasmic strands. They are considered as
specialized primary pit-fields. In mature and active conducting
sieve elements, callose is relatively small. Callus gradually
increases and its amount lining the pores of sieve areas is larger
and accordingly the pores become narrower. The wall parts bearing
the more highly differentiated sieve areas are called sieve plates.
Two types of sieve plates are distinguished:

114
1- Simple sieve plates with one sieve area, the pores tend to be
very large and the plate occupies all or nearly all the transverse end
wall.
2- Compound sieve plates with several sieve areas, the pores tend
to be smaller than that of simple type and the plate occupies only
part of the oblique end.

b) Companion cells
The sieve tube members of angiosperms are commonly associated
with specialized parenchyma cells called companion cells. These
cells arise from the same meristematic cells as the associated sieve
tube members. In the formation of the companion cells, the
meristematic mother cell divides longitudinally one or more times.
One of the resulting cells, usually the larger, differentiates into
sieve-tube member while the other becomes a companion cell.
The number of companion cells varies from one to several in
different species and may also be variable in the same plant. They
also vary in size; some are as long as the sieve tube member, others
are shorter than the sieve tube member. The companion cells of a
given sieve tube element may occur on various sides of this

115
element or they may form continuous longitudinal series on one
side of it.
In contrast to the sieve element, the companion cell retains its
nucleus at maturity.
The sieve tube and its companion cell appear to be closely
associated not only ontogenetically and morphologically but also
physiologically. When the sieve tube protoplasts are disorganized
at the end of its activity, the associated companion cells also die.
Companion cells are lacking in the earlier parts of primary phloem
(protophloem), while are present in metaphloem.
Solitary long companion cells are common in primary phloem in
herbaceous plants, while short and numerous companion cells
appear to be characteristic of secondary phloem of woody plants.
Companion cells occur in highly specialized phloem (monocot)
where they are abundant together with sieve tubes making up the
entire tissue.
In transverse section, companion cells are seen as small triangular,
rounded, or rectangular cells beside the sieve tube elements.
Companion cells are responsible for the transfer of sugars and
other solutes from and to sieve elements and contiguous cells.

116
II) Phloem parenchyma

Phloem contains variable types of parenchyma cells other than

companion cells. These are concerned with many other activities
characteristic of living parenchyma cells such as storage of starch,
fat and other organic food materials and accumulations of tannins
and resins.
In shape, these cells range from elongate and tapering to broadly
cylindrical, sub-spherical, and polyhedral. The parenchyma cells of
the primary phloem are elongated and their axes parallel with the
longitudinal extent of the vascular tissue.
In the secondary phloem, parenchyma occurs in two systems; the
axial and the ray systems, which are called axial phloem
parenchyma and phloem rays successively.
Pit fields occur between parenchyma cells and companion cells and
sieve cells. After phloem ceases to conduct, the parenchyma cells
may remain relatively unchanged or they may become sclarified
(by deposition of lignin). Phloem parenchyma and ray parenchyma
produce and form cork. Phloem parenchyma cells are absent in
monocots.

117
III) Phloem fibers and sclereids

Fibers are common components of both primary and secondary

phloem.
In the primary phloem, fibers occur in the outermost part of the
tissue; in the secondary, in various distributional patterns among
the other phloem cells of the axial system.
The fibers may be septate or non-septate and may be living or non-
living at maturity. Living fibers serves as storage cells as they do
in the xylem.
In many species, primary and secondary fibers are long cells with
thick walls and are used as a commercial source of fiber (Linum,
Cannabis, and Hibiscus).
Sclereids are also frequently found in the phloem. They may occur
in combination with fibers or alone, and may be present in both
axial and radial systems of the secondary phloem. Sclereids
develop from parenchyma cells as the tissue ages and the sieve
tubes cease to function.

118
 Structure of the primary body in stems

The stem consists of three tissue systems, the dermal, the

fundamental, and the vascular. The variation of the primary
structure in stems of different plants is based chiefly on differences
in the relative distribution of the fundamental and vascular tissues.

1- Dicotyledonous stem
In conifers and dicotyledons, the vascular system of the internodes
commonly appears as a hollow cylinder delimiting an outer and an
inner region of ground tissue, the cortex and the pith,
respectively.
The vascular bundles are separated from each other by more or less
wide panels of ground parenchyma (interfascicular
parenchyma), that interconnects the pith and the cortex. This
tissue called interfascicular because it occurs between the bundles
or fascicles.
A panel of interfascicular parenchyma is often called the
medullary (pith) ray.
In the following we will discuss the internal structure of the
primary body in stems.

