MOLECULAR BIOLOGY ASSIGNMENT

on

PLASMA MEMBRANE IN PLANTS

SUBMITTED BY:
Aditya Kanwal 2012BT09 M.Tech Biotechnology MNNIT

INTRODUCTION
We all know that cells are the basic functional and structural unit of life. For these cells to survive and perform their characteristic functions, they need to create a specific micro-environment. This micro-environment is provided by the cytoplasm which is separated from the surroundings with the help of a barrier. Similar is the case for each organelle (in eukaryotes), which in turn needs to compartmentalize and create a micro-environment within themselves. These barriers that separate the internal cell fluids (like cytoplasm, nucleoplasm etc.) from the surroundings by compartmentalization are known as Biomembranes. All biomembranes are composed of a lipid bi-layer and proteins. Lipid bilayer forms the permeability barrier whereas proteins perform all other functions from active transport of molecules to cell signalling. Its these proteins that decide the function of cells because the lipid components are almost the same in every membrane. One of the most important feature of a biomembrane is its selective permeablility. It allows only small and uncharged molecules to pass through it via diffusion. For larger charged molecules, involvement of membrane proteins is required and transport is active i.e. energy expenditure is done via ATP consumption. This property is what makes this structure so appropriate to play its role. All membrane molecules, let it be lipids or proteins, are able to diffuse freely within the plane of the membrane i.e. laterally. This property provides a clue for the elucidation of membrane model. The fluid-mosaic membrane model is based on this concept and will be discussed later. A single plant cell can have 17-20 different types of biomembranes like plasma membrane, mitochondrial membrane, chloroplast membrane, nuclear membrane, golgi cisternae, peroxisomal membrane, thylakoid membrane, Endoplasmic reticulum membrane, tonoplast, endosomal membrane etc. All cells inherit the complete set of these membranes from the pre-existing cells and these membranes cannot arise de novo. This helps in the maintainence of integrity and continuity of composition from one generation to next.

The Fluid Mosaic Model

The fluid mosaic model was given by S.J. Singer and G.L. Nicolson in 1972. It considers biological membranes as two-dimensional fluids in which lipids and proteins diffuse and are laterally mobile in it. This model combines various inherent properties of cellular membranes like molecular and physicochemical characteristics of its components i.e. lipids and proteins, their assembly and their association with each other. LIPIDS: Lipids are a large and diverse group of organic compounds involved in storage, structure, signalling and nutrition. Here, we are concerned only with its structural roles. Phospholipids, sphingolipids and sterols are the ones involved in cellular compartmentalization. These molecules are amphipathic in nature i.e. they have two domains: one is hydrophillic and the other is hydrophobic. This property plays a major role in assembly of these lipids. In a two phase system having water and an organic solvent, these molecules will arrange themselves in the interface with hydrophillic ends in water and hydrophobic ends in organic solvent. However, if we add these amphipathic molecules in water, in the absence of organic solvent, they will self assemble so as to hide their hydrophobic end from water. This leads to the formation of bilayers as they are the most stable conformation in such a situation. Phospholipids are the most common type of membrane lipid, possesing a charged polar head that interacts with aqueous environments and two hydrophobic tails 14-24 carbon atom long. These tails can have double bonds, leading to the formation of kinks which help in the packaging of bilayer and maintaining its fluidity. These are derivatives of glycerol 3-phosphate with polar head group attached to the phosphate group. Another class is of sphingolipids which are derived from sphingosine, an amino alcohol with a long hydrocarbon chain, and contain a long-chain fatty acid attached to the sphingosine amino group. These sphingolipids are asymmetrically distributed in the plasma membrane both laterally and in the exoplasmic and cytoplasmic leaflet.

Sterols are another class of lipids that occur in membranes and differ from cholesterol that occurs in animal cells. In plant membranes generally three kinds of sterols are found. They are: Beta-sitosterol Campesterol Stigmasterol All these are abundant in plant cell membranes and provides composure to the membranes. As the individual lipid molecules in the lipid bilayer are not bonded to each other covalently, they can move freely within the plane of bilayer. This creates the characteristic fluidity of membranes. These phospholipids can diffuse laterally, rotate, bob up and down, and flip- flop. Rotation and bobbing occurs within a single position or axis, lateral diffusion occurs within the monolayer and flip-flop occurs across the bilayer. Flip-flop is seen to take place mainly in the presence of flippases which mediate the movement of newly synthesized lipids across the bilayer and is rare otherwise. Sterols, however, can diffuse rapidly and flip-flop without any additional help because of their large hydrophobic domains. Lipids have two major functions to perform: 1) To create a barrier: It does so by providing a hydrophobic core in the centre of bilayer that prevents the diffusion of hydrophilic solutes across the membrane. Proteins selectively help in the transport of those hydrophilic solutes that are required by the cell. 2) To maintain its composure: Stabilization and maintainence of the bilayer structure is mainly through hydrophobic and van der Waals interactions within the lipid chains. This helps it in maintaining cell structure, prevent lysis and repairing itself in case of damage.

