Cell membranes amphipathic molecules

CTC, used extensively to monitor calcium release in both whole cells and isolated organelles (28-33), is an amphipathic molecule that easily passes through cell membranes (see Figure 1). The fluorescence of this probe is enhanced more than fiftyfold by binding of calcium when the dye is intercalated into biological membranes. 

The nonpolar lipid core consists of mainly triacylglycerol and cholesteryl ester and is surrounded by a single surface layer of amphipathic phospholipid and cholesterol molecules (Figure 25-1). These are oriented so that their polar groups face outward to the aqueous medium, as in the cell membrane (Chapter 14). The protein moiety of a lipoprotein is known as an apo-lipoprotein or apoprotein, constituting nearly 70% of some HDL and as litde as 1% of chylomicrons. Some apolipoproteins are integral and cannot be removed, whereas others are free to transfer to other hpoproteins. 

The cells of all contemporary living organisms are surrounded by cell membranes, which normally consist of a phospholipid bilayer, consisting of two layers of lipid molecules, into which various amounts of proteins are incorporated. The basis for the formation of mono- or bilayers is the physicochemical character of the molecules involved these are amphipathic (bifunctional) molecules, i.e., molecules which have both a polar and also a non-polar group of atoms. Examples are the amino acid phenylalanine (a) or the phospholipid phosphatidylcholine (b), which is important in membrane formation. In each case, the polar group leads to hydrophilic, and the non-polar group to hydrophobic character. 

Phospholipids are the most important of these liposomal constituents. Being the major component of cell membranes, phospholipids are composed of a hydrophobic, fatty acid tail, and a hydrophilic head group. The amphipathic nature of these molecules is the primary force that drives the spontaneous formation of bilayers in aqueous solution and holds the vesicles together. 

Although the notion of monomolecular surface layers is of fundamental importance to all phases of surface science, surfactant monolayers at the aqueous surface are so unique as virtually to constitute a special state of matter. For the many types of amphipathic molecules that meet the simple requirements for monolayer formation it is possible, using quite simple but elegant techniques over a century old, to obtain quantitative information on intermolecular forces and, furthermore, to manipulate them at will. The special driving force for self-assembly of surfactant molecules as monolayers, micelles, vesicles, or cell membranes (Fendler, 1982) when brought into contact with water is the hydrophobic effect. 

Amphipathic molecules can form bilayered lamellar structures spontaneously if they have an appropriate geometry. Most of the major cell membrane lipids have a polar head, most commonly a glycerophosphorylester moiety, and a hydrocarbon tail, usually consisting of two... 

We have encountered examples of simple lipid bilayers earlier. These bilayers are composed largely of amphipathic molecules. In water, they have their hydrophobic parts occupying the center of the bilayer and their hydrophilic parts occupying the bilayer surface. Such bilayers form a continuous and essential structural feature of virtually all biological membranes. We need to distinguish between that layer which faces out from the cell and is in contact with the external environment, the exoplasmic leaflet, and that which faces in and is in contact with the cellular contents, the cytoplasmic leaflet. As we shall see, these two aspects of the lipid bilayer are quite distinct. 

Cell membranes are bilayers of amphipathic acids, for example phospholipids and sterols, which contain globular proteins. The structure is governed by the essential requirement for stability in an aqueous environment, that is, the hydrophobic tails of the lipid molecules point towards each other, leaving the outer surfaces composed of polar, hydrophilic groups. 

Why are membranes so important to cells Besides their obvious importance in enclosing and defining the limits of cells, membranes are the result of a natural aggregation of amphipathic molecules. They also represent a natural arrangement for boundaries between different aqueous phases within a cell. In addition, membranes are the "habitat" for many relatively nonpolar molecules formed by metabolism. These include proteins with hydrophobic surfaces and... 

Phospholipids are ideal compounds for making membranes because of their amphipathic nature (see chapter 17). The polar head-groups of phospholipids prefer an aqueous environment, whereas the nonpolar acyl substituents do not. As a result, phospholipids spontaneously form bilayer structures (see fig. 17.6), which are a dominant feature of most membranes. The phospholipid bilayer is the barrier of the cell membrane that prevents the unrestricted transport of most molecules other than water into the cell. Entry of other molecules is allowed if a specific transport protein is present in the cell membrane. Similarly, the phospholipid bilayer prevents leakage of metabolites from the cell. The amphipathic nature of phospholipids has a great influence on the mode of their biosynthesis. Thus, most of the reactions involved in lipid synthesis occur on the surface of membrane structures catalyzed by enzymes that are themselves amphipathic. 

Fatty acids are an essential component of all living cells [28] and are used in several major cellular functions. Their structural roles include use in biological membranes, facilitating the compartmentalization that is a major feature of cellular organisms. Here, hydrophilic alcohols including choline, ethanolamine, and serine are esterified with fatty acids to create amphipathic molecules which are the primary components of biological membranes. Proteins may be acylated with fatty acids as a means of targeting these proteins to membrane-bound loca-... 

At some stage in evolution, sufficient quantities of appropriate amphipathic molecules must have accumulated from biosynthetic or other processes to allow some nucleic acids to become entrapped and cell-like organisms to form. Such compartmentalization has many advantages. When the components of a cell are enclosed in a membrane, the products of enzymatic reactions do not simply diffuse away into the environment but instead are contained where they can be used by the cell that produced them. The containment is aided by the fact that nearly all biosynthetic intermediates and other hiochemicals include one or more charged groups such as phosphates or carboxylates. Unlike more nonpolar or neutral... 

However, the most important use for lipids in the cell is in the formation of membranes, membranes contain amphipathic molecules, i.e., ones with a hydrophobic end and a hydrophilic end. 

Phospholipids and sphingolipids are major components of cell membranes. They are amphipathic molecules that is, one portion of the molecule is hydrophilic and associates with H20, and another portion contains the hydrocarbon chains derived from fatty acids, which are hydrophobic and associate with lipids (see Figure 6-4). 

Generalized structure of a lipoprotein molecule showing the distribution of polar components in an outer shell composed of free cholesterol, phospholipids, and amphipathic proteins and in an inner core composed of neutral triacylglycerols and cholesteryl esters. Phospholipids are oriented with polar head groups toward the aqueous environment and hydrophobic tails toward the neutral core, analogous to their positioning in the outer leaflet of the typical cell membrane. 

There are other chemical questions surrounding the emergence of life. For example, membranes are needed to confine the contents of cells and provide a stable, controlled environment for biochemical reactions. Some fairly simple organic molecules (amphipathic molecules), in which one end has an affinity for water (i.e. it is hydrophilic) but the other does not (i.e. it is hydrophobic, like oil), can naturally form membranelike structures, which, in the presence of water, form cell-like spheres (see Box 1.7). Self-replication is another vital factor, which needs to be accomplished with minimal error.The information necessary for con-... 

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