119
*Epidermis

The stem epidermis commonly consists of one layer of cells (in

few plants biseriate or multiseriate) and has a cuticle and
cutinized walls. It is a living tissue capable of mitotic activity, an
important characteristic in view of the stresses to which the tissue
is subjected during the primary and secondary increase in thickness
of the stem.

Stomata constitute a less prominent epidermal component in the

stem than in the leaves.

The function of the epidermis is mainly to protect the delicate

internal tissue from rapid loss of water and mechanical injury. It
may serve secondarily in photosynthesis and secretion.

*The cortex

The cortex is the portion of an axis that surrounds the central

cylinder and is separated from the vascular cylinder by the
endodermis.

It is essentially parenchymatous in nature and may contain many

kinds of cells arranged in many ways. Parenchyma usually has

120
chloroplasts. Intercellular spaces are prominent but sometimes are
largely restricted to the median part of the cortex.

In many aquatic plants, the cortex develops as an aerenchyma

with a system of large intercellular spaces.

The peripheral part of the cortex frequently contains collenchyma

in strands or in more or less continuous layer. In some plants,
notably grasses (monocotyledons), sclerenchyma rather than
collenchyma develops as the primary supporting tissue in the outer
region of the stem. The function of the cortex is primarily a
protective layer and secondarily supports photosynthesis and
storage.

* Starch sheath

In young stems, the innermost layer of cortex is represented by

compactly arranged barrel shaped cells, without any intercellular
spaces. It is wavy in appearance. The cells are richly deposited
with starch grains and hence, are commonly described as starch
sheath. When no starch accumulates and no special wall
characteristics develop in the innermost cortical layer, the
delimitation of the cortex from the vascular region may be
problematic.

121
*Pericycle

The pericycle in stems of hydrophytes is a thin cylinder of tissue

limited internally by the vascular bundles and externally by the
starch sheath and is composed of one layer as in roots.

The pericycle is often occupied by a zone of pericyclic fibers,

which is sometimes continuous or sometimes broken up into
groups alternating with parenchyma. The groups of pericyclic
fibers lie outside the vascular bundles. The pericyclic fibers in
angiospermous stems have been shown to arise from primary
phloem (protophloem fibers) after the obliteration ‫ محو‬of the
sieve tubes.

*Vascular cylinder

The primary vascular system of seed plants consists of strands

variable in size and degree of distinctness. Discrete individual
strands commonly referred to as vascular bundles.

The prevalent arrangement of the phloem and xylem in stems is

collateral, bicollateral or concentric.

122
The vascular cylinder is composed of the primary conducting
tissues (primary phloem and primary xylem) including in between
the cambium.

►The primary phloem is classified into protophloem and

metaphloem. The protophloem matures in plant parts that are still
undergoing extension growth. The metaphloem differentiates later.

The protophloem sieve elements are usually narrow and

inconspicuous, but they are enucleate and have sieve areas with
callose. They may or may not have companion cells.

While the sieve elements of the protophloem cease to function and

are obliterated, the fiber primordia increase in length, develop
secondary walls and mature as fibers. Such fibers are found on the
periphery of the phloem region in numerous dicotyledonous stems
and are often called pericyclic fibers.

The metaphloem has more and commonly wider sieve elements

than the protophloem. Companion cells are regularly present in
the metaphloem of angiosperms, but fibers are usually absent.
Parenchyma cells may become sclerified after the phloem ceases to
conduct.

123
►The procambium: The strands of meristematic tissue, which
developed from the promeristem by several longitudinal
divisions, are known as procambium. This tissue is formed of
elongate cylinder cells with dense cytoplasm and is responsible for
formation of primary phloem and primary xylem.

In dicotyledonous plants, procambium strips are present in

vascular bundles and so the vascular bundle is said to be an open
vascular bundle. This cambium remains meristematic and
becomes active only in older stems, forming additional xylem and
phloem in radial rows through the secondary thickening.

In monocot stems all the procambium differentiated into primary

phloem and xylem and so the vascular bundle is said to be closed.

►Primary xylem contains tracheary elements, fibers and

parenchyma cells. Developmentally, the primary xylem usually
consists of an earlier part, the protoxylem, and a later part, the
metaxylem.