 PROTEINS: Membrane proteins are the molecules present in the membrane of the cells and their organelles. Every biological membrane has the same basic bilayer structure but its the proteins associated with a particular membrane that makes it different from others . They play a variety of structural and functional roles depending on the cell type and subcellular location. The extracellular domains of proteins generally bind to other molecules like ions, signalling messengers, carbohydrates and to certain adhesion molecules on other cells. Domains within the plasma membrane help mainly in transport of molecules across the membrane and in interaction with hydrophobic region of lipid bilayer. Domains towards the cytosolic face are involved in intracellular signaling pathways, anchoring cytoskeletal proteins and provide surface for various cytosolic reactions to occur. These membrane proteins can be divided into three categories: -Integral membrane proteins: These are permanently bound to the lipid bilayer. They have atleast one hydrophobic domain that associates with the hydrophobic interior of the bilayer. This also makes them water insoluble. Their removal requires the degradation of bilayer. -Peripheral membrane proteins: These are temporarily associated with the bilayer or with the integral proteins with the help of weak interactions. These can be removed from the bilayer without damaging it. -Lipid anchored proteins: These were not classified in the original fluid mosaic model and were classified later. These proteins are bound to lipid bilayer with the help of lipidated amino acid residues. In this, some short chain lipids like fatty acids, prenyl lipids, etc. act as connectors between the lipid bilayer and the proteins. They can be divided further into four classes based on the kind of group it is linked with: -Fatty acid linked proteins -Prenyl group linked proteins -Phosphatidylinositol anchored proteins -Cholesterol linked proteins. The two most common structural classes of transmembrane proteins are alpha-helical bundles and beta-barrels. These two structural motifs have been seen to be conserved for transmembrane proteins.

THE PLASMA MEMBRANE

All living cells are surrounded by a common architecture of lipid bilayer called the plasma membrane or the cell membrane. They act as an active interface between the cell and its surroundings by: mediating transport of molecules in and out of the cell maintaining membrane potential mediating cell signalling by interacting with small chemical messengers associating with elements of cytoskeleton to provide shape and structure synthesis and assembly of cell wall molecules. formation of plasmodesmata in conjunction with specialized domains of endoplasmic reticulum. As discussed already, plasma membranes help in the transport of molecules in and out of the cells. Smaller and uncharged molecules can pass the bilayer passively in the presence of concentration gradient via diffusion whereas large and charged molecules pass actively with the help of membrane proteins that utilize energy for transporting these molecules across the membrane. Membrane potential also allows the cells to transport various ions and metabolites into the cell. Cell signalling is another major property mediated by membrane proteins. This cellular signalling controls development, homeostasis and response to damage and infection. This signalling can be either over short distances (between two adjacent cells) or over long distances (between two different tissues located far apart). By interacting with cytoskeletal elements like microfilaments, microtubules and intermediate filaments, plasma membrane provides shape and structure to the cell. Also, the plasmamembrane is the site for the synthesis of cellulose which is the most abundant biopolymer on earth. It has been seen that the plant cell membranes contain terminal complexes or particle rosettes that are complexes of many cellulose synthases and are the sites of cellulose synthesis. Two transferase molecules can act simultaneously from opposite sides and add two glucoses at a time to the growing chain. This explains the rotation of alternating glucoses in cellulose molecules. Plasmodesmata are another characteristic structural feature of plant membranes that cross the cell walls and connect adjacent cells enabling transport and communication. These are made up of three main layers formed by plasma membrane, cytoplasmic sleeve and desmotubule. The plasma membrane involved is the continous extension of cell membrane or the plasmolemma and is similar in structure to that of the bilayer.

The ratio between various types of lipids (i.e. phospholoipids, sphingolipids and sterols) present in plant cell membranes is very variable amongst cells of different organs of same plant and cells of same organs in different plants. However the reason behind this phenominan is not clearly understood.
Table showing lipid composition of plasma membranes from various tissues (mole %) [from - Buchanan B.B, Gruissem W, Jones R.L, Biochemistry and Molecular Biology of Plants(2002), Wiley]

LIPID TYPE Phospholipids Free Sterols Steryl glucosides Acylated steryl glucosides Glucocerebosides

BARLEY ROOT 26 57 7 9

BARLEY LEAF 44 35 16

SPINACH LEAF 64 7 13 14

Almost all plants are able to withstand cold temperatures. To survive, plants undergo a process called cold acclimation. This process involves changes in physiological properties and the chemical composition of membranes, cytoplasm and the cell wall. It has been seen that the composition of phospholipids increases whereas the composition of glucocerebosides decreases. Plasma membrane proteins on the other hands play variety of roles as already discussed.

REFERENCES
Buchanan B. , Gruissem W. , Jones R. , Biochemistry & Molecular Biology of Plants(2002) John Wiley & Sons. Lodish H., Berk A., Kiaser C.A.,Krieger M.,Scott M.P., Molecular Cell Biology, (2007) Macmillan Higher Education.