STUDY OF CELL WALL CONSTITUENTS

AND CELL INCLUSIONS

 CELL WALL
The cells of all plants, bacteria and fungi have a rigid, protective covering outside
the plasma membrane called cell wall.

The presence of cell wall in plant cells distinguishes them from animal cells.
Among the vascular plants only certain cells connected with the reproductive
processes, are naked, all other cells have walls.

The cell wall was first observed by Hooke in the year 1865 in cork cells.

 Originally it was thought that the cell wall was a non-living secretion of the
protoplasm, but now it is known to be metabolically active and is capable of growth
and at least during its growth, contains protoplasmic material.
DEFINITION
Cell wall is the outer, rigid, protective, semi-elastic, semi transparent, supportive
covering of plant cells. Its thickness varies from 0.1μm to 10 μm.

Chemical composition:
1. Matrix: water 30-60%, Hemicellulose-5-15%, Pectin substances-2-8%, Lipids-0.5-
3%, Proteins-1-2%

2. Microfibrils: cellulose, microfibrils-20-40%

3. Other Ingredients: Lignin, cutin, suberin, silica, minerals, waxes, tannins, resins, gums
etc. 0-25%
 STRUCTURE OF CELL WALL
 Plant cell walls consists predominantly of cellulose microfibrils enmeshed in

a complex network of branched polysaccharides and glycoproteins called

extensins. The two main polysaccharides are hemicellulose and pectins.

Cellulose, hemicellulose and glycoproteins are linked together to form a

rigid interconnected network that is embedded in a pectin matrix.

A typical plant cell has the following three parts.
1. Middle lamella 2.Primary wall 3. Secondary Wall
Middle lamella( thickness 0.2-1μm)

It is a thin amorphous cement like layer between two adjacent cells. Middle
lamella is the first layer, which is deposited at the time of cytokinesis. It is
optically inactive (isotropic). It is made up of calcium and magnesium pectates.
In addition to these substances proteins are also present.
Primary wall ( thickness 0.1-3μm)

It is the first formed wall of the cell which is produced inner to the middle
lamella. It is thin, elastic and extensible in growing cells.

It is optically active (anisotropic). It grows by addition of more wall material

within the existing one. Such a growth is termed as intussusception.

Some cells like the parenchymatous cells and meristematic cells have only the
primary wall.

The primary wall consists of a loose network of cellulose microfibrils

embedded in a gel like matrix or ground substances. In most of the plants the
micro fibrils are made up of cellulose.
Structure of primary cell wall
The micro fibrils are oriented variously according to shape and thickness of
the wall.

The matrix of the primary wall in which the micro fibrils are embedded is
mainly composed of water, hemi-cellulose, pectin and glycoprotein. Pectin is
the filling material of the matrix.

Hemi-cellulose binds the microfibrils with the matrix and the glycoprotein
control the orientation of the microfibrils.
Secondary Wall ( thickness 3-10μm)

A thick secondary wall is laid inner to the primary wall after the cell has
reached maturity. It is laid down in succession of at least three layers often
named s1, s2 and s3 .

 It grows in thickness by accretion (apposition) i.e deposition of materials over

the existing structures. The central layer (S2) is usually the thickest layer. In some
cells however, the number of layers may be more than three.

The formation of secondary wall is not uniform in all the cells. This results in the
differentiation of various types of cells, such as parenchyma, collenchyma,
sclerenchyma, fibres and tracheids.
The micro fibrils of secondary wall are compactly arranged with different
orientation in different layers embedded in a matrix of pectin and hemicellulose.

 Substances like lignin, suberin, minerals, waxes, tannins, resins, gums, inorganic
salts such as calcium carbonate, and calcium oxalate, silica etc may be
deposited in the secondary wall.

The secondary wall is very strongly anisotropic and layering can be observed
in it.
Structure of secondary cell wall

3-10 μm

0.1-3μm
The secondary wall consists of two continuous interpenetrating systems one of
which is the cellulose micro fibrils and the other, the continuous system of
microcapillary spaces.

These spaces may be filled with lignin, cutin, suberin, hemi-cellulose and
other organic substances and sometimes even some mineral crystals.
NATURE OF THICKENING OF SECONDARY CELL WALL

Annular or ringlike: thickening is noticed in the protoxylem elements where

secondary matters are placed centripetally in form of rings at regular
intervals

Spiral: thickening is also found in protoxylem elements, secondary wall being

deposited in form of spiral

Scalariform: Secondary matters took like the rungs of a ladder here

Reticulate: The secondary matters here assume the form of a network

Pitted: In this case secondary cell wall materials are deposited practically all over
the primary wall, only leaving some small thin areas here and there. These
unthickened areas are the pits

A & B Annular, C&D. Spiral, E&F. Scaliform. G. Reticulate.