The protoxylem differentiates in the parts of the primary body that

have not completed their growth and differentiation.

124
The protoxylem usually contains only tracheary elements
embedded in parenchyma that is concerned to be part of the
protoxylem. When the tracheary elements are destroyed they may
become completely obliterated by the surrounding parenchyma
cells.

The metaxylem is commonly initiated in the still growing primary

plant body, but it matures largely after the elongation is completed.
It is, therefore, less affected by the primary extension of the
surrounding tissues than the protoxylem.

The metaxylem is somewhat more complex than the protoxylem

and may contain fibers in addition to tracheary elements and
parenchyma cells. The tracheary elements of the metaxylem are
retained after the primary growth is completed but become non-
functioning after some secondary xylem is produced. In plants
lacking secondary growth, the metaxylem remains functional in
mature plant organs.

125
*Pith

It is the innermost tissue, which is surrounded by the vascular

tissue. The pith usually formed of parenchyma cells with
intercellular spaces, having storage function.

Frequently certain pith cells contain crystals, tannins or mucilage

(e.g. Tilia and Hibiscus).

Pith may be destroyed during the growth of the stem and

accordingly the internodes are frequently hollow while the nodes
are solid, (e.g. Trifolium).

Pith cells may develop thick lignified walls to increase mechanical

support of the plants, e.g. pith of Retama stem. Pith is distinct in
dicotyledonous stems and few monocots.

2- Monocotyledonous stem

The fundamental difference between the stems of a typical

monocotyledon and dicotyledon is that in monocotyledon all the
tissues of the growing point become differentiated into permanent
tissues, no meristematic tissue remains, nor does any secondary
meristem arise (with few exceptions), and the stem remains
unaltered in structure.

126
1- The stele is broken up into bundles, which are commonly
distributed throughout the fundamental tissue.
2- The endodermis is lacking and the limits of cortex, pericycle
and pith are often indistinguishable. However, in some herbaceous
monocotyledons, as in most grasses (e.g. Triticum) there is present
a central region in which no bundles occur; this region may
represent the pith.
3- In monocotyledonous stems, the epidermis is often followed
internally by a cylinder of a hypodermal strengthening tissue of
sclerenchyma interrupted by parenchyma.
4- The vascular bundles which are numerous and scattered through
the whole of the ground tissue are enclosed in a sheath of
sclerenchyma. This sheath of mechanical tissue is thickest on the
inner and outer sides of the bundle.

Protoxylem lacunae

In the shoot xylem of many monocotyledons, the stretched non-

functioning elements are partly collapsed ‫ تضعف أو تنهار‬but not
obliterated, instead, open canals, the so called, protoxylem lacunae,
surrounded by parenchyma cells, appear in their places.

127
Structure of primary body in roots

The first root of a seed plant develops from the apical meristem at
the root end of the embryo. This root is called taproot, or primary
root.

In the gymnosperms and dicotyledons, the taproot and its branched

lateral roots comprise the root system.

In the monocotyledons, the first root commonly lives a relatively

short time and the root system is formed by adventitious roots
arising on the shoot, often in connection with axillary buds.
Although these roots become branched they form a rather fibrous
root system.

The internal organization of the root is variable but generally

simpler and more primitive than that of the stem. The root is an
axial structure with no leaf-like organs and with no division into
nodes and internodes.

A transverse section through a root in the primary state of growth

shows a clear separation between the usual three tissue systems,
the epidermis (dermal tissue system), the cortex (ground tissue
system), and the vascular tissue system. The root cap, which

128
covers the apical meristem of the root, is also a part of the primary
body.

* Epidermis or piliferous layer

It is the outermost covering of the root formed by single layer of

compactly arranged, barrel-shaped, parenchyma cells. The cells
are characteristically thin-walled since they are involved in
absorption of water. A cuticle and stomata are absent.

Some of the cells are produced into long unicellular projections

called root hairs. Hence, the epidermis is also known as piliferous
layer. This single layer of cells is derived from the dermatogen.
Root hairs are confined to the zone of root hairs near the tip.

Each root hair arises from a separate epidermal cell, as a lateral

outgrowth occupied by a large vacuole continuous with that of the
dermal cell and filled with sap.

The epidermal cell walls composed of cellulose and pectin

substances. The root hair and the epidermal cell constitute a single
cell having a thin living layer of cytoplasm. The nucleus is usually
lies near the end of the hair.

