Endoplasmic Reticulum

Reticular network of membranes within cytoplasm is called endoplasmic

reticulum (ER). The labyrinthine network of ER pervades the entire

cytoplasmic fluid and provides continuous but a dynamic

compartmentalization and fluidity. In fact plasma membrane and nuclear

membrane are interconnected through various forms of endoplasmic

reticulum.

Endoplasmic Reticulum exhibits various forms and shapes. They also show

variations in their chemical composition and structural organization. Some

of the ER membranes are found as flat membranous sheets and some are in

the form of branched tubular structures. They also exist as membranes

vesicles. Nevertheless such forms are not constant and they undergo rapid

changes from one from to another form and vice versa.

Though ER is basically made up of two unit membranes containing lumen in

between which is filled with fluid. The extent and magnitude of ramification

of this membrane system varies from cell type to cell type. The cell that is

actively engaged in protein synthesis and metabolically active contains

greater amount of ER occupying more than ¾ th of the cytoplasmic space.

Endoplasmic membranes are further classified into rough endoplasmic

reticulum (RER) and smooth endoplasmic reticulum (SER). The former is

studded with a large number of ribosomes on its cytosolic surface. The

bound ribosomes are active in protein synthesis; on the contrary, SER is free
from ribosomes. In RER, ribosomes are bound to the membrane surface by

its larger subunit. The binding is attributed to the presence of two specific

transmembrane proteins called ribophorins of mol wt. 65 KD and 63 KD

respectively. These proteins are absent in SER. Most of the ribosomes

found on RER membranes are in the form of polysome complexes actively

engaged in protein synthesis. The nascent polypeptides thus synthesized

enter into the lumen by its NH3 end through a groove found in the large

ribosomal subunit. The first 20-40 amino acid recidues found at the NH2

end of the polypeptide chain act as signal polypeptides and they facilitate

the translocation of proteins into the lumen of the RER. The proteins thus

packed into RER lumen are transported towards SER to reach specific

destinations. Some of the ER membranes carrying proteins either reach

Golgi bodies or pinch off into vesicles which ultimately find their way into

respective organelles. The ER membranes are supported by microtubular

cytoskeleton structures. That is the reason they remain intact and also

mobile. At the time of cell division at end of prophase MTs undergo

depolymerization, with that ER collapse into fine vesicles. They reappear at

the return of telophase, and MTs help in organizing the membranes.

Chemical Composition:

Endoplasmic reticular membranes contain high lipid content in relation to its

protein partners. The common lipids found are phospholipids, phosphotidyl

inositol, neutral lipids, sulfolipids, cholesterol and some phytosterols. Nearly

30-40 different types of ER membrane proteins have been isolated, of

which many are enzymes like cytochrome P 450 and its subgroups, electron

transport protein complexes like Cytochrome C reductase, Cytochrome b5

reductase. Gulcose 6 phosphotases are also common in ER. Furthermore,

though ER shows similarity in their structural appearances they exhibit

chemical heterogeneity at their cytoplasmic and lumen surfaces. They do

contain a variety of resident proteins, which perform functions like, protein

folding, protein modification and protein transport.

Biogenesis:

There are many views with regard to the biosynthesis of endomenbranes.

According to one view, the membranes develop from plasma membranes.

According to one view, the membranes develop from plasma membranes by

inward invagination and growth. But other view points out that ER develop

from the outer nuclear membrane. Both these views point out that ER

develops from both plasma membranes and outer nuclear membrane. It is

also true that both plasma membranes and nuclear membrane are also

derived from endomenbranes. In protein synthesis and other cell membrane

components are involved in the synthesis of various liquid components, thus

they get self assembled by incorporating required proteins and lipid and

other fatty acid derivatives.

Functions:

Endoplasmic membranes found in the cytoplasmic fluid are highly dynamic

and show rapid but sweeping movements. In fact they exhibit high fluidity

because the membranes are always in constant breakdown and

reorganization. Some of the important functions of it are listed below RER

is engaged in the synthesis of a wide variety of proteins at different site.

SER is involved in glycogenolysis. In plant cells SER is responsible for the

secretion of vesicles containing raw materials for cell wall formation. SER is

also responsible for the synthesis of various kinds of lipids and its

derivatives. ER enzymes like Cyt. P450 and its allied enzymes are capable of

detoxification of drugs. Alcohols and other metabolically harmful

components are detoxified; but the same can also cause damage to the liver.