SBN 222: PLANT STRUCTURE & FUNCTION @2019

LECTURE ONE
PLANT CELL STRUCTURE AND TISSUE SYSTEMS
Most organelles are common to both animal and plant cells. However, plant cells also have
features that animal cells do not have. Many plant cells are green. Why? Plant cells also usually
have a distinct shape. The rigid exterior around the cells is necessary to allow the plants to grow
upright. Animal cells do not have these rigid exteriors. There are other distinct differences
between plant and animal cells.
Plant Cell Structure
Plants are unique amongst the eukaryotes, organisms whose cells have membrane-enclosed
nuclei and organelles, because they can manufacture their own food. Chlorophyll, which gives
plants their green colour, enables them to use sunlight to convert water and carbon dioxide into
sugars and carbohydrates; chemicals the cell uses for fuel.
Like the fungi, another kingdom of eukaryotes, plant cells have retained the protective cell wall
structure of their prokaryotic ancestors. The basic plant cell has the basic construction of a
typical eukaryote cell, but does not have centrioles, lysosomes, cilia, or flagella as does the
animal cell. Plant cells do have specialized structures: a rigid cell wall, central vacuole,
plasmodesmata, and chloroplasts (figure 1).

Figure 1: A diagram showing parts of the Plant cell

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 Cell Wall - Like their prokaryotic ancestors, plant cells have a rigid wall surrounding the
plasma membrane. It is a far more complex structure, however, and serves a variety of
functions, from protecting the cell to regulating the life cycle of the plant organism.
 Chloroplast - The most important characteristic of plants is their ability to
photosynthesize, i.e. make their own food by converting light energy into chemical
energy. This process is carried out in specialized organelles called chloroplasts.
 Endoplasmic Reticulum - The endoplasmic reticulum is a network of sacs that
manufactures, processes, and transports chemical compounds for use inside and outside
of the cell. It is attached to the double-layered nuclear envelope, providing a connection
between the nucleus and the cytoplasm. In plants, it also connects between cells via the
plasmodesmata.
 Golgi Apparatus - The Golgi apparatus is the distribution and shipping department for
the cell's chemical products. It modifies proteins and fats built in the endoplasmic
reticulum and prepares them for export as outside of the cell.
 Microfilaments - Microfilaments are solid rods made of globular proteins called actin.
These filaments are primarily structural in function and are an important component of
the cytoskeleton.
 Microtubules - These straight, hollow cylinders, composed of tubulin protein, are found
throughout the cytoplasm of all eukaryotic cells and perform a number of functions.
 Mitochondria - Mitochondria are oblong shaped organelles found in the cytoplasm of all
eukaryotic cells. In plant cells, they break down carbohydrate and a sugar molecule to
provide energy, particularly when light isn’t available for the chloroplasts to produce
energy.
 Nucleus - The nucleus is a highly specialized organelle that serves as the information and
administrative center of the cell.
 Peroxisomes - Microbodies are a diverse group of organelles that are found in the
cytoplasm, roughly spherical and bound by a single membrane. There are several types of
microbodies but peroxisomes are the most common. Peroxisomes have catalyse enzymes
which breaks down hydrogen peroxide into water and oxygen.
 Plasmodesmata - Plasmodesmata are small tubes that connect plant cells to each other,
providing living bridges between cells.
 Glyoxysomes – Contain isocitrase enzymes important in Citric Acid Cycle/Krebs cycle
 Plasma Membrane - All living cells have a plasma membrane that encloses their
contents. In prokaryotes and plants, the membrane is the inner layer of protection
surrounded by a rigid cell wall. These membranes also regulate the passage of molecules
in and out of the cells.
 Ribosomes- All living cells contain ribosomes, tiny organelles composed of
approximately 60 percent RNA and 40 percent protein. In eukaryotes, ribosomes are
made of four strands of RNA. In prokaryotes, they consist of three strands of RNA.
 Vacuole - Each plant cell has a large, single vacuole that stores compounds, helps in
plant growth, and plays an important structural role for the plant.

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Special Structures in Plant Cells

a) The Cell Wall
A cell wall is a rigid layer that is found outside the cell membrane and surrounds the cell. The
cell wall contains not only cellulose and protein, but other polysaccharides as well. The cell wall
provides structural support and protection. It is the outermost part of the cell. It is made up of
cellulose, pectin, lignin, lipids and proteins. It is the non-living part of the cell. Pores in the cell
wall allow water and nutrients to move into and out of the cell. The cell wall also prevents the
plant cell from bursting when water enters the cell.
The cell wall can be differentiated into three divisions/components:
i) Middle lamella- It is formed during cell division. It is formed as a basic cell wall
ii) Primary wall- Deposition of lignin to the middle lamella forms the primary cell wall.
It is made up of cellulose, unicellular, pectin and some protein and lipids
iii) Secondary wall- It is deposited around the primary cell wall. It is usually hard, tough
and has deposition of lignin.
When a plant matures, the cell wall undergoes some changes/modifications as follows:
i) Lignification- This is the deposition of lignin or the conversion of cell wall into
lignin. It is usually found in the tracheids and vessels in sclerenchyma cells. There are
different patterns of lignification:
i) Annular/ring
ii) Spiral lignification- in a spiral manner
iii) Scalariform/ladder- looks like a ladder
iv) Reticulate- almost like a net
ii) Suberization- This is the deposition of suberin. It is found mainly in cork cells-
outermost part of the epidermis. (Endodermis-innermost part of the cork cells).
Casparian strip in the endodermis also under suberization.
iii) Cutinization- deposition of cutin. Cutin is found in the outermost layer of the
epidermis especially leaves and stems forming a cuticle layer. It helps to regulate
transpiration, gaseous exchange and regulate amount of light absorbed.
iv) Mucilagenous substances- Conversion of cellulose into mucilaginous substances- a
thick glue like substance produced by nealy all plants. It helps to absorb and retain
water e.g. in ladies’ fingers – Okia
v) Deposition of minerals- e.g. Silica (silification) and calcium oxalate (calcification).
Microtubules guide the formation of the plant cell wall. Cellulose is laid down by enzymes to
form the primary cell wall. Some plants also have a secondary cell wall. The secondary wall
contains a lignin, a secondary cell component in plant cells that have completed cell
growth/expansion.
b) The Central Vacuole
Most mature plant cells have a central vacuole that occupies more than 30% of the cell's
volume. The central vacuole can occupy as much as 90% of the volume of certain cells. The
central vacuole is surrounded by a membrane called the tonoplast. The central vacuole has many
functions. Aside from storage, the main role of the vacuole is to maintain turgor pressure against
the cell wall. Proteins found in the tonoplast control the flow of water into and out of the
vacuole. The central vacuole also stores the pigments that colour flowers.
The central vacuole contains large amounts of a liquid called cell sap, which differs in
composition to the cell cytosol. Cell sap is a mixture of water, enzymes, ions, salts, and other

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substances. Cell sap may also contain toxic by-products that have been removed from the
cytosol. Toxins in the vacuole may help to protect some plants from being eaten.
c) Plastids
Plant plastids are a group of closely related membrane-bound organelles that carry out many
functions. They are responsible for photosynthesis, for storage of products such as starch, and for
the synthesis of many types of molecules that are needed as cellular building blocks. Plastids
have the ability to change their function between these and other forms. Plastids contain their
own DNA and some ribosomes, and scientists think that plastids are descended from
photosynthetic bacteria that allowed the first eukaryotes to make oxygen. The main types of
plastids and their functions are:
i) Chloroplasts are the organelle of photosynthesis. They capture light energy from the sun
and use it with water and carbon dioxide to make food (sugar) for the plant. The
arrangement of chloroplasts in a plant’s cells (Figure 2).

Figure 2: A diagram of the chloroplast

The chloroplast is enclosed by an inner and an outer phospholipid membrane. Between

these two layers is the inter-membrane space. The fluid within the chloroplast is called
the stroma, and it contains one or more molecules of small, circular DNA. The stroma
also has ribosomes. Within the stroma are stacks of thylakoids, sub-organelles that are
the site of photosynthesis. The thylakoids are arranged in stacks called grana (singular:
granum). A thylakoid has a flattened disk shape. Inside it is an empty area called the
thylakoid space or lumen. Photosynthesis takes place on the thylakoid membrane.
Within the thylakoid membrane is the complex of proteins and light-absorbing pigments,
such as chlorophyll and carotenoids.
ii) Chromoplasts make and store pigments that give petals and fruit their orange and yellow
colors.
iii) Leucoplasts do not contain pigments and are located in roots and non-photosynthetic
tissues of plants. They may become specialized for bulk storage of starch, lipid, or
protein. However, in many cells, leucoplasts do not have a major storage function.
Instead, they make molecules such as fatty acids and many amino acids.

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LECTURE TWO
PLANT TISSUES
Meristematic Cells
Meristem is derived from a greek word ‘meristos’ which means divisible. These are immature
cells from plants actively dividing i.e. undifferentiated. They are responsible for growth. They do
not have specific shape i.e. irregular. They are found at the tip of the shoot and roots. They have
thin cell wall. They do not store any food. The can also be found in mature tissues for repair of
wounds (figure 3).

Figure 3: Meristematic cells

Characteristic features of meristematic cells

i) Cells may be spherical, oval polygon or rectangular in shape i.e. don’t have a specific
shape
ii) Cell wall is thin, uniform and is made up of cellulose
iii) Cells are compactly arranged without any inter-cellular spaces
iv) Each cell has large prominent nucleus and dense cytoplasm usually without vacuole
v) Cells are in active state of metabolism and are capable of undergoing rapid division
vi) Cells do not store food materials.
vii) Large nucleus
viii) Small or no vacuole at all

They are classified based on:

a) Position/location in the plant body
i) Apical meristem – found at the shoot and the roots for growth in length. They
continuously differentiate. They increase the length of plant i.e. Primary growth
ii) Intercalary – found at the tip and base stem of the leafs/nodes/internodes. They
regenerate buds by increasing length between nodes. E.g. leafs regrowth in
grasses after herbivory and lawn mower allows them to survive
iii) Lateral meristem/cambium- Found on the radial portion/sides of the roots or stem.
They bring about secondary growth in plants i.e. increase in width/girth
b) Function
i) Protoderm – gives rise to the epidermis
ii) Procambium- gives rise to vascular tissues
iii) Ground meristem – gives rise to the cortex, pith, medullary rays and pericycle.

