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    Bio-membranes

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    Eukaryotes.. Eukaryotic cell has an efficient membrane functions due to increase in the surface

    area. This is attributed to the presence of many internal membranes with

    numerous convolutions and infoldings.

    Membranes surround the nucleus, mitochondria, and (in plant cells) the

    chloroplasts as well as Endoplasmic Reticulum. Membranes form stacks of sacs

    with Golgi apparatus, surround lysosomes responsible for intracellular digestion,

    peroxisomes as well as form small vesicles and, in plants, a large liquid-filledvacuole.

    Each organelle is surrounded by one or more biomembranes, with unique set of

    proteins specific for its characteristic function.

    All these membrane-bounded structures correspond to distinct internal

    compartments within the cytoplasm. In a typical animal cell these organelles

    occupy nearly half the total cell volume and the remaining is the cytosol.

    Keeping all the organelles in proper shape and in a controlled movement

    eukaryotic cells have a tough cytoskeleton.

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    Functions ofbiomembranes

    Cell membranes are crucial to the life of the cell.

    The plasma membrane encloses the cell, defines its boundaries, andmaintains the essential differences between the cytosol and the extracellular

    environment.

    The membranes of the ER, Golgi apparatus, mitochondria, and other

    membrane-bounded organelles in eukaryotic cells maintain the characteristicdifferences between the contents of each organelle and the cytosol.

    Responsible for maintaining ion gradients across membranes (function of

    specialized membrane proteins) which can be used to synthesize ATP, to drive

    the transport of selected solutes, or, in production and transmission of

    electrical signals in nerve and muscle cells.

    Plasma membrane also contains proteins that act as sensors of external

    signals, allowing the cell to change its behavior in response to environmental

    changes; These are called receptors , that transfer information rather than

    ions or molecules across the membrane.

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    Membrane structure

    Inspite of diverse functions, all the cell membranes have a common

    basic molecular structure.

    Each membrane is a thin film of lipid and protein molecules, held

    together mainly by non-covalent interactions. Carbohydrates are

    present in covalent linkage with lipids or proteins.

    The lipid molecules are arranged as a continuous double layer (~5 nm

    thick). This lipid bilayerprovides the basic structure of the membrane

    and is a relatively impermeable barrier to the passage of most water-

    soluble molecules.

    Cell membranes are asymmetrical structures:

    The lipid and protein compositions of the outside and inside faces

    differ from one another in ways that reflect the different functions

    performed at the two surfaces of the membrane.

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    (a) Electron micrograph of plasma membrane (b, c) Schematic representations of lipid bilayer

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    The Lipid bilayer-Composition

    The lipid bilayer has been firmly established as the universal basisfor cell-membrane structure.

    It is easily seen by ordinary electron microscopy, although

    specialized techniques, such as x-ray diffraction and freeze-fracture

    electron microscopy, are needed to reveal the details of itsorganization.

    The bilayer structure is attributed to the special properties of the

    lipid molecules i.e. the amphipathic nature, due to which it

    assembles spontaneously to form bilayers.

    The most abundant are the phospholipids consist of two long-chain,

    nonpolar fatty acid groups linked (usually by an ester bond) to small,

    highly polar groups, including a phosphate.

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    The tails are usually fatty acids which may vary in length (14-24 C atoms).

    One tail usually has one or more cis-double bonds (that is , it is unsaturated),

    while the other tail does not (that is, it is saturated). Each double bondcreates

    a small kinkin the tail.

    Differences in the length and saturation of the fatty acid tails influence the

    ability of phospholipid molecules to pack against one another, and for this

    reason they affect the fluidity of the membrane.

    Major phosphoglyceride, phopshatidylcholine

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    Types of fatty acids in membrane

    14:0 myristic acid

    16:0 palmitic acid

    18:0 stearic acid

    18:1 cis 9 oleic acid

    18:2 cis 9,12 linoleic acid

    18:3 cis 9,12,15 linonenic acid

    20:4 cis 5,8,11,14 arachidonic acid 20:5 cis 5,8,11,14,17 eicosapentaenoic acid

    (an omega-3 fatty acid because of

    double bond 3 C from distal end)

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    Three Classes of Lipids Are Foundin Biomembranes

    A typical biomembrane is assembled from phosphoglycerides or

    glycerophospholipids (major class), sphingolipids, and steroids.

    Phosphoglycerides contain two fatty acyl chains esterified with glycerol-3-P and a polar head group

    attached to the P group; nature of the polar group decides the classification. For eg. In

    Phosphatidylcholine head group consists of choline, a positively charged alcohol, esterified to the

    negatively charged phosphate.

    Esterificn of C1 and

    C2 hydroxylswith fatty acids and

    C3 hydroxyl with

    phosphate

    Phosphate esterified to an

    alcohol of a polar headgrp: choline, Inositol,

    ethanolamine, serine

    X=OR

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    Sphingolipids andCholesterol are other two classes of membrane lipids

    Sphingolipids are derivatives

    of sphingosine (red), anamino alcohol with a long

    hydrocarbon chain.

    Various fatty acyl chains are

    connected to sphingosine byan amide bond to yield

    ceramide inolved in forming

    complex sphingolipids.

    The sphingomyelins (SM),which contain a phospho-

    choline or phospho-

    ethanolamine head group.

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    Cholesterol.Like other membrane lipids, the steroid

    cholesterol is amphipathic. Its single

    hydroxyl group is equivalent to the polar

    head group in other lipids; the

    conjugated ring and short hydrocarbon

    chain form the hydrophobic tail.

    Other sphingolipids are glycosphingolipids

    with single sugar residue or branched

    oligosaccharide attached to the sphingosine

    backbone. eg A cerebroside is a

    sphingolipid (ceramide) with a

    monosaccharide such as glucose or

    galactose as polar head group.

    A ganglioside is a ceramide with a polar

    head group that is a complexoligosaccharide, including the acidic sugar

    derivative sialic acid

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    In aqueous solution, when lipid molecules are surrounded on all sides by

    water, they tend to aggregate so that their hydrophobic tails are buried in the

    interior and their hydrophilic heads are exposed to water.

    Depending on their shape, they can do this in either of two ways: they form

    spherical micelles, with the tails inward, or they can form bimolecular sheets,

    or bilayers , with the hydrophobic tails sandwiched between the hydrophilic

    head groups.

    The hydrophobic effect and van der Waalsinteractions, cause the tail groups to self-

    associate into a bilayer with the polar head

    groups oriented toward water.

    Because of their cylindrical shape,

    membrane phospholipid moleculesspontaneously form bilayers in aqueous

    environments and tend to seal on

    themselves to form compartments.

    This is one property that makes it an

    ideal structure to form membranes.

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    Liposomes: artificial micelles ofphospholipids