129
In some herbaceous perennials the epidermis remains for a long
time or permanently, as a protective tissue. Its walls increase in
thickness, and the cell lumen sometimes become filled with deeply
coloured substances.

* Cortex

The cortex of the root is relatively wider than that of the stem. It is
derived from the periblem. Cortex is a major component of the
ground tissue of root. It is represented by several layers of loosely
arranged parenchyma cells. Intercellular spaces are prominent.
The cortical cells are thin walled, rounded with well-developed
intercellular spaces, free from chlorophyll serving the storage
function. Aerenchymatic root cortex is common in hydrophytes
and plants of moist habitats.

In many monocotyledons abundant sclerenchyma may develop

in addition to parenchyma.

The cortex is mainly meant for storage of water. The cells also
allow a free movement of water into the xylem vessels.

130
* Endodermis

It is the innermost layer of cortex formed by compactly arranged

barrel-shaped cells. The cells in the endodermis are characterized
by the presence of thickening on their radial walls. These
thickenings are known as casparian thickenings. They are formed
by the deposition of a waxy substance called suberin. The
casperian thickenings play an important role in creating and
maintaining a physical force called root pressure.

Some of the cells in the endodermis are thin-walled and are

known as passage cells. The passage cells allow water to pass into
the xylem vessels.

In the absence of secondary growth (most monocotyledons and

few dicotyledons) the endodermis commonly undergoes certain
wall modifications result in the occurrence of thick-walled
endodermis. Thickening may confine to radial and inner tangential
walls “Zea root” or occurs on all the walls of endodermal cells
facing the phloem. Other endodermal cells facing the xylem
without casperian strips are called the passage cells, which allow a
limited transfer of materials between the cortex and the vascular
cylinder.

131
* Pericycle
The layer next to the endodermis is commonly known as
pericycle. The pericycle of relatively young roots consists of thin-
walled parenchyma. It makes the outer boundary of the primary
vascular cylinder of the dicotyledonous roots. It may be uniseriate
or multiseriate.

The lateral roots in dicots arise in this tissue. The phellogen and
part of vascular cambium originate in the pericycle.

* Vascular system or cylinder

The vascular cylinder (stele) comprises the vascular tissues in

radial arrangement.

The xylem frequently forms a solid core with ridge-like

projections, seen as radial files of cells in transverse sections,
extending towards the pericycle. Generally the xylem forms
discrete strands, alternating with the phloem strands. Sometimes
the xylem occupies the centre, with the strand-like parts projecting
from the central core like ridges. If xylem is not differentiated in
the centre, the centre is occupied by pith. The root typically shows
an exarch xylem, i.e., the protoxylem is located near the periphery

132
of the vascular cylinder, the metaxylem farther inward. The
protoxylem consists of annular and spiral vessels whereas
metaxylem of reticulate and pitted vessels.

The number of xylem ridges varies in different species and among

roots of the same plant, and in relation to this variation the roots
are called diarch, triarch, tetrarch, etc., or polyarch.

The phloem strand consists of sieve tubes, companion cells and

phloem parenchyma. The parenchymatous conjunctive tissue
occurs in between xylem and phloem strands.

The first mature sieve elements, those of the protophloem, occupy

a peripheral position in the vascular cylinder; the metaphloem
occurs further inward. Thus, the primary phloem show a
centripetal order of differentiation as does the xylem.

Companion cells are characteristic of the metaphloem but may be

absent in the protophloem.

In roots having secondary growth, the cells located between the

xylem and the phloem; eventually function as a vascular cambium.
In roots without secondary growth the cells in the same location
mature as parenchyma or sclerenchyma cells.

133
* Pith

At the centre there is small parenchymatous pith. It may be even

absent. It is frequent in the roots of herbaceous dicotyledons and
monocotyledons, but in woody dicotyledons and conifers the
xylem groups often meet in the center.

Monocoteledonous root
1. Peliferous layer is the outermost single layer made from
compactly arranged parenchymatous cells without intercellular
space.

2. Cortex is a multi-layered well developed and made from oval

parenchymatous cells with intercellular spaces. The intercellular
spaces usually help in gaseous exchanges, storage of starch, etc. In
monocots and several old roots, few layers of cortex just below
Peliferous layer give rise to a single or multilayered cuticularized
sclerenchymatous region called exodermis. Cortex helps in
mechanical support to the roots (like hypodermis to stem).