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c) Origin and development

i) Promeristem – arise from the embryo
ii) Primary meristem- arise from the promeristem
iii) Secondary meristem- involved in secondary growth and arise from primary
meristem
d) How they divide/plane of division
i) Mass- cells divide in all different planes
ii) Plate- divide in one single plane e.g. in a leaf
iii) Rib- divide at right angles resulting to growth in girth
Permanent Tissues
They are permanent and do not change the structure. The meristematic cells keep on dividing and
form permanent tissues through a process known as differentiation. They take up a specific role
and loss the ability to divide hence becoming permanent tissues. The vacuoles grow to many
different shapes and size depending on the need of the cell. It is possible that the vacuole may
take up to 90% of the cell total volume in permanent tissues.
i) Simple Permanent Tissues
a) Parenchyma cells
o They are living cells inside the plants.
o They are found in all parts of the plant.
o They have a thin cell wall.
o They are isodiametric in shape.
o They are found in most of the edible fruits
o They are divide according to their functions into:
i) Chlorenchyma cells- they have chloroplasts.
ii) Aerenchyma cells- they have large intercellular space. They are loosely packed
which help some cell to increase buoyancy and help them to float in water e.g. in
water lilies
iii) Idioblasts – they store food in form of starch and water.
b) Collenchyma cells
o They are living cells.
o Formed from parenchyma cells
o Their cell wall is thick because of a thick layer of cellulose.
o They have very less intercellular spaces
o They are found especially just below the epidermis hence are referred to as
hypodermis.
o They have uneven walls
o They provide flexibility and support for the plants like the leaves and flower buds.
c) Sclerenchyma cells
o They are dead cells.
o They have thick tough secondary walls
o They have lignin which makes them super tough e.g. the husk of a coconut is made
up of sclerenchyma cell
o They give mechanical support to the plants.

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o They are found below the surface of the epidermis e.g. in the veins of the flowers,
hard covering of nuts
o They can be elongated i.e. fibers.
o They can be round e.g. sclerenchyma stones.
o They are of two types:
i) Osteosclerenchyma- bone shaped
ii) Astrosclerenchyma- Star shaped.
iii) Complex Permanent Tissues
The complex tissues are heterogeneous in nature, being composed of different types of cells
performing diverse functions. Xylem and phloem are complex tissues which constitute the major
part of the vascular bundles and hence are also called vascular tissues. Vascular bundles form a
continuous and interconnected system of different organs of the plant.
a) Xylem
Xylem is derived from the Greek work xylon. It is the “plumbing system of the plant”.
Structurally, xylem consists of both living and non-living cells. The most important constituent
of it are the tracheary elements. However, xylem consists of four elements, (i) tracheids, (ii)
vessels, (iv) xylem fibres and (iv) xylem parenchyma (Figure 4). The tracheids and vessels does
the actual transportation of water and mineral salt and together they are called tracheary
elements.
i) Tracheids
A tracheid is very much elongated or tube-like cell with hard, thick and lignified walls and a
large cavity. It is devoid of protoplast and hence dead. Their ends are tapering, blunt or chisel –
like. In transverse section these appear circular, polygonal or polyhedral in shape. The cell wall
is hard, moderately thick and lignified. They are shorter in length and have no perforations.
Because of structural specializations, tracheids are well adapted for transport of water and solutes
from the roots to the stem and leaves. They are found in all vascular plants. In gymnosperms
they are the chief water transporting elements. Besides this they also provide mechanical
support.

Figure 4: A-Tracheids and B-Vessel elements

ii) Vessels/Trachea
A vessel is an elongated long, cylindrical, tube-like structure with lignified walls and a wide
central lumen. The cells are dead as these do not have the protoplasm. Because of structural

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peculiarities, vessels are more efficient in transport of water and minerals as compared to
tracheids. Their efficiency is due to presence of perforation plates. Besides these they also give
mechanical strength. They are longer in length. They are found in angiosperms.