H. Pitted (Simple). I. Pitted (Bordered)
Plasmodesmata

The cell wall is not totally complete around the cell. It is interrupted by narrow pores carrying
fine strands of cytoplasm, which interlink the contents of the cells. They are called
plasmodesmata.

They form a protoplasmic continuum called symplast. It consists of a canal, lined by plasma
membrane. It has a simple or branched tubule known as desmotubule.

Desmotubule is an extension of endoplasmic reticulum. Plasmodesmata serves as a passage

for many substances to pass through. It is also believed that they have a role in the relay of
stimuli.
Pits

Pits are the areas on the cell wall on which the secondary wall is not laid down.
The pits of adjacent cells are opposite to each other. Each pit has a pit
chamber and a pit membrane. The pit membrane consists of middle lamella
and primary wall. Pit membrane has many minute pores and thus they are
permeable.

Pits are of two types

1. Simple pits

2. Bordered pits.
In simple pits the width of the pit chamber is uniform. There is no secondary wall in
the simple pit .

In bordered pit the secondary wall partly overhangs the pit.

Pits help in the translocation of substances between two adjacent cells. Generally
each pit has a complementary pit lying exactly opposite to it in the wall of the
neighbouring cell. Such pits form a morphological and functional unit called the pit
pair.
Diagrammatic representation of pit
with torus

A. A vessel with bordered pits in front view. B. Same in sectional view

C. Perspective diagram of the same D. sectional view of bordered with
changed position of torus

The pit membrane usually has a thickening called torus.

FUNCTIONS OF CELL WALL
1. It gives definite shape to the cell.

2. It protects the internal protoplasm against injury.

3. It gives rigidity to the cell 4. It prevents the bursting of plant cells due to
endosmosis.

5. The walls of xylem vessels, tracheids and sieve tubes are specialized for long
distance transport.

6. In many cases, the cell wall takes part in offense and defense.
Structure of cellulose
CELLULOSE (POLYSACCHARIDE- HOMO)
Cellulose, the single most abundant organic macromolecule on earth is an unbranched
polymer that consists of thousands of β-D-glucose units liked together by β(1-4) bonds.

Cellulose molecules are long, ribbon like structures that are stabilized by
intramolecular hydrogen bonds.

Many such molecules(about 50-60) associated laterally to form the microfibrils

found in cellwalls.

Microfibrils are 25 nm in dia and are often twisted in a rope like fashion to generate
even larger structure called macrofibrils.
 HEMICELLULOSE (POLYSACCHARIDE- HETERO)
 Each occurs as a linear chain of a single kind of sugar (glucose /
xylose) with short side chains with different kinds of sugars
(hexoses like glucose, galactose and mannoses; pentoses like
xylose and arabinose).

 The sugar units in the hemicellulose backbone form hydrogen

bonds along the surface of the cellulose, creating a coating.

 The coating helps to bind the fibrils together into a rigid

interconnected network
PECTINS (POLYSACCHARIDE- HETERO)
They are branched polysaccharides but with backbone called
rhamnogalacturonns that consists mainly of negatively charged galacturonic acid
and rhanmnose.

The side chains attached are similar to that of hemicellulose. Pectin molecule form
the matrix in which cellulose fibrils are embedded.

In addition they bind adjacent cellwalls together. Because of their highly branched
structure and their negative charge, pectin traps and bind water molecules. As a
result pectin have gel like consistency.
EXTENSINS (GLYCOPROTEIN)
б Their polypeptide backbone is rich in aminoacids serine, hydroxyproline and lysine.
Due to lysine they have net positive charged has high affinity for negatively charged
pectin molecule.

б Extensins have numerous oligosaccharides usually attached to the hydroxyl group of

serine and hydroxyproline.

б They are rigid, rod like molecules that are tightly woven into complex polysaccharide
network of the cell wall.

б Extensins are least abundant in the cellwalls of actively growing tissue and most
abundant in the cell walls of tissues that provide support to the plant.
LIGNINS
 They are very insoluble polymers of aromatic alcohols that occur mainly in woody tissues.

 Lignin molecules are localized mainly between the cellulose fibril, where they function to
resist compression forces.
CELL INCLUSIONS (ERGASTIC SUBSTANCES OF PLANTS)

The non-living substances of plant metabolism are known as ergastic

substances. They may be reserve foods, secretory and excretory or end

products of metabolism.

(a) Reserve foods : The materials which occur as reserve food in a cell

are the carbohydrates, proteins and lipids. They are present in insoluble

forms. Their conversion to soluble form is covered by enzymatic processes.