3. Endodermis is innermost layer of cortex made from barrel

shaped parenchyma. It forms a definite ring around the stele. These

134
cells are characterized by the presence of casparian stripes. It is
deposition of suberin and lignin, and their radial and tangential
walls.

4. Due to presence of casparian stripes, endodermis forms water

tight jacket around the vascular tissues, hence it is also called
biological barrier.
It regulates the inward and outward flow of water and minerals and
prevents diffusion of air into xylem elements.

5. Pericycle is uniseriate and made from thin walled

parenchymatous cells. Several lateral roots arise from this
layer. Hence, lateral roots are endogenous in origin.

6. Vascular bundle is radial, arranged in a

ring, polyarch (presence of many alternating xylem and phloem
bundles).

Xylem and phloem are found at different radii alternating with

each other. The number of xylem and phloem vary from, 8 to 46.
The xylem is exarch, i.e. the protoxylem lies towards periphery
and metaxylem toward center.

135
The phloem is also exarch (protophloem towards the periphery and
metaphloem towards the center).
Secondary growth is absent in monocot roots due to lack of
vascular and cork cambium. Conjunctive tissue is
parenchymatous tissues which separates xylem and phloem
bundles.

Pith is large, well developed portion of monocot root. It occupies

the central portion and made from thin walled parenchymatou
tissue with intercellular spaces. It contains abundant amount of
starch grains.
Monocot root have bands of vascular bundles. Vessels are not
found in linear rows, but arranged in V-shaped structure.

136
Comparison between the structures of stem and root:
Stem Root
1. Cuticle Present, thick / thin Usually absent
2. Trichomes When present, is multi- Root hairs are unicellular
cellular or unicellular
3. Stomata Present Absent
4. Hypodermis collenchymatous sclerenchymatous.
5. Cortex heterogeneous, with Homogenous with cells
collenchyma, and of parenchyma.
parenchyma
6. Endodermis Usually absent Generally present
7. Pericycle absent One layered
8.Vascular Many, conjoint, 2-12, radial, exarch, form
bundles collateral or bicollateral, a broken ring.
regularly arranged
forming a broken ring
or scattered in ground
tissue, endarch.
9. Pith Well marked and large Relatively small or
or not differentiated absent.
10. Branching Exogenous, originating Endogenous, arising form
from cortex pericycle.

137
The leaf

Foliage leaves which, among the Angiosperm, exhibit great

variation in anatomical and morphological structure. The foliage
leaves in certain angiosperm genera are sessile and consist almost
entirely of lamina, but in most genera several distinct parts can be
distinguished, i.e. the leaf base, the petiole, and the lamina.
Histologically the leaf is composed of three types of tissue
systems: epidermis, mesophyll, and vascular tissue.

The epidermis

The epidermis of leaves of different plants varies in the number of

layers, its shape, structure, arrangement of stomata, appearance and
arrangement of trichomes, and occurrence of specialized cells.
Because of the usually flat structure of the leaf, a distinction is
made between the epidermal tissues of the two surfaces of the leaf;
that surface of the leaf that is closer to the internodes above it and
which usually faces upwards is referred to as the adaxial surface
(upper epidermis), and the other surface as the abaxial surface
(lower epidermis).

138
The mesophyll

The mesophyll comprises the parenchymatous tissue internal to the

epidermis. It is usually undergoes differentiation to form the
photosynthetic tissues and so contains chloroplasts.
In many plants especially among dicotyledons, two types of
parenchyma can be distinguished in the mesophyll: palisade
parenchyma and spongy parenchyma.
The cells of typical palisade parenchyma are elongated, and in
cross section of the leaf they are rod-shaped and appear to be
arranged in rows, while in section parallel to the leaf surface these
cells are seen to be rounded and separated or only slightly attached
to one another.
The palisade cells are found immediately below the epidermis, but
sometimes a hypodermis may be present between the epidermis
and the palisade tissue.
The cells of the palisade parenchyma may be arranged in one or
more layers and in the latter case the length of cells in the different
rows may be equal, or they may become shorter towards the center
of the mesophyll. The palisade tissue is usually found on the
adaxial surface of the leaf.

