6.1 The Tissues
6.2 The Tissue
System
6.3 Anatomy of
Dicotyledonous
and
Monocotyledonous
Plants
6.4 Secondary
Growth
You can very easily see the structural similarities and variations in the
external morphology of the larger living organism, both plants and
animals. Similarly, if we were to study the internal structure, one also
finds several similarities as well as differences. This chapter introduces
you to the internal structure and functional organisation of higher plants.
Study of internal structure of plants is called anatomy. Plants have cells
as the basic unit, cells are organised into tissues and in turn the tissues
are organised into organs. Different organs in a plant show differences in
their internal structure. Within angiosperms, the monocots and dicots
are also seen to be anatomically different. Internal structures also show
adaptations to diverse environments.
6.1 THE TISSUES
A tissue is a group of cells having a common origin and usually performing
a common function. A plant is made up of different kinds of tissues. Tissues
are classified into two main groups, namely, meristematic and permanent
tissues based on whether the cells being formed are capable of dividing
or not.
6.1.1 Meristematic Tissues
Growth in plants is largely restricted to specialised regions of active cell division
called meristems (Gk. meristos: divided). Plants have different kinds of
meristems. The meristems which occur at the tips of roots and shoots and
produce primary tissues are called apical meristems (Figure 6.1).
Root apical meristem occupies the tip of a root while the shoot apical
meristem occupies the distant most region of the stem axis. During the
formation of leaves and elongation of stem, some cells ‘left behind’ from
shoot apical meristem, constitute the axillary bud. Such buds are present
in the axils of leaves and are capable of forming a branch or a flower. The
meristem which occurs between mature tissues is known as intercalary
meristem. They occur in grasses and regenerate parts removed by the
grazing herbivores. Both apical meristems and intercalary meristems are
primary meristems because they appear early in life of a plant and
contribute to the formation of the primary plant body.
The meristem that occurs in the mature regions of roots and shoots of
many plants, particularly those that produce woody axis and appear
later than primary meristem is called the secondary or lateral meristem.
They are cylindrical meristems. Fascicular vascular cambium,
interfascicular cambium and cork-cambium are examples of lateral
meristems. These are responsible for producing the secondary tissues.
Following divisions of cells in both primary and as well as secondary
meristems, the newly formed cells become structurally and functionally
specialised and lose the ability to divide. Such cells are termed permanent
or mature cells and constitute the permanent tissues. During the
formation of the primary plant body, specific regions of the apical meristem
produce dermal tissues, ground tissues and vascular tissues.
6.1.2 Permanent Tissues
The cells of the permanent tissues do not generally
divide further. Permanent tissues having all cells
similar in structure and function are called simple
tissues. Permanent tissues having many different
types of cells are called complex tissues.
6.1.2.1 Simple Tissues
A simple tissue is made of only one type of cells.
The various simple tissues in plants are
parenchyma, collenchyma and sclerenchyma
(Figure 6.2). Parenchyma forms the major
component within organs. The cells of the
parenchyma are generally isodiametric. They
may be spherical, oval, round, polygonal or
elongated in shape. Their walls are thin and made
up of cellulose. They may either be closely packed
or have small intercellular spaces. The
parenchyma performs various functions like
photosynthesis, storage, secretion.
The collenchyma occurs in layers below the
epidermis in most of the dicotyledonous plants. It is
found either as a homogeneous layer or in patches.
It consists of cells which are much thickened at the
corners due to a deposition of cellulose,
hemicellulose and pectin. Collenchymatous cells
may be oval, spherical or polygonal and often
contain chloroplasts. These cells assimilate food
when they contain chloroplasts. Intercellular spaces
are absent. They provide mechanical support to the
growing parts of the plant such as young stem and
petiole of a leaf.