(b) Excretory products : Animals are able to get rid of their excretory
products in liquid or solid form. Plants being unable to do so, their
wastes are excreted in the form of insoluble products and are stored
inside the cells only. Excretory products of the plants are named as
tannins, resins, latex, volatile oils, chemicals like alkaloids, glycosides
and mineral crystals, such as calcium oxalate, calcium carbonate, silica,
etc.
 A) RESERVE FOODS
(i) Carbohydrates : Many water soluble, as well as, water in-soluble carbohydrates are
found in plants. Glucose, fructose and sucrose are all water-soluble. The water-insoluble
contents are normally poly-saccharides.

lnulin : It is the only polysaccharide soluble in water and is found in tubers of Dahlia and
Dandelion. In alcohol, it forms sphere-crystals.

Starch : it is another polysaccharide mostly found in tubers, food-grains and seeds of the
plant. It is found in the form like granules, concentric or eccentric. Starch gives blue colour
with dilute solution of iodine.
Cellulose : A complex polysaccharide forming cell wall and is structural material
of plants. Mucilage in senna, isapgol, and linseed, forming gummy solution in
epidermal cells is another example of poly-saccharides.

(ii) Proteins : These are the nitrogenous substances either soluble or insoluble in
water. They are of great importance in the structure and functioning of living
cells. They are produced by and associated with living matter. They may be
amorphous or crystalline. In maize, they are amorphous found in the form of
layer as aleurone layer or crystalline as aleurone grain in castor seeds. They are
converted to amino-acids by proteolytic enzymes.
(iii) lipids : These are reserves of food materials of plants. Those which are liquid at room-
temperature are called oils and solids are called as fats. They are made up of fatty acids
andglycerin. Oils are found in endosperm of the following seeds of Castor, pea-nut, and sesame.
(B) EXCRETORY PRODUCTS:

(i) Alkaloids : These are the basic nitrogenous secondary metabolites having
marked physiological action, if taken internally. They occur in plants as salt of
nicotinic, quinic, citric or oxalic acid. They are found in seeds, barks, leaves,
roots, etc. Alkaloids are used as highly potent medicaments and possess curative
properties. They are protective materials of plants discouraging animal or insect
attacks. Alkaloids are detoxicating agents of plants "and reservoirs. for protein
synthesis e.g. ergot, vinca, datura, etc.
(ii) Glycosides : These are the condensation products of sugar and aglycon. Glycosides
are secondary metabolites of pharmaceutical significance. These are soluble in water,
as well as, alcohol. e.g. digitalis, senna, rhubarb, etc.

(iii) Tannins : Tannins are present in cell sap, soluble in water and alcohol. They give
blue- black or green colour with iron compounds. They have several medicinal
properties and hence used as drugs. Nut-galls, myrobolan are the examples of tannin.

(iv) Resins : These are found in abundance in the resin ducts of the trunk of the trees or
in rhizomes (ginger), fruits (capsicum) and other parts of the plants. They are insoluble
in water, may be semi-solid or solid. Many a times, they are associated with gums or
volatile oils.
(v) Latex : It is a white suspension wherein microscopically small particles or oil globules
are suspended. They contain proteins, sugars, minerals and alkaloidal salts in true
solution, while gums, starch, resins are in suspended form. Latex is present in
lateciferous tissue, characteristic to certain families like Euphorbiaceae, Caricaceae,
Papaveraceae, etc. Banyan, madar, papaya, poppy, ficus are examples.

(vi) Volatile oils : These are the fragrant liquids found in plant. They are volatile, liquid
and aromatic and are present in roots, leaves, barks, fruits, etc. They are insoluble in
water and are soluble in alcohol. Normally volatile oils are carminative stimulants and
antiseptic. Orange, lemon, coriander, cinnamon, ginger, eucalyptus and mentha are
examples of volatile oil containing drugs.
(vii) Mineral crystals : These occur in cell-wall or cell cavity. They are all insoluble in
water. Common crystals are calcium oxalate, calcium-carbonate and silica. They are
found in roots, stems, leaves, fruits, etc. Various types of crystals of calcium oxalate are
found like prisms, acicular, raphides, clusters, rosettes, etc.
(C) SECRETORY PRODUCTS
Various other products, not covered under any one of the above types, are
formed in the plants and serve some important function in the plants and are the
secretory products.

Enzymes, colouring substances, nectar are the examples of this type.

Enzymes are nitrogenous water soluble compounds carrying out hydrolysis of

carbohydrates and proteins.

Colouring substances’or pigments like chlorophyll are essential for photosynthesis,

anthocyanin and several other. Flavonoid glycosides giving attractive colour to
various flowers and leaves are the examples of this type.
Nectar is the sugary solution secreted by many flowers in special cells or glands to attract the
insects for effective pollination.