139
In certain plants, including many xeromorphic species palisade
parenchyma is present on both sides of the leaf with the result that
only a small strip of spongy parenchyma is present in the central
portion of the lamina.
A leaf in which the palisade parenchyma occurs on one side of the
leaf and the spongy parenchyma on the other is termed
dorsiventral or bifacial. When the palisade parenchyma is present
on both sides of the leaf, the leaf is said to be isolateral or
isobilateral.
The cells of spongy parenchyma are variously shaped. They may
resemble the palisade cells, or have equal diameters, or be
elongated in a direction parallel to the leaf surface.
However, a characteristic of all spongy parenchyma cells is the
presence of lobes by which the neighboring cells are connected.
In certain plants, such as Zea and many other grasses
(monocotyledons), the mesophyll cells are more or less uniform in
shape. In certain species of dicotyledons (e.g. Eucalyptus and
Atriplex), it is not possible to distinguish between the two types of
parenchyma, and the mesophyll is entirely composed of palisade
cells.

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The palisade tissue has become specialized in such a way that the
efficiency of photosynthesis has been increased.
In mesophyll that can be clearly divided into palisade and spongy
parenchyma the large majority of the chloroplasts are found in the
palisade cells.
The specialization of the palisade tissue that result in more
efficient photosynthesis is brought about not only by the increased
number of chloroplasts in the cells but also by the dimensions of its
free surface area resulting from the well-developed system of
intercellular spaces which facilitates rapid gas exchange.

Vascular system of the leaf

In angiosperms, two main types of venation are usually

distinguished – reticulate and parallel venation. In leaves with
reticulate venation, which is the commonest type among the
dicotyledons, the veins are of different size depending on the
degree of branching; central vein (or midrib), secondary veins
and so on.
In leaves with parallel venation, which is the commonest type
among the monocotyledons, the main vein continue throughout
the entire leaf and are almost parallel for most of their length but

141
approach one another and fuse at the leaf tip or both at the leaf tip
and base. These parallel veins are interconnected by very thin
commissural bundles, which are scattered throughout the lamina.
The large veins in dicotyledonous leaves may consist of both
primary and secondary tissues, while the smaller veins consist of
primary tissues only. The large and medium-sized veins contain
vessels and sieve tubes.
In the smallest veins, the tracheary elements are tracheids with
annular and spiral wall thickenings.
The structure of vascular bundle corresponds to that in the stem.
The collateral bundles of the leaf occupy such a position that the
xylem is directed towards the upper, and the phloem towards the
lower surface of the leaf.
A sheath of parenchyma surrounds the large vascular bundles of
the midrib as well as the lateral vascular bundles, which, is in close
connection with the mesophyll cells. The cells of these sheaths are
usually elongated and have thin cell walls.
Strands of supporting tissue, usually collenchyma in
dicotyledonous leaf, are frequently present on one or both faces of
the bundle, especially on the phloem side.

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Monocotyledonous leaves develop relatively large amounts of
sclerenchyma in association with the vascular bundles or in
separate strands.
In grasses, strands of fibers occur on one or both sides of vascular
bundles, and are connected to the bundle sheaths as well as to the
epidermis.

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I: What are the differences between each of the following?
1- Epigeal and hypogeal germination
2- Viability and dormancy
3- Fibrous roots and prop roots

II: Summarize the types of subterranean (underground) stems

III: Enumerate the main characteristics of permanent tissue

IV: Enumerate the anatomical features of the dicotyledonous root,

indicating your answer with a labeled diagram.

V: : Read the following sentences and write the scientific concepts:

1- The unit of construction of the body of living organisms
2- An essential non- living constituent delimiting the plant cells
3- The second stage of the cell division

144
References

Beck, CB (2010). An introduction to plant structure and

development: plant anatomy for the twenty-first century. Cambridge
university press
Esau, K. (1965). Plant anatomy, 2nd ed. John wiley&Sons, Inc. New
York, London, Sydney.

Evert, RF (2006). Esau's plant anatomy: meristems, cells, and tissues of

the plant body: their structure, function, and development. John
wiley&Sons, Inc.
Fahn, A. (1967). Plant anatomy, Pergamon press Ltd Oxford.

Metcalfe, CR., Chalk, L (1950) - Anatomy of the dicotyledons, Vols. 1 &

2, cabdirect.org

Tomlinson, P.B. (1970). Monocotyledons — Towards an Understanding

of their Morphology and Anatomy. Academic Press Inc. (London) Ltd.
Published by Elsevier Ltd.
Tomlinson, PB. (1977). Plant morphology and Anatomy in the Tropics-
The Need for Integrated Approaches. Annals of the Missouri Botanical
Garden, JSTOR

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