Sclerenchyma consists of long, narrow cells
with thick and lignified cell walls having a few or
numerous pits. They are usually dead and without
protoplasts. On the basis of variation in form,
structure, origin and development, sclerenchyma
may be either fibres or sclereids. The fibres are
thick-walled, elongated and pointed cells,
generally occuring in groups, in various parts of
the plant. The sclereids are spherical, oval or
cylindrical, highly thickened dead cells with very
narrow cavities (lumen). These are commonly found in the fruit
walls of nuts; pulp of fruits like guava, pear and sapota; seed
coats of legumes and leaves of tea. Sclerenchyma provides
mechanical support to organs.
6.1.2.2 Complex Tissues
The complex tissues are made of more than one type of cells
and these work together as a unit. Xylem and phloem constitute
the complex tissues in plants (Figure 6.3).
Xylem functions as a conducting tissue for water and
minerals from roots to the stem and leaves. It also provides
mechanical strength to the plant parts. It is composed of four
different kinds of elements, namely, tracheids, vessels, xylem
fibres and xylem parenchyma. Gymnosperms lack vessels in
their xylem. Tracheids are elongated or tube like cells with
thick and lignified walls and tapering ends. These are dead and
are without protoplasm. The inner layers of the cell walls have
thickenings which vary in form. In flowering plants, tracheids
and vessels are the main water transporting elements. Vessel is
a long cylindrical tube-like structure made up of many cells
called vessel members, each with lignified walls and a large
central cavity. The vessel cells are also devoid of protoplasm.
Vessel members are interconnected through perforations in their
common walls. The presence of vessels is a characteristic feature
of angiosperms. Xylem fibres have highly thickened walls and
obliterated central lumens. These may either be septate or
aseptate. Xylem parenchyma cells are living and thin-walled,
and their cell walls are made up of cellulose. They store food
materials in the form of starch or fat, and other substances like
tannins. The radial conduction of water takes place by the ray
parenchymatous cells.
Primary xylem is of two types – protoxylem and metaxylem.
The first formed primary xylem elements are called protoxylem
and the later formed primary xylem is called metaxylem. In
stems, the protoxylem lies towards the centre (pith) and the
metaxylem lies towards the periphery of the organ. This type
of primary xylem is called endarch. In roots, the protoxylem
lies towards periphery and metaxylem lies towards the centre.
Such arrangement of primary xylem is called exarch.
Phloem transports food materials, usually from leaves to
other parts of the plant. Phloem in angiosperms is composed
of sieve tube elements, companion cells, phloem parenchyma
and phloem fibres. Gymnosperms have albuminous cells and sieve cells.
They lack sieve tubes and companion cells. Sieve tube elements are
also long, tube-like structures, arranged longitudinally and are
associated with the companion cells. Their end walls are perforated in a
sieve-like manner to form the sieve plates. A mature sieve element
possesses a peripheral cytoplasm and a large vacuole but lacks a nucleus.
The functions of sieve tubes are controlled by the nucleus of companion
cells. The companion cells are specialised parenchymatous cells, which
are closely associated with sieve tube elements. The sieve tube elements
and companion cells are connected by pit fields present between their
common longitudinal walls. The companion cells help in maintaining the
pressure gradient in the sieve tubes. Phloem parenchyma is made up
of elongated, tapering cylindrical cells which have dense cytoplasm and
nucleus. The cell wall is composed of cellulose and has pits through which
plasmodesmatal connections exist between the cells. The phloem
parenchyma stores food material and other substances like resins, latex
and mucilage. Phloem parenchyma is absent in most of the
monocotyledons. Phloem fibres (bast fibres) are made up of
sclerenchymatous cells. These are generally absent in the primary phloem
but are found in the secondary phloem. These are much elongated,
unbranched and have pointed, needle like apices. The cell wall of phloem
fibres is quite thick. At maturity, these fibres lose their protoplasm and
become dead. Phloem fibres of jute, flax and hemp are used commercially.
The first formed primary phloem consists of narrow sieve tubes and is
referred to as protophloem and the later formed phloem has bigger sieve
tubes and is referred to as metaphloem.
