Cell Biology Chapter 11 Membrane Structure and Function Membrane Functions o Cells have membranes so they can: Compartmentalize Regulate flow of materials and information o Some membranes have enzymatic, signal transduction, and energy production systems localized within or upon them. Comprised primarily of proteins. o Membrane model is the Fluid Mosaic Model; membrane comprised of a mixture of lipids, and proteins, free to move laterally so the membrane is a dynamic structure. Mobility is a critical membrane characteristic: Without it, the membrane is unable to function Cellular and Intracellular Membranes o Plasma Membrane: Hydrophobic core barrier to polar compounds, ions Keeps desirable substances in, undesirable substances out Allows gradients to be built Proteins form channels or pumps to regulate transit of polar molecules, ions across membrane o Intracellular Membrane: Compartmentalize functions Enzymes and substrates of those functions isolated, colocalized Facilitate locally high concentrations of enzymes and substrates so reactions more efficient and faster Allows gradients to be built Transport Regulation o Lipophilic compounds cross w/o special transporters o Polar/ionic molecules require transport proteins o Transport may be passive, crossing membrane from high concentration to low concentration regions o Transport may be active, using energy to pump ions, sugars, amino acids across membranes against electrical and/or concentration gradients o Direct Communication: Plasma membranes of cells may be fused together forming gap junctions between animal cells across which signals and low molecular weight substances freely pass. Cardiac cells interconnected by gap junctions. Plasmodesmata: Intercellular bridges in plants Signal Transduction o Receptors: First step in signal transduction. Proteins. o Signal from outside received on outer plasma membrane surface by receptor proteins, causing changes in receptors that start signal transduction cascade resulting in release of secondary messengers in cytoplasm o 2md messenger events on inside transmit information that a signal has been received to internal cellular systems o Steroid receptors in cytoplasm or nucleus, not on cellular surface, as steroids are lipid-soluble and diffuse freely across plasma and nuclear membranes. Membrane Lipids o Phospholipids o Sphingolipids Mostly neurons o Gycolipids Neurons, chloroplasts o Sterols: Cholesterol; phytosterols. Hopanoids-not sterols Similar functions: Enhance stability and fluidity of membrane, also make it a better hydrophobic barrier Differing Membrane Compositions o Different cellular membranes have different lipid and protein compositions o Different faces of the same membrane can have different lipid compositions, different proteins/protein orientations o These differences relate to specific functions of each membrane Phospholipids o Amphipathic: Polar heads, hydrophobic tails o Fatty acid tails 12-20 carbons long: Usually 16 or 18 Chains less than 12 C result in unstable bilayer. Tail length typically an even number of C’s, some exceptions o Chain length determines membrane stability and thickness Shorter chains, thinner, more fluid, less stable membranes o Length and ° of saturation varies, often between chains in the same phospholipids molecule o All double bonds in cis conformation. Living organisms do not make trans fatty acids. Cis conformation puts kink in chain at double bond, loosening packing which makes the membrane more fluid but less stable. Cholesterol o Up to ½ of animal membrane lipid cholesterol o Cholesterol has 1 hydrophilic –OH group, 4 hydrophobic rings and a hydrophobic side chain: Extremely hydrophobic o Effect of these characteristics: At high temperatures cholesterol reduces membrane fluidity, stabilizing membrane. Low temperatures: Reduces fatty acid chain packing, increasing membrane fluidity o Cholesterol also blocks ions and small molecules from crossing membrane by plugging gaps and improving membrane stability and flexibility: Better insulation o Phytosterols in plants, hopaoids in bacteria have structures similar to cholesterol and serve similar functions, though synthesis pathways are quite different Importance of Membrane Fluidity o Fluidity measure of how readily phospholipids and mobile proteins move laterally. Transverse motion rare o ΔT means potential change in fluidity: Membranes can “freeze”, becoming nonfluid. Phase transition in membranes is a change in phase from liquid to solid or vice versa. Fluid state is functional state o Lipid composition regulated by cell, influencing phase transition temperature. Kinds/percent of lipid composition, chain length, ° of saturation, percent cholesterol all regulate membrane fluidity Factors Influence Fluidity o Longer chains more rigid, freeze at higher temperatures than shorter chains but more stable at higher temperatures o Shorter chains less rigid, fluid at lower temperatures than longer chains but less stable at higher temperatures o More double bonds, less saturated, less rigid, more fluid at lower temperatures, less stable at higher temperatures o Sterols stabilize membranes at high temperatures, keep membranes fluid as temperatures drop o Cell trying to maintain constant fluidity under changing temperature conditions o Cell may do this by altering lipid composition of membrane, percentages of different lipids, or ° of saturation of lipid chains Homeoviscous Adaptation o Alteration of lipid content of membranes in response to temperature fluctuations is homeoviscous adaptation o Enzyme activation/deactivation as temperature changes involved in homeoviscous adaptation o Acetyl CoA tranferase +/- 2 C’s at a time to fatty acyl chains o Desaturase removes H’s in fatty chains, replaces them with double bonds between C atoms of backbone. Loosens packing, increasing membrane fluidity o Critically important for hibernating/estivating animals, which experience drop in body temperature, as well as poikilotherms which do not regulate temperature Motion of Lipids in Membranes o Transverse Diffusion not thermodynamically favored; rare but does happen when facilitated by specific proteins called flippases. Neither phospholipids nor proteins generally flip orientation without aid. Enables membrane to maintain asymmetric distributions of different membrane components on the 2 faces o Rotation and lateral diffusion common for lipids and proteins o Model of membrane structure is the Fluid Mosaic Model Membrane Proteins o Some proteins have sugars covalently bound: Glycoproteins o Found on outer face of plasma membrane o Glycosylation occurs in lumen of RER, further processing in the Golgi o 2 types of covalent linkage: N-linked sugars: NH2 on the side group of asparagines is used for the sugar linkage O-linked sugars: OH groups of serine, threonine, hydroxylysine, or hydroxyproline used for sugar linkage o 2-60 sugars, found on proteins on plasma membrane surface facing out from cell o Glycoproteins involved in cellular adhesion, signal recognition, immune response, tissue lining/protection, and membrane stability