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.