6.2 THE TISSUE SYSTEM
We were discussing types of tissues based on the types of cells present.
Let us now consider how tissues vary depending on their location in the
plant body. Their structure and function would also be dependent on
location. On the basis of their structure and location, there are three types
of tissue systems. These are the epidermal tissue system, the ground or
fundamental tissue system and the vascular or conducting tissue system.
6.2.1 Epidermal Tissue System
The epidermal tissue system forms the outer-most covering of the whole
plant body and comprises epidermal cells, stomata and the epidermal
appendages – the trichomes and hairs. The epidermis is the outermost
layer of the primary plant body. It is made up of elongated, compactly
arranged cells, which form a continuous layer. Epidermis is usually single-
layered. Epidermal cells are parenchymatous with a small amount of
cytoplasm lining the cell wall and a large vacuole. The outside of the
epidermis is often covered with a waxy thick layer called the cuticle which
prevents the loss of water. Cuticle is absent in roots. Stomata are structures
present in the epidermis of leaves. Stomata regulate the process of
transpiration and gaseous exchange. Each stoma is composed of two bean-
shaped cells known as guard cells which enclose stomatal pore. In grasses,
the guard cells are dumb-bell shaped. The outer walls of guard cells (away
from the stomatal pore) are thin and the inner walls (towards the stomatal
pore) are highly thickened. The guard cells possess chloroplasts and
regulate the opening and closing of stomata. Sometimes, a few epidermal
cells, in the vicinity of the guard cells become specialised in their shape and
size and are known as subsidiary cells. The stomatal aperture, guard cells and
the surrounding subsidiary cells are together called stomatal apparatus (Figure 6.4).
The cells of epidermis bear a number of hairs. The root hairs are
unicellular elongations of the epidermal cells and help absorb water and
minerals from the soil. On the stem the epidermal hairs are called
trichomes. The trichomes in the shoot system are usually multicellular.
They may be branched or unbranched and soft or stiff. They may even
be secretory. The trichomes help in preventing water loss due to
transpiration.
6.2.2 The Ground Tissue System
All tissues except epidermis and vascular bundles constitute the ground
tissue. It consists of simple tissues such as parenchyma, collenchyma
and sclerenchyma. Parenchymatous cells are usually present in cortex,
pericycle, pith and medullary rays, in the primary stems and roots. In
leaves, the ground tissue consists of thin-walled chloroplast containing
cells and is called mesophyll.
6.2.3 The Vascular Tissue System
The vascular system consists of complex tissues,
the phloem and the xylem.The xylem and
phloem together constitute vascular bundles
(Figure 6.5). In dicotyledonous stems, cambium
is present between phloem and xylem. Such
vascular bundles because of the presence of
cambium possess the ability to form secondary
xylem and phloem tissues, and hence are called
open vascular bundles. In the monocotyledons,
the vascular bundles have no cambium present
in them. Hence, since they do not form secondary
tissues they are referred to as closed. When
xylem and phloem within a vascular bundle are
arranged in an alternate manner along the
different radii, the arrangement is called radial
such as in roots. In conjoint type of vascular
bundles, the xylem and phloem are jointly
situated along the same radius of vascular
bundles. Such vascular bundles are common
in stems and leaves. The conjoint vascular
bundles usually have the phloem located only
on the outer side of xylem.
6.3 ANATOMY OF DICOTYLEDONOUS AND
MONOCOTYLEDONOUS PLANTS
For a better understanding of tissue
organisation of roots, stems and leaves, it is
convenient to study the transverse sections of
the mature zones of these organs.
6.3.1 Dicotyledonous Root
Look at Figure 6.6 (a), it shows the transverse
section of the sunflower root. The internal tissue
organisation is as follows:
The outermost layer is epiblema. Many of
the cells of epiblema protrude in the form of
unicellular root hairs. The cortex consists of
several layers of thin-walled parenchyma cells
with intercellular spaces. The innermost
layer of the cortex is called endodermis.