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iii) Xylem fibres

Xylem fibers are found both in primary and secondary xylem and are basically of two types, the
fibre tracheids and the libriform fibres. Fibre tracheids are intermediate forms between typical
fibres and tracheids which possesses bordered pits, though borders are less developed. The
libriform fibres are narrow with highly thickened secondary wall, obliterated central lumen and
simple pits. Their main function is support of the plant.
iv) Xylem parenchyma
The parenchymatous cells are found both in primary and secondary xylem but more commonly
in primary xylem. Their main function is the storage of food and help in the sideways conduction
of water. The parenchyma found in secondary xylem is classified into two types.
(i) Axial or wood parenchyma. These are derived from the fusiform cambium and occur in
radial transverse series.
(ii) Ray parenchyma- these originate from ray initials of fusiform cambium and occur in
radial transverse series.
b) Phloem
It is another type of complex tissue found in the vascular system which may also be called bast
or leptome. Always think of the back when we talk of the phloem. Its main function is the
transportation of dissolved food throughout the plant. As such phloem consists of (i) sieve
elements (sieve cells and sieve tubes) (ii) companion cells, (iii) phloem parenchyma and (iv)
phloem fibres.

Figure 5: Phloem tissue

The different elements of phloem originate from the procambium or vascular cambium. The
phloem formed from pro cambium is called primary phloem and that formed from vascular
cambium is secondary phloem. The phloem is typically found outside the vascular cambium.
However, in the stem of some dicotyledonous, parches of primary phloem are found in the pith,
i.e. inner to the xylem.
i) Sieve elements
They are the conducting elements. Basically sieve elements include two types of cells, the less
specialized sieve cells and more specialized sieve tubes.
a) Sieve cells
These are narrow elongated cells without conspicuous sieve areas, and taper at the ends or have
greater inclined walls which overlap in the tissues. These are found among members of lower
vascular plants and gymnosperms.

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b) Sieve tubes
These are long tube-like slender bodies arranged in longitudinal series where the end walls are
perforated in a sieve like manner and referred to as sieve plates. The sieve plate is the more
specialised area found usually in or near an end wall. The main function of the sieve tubes is the
longitudinal translocation of prepared food materials, from leaves to the storage organs in the
downward direction and then to the growing regions in the upward direction. The perforations of
the tubular walls make the process more efficient. The food moves mainly in the form of sucrose
followed by other types of sugars. They are only found in angiosperms.
ii) Companion cells
These are specialised parencymatous cells found closely associated with sieve elements in origin,
position and function. These develop from same meristematic cells that give rise to the sieve
element. Such a meristematic cell divides longitudinally and one of them differentiate into sieve
tube and other into companion cell. These are living cells and have dense cytoplasm, prominent
nucleus, several vacuoles, and other cytoplasmic organelles. Starch grains are never found in
companion cells. Companion cells play an important role in maintenance of a pressure gradient
in the sieve tubes. They also give support to the sieve tubes.
iii) Phloem parenchyma
The phloem parenchyma found associated with phloem are elongated tapering to broadly
cylindrical, sub-spherical and polyhedral in shape. These are living cells which have dense
cytoplasm and nucleus. The cell wall is composed of cellulose, having primary pit fields,
interconnecting, axial parenchyma cells and ray cells, and sometimes companion cell and sieve
tubes. Lignified cell wall has been found in inactive phloem parenchyma. Phloem parenchyma is
characteristically not found in monocots. The phloem parenchyma is mainly associated with the
storage of organic food materials and other substances such as resins, tannins, mucilage, latex
etc.
iv) Phloem fibres or bast fibres
Sclerencymatous fibres are found both in primary and secondary phloem and are commonly
called phloem fibres. They are used for support. Used commercially e.g. Sisal materials
Other specialised tissues: Secretory tissues are structurally modified and specially organized
for secretory functions. These may occur either as a single cell or in isolated patches or in a mass
and likewise may be external or internal in position.
External secretory structures: Trichomes and glands: These may also be called glandular
Trichomes. These are the simplest kind of excretory structures, though well organised and
composed of different types of cells. These are the following types:
(i) Digestive glands: Found in the body of insectivorous plants. They secret digestive enzymes as well as
poisonous substances for paralysing the insects.
(ii) Stinging hairs: these are found in Urtica all along the body. These hairs secret poisonous and muscle
irritating substances
(iii) Colleters: these are commonly found on the bud scales of roses and secret sticky excretory substances rich
in terpenes and mucilage
(iv) Nectaries: Secrets sugary substances called nectar and or honey composed of sucrose, glucose and fructose
which are derived from the phloem. E.g. Hydathodes
Internal secretory structures: Secretory cells: Found in the ground tissue and are called
Idioblasts which appear distinct from other neighboring cells. These have been named on the
basis of their secretory materials such as oils, resins, tannins, gums, mucilage and crystals of
various types

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i) Oil glands: Many plants have characteristic oil glands composed of small groups of thin walled, densely
cytoplasmic cells secretory in function.
ii) Laticiferous tissues: These are specialised parencymatous cells which secrete vicious fluid called latex.
LECTURE THREE
TISSUE SYSTEMS IN PLANTS
Structural tissues on the basis of physiological similarities or topographical continuity, in
reference to division of labour, are organised into tissue system. The precursors of these tissue
systems are meristems which may be outlined as follows (Figure 6).

Figure 6: A figure showing the origin and composition of tissue systems in plants

Sach (1875) recognized three basic tissue systems found in the primary structure of the plants;
Epidermal tissue system, Ground or fundamental tissue system and Vascular tissue system
(Figure 7).

