Membrane Structure and Dynamics

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CHAPTER 7 Membrane Structure and Dynamics

Membranes composed of lipids and proteins form the barrier between each cell and its environment. Membranes also partition the cytoplasm of eukaryotes into compartments, including the nucleus and membrane-bounded organelles. Each type of membrane is specialized for its various functions, but all biological membranes have much in common: a planar fluid bilayer of lipid molecules, integral membrane proteins that cross the lipid bilayer, and peripheral membrane proteins on both surfaces.

This chapter opens with a discussion of the lipid bilayer. It then considers examples of integral and peripheral membrane proteins before concluding with a discussion of the dynamics of both lipids and proteins. The following three chapters introduce three large families of membrane proteins: pumps, carriers, and channels. Chapter 11 explains how pumps, carriers, and channels cooperate in a variety of physiological processes. Chapters 24 and 30 cover plasma membrane receptor proteins.

Development of Ideas about Membrane Structure

Our current understanding of membrane structure began with E. Overton’s proposal in 1895 that cellular membranes consist of lipid bilayers (Fig. 7-1A). Biochemical experiments in the 1920s supported the bilayer hypothesis. It was found that the lipids extracted from the plasma membrane of red blood cells spread out in a monolayer on the surface of a tray of water to cover an area sufficient to surround the cell twice. (Actually, offsetting errors—incomplete lipid extraction and an underestimation of the membrane area—led to the correct answer!) X-ray diffraction experiments in the early 1970s established definitely that membrane lipids are arranged in a bilayer.

During the 1930s, cell physiologists realized that a simple lipid bilayer could not explain the mechanical properties of the plasma membrane, so they postulated a surface coating of proteins to reinforce the bilayer (Fig. 7-1B). Early electron micrographs strengthened this view, since when viewed in cross sections, all membranes appeared as a pair of dark lines (interpreted as surface proteins and carbohydrates) separated by a lucent area (interpreted as the lipid bilayer). By the early 1970s, two complementary approaches showed that proteins cross the lipid bilayer. First, electron micrographs of membranes that are split in two while frozen (a technique called freeze-fracturing; see Fig. 6-4D) revealed protein particles embedded in the lipid bilayer. Later, chemical labeling showed that many membrane proteins traverse the bilayer, exposing different regions of the polypeptide to the aqueous phase on the two sides. Light microscopy with fluorescent tags demonstrated that membrane lipids and some membrane proteins diffuse in the plane of the membrane. Quantitative spectroscopic studies showed that lateral diffusion of lipids is a rapid process but that flipping from one side of a bilayer to the other is a slow one. The fluid mosaic model of membranes (Fig. 7-1C) incorporated this information, showing transmembrane proteins floating in a fluid lipid bilayer. Subsequent work revealed structures of many proteins that span the lipid bilayer, the existence of lipid anchors on some membrane proteins, and a network of cytoplasmic proteins that restricts the motion of many integral membrane proteins (Fig. 7-1D).


Lipids form the framework of biological membranes, anchor soluble proteins to the surfaces of membranes, store energy, and carry information as extracellular hormones and as intracellular second messengers. Lipids are organic molecules generally less than 1000 D in size that are much more soluble in organic solvents than in water. They consist predominantly of aliphatic or aromatic hydrocarbons.

This chapter concentrates on major lipids found in biological membranes. After an introduction to their structures, the following section explains how the hydrophobic effect drives lipids to self-assemble stable bilayers. Membranes also contain hundreds of minor lipids, some of which might have important biological functions that are not yet appreciated. For example, during the 1980s, a minor class of lipids with phosphorylated inositol head groups first attracted attention when investigators found that they had a major role in signaling (see Fig. 26-7).


Phosphoglycerides (also called glycerolphospholipids) are the main constituents of membrane bilayers (Fig. 7-2). (These lipids are often called phospholipids, an imprecise term, as other lipids contain phosphate.) Phosphoglycerides have three parts: a three-carbon back-bone of glycerol, two long-chain fatty acids esterified to carbons 1 and 2 (C1 and C2) of the glycerol, and phosphoric acid esterified to C3 of the glycerol. Fatty acids have a carboxyl group at one end of an aliphatic chain of 13 to 19 additional carbons (Table 7-1). More than half of the fatty acids in membranes have one or more double bonds, which create a bend in the aliphatic chain. These bends contribute to the fluidity of the bilayer. Fatty acids and phosphoglycerides are amphiphilic, since they have both hydrophobic (fears water) and hydrophilic (loves water) parts. The aliphatic chains of fatty acids are hydrophobic. The carboxyl groups of fatty acids and the head groups of phosphoglycerides are hydrophilic. The cross-sectional areas of the head groups and the aliphatic tails are similar, so a phosphoglyceride is shaped approximately like a cylinder—an important factor in membrane structure. The hydrophobic effect (see Fig. 4-5) drives amphiphilic phosphoglycerides to assemble bilayers (see later).