It comprises a single layer of barrel-shaped
cells without any intercellular spaces. The
tangential as well as radial walls of the
endodermal cells have a deposition of
water-impermeable, waxy material suberin
in the form of casparian strips. Next to
endodermis lies a few layers of thick-walled
parenchyomatous cells referred to as
pericycle. Initiation of lateral roots and
vascular cambium during the secondary
growth takes place in these cells. The pith
is small or inconspicuous. The
parenchymatous cells which lie between
the xylem and the phloem are called
conjuctive tissue. There are usually two
to four xylem and phloem patches. Later,
a cambium ring develops between the
xylem and phloem. All tissues on the
innerside of the endodermis such as
pericycle, vascular bundles and pith
constitute the stele.
6.3.2 Monocotyledonous Root
The anatomy of the monocot root is similar
to the dicot root in many respects (Figure
6.6 b). It has epidermis, cortex, endodermis,
pericycle, vascular bundles and pith. As
compared to the dicot root which have fewer
xylem bundles, there are usually more than
six (polyarch) xylem bundles in the monocot
root. Pith is large and well developed.
Monocotyledonous roots do not undergo
any secondary growth.
6.3.3 Dicotyledonous Stem
The transverse section of a typical young
dicotyledonous stem shows that the epidermis
is the outermost protective layer of the stem
(Figure 6.7 a). Covered with a thin layer of cuticle, it may bear trichomes and
a few stomata. The cells arranged in multiple layers between epidermis and
pericycle constitute the cortex. It consists of three sub-zones. The outer
hypodermis, consists of a few layers of collenchymatous cells just below the
epidermis, which provide mechanical strength to the young stem. Cortical
layers below hypodermis consist of rounded thin walled parenchymatous
cells with conspicuous intercellular spaces. The innermost layer of the cortex
is called the endodermis. The cells of the endodermis are rich in starch
grains and the layer is also referred to as the starch sheath. Pericycle is
present on the inner side of the endodermis and above the phloem in the
form of semi-lunar patches of sclerenchyma. In between the vascular bundles
there are a few layers of radially placed parenchymatous cells, which constitute
medullary rays. A large number of vascular bundles are arranged in a ring ;
the ‘ring’ arrangement of vascular bundles is a characteristic of dicot stem.
Each vascular bundle is conjoint, open, and with endarch protoxylem. A
large number of rounded, parenchymatous cells with large intercellular
spaces which occupy the central portion of the stem constitute the pith.
6.3.4 Monocotyledonous Stem
The monocot stem has a sclerenchymatous hypodermis, a large number
of scattered vascular bundles, each surrounded by a sclerenchymatous
bundle sheath, and a large, conspicuous parenchymatous ground tissue
(Figure 6.7b). Vascular bundles are conjoint and closed. Peripheral
vascular bundles are generally smaller than the centrally located ones.
The phloem parenchyma is absent, and water-containing cavities are
present within the vascular bundles.
6.3.5 Dorsiventral (Dicotyledonous) Leaf
The vertical section of a dorsiventral leaf through the lamina shows three
main parts, namely, epidermis, mesophyll and vascular system. The
epidermis which covers both the upper surface (adaxial epidermis) and
lower surface (abaxial epidermis) of the leaf has a conspicuous cuticle.
The abaxial epidermis generally bears more stomata than the adaxial
epidermis. The latter may even lack stomata. The tissue between the upper
and the lower epidermis is called the mesophyll. Mesophyll, which
possesses chloroplasts and carry out photosynthesis, is made up of
parenchyma. It has two types of cells – the palisade parenchyma and
the spongy parenchyma. The adaxially placed palisade parenchyma is
made up of elongated cells, which are arranged vertically and parallel to
each other. The oval or round and loosely arranged spongy parenchyma
is situated below the palisade cells and extends to the lower epidermis.
There are numerous large spaces and air cavities between these cells.