Figure 7: A figure showing the three (3) types of tissue systems and their functions in plants

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A. The Epidermal Tissue System

Epidermal tissue system forms the outermost skin layer which is basically a protective layer. It is
subjected to a number of adverse factors such as evaporation of water, risk of mechanical injury
by rains, wind and biological agencies, exposure to weak and intense light and varied thermal
fluctuations. As such several adaptations and variations are found in epidermal tissue system.
Epidermal cells are parencymatous in nature and are compactly arranged without intercellular
spaces. The epidermis consists of living cells. Cells have a large central vacuole and thin
peripheral uninucleate cytoplasm which contains chromoplasts and leucoplast and occasionally
chloroplasts. In general, they have unevenly thick walls, the outer and radial walls being much
thicker than the inner walls.
The walls are strongly cutinised- important for protection against mechanical injury and
prevention of loss of water. It forms a layer on epidermis called cuticle which is present all over
the plant body except in roots and some submerged aquatic plants. Epidermis of root without
cuticle is called epiblema. The thickness of cuticle is variable which may be smooth as in
sunflower stem or have protrusions and ridges of various shapes. In cork cells, suberin is found
deposited and the dermal system is referred to as periderm.
Functions:
(i) It is protective layer that protects internal tissues against mechanical injury, high light
intensities, injurious thermal changes, attack of microbial parasites, high wind velocity,
acid rain effect etc. This is due to deposition of cutin, wax, oil, resin, hairs, lignin etc.
(ii) It reduces loss of water due to excessive transpiration.
(iii) the presence of sharp and stiff hairs on epidermis in certain cases protects plants from
grazing.
(iv) Epidermis of some xerophytic plants store water and food materials in the form of
mucilage
(v) In Bryophytes, Pteridophytes and certain hydrophytes, epidermal cells contain
chloroplasts and thus manufacture the food.
Specialised epidermal structures
a. Lithocytes: Contain deposition of calcium carbonate crystals, arranged in the form of
bunches, like those of grapes. Such deposits are called cystoliths and cells containing
them are called Lithocytes.
b. Myrosin cells: Contain sac-like cells scattered in the epidermis. These remains filled
with an enzyme, myrosin, and are called myrosin cells. These are idioblastic cells
resembling the laticifers in nature.
c. Sclerotic cells: The epidermis of seed coats of some plants of leguminosae family is
often made up of a layer of sclereids
d. Bulliform cells: Some epidermal cells found in groups in the leaves of certain monocots,
especially grasses are comparatively larger, thin walled and highly vacuolated. These are
called bulliform cells. They are mainly water containing cells, poor in solid and
chlorophyll contents.
e. Stomata: Stomata are narrow openings found in the epidermis of green aerial parts,
especially leaves of plant. A pore the stoma, remains surrounded by two specialised
epidermal cells called guard cells. The guard cells are kidney shaped in dicotyledons but
dumble-shaped in members of graminae (monocots). Their wall is unevenly thickened.
The wall surrounding the pore is thickened and inelastic due to presence of secondary
layer of cellulose but remaining part of the wall remains thin, elastic and permeable. The

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epidermal cells surrounding guard cells are specialised and called subsidiary cells or
accessory cells which give support in the movement of guard cells. Functions of
stomata: Stomata are very important structures from physiology point of view.
Physiological functional like photosynthesis, respiration and transpiration takes place
with the help of stomata, as it is through them interchange of gases takes place between
the intercellular space system of the internal tissues and the outer atmosphere.
f. Trichomes: the term trichomes include different types of unicellular or multicellular
appendages of the epidermis. Trichomes are found both on vegetative as well as
reproductive parts and may be living or dead, ephemeral or persistent and variable in
structure, form and function. For instance,
 Foliar trichomes are important in the regulation of moisture exchange with the
atmosphere
 Covering hairs for plant defence against phytophagous insects
 Secretory trichomes for providing chemical defence against insects and
 Stinging hairs for protecting plants from animals.
B. The Ground Tissue System
The tissue system found in the region between epidermis and vascular system are collectively
referred to as ground meristem. It is heterogeneous in nature composed of different types of
cells concerned with different functions. This system forms the major mass of the primary body.
In a stem, it basically includes cortex, pericycle, pith and modular rays and in a leaf,
mesophyll tissues are included in this system. It is partly derived from periblem and partly from
plerome.
i) Cortex
Cortex is the extra-stelar zone, which forms that part of the axis which spreads from the layer
next to epidermis to the layers enveloping the stele. In a dicotyledonous stem, cortex is generally
differentiated into hypodermis, general cortex, endodermis etc.
(a) hypodermis: It is situated just below the epidermis. In a dicot stem, hypodermis is
characteristically composed of collenchymatous cells while in a monocot stem, it consists
of Sclerencymatous cells.
(b) General cortex: The region between the hypodermis and endodermis is called general
cortex which is usually composed of loosely arranged parenchyma cells with intercellular
spaces. The cells are isodiametric in shape. Starch grains and different types of crystals
are found in these cells. Chloroplast may or may not be present. Sclereids, resin ducts,
tannin cells, oil cavities and lactiferous tissues may also occur in this region.
(c) Endodermis: the innermost layer of the cortex is referred to as endodermis. Some
scientists consider it as the limiting layer of stele. So it is origin remains controversial. It
is a single layer composed of compactly arranged modified parenchyma cells lacking
intercellular spaces. Cells are vertically elongated which appears tubular or barrel-shaped
in cross section. Endodermal cells are characterised by presence of waxy substances, like
suberin, in the form of bands or strips on their radial and transverse walls. These are
commonly called casparian strips or bands named after its discoverer, Caspary (1865).
Lignin may also be found present in these thickening strips.
The exact and basic functions of the cortex on the basis of its relation to water and vascular
tissues have been attributed to it. This includes the following functions:
(i) It is considered as a water tight jacket between xylem and cortex.

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(ii) It acts like an air dam that prevents diffusion of air into xylem, which would
otherwise get clogged.
(iii) It is a protective layer, sometimes referred to as accessory epidermis.
(iv) It regulates and maintains root pressure,
(v) It acts as a water dam due to the presence of compactly set thickened cells.
(vi) It provides regulated and systematised flow of water through its passage cells.
(vii) The thin walled cells retain power of division, as such; the lateral branch and
adventitious buds often originate through this layer.
ii) Pericycle
Pericycle may consist of one or several layers of cells found between the endodermis and
vascular bundles. Pericle region surrounds the vascular tissue and is regarded as the limiting
layer of the stele. It forms the outermost part of the intrastelar ground tissue, the other part being
the pith and medullary rays.
Function: In roots pericycle gives rise to lateral roots and since it is meristematic in nature (in
this case), it has also been referred to as pericambium. In stems, during secondary growth,
phellogen or cork cambium arises from it and in certain cases where anomalous secondary
growth is found, it forms the secondary cambium. In monocots where secondary growth is
absent, pericycle becomes Sclerencymatous in nature and thus provides mechanical strength to
the stem. In normal cases, the parencymatous cells of the pericycle serve for storage of food
materials, and only in certain cases, these may contain laticiferous tissues and secondary ducts.
iii) Pith
Pith is the main internal, more or less cylindrical ground tissue which forms the central core of
the stem and which externally remains bounded by the vascular bundles. Normally it consists of
long isodiametric and loosely arranged normal parencymatous cells with large number of
intercellular spaces.
Function: Pith, where it is presents basically serves as a region for storage of various substances
such as starch, fatty substances, mucilage, tannins etc.
iv) Medullary rays
The vascular bundles remain separated due to the presence of the long strips of parencymatous
tissues passing through in between them. Such strips are called medullary rays or pith rays. The
chief function is help in the conduction of food and water from critical region to pith or central
region.
v) Mesophyll tissues/Ground tissues of leaves
The ground tissue of leaves is commonly called mesophyll tissues, which is remarkably
different from that of stem and roots. It shows variation in different types of leaves.
In isobilateral leaves (monocotyledonous leaves with parallel venation), mesophyll tissues
consist of more or less isodiametric, parencymatous cells with intercellular spaces. In
dorsiventral (dicotyledonous leaves) and concentric leaves, having reticulated venation,
mesophyll tissues are found differentiated into two types of cells, as follows:
(a) Palisade cells: These are usually found beneath the upper epidermis (adaxial surface) and are
elongated or columnar shaped, arranged more or less at right angles to the epidermis.
However, in certain cases, palisade parenchyma is present below the upper and lower
epidermis.
(b) Spongy cells: These occur towards lower epidermis (abaxial surface) and are characterised
by isodiametric or irregular shaped cells with large number of intercellular spaces. The
mesophyll cells contain chloroplast and perform photosynthetic functions. In certain

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xerophytic or succulent plants these cells store water and food substances in soluble forms. In
some cases, sclereids and secretory cells are also found in this region. In aquatic and marshy
plants, mesophyll tissues consist of parenchyma which in addition to photosynthesis provides
buoyancy to the leaves.
C. The Vascular Tissue System
Composed of two complex tissues. Xylem and phloem occur together forming clusters or
bundles. In basal angiosperms and eudicots, vascular bundles are arranged in one ring
surrounding the pith. In monocots, vascular bundles are scattered throughout the ground tissue.
Vascular bundles are collateral: they contain both xylem and phloem. The xylem and phloem of
a bundle are both primary tissue, that is, they are both derived from the activity of the apical
meristem. Parenchyma and sclerenchyma cells are also found in the bundles but their amounts
vary with the function of the stem. Vascular tissue system conducts materials throughout the
plant body and provides support.

Figure 8: A diagram showing the vascular bundles

i) Xylem conducts water and minerals from the roots to all parts of the plant; it also
supports the plant and stores food and other organic substances. Together with the phloem, the
xylem forms a continuous system of vascular tissue extending throughout the body.

