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Preface There is now general acceptance of the view that the plasma membranes of most, if not all, cells are dynamic assemblies of molecules that are able to undergo rapid and reversible structural rearrangements in response to both intra- and extracellular stimuli. This concept is, however, of relatively recent origin. Until the mid-nineteen sixties, experimental observations on the mor phology, chemical composition and function of biological membranes were in terpreted against a conceptual background in which membrane components were thought to exist largely in rather rigid, ordered and static structural arrangements. In the late nineteen sixties and early seventies information began to accumulate from studies in a variety of disciplines which indicated that certain membrane components were free to move within the membrane and thus undergo topographic rearrangement. These findings were difficult to reconcile with the existing static models for membrane structure and prompted the formulation of new generalized concepts for membrane organization such as the "liquid crystalline,, and "fluid mosaic" models which, for the first time, appeared to fit the growing number of observations on the dynamic properties of biological membranes. From these initial experimental observations less than a decade ago, information on membrane dynamics has grown at a remarkable pace so that today the literature on the subject is of voluminous proportion. Although many fundamental questions remain to be answered, and speculation currently surrounds interpretation of several aspects of the subject, sufficient information is already available to support a functional relationship between the topography and dynamics of plasma membrane macromolecules and the control of cell surface properties. In its most simple form, the plasma membrane can be thought of as a two-dimensional solution of a mosaic of lipids and proteins. The lipids, ar ranged predominantly as a bilayer, exist in a "fluid'' state while the proteins (and glycoproteins) are either inserted to varying depths into the lipid bilayer (integral proteins) or bound loosely to the surfaces of the bilayer (peripheral proteins). Both proteins and lipids can be organized asymmetrically within this xii type of structure, thus allowing specific classes of membrane components to be localized exclusively or predominantly at the inner or outer membrane surface. Another important feature is that the fluid state of the membrane lipids permits lateral diffusion of macromolecules within the plane of the membrane, thereby creating opportunities for rapid and reversible changes in membrane topography. The distinctive lateral mobilities of different membrane com ponents, ranging from extremely rapid to relatively immobile, permits cells to maintain certain surface molecules in relatively ordered topographic arrays or "patterns", while allowing others to diffuse randomly and avoid ordered topographic restraint. The fluid nature of cell membranