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Structure and functions, Biogenesis, growth.


CELL WALL

PRIMARY STRUCTURE


Cell wall is complex in nature .It remains differentiated in the following layers:
                                                                                                                                       
(i) Middle Lamella         
(ii)Primary cell wall    
(iii)Secondary cell wall 
(iv)Tertiary cell wall

(i) Middle Lamella-The cells of plant tissue generally remain cemented together by inter cellular matrix known as middle lamella. It is composed of pectin, lignin & some other proteins. They are dissolved in strong acids.

(ii) Primary Cell Wall- It first formed cell wall. Its thickness is about 100-200 NM. It is the outer most layer of the cell & it forms the only cell wall in the immature meristamatic & parenchymatous cells. It is comparatively thin & permeable. It is composed of loosely organised network of cellulose microfibrils associated with hemicellulose pectins & glycoproteins. Pectins are important in imparting flexibility & makes primary cell wall to expand during cell growth. The cellulose synthesizing enzyme that synthesize cellulose microfibrils are localized within plasma membrane. These are called rosettes; add glucose molecules to growing microfibrils. It is composed of polysaccharide cellulose but other substance may be incorporated in it. Lignin or suberin may also present in it & epidermal cells of leaves & stem also posses cutin & cuter waxes & are impermeable to water. Primary cell wall of yeast and fungi is composed of chitn.

(iii) Secondary Cell Wall- It lies after primary cell wall. It is thick, permeable & lies close to tertiary cell wall or lies close to plasma membrane .It consist of bulk of many layers and at maturity  cell wall consist  It gives shape & mechanical strength to the cell .It is chemically composed of compactly arranged cellulose & lignins.

(iv) Tertiary Cell Wall- It is found beneath the secondary cell wall. It differs form primary and secondary cell wall, and contains xylans. 




CHEMICAL NATURE

Plant's cell wall are composed of carbohydrates known as cellulose. Besides cellulose various chemical substances as hemicelluloses, pectin, lignin, cutin & chitn. The cell wall also contains certain minerals like calcium & magnesium in the form of carbonates & silicates. The cellulose is a polysaccharide & it is the most abundantly occurring chemical substance of most plant cell. Chemically cellulose contains long chains of glucose molecules. Glucose is the structural unit of cellulose & about 3000 glucose molecule are linked together to form long chain of cellulose molecule. A bundle of 100 chain molecules of cellulose forces the elementary fibril known as micelle. The 20 micelle when get arranged parallel form the fibrils of 25nm thick known as microfibrils (5-15nm in diametre).These microfibrils form large sized bundles 0.5micrometre thick of cellulose fibre to form the macrofibrils. The macrofibrils consisting of many cellulose fibrils. The hemicellulose is composed of monosaccharide units such as arabinose, xylose, mannos& galactose. It occurs sometimes in between the macrofibrils of cellulose.

HEMICELLULOSE

They are heterogeneous groups of carbohydrates polymers constructed from various 5&6 carbons sugars including xylose, arabinose, mannose & galactose. The hemicellulose xylan which utilizes the pentose xylose as its main building block, accounts as much as 50% of the cell wall in woody tissues. Hemicellulose molecule binds to the cellulose microfibrils &to each other creating a coating that helps to bond these microfibrils together into a rigid interconnected network of cellulose & hemicellulose.


PECTINS
These are the polymers of carbohydrates D-galactouronic acid & D-glucouronic acid units .Like the glycosoaminoglycane of animal cell, they readily form hydrated gels. This property of pectin molecules is responsible for gelatin that occurs during the process of making jams and jellies .Pectin molecules are involved in binding adjacent cell wall together & in forming the matrix in which cellulose microfibrils are embedded. (FIG II)

LIGNINS
They are group of polymerized aromatic phenols. They occurs mainly in woody tissues, as cross linked network that contributes to the hardening of the cell wall. Lignin molecules are localized between the cellulose fibrils, where they function to rest compression forces . 


GLYCOPROTEINS
They are important constituent of plant cell wall, accounting as much as 10% of the total mass. Among the wall glycoprotein a group of common glycoprotein are called extensins. Extensins & other glycoprotein forms cross linked networks with each other as well as with cellulose microfibrils 

OTHER COMPONENT
In addition to the preceding components a small percent of the total mass of the cell wall is accounted for the lipids, including waxes & other complex polymers .Minerals such as Ca & K also occurs in the plant cell wall in the form of inorganic salts.


The cell wall originates in the developing cell plate. During telophase of the mitotic cell cycle, the phragmosome, a flattened membranous vesicle containing cell wall components,froms across the cell within a cytoskeletal array called the phragmoplast. 

The noncellulosic cell wall polysaccharides synthesized in the golgi apparatus and packaged in the vesicles fuse with the growing cell plate. The plate grows outward until the edges of the membranous vesicle fuse with the plasma membrane, creating two cells. Finally the new cell wall fuses with the existing primary cell wall, 

The plant golgi apparatus is a factory for the synthesis, processing and targeting of glycoproteins. Thus except for cellulose the polysaccharides, the structural proteins, and a broad spectrum of enzymes are coordinately secreted in golgi derived vesicles and targeted to the cell wall.

Cell enlargement depends on the activities of endoglycosidase, endotransglycosylase or expansin or some combination of these. But cell shape is largely governed by the pattern of cellulose deposition. Termination of cell growth is accompanied by cross linking reaction involving proteins and aromatic substances.

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