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The xylem is derived from the procambium in the primary plant body. During secondary growth,
xylem is derived from the cambium. There are two tracheary elements that make the xylem:
tracheids and vessel elements. Tracheids and vessel elements are the conducting cells and are
dead at maturity; both have wall pits on their sidewalls for lateral transport. A stack of vessel
elements end-to-end forms a vessel. Xylem vessels arise from individual cylindrical cells
oriented end to end. At maturity the cytoplasmic contents die.
The secondary walls of the xylem vessels are deposited in spirals and rings and are usually
perforated by pits. The tracheary elements of the primary xylem have a variety of secondary wall
thickenings. Secondary wall is deposited in the tracheary elements during the period of cell
elongation in the procambium. Secondary wall of the first-formed tracheary elements of the
early-formed primary xylem called protoxylem, are deposited in the form of rings or spirals.
These rings allow the cell to elongate after the cells have differentiated. Elements with annular
and helical thickenings will develop in the late-formed primary xylem, called metaxylem, and
in the secondary xylem. Scalariform thickenings consists of annular thickening interconnected,
and reticulate secondary wall is form by a network of thickenings. Circular bordered pits
interconnect tracheary elements. These are the most derived and strongest of the tracheary
elements. In these tracheary elements, the entire primary wall is covered with secondary wall.
Pits are openings in the secondary wall that are surrounded by a thickened border.
Circular bordered pits are aligned with those of adjacent cells and form pit-pairs. The primary
cell walls of both adjacent cells and the middle lamella forms the pit membrane that separated
both cells. The pit membrane is very permeable to water. Only tracheary elements have bordered
pits. Fibers and sclereids do not have bordered pits, just simple pits. Tracheary elements with
annular thickenings are weak, but a large percentage of the primary wall is not covered with
secondary wall and is available for water movement between adjacent cells. In the metaxylem
and in the secondary xylem, the secondary cell walls of the tracheids and vessels cover the entire
primary walls, except the pit membranes and at the perforation of the vessel elements. At
maturity, vessel elements have perforations, which are areas lacking primary and secondary
walls. These perforations or holes occur on the end walls of the vessel elements. The result is the
xylem vessel, a continuous nonliving duct.
Xylem also contains tracheids. These are individual cells tapered at each end so the tapered end
of one cell overlaps that of the adjacent cell. Like xylem vessels, tracheids have thick, lignified
walls and, at maturity, no cytoplasm. Their walls are perforated so that water can flow from one
tracheid to the next. There are thin membranes in the pit that prevent air bubbles from passing to
the adjacent tracheid. Vessels are the principal water-conducting cell in angiosperms. Many
angiosperms also have tracheids in addition to vessels. The xylem of ferns and conifers contains
only tracheids. Water flowing from tracheid to tracheid must pass through the pit membrane –
the thin modified primary walls - of the pit pairs. Tracheids lack perforations and have only pits.
Perforations greatly reduce friction and allow the free flow of water, while pits have the pit
membrane that slows down the passage of water. Water can flow relatively unimpeded from
vessel element to vessel element through the perforation. Tracheids evolved over 420 million
years ago and virtually all plants with vascular tissue have them. Vessels evolved more recently
and occur mostly in flowering plants. Air bubbles formed during the freezing and thawing can
potentially obstruct the flow of water for the entire length of the vessel. Programmed cell death
(apoptosis) results in the elimination of the protoplast in the tracheary elements. The xylem
tissue also contains parenchyma cells that store various substances. Xylem parenchyma
commonly occurs in vertical strands, but in the secondary xylem, they are also found in the rays.

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Xylem may also contain fibers some of which are living at maturity and serve a dual function of
storage and support. Sclereids are sometimes found in the xylem.

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ii) Phloem

It is the principal food conducting tissue in vascular plants. In addition to sugars, phloem
transports many other substances including amino acids, lipids, micronutrients, hormones,
proteins, and RNA, some of which act as signaling molecules.

There is primary and secondary phloem. The first formed primary phloem, the protophloem, is
often stretched and destroyed during elongation of the organ. The principal conducting cells of
the phloem are the sieve elements. The protoplasts of adjacent sieve elements are interconnected
through sieve areas.
Two types of sieve elements are recognized: sieve cells and sieve tube elements.
 Sieve cells are found only in gymnosperms.
 Sieve tube cells are found only in angiosperms.
 Sieve elements are variable in seedless vascular plants and are simply called sieve
elements.
In sieve cell
 Cells are long and narrow with pointed ends.
 The pores are narrow.
 The sieve areas are located over all the cell surface.
 Sieve areas are concentrated on the overlapping walls of the long and slender sieve cells.
 Albuminous cells are associated with them.
 Found in all non-angiosperm vascular plants.
In sieve tube elements
 Cells are short and wide with flat ends.
 The sieve areas in some walls have larger pores than those in other walls.
 The part of the sieve area bearing the larger pores is called the sieve plate or sieve area.
 Sieve plates are generally located on the end walls but they may occur anywhere in the
cell.
 The sieve tube elements are arranged end to end in longitudinal series called sieve tubes.
 The presence of sieve plates is a distinguishing characteristic of the sieve tube elements.
 Sieve tube elements are aligned end-to-end and form a sieve tube.
 Companion cells are associated with them.
 Found in angiosperms only.
Sieve elements have only primary wall. Sieve elements remain alive at maturity; sieve elements
lack nucleus, vacuoles, Golgi complex, ribosomes and cytoskeleton. Plasma membrane and
endoplasmic reticulum remain. ER is particularly abundant near the sieve plates. At maturity, the
plasma membrane, ER, some plastids and mitochondria remain distributed along the wall of the
sieve elements. Sieve tube members are the conducting cells; companion cells regulate the
metabolism of the sieve tube members. Both are derived from the same mother cell. Companion
cells contain all the organelles found in plant cells. Companion cells move sugars, amino acids,
informational molecules, ATP and other substances into and out of the sieve elements. There are
numerous cytoplasmic connections (plasmodesmata) between the companion cells and the sieve
tube members. Albuminous cells are parenchyma cells found in the phloem of gymnosperms.

