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Fig. 6.7 A scheme of an animal cell
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bimolecular layer (bilayer) of phospholipids, with their long alkyl chains oriented inwards (Fig. 6.9). Electron microscope studies have revealed that the central layer corresponds to this largely hydrocarbon region. In fact, biological membranes are still more complex as will be discussed in Section 6.4.2.
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6.4.1 Composition of biological membranes Biological membranes consist of lipids, proteins and also sugars, sometimes mutually bonded in the form of lipoproteins, glycolipids and glycoproteins. They are highly hydrated water forms up to 25 per cent of the dry weight of the membrane. The content of the various protein and lipid components varies with the type of biological membrane. Thus, in
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Fig. 6.8 Electron photomicrograph of mouse kidney mitochondria. The structure of both the cytoplasmatic membrane (centre) and the mitochondrial membranes is visible on the ultrathin section. Magnification 70,000x. (By courtesy of J. Ludvik)
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Fig. 6.9 Characteristic structures of biological membranes. (A) The fluid mosaic model (S. J. Singer and G. L. Nicholson) where the phospholipid component is predominant. (B) The mitochondrial membrane where the proteins prevail over the phospholipids
myelin cell membranes (myelin forms the sheath of nerve fibres) the ratio protein: lipid is 1:4 while in the inner membrane of a mitochondrion (cf. page 464) 10:3. The lipid component consists primarily of phospholipids and cholesterol. The most important group of phospholipids are phosphoglycerides, based on phosphatidic acid (where X = H), with the formula O O H2C O C R R C O CH
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where R and R' are alkyl or alkenyl groups with long chains. Thus, glycerol is esterified at two sites by higher fatty acids such as palmitic, stearic, oleic, linolic, etc., acids and phosphoric acid is bound to the remaining alcohol group. In phospholipids, the phosphoric acid is usually bound to nitrogen-
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substituted ethanolamines. For example, the phospholipid lecithin is formed by combination with choline (X = CH2CH2N+(CH3)3). Other molecules can also be bound, such as serin, inositol or glycerol. Some phospholipids contain ceramide in place of glycerol: CH(OH)CH=CH(CH2)12CH3 CHNHCOR CH2OH where R is a long-chain alkyl group. The ester of ceramide with phosphoric acid bound to choline is sphingomyelin. When the ceramide is bound to a sugar (such as glucose or galactose) through a /3-glycosidic bond, cerebrosides are formed. The representation of various lipidic species strongly varies among biological membranes. Thus, the predominant component of the cytoplasmatic membrane of bacterium Bacillus subtilis is phosphatidyl glycerol (78 wt.%) while the main components of the inner membrane from rat liver mitochondrion are phosphatidylcholine = lecithin (40 wt.%) and phophatidylethanolamine = cephalin (35 wt.%). Cholesterol contributes to greater ordering of the lipids and thus decreases the fluidity of the membrane. Proteins either strengthen the membrane structure (building proteins) or fulfil various transport or catalytic functions (functional proteins). They are often only electrostatically bound to the membrane surface (extrinsic proteins) or are covalently bound to the lipoprotein complexes (intrinsic or integral proteins). They are usually present in the form of an or-helix or random coil. Some integral proteins penetrate through the membrane (see Section 6.4.2). Saccharides constitute 1-8 per cent of the dry membrane weight in mammals, for example, while this content increases to up to 25 per cent in amoebae. They are arranged in heteropolysaccharide (glycoprotein or glycolipid) chains and are covalently bound to the proteins or lipids. The main sugar components are L-fucose, galactose, manose and sometimes glucose, N-acetylgalactosamine and N-acetylglucosamine. Sialic acid is an important membrane component: CH2OH CHOH CHOH
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OH This substance often constitutes the terminal unit in heteropolysaccharide chains, and contributes greatly to the surface charge of the membrane.