Name Carbons Double Bonds (Positions)
Myristate 14 0
Palmitate 16 0
Palmitoleate 16 1 (Δ9)
Stearate 18 0
Oleate 18 1 (Δ9)
Linoleate 18 2 (Δ9, Δ12)
Linolenate 18 3 (Δ9, Δ12, Δ15)
Arachidonate 20 4 (Δ5, Δ8, Δ11, Δ14)

Cells make more than 100 major phosphoglycerides by using several different fatty acids and by esterifying one of five different alcohols to the phosphate. In general, the fatty acids on C1 have no or one double bond, whereas the fatty acids on C2 have two or more double bonds. Each double bond creates a permanent bend in the hydrocarbon chain. The alcohol head groups, rather than the fatty acids, give phosphoglycerides their names:

The several head groups confer distinctive properties to the various phosphoglycerides. All have a negative charge on the phosphate esterified to glycerol. Neutral phosphoglycerides—PE and PC—have a positive charge on their nitrogens, giving them a net charge of zero. PS has extra positive and negative charges, giving it a net negative charge like the other acidic phosphoglycerides (PA, PG, and PI). PI can be modified by esterifying one to five phosphates to the hexane ring hydroxyls. These polyphosphoinositides are highly negatively charged.

The complicated metabolism of phosphoglyce-rides can be simplified as follows: Enzymes can interconvert all phosphoglyceride head groups and remod-el fatty acid chains. For example, three successive enzymatic methylation reactions convert PE to PC, whereas another enzyme exchanges serine for ethanolamine, converting PS to PE. Other enzymes exchange fatty acid chains after the initial synthesis of a phosphoglyceride. These enzymes are located on the cytoplasmic surface of the smooth endoplasmic reticulum. Biochemistry texts provide more details of these pathways.

Several minor membrane phospholipids are variations on this general theme. Plasmalogens have a fatty acid linked to carbon 1 of glycerol by an ether bond rather than an ester bond. They serve as sources of arachidonic acid for signaling reactions (see Fig. 26-9). Cardiolipin has two glycerols esterified to the phosphate of PA.


Most sugar-containing lipids of biological membranes are sphingolipids. Sphingolipids get their name from sphingosine, a nitrogen-containing base (Fig. 7-3) that is the structural counterpart of glycerol and one fatty acid of phosphoglycerides. Sphingosine carbons 1 to 3 have polar substituents. A double bond between C4 and C5 begins the hydrocarbon tail. Two variable features distinguish the various sphingolipids: the fatty acid (often lacking double bonds) attached by an amide bond to C2 and the nature of the polar head groups esterified to the hydroxyl on C1.

The head groups of glycosphingolipids consist of one or more sugars. Some are neutral; others are negatively charged. Note the absence of phosphate. Sugar head groups of some glycosphingolipids serve as receptors for viruses. Alternatively, a phosphate ester can link a base to C1. These so-called sphingomyelins have phosphorylcholine or phosphoethanolamine head groups just like PC and PE. Receptor-activated enzymes remove phosphorylcholine from sphingomyelin to produce the second messenger ceramide (see Fig. 26-11). Sphingolipids are much more abundant in the plasma membrane than in membranes inside cells. The hydrocarbon tails of sphingosine and the fatty acid contribute to the hydrophobic bilayer, and polar head groups are on the surface.


Sterols are the third major class of membrane lipids. Cholesterol (Fig. 7-4) is the major sterol in animal plasma membranes, with lower concentrations in internal membranes. Plants, lower eukaryotes, and bacteria have other sterols in their membranes. The rigid four-ring structure of cholesterol is apolar, so it inserts into the core of bilayers with the hydroxyl on C3 oriented toward the surface.

Cholesterol is vital to metabolism, being situated at the crossroads of several metabolic pathways, including those that synthesize steroid hormones (such as estrogen, testosterone, and cortisol), vitamin D, and bile salts secreted by the liver. Cholesterol itself is synthesized (see Fig. 20-13) from isopentyl (5-carbon) building blocks that form 10-carbon (geranyl), 15-carbon (farnesyl), and 20-carbon (geranylgeranyl) isoprenoids. As is described later, these isoprenoids are used as hydrocarbon anchors for many important membrane-associated proteins. Isoprenoids are also precursors of natural rubber and of cofactors present in visual pigments.

Physical Structure of the Fluid Membrane Bilayer

In an aqueous environment, amphiphilic lipids spontaneously self-assemble into ordered structures in microseconds. The cylindrical shapes and amphiphilic nature of phosphoglycerides and sphingolipids favor formation of lamellar bilayers, planar structures with fatty acid chains lined up more or less normal to the surface and polar head groups on the surfaces exposed to water (Fig. 7-1D). Bilayer formation is energetically favorable, owing to the increase in entropy when the hydrophobic acyl chains interact with each other and exclude water from the core of the bilayer. This hydrophobic effect increases the entropy of the system and drives the assembly process.

An atomic model of a phosphoglyceride bilayer (Fig. 7-5

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