Vascular system includes vascular bundles, which can be seen in the
veins and the midrib. The size of the vascular bundles are dependent on
the size of the veins. The veins vary in thickness in the reticulate venation
of the dicot leaves. The vascular bundles are surrounded by a layer of
thick walled bundle sheath cells. Look at Figure 6.8 (a) and find the
position of xylem in the vascular bundle.
6.3.6 Isobilateral (Monocotyledonous) Leaf
The anatomy of isobilateral leaf is similar to that of the dorsiventral leaf in
many ways. It shows the following characteristic differences. In an
6.4 SECONDARY GROWTH
The growth of the roots and stems in
length with the help of apical meristem is
called the primary growth. Apart from
primary growth most dicotyledonous
plants exhibit an increase in girth. This
increase is called the secondary growth.
The tissues involved in secondary growth
are the two lateral meristems: vascular
cambium and cork cambium.
6.4.1 Vascular Cambium
The meristematic layer that is responsible
for cutting off vascular tissues – xylem and
pholem – is called vascular cambium. In
the young stem it is present in patches as
a single layer between the xylem and
phloem. Later it forms a complete ring.
6.4.1.1 Formation of cambial ring
In dicot stems, the cells of cambium present
between primary xylem and primary
phloem is the intrafascicular cambium.
The cells of medullary rays, adjoining these intrafascicular cambium become
meristematic and form the interfascicular cambium. Thus, a continuous
ring of cambium is formed.
6.4.1.2 Activity of the cambial ring
The cambial ring becomes active and begins to cut off new cells, both
towards the inner and the outer sides. The cells cut off towards pith,
mature into secondary xylem and the cells cut off towards periphery
mature into secondary phloem. The cambium is generally more active
on the inner side than on the outer. As a result, the amount of secondary
xylem produced is more than secondary phloem and soon forms a
compact mass. The primary and secondary phloems get gradually
crushed due to the continued formation and accumulation of secondary
xylem. The primary xylem however remains more or less intact, in or
around the centre. At some places, the cambium forms a narrow band of
parenchyma, which passes through the secondary xylem and the
secondary phloem in the radial directions. These are the secondary
medullary rays (Figure 6.9).
6.4.1.3 Spring wood and autumn wood
The activity of cambium is under the control of many physiological and
environmental factors. In temperate regions, the climatic conditions are
not uniform through the year. In the spring season, cambium is very
active and produces a large number of xylary elements having vessels
with wider cavities. The wood formed during this season is called spring
wood or early wood. In winter, the cambium is less active and forms
fewer xylary elements that have narrow vessels, and this wood is called
autumn wood or late wood.
The spring wood is lighter in colour and has a lower density whereas
the autumn wood is darker and has a higher density. The two kinds of
woods that appear as alternate concentric rings, constitute an annual ring.
Annual rings seen in a cut stem give an estimate of the age of the tree.
6.4.1.4 Heartwood and sapwood
In old trees, the greater part of secondary xylem is dark brown due to
deposition of organic compounds like tannins, resins, oils, gums, aromatic
substances and essential oils in the central or innermost layers of the stem.
These substances make it hard, durable and resistant to the attacks of micro-
organisms and insects. This region comprises dead elements with highly
lignified walls and is called heartwood. The heartwood does not conduct
water but it gives mechanical support to the stem. The peripheral region of
the secondary xylem, is lighter in colour and is known as the sapwood. It is
involved in the conduction of water and minerals from root to leaf.
6.4.2 Cork Cambium
As the stem continues to increase in girth due to the activity of vascular
cambium, the outer cortical and epidermis layers get broken and need to
be replaced to provide new protective cell layers. Hence, sooner or later,
another meristematic tissue called cork cambium or phellogen develops,
usually in the cortex region. Phellogen is a couple of layers thick. It is
made of narrow, thin-walled and nearly rectangular cells. Phellogen cuts
off cells on both sides. The outer cells differentiate into cork or phellem
while the inner cells differentiate into secondary cortex or phelloderm.