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438 6.4.2 The structure of biological membranes
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Following the original simple concepts of Gorter and Grendel, a large number of membrane models have been developed over the subsequent half a century; the two most contrasting are shown in Fig. 6.9. The basic characteristic of the membrane structure is its asymmetry, reflected not only in variously arranged proteins, but also in the fact that, for example, the outside of cytoplasmatic (cellular) membranes contains uncharged lecithin-type phospholipids, while the polar heads of strongly charged phospholipids are directed into the inside of the cell (into the cytosol). Phospholipids, which are one of the main structural components of the membrane, are present primarily as bilayers, as shown by molecular spectroscopy, electron microscopy and membrane transport studies (see Section 6.4.4). Phospholipid mobility in the membrane is limited. Rotational and vibrational motion is very rapid (the amplitude of the vibration of the alkyl chains increases with increasing distance from the polar head). Lateral diffusion is also fast (in the direction parallel to the membrane surface). In contrast, transport of the phospholipid from one side of the membrane to the other (flip-flop) is very slow. These properties are typical for the liquid-crystal type of membranes, characterized chiefly by ordering along a single coordinate. When decreasing the temperature (passing the transition or Kraft point, characteristic for various phospholipids), the liquid-crystalline bilayer is converted into the crystalline (gel) structure, where movement in the plane is impossible. The cells of the higher plants, algae, fungi and higher bacteria have a cell wall in addition to the cell membrane, protecting them from mechanical damage (e.g. membrane rupture if the cell is in a hypotonic solution). The cell wall of green plants is built from polysaccharides, such as cellulose or hemicelluloses (water-insoluble polysaccharides usually with branched structure), and occasionally from a small amount of glycoprotein (sugar-protein complex). Because of acid groups often present in the pores of the cell wall it can behave as an ion-exchanger membrane (Section 6.2). Gram-positive bacteria (which stain blue in the procedure suggested by H. Ch. J. Gram in 1884) have their cell walls built of cross-linked polymers of amino acids and sugars (peptidoglycans). The actual surface of the bacterium is formed by teichoic acid (a polymer consisting of a glycerolphosphate backbone with linked glucose molecules) which makes it hydrophilic and negatively charged. In this case the cell wall is a sort of giant macromolecule a bag enclosing the whole cell. The surface structure of gram-negative bacteria (these are not stained by Gram's method and must be stained red with carbol fuchsin) is more diversified. It consists of an outer membrane whose main building unit is a lipopolysaccharide together with phospholipids and proteins. The actual cell
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439 wall is made of a peptidoglycan. A loosely bounded polysaccharide layer termed a glycocalyx is formed on the surface of some animal cells. 6.4.3 Experimental models of biological membranes Phospholipids are amphiphilic substances; i.e. their molecules contain both hydrophilic and hydrophobic groups. Above a certain concentration level, amphiphilic substances with one ionized or polar and one strongly hydrophobic group (e.g. the dodecylsulphate or cetyltrimethylammonium ions) form micelles in solution; these are, as a rule, spherical structures with hydrophilic groups on the surface and the inside filled with the hydrophobic parts of the molecules (usually long alkyl chains directed radially into the centre of the sphere). Amphiphilic substances with two hydrophobic groups have a tendency to form bilayer films under suitable conditions, with hydrophobic chains facing one another. Various methods of preparation of these bilayer lipid membranes (BLMs) are demonstrated in Fig. 6.10. If a dilute electrolyte solution is divided by a Teflon foil with a small window to which a drop of a solution of a lipid in a suitable solvent (e.g. octane) is applied, the following phenomenon is observed (Fig. 6.10A). The layer of lipid solution gradually becomes thinner, interference rainbow bands appear on it, followed by black spots and finally the whole layer becomes black. This process involves conversion of the lipid layer from a multimolecular thickness to form a bilayer lipid membrane. Similar effects were observed on soap bubbles by R. Hooke in 1672 and I. Newton in 1702. Membrane thinning is a result of capillary forces and dissolving of components of the membrane in the aqueous solution with which it is in contact. The thick layer at the edge of the membrane is termed the Plateau-Gibbs boundary. Membrane blackening is a result of interference between incident and reflected light. If the membrane thickness is much less than one quarter of the wavelength of the incident light, then the waves of the incident and reflected light interfere and the membrane appears black against a dark background. The membrane thickness can be found from the reflectance of light with a low angle of incidence, from measurements of the membrane capacity and from electron micrographs (application of a metal coating to the membrane can, however, lead to artefact formation). The thickness of a BLM prepared from different materials lies in the range between 4 and 13 nm. A BLM can even be prepared from phospholipid monolayers at the water-air interface (Fig. 6.10B) and often does not then contain unfavourable organic solvent impurities. An asymmetric BLM can even be prepared containing different phospholipids on the two sides of the membrane. A method used for preparation of tiny segments of biological membranes (patch-clamp) is also applied to BLM preparation (Fig. 6.IOC).
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