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Parenchyma cells are also found in the phloem and are associated with the storage of a variety
of substances.
Fibers and sclereids may also be present in the phloem and help in supporting the plant body.
The Vascular Tissue System (additional information)
It consists of the complex tissues, the xylem and the phloem which together make a unit strand
called vascular bundle. Vascular bundles remain distributed in the stelar region. A stele is
considered as a central cylinder of root, stem or leaf which consist of vascular bundles, pith and
pith rays enclosed in a pericycle. The pericycle remains surrounded by endodermis.
Origin. The elements of vascular bundle arise from procambium of apical meristem. During
longitudinal division in the apical meristem, some cells are distinguished from other cells due to
presence of dense cytoplasm in them. These cells are collectively called procambium which soon
gets differentiated into two parts, the phloic procambium which forms phloem and the xyloic
procambium which forms xylem.
The phloem elements mature earlier and these develop only in acropetal manner but xylem
elements develop both in acropetal and basipetal directions. The phloem always differentiates
centripetally, i.e. towards the centre of the axis. However, differentiation in xylem shows three
different patterns:
(a) Exarch- in this pattern, the course of development is centripetal or towards the axis and
thus, the initial xylem elements are located far from the axis. Exarch xylem is
characteristic of roots.
(b) Endarch- in this pattern, the course of development is centrifugal and thus, the initial
xylem elements are towards the centre. Endarch xylem elements are characteristic of the
stem of phanerogams.
(c) Mesarch- In this pattern, the course of development proceeds in two directions, i.e. both
centrifugally and centripetally, and thus, it leads to a condition where protoxylem is
found in the centre and metaxylem towards its periphery. Mesarch xylem is characteristic
of rhizome and rachis of certain ferns.
Elements of Vascular bundles
A typical primary vascular bundle of the dicot stem consists of three types of elements, (i)
primary xylem, (ii) primary phloem and (iii) cambium.
Primary xylem. Primary xylem is composed of protoxylem and metaxylem elements.
Protoxylem are the first formed xylem and metaxylem are lately formed.
Primary phloem. Primary phloem consists of protophloem and metaphloem both recognized on
basis of differentiation. The first formed phloem elements are called protophloem while lately
formed are called metaphloem.
Cambium. Cambium occurs in the form of a thin strip of cells between xylem and phloem, in
vascular bundles of stem of dicots and gymnosperms.
Types of Vascular bundles
According to the mode of occurrence of vascular elements, vascular bundles are classified as
follows:
1. Radial. In this type, the xylem and the phloem are found as separate parches on alternate
radii. These are most primitive in nature and are characteristic of roots. (Fig 6.12, F and
6).
2. Conjoint. In this type the two complex tissues (xylem and phloem) occur side by side on
the same radius. According to the position of the phloem, these are recognized into two
types, (a) collateral and (b) Bicollateral. The collateral V.B. are the most common type

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found in stem and leaves of phanerograms. In this case, xylem and phloem lie together on
the same radius, the phloem being externally and the xylem being internally (towards the
pith) in position. The collateral V.B. are referred to as open type if a strip of cambium
between xylem and phloem is present as in the case of dicot stems, and closed type, if
cambium is not found, as in monocots stems. In bicollateral V.B., two phloem patches,
one on outer side (external) and other on inner side (internal) of the xylem, are found, as
in stems of members of cucurbitaceae. In these vascular bundles, the sequence would be
the outer phloem, outercambium, xylem, inner cambium and inner phloem. These are
always open (Fig. 6.12)
3. Concentric. In this type, one complex tissue remains completely surrounded by the
other. These are of two types, (i) amphivasal or leptocentric bundles, where phloem is
surrounded by xylem, such as secondary vascular bundles of Dracaena, Yucca and
medullary bundles in dicotyledons and (ii) amphicribal or hadrocentric, where xylem is
surrounded by phloem, as in rhizomes of ferns, stems of Lycopodium and Delaginella
and small bundles of flower, fruits and some leaves of dicotyledons. The concentric
bundles are always closed.

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