The cork is impervious to water due to suberin deposition in the cell wall.
The cells of secondary cortex are parenchymatous. Phellogen, phellem,
and phelloderm are collectively known as periderm. Due to activity of
the cork cambium, pressure builds up on the remaining layers peripheral
to phellogen and ultimately these
layers die and slough off. Bark is a
non-technical term that refers to all
tissues exterior to the vascular
cambium, therefore including
secondary phloem. Bark refers to a
number of tissue types, viz.,
periderm and secondary phloem.
Bark that is formed early in the
season is called early or soft bark.
Towards the end of the season, late
or hard bark is formed. Name the
various kinds of cell layers which
constitute the bark.
At certain regions, the phellogen
cuts off closely arranged
parenchymatous cells on the outer
side instead of cork cells. These
parenchymatous cells soon rupture
the epidermis, forming a lens-
shaped openings called lenticels.
Lenticels permit the exchange of
gases between the outer atmosphere
and the internal tissue of the stem.
These occur in most woody trees
(Figure 6.10).
6.4.3 Secondary Growth in
Roots
In the dicot root, the vascular
cambium is completely secondary in
origin. It originates from the tissue
located just below the phloem
bundles, a portion of pericycle tissue,
above the protoxylem forming a
complete and continuous wavy ring,
which later becomes circular (Figure
6.11). Further events are similar to
those already described above for a
dicotyledon stem.
Secondary growth also occurs in stems and roots of gymnosperms.
However, secondary growth does not occur in monocotyledons.
SUMMARY
Anatomically, a plant is made of different kinds of tissues. The plant tissues are
broadly classified into meristematic (apical, lateral and intercalary) and permanent
(simple and complex). Assimilation of food and its storage, transportation of water,
minerals and photosynthates, and mechanical support are the main functions of
tissues. There are three types of tissue systems – epidermal, ground and vascular.
The epidermal tissue systems are made of epidermal cells, stomata and the
epidermal appendages. The ground tissue system forms the main bulk of the
plant. It is divided into three zones – cortex, pericycle and pith. The vascular
tissue system is formed by the xylem and phloem. On the basis of presence of
cambium, location of xylem and phloem, the vascular bundles are of different
types. The vascular bundles form the conducting tissue and translocate water,
minerals and food material.
Monocotyledonous and dicotyledonous plants show marked variation in their
internal structures. They differ in type, number and location of vascular bundles.
The secondary growth occurs in most of the dicotyledonous roots and stems and
it increases the girth (diameter) of the organs by the activity of the vascular cambium
and the cork cambium. The wood is actually a secondary xylem. There are different
types of wood on the basis of their composition and time of production.
EXERCISES
- State the location and function of different types of meristems.
- Cork cambium forms tissues that form the cork. Do you agree with this
statement? Explain. - Explain the process of secondary growth in the stems of woody angiosperms
with the help of schematic diagrams. What is its significance? - Draw illustrations to bring out the anatomical difference between
(a) Monocot root and Dicot root
(b) Monocot stem and Dicot stem - Cut a transverse section of young stem of a plant from your school garden and
observe it under the microscope. How would you ascertain whether it is a
monocot stem or a dicot stem? Give reasons. - The transverse section of a plant material shows the following anatomical
features – (a) the vascular bundles are conjoint, scattered and surrounded by a
sclerenchymatous bundle sheaths. (b) phloem parenchyma is absent. What
will you identify it as? - Why are xylem and phloem called complex tissues?
- What is stomatal apparatus? Explain the structure of stomata with a labelled
diagram. - Name the three basic tissue systems in the flowering plants. Give the tissue
names under each system. - How is the study of plant anatomy useful to us?
11, What is periderm? How does periderm formation take place in the dicot stems? - Describe the internal structure of a dorsiventral leaf with the help of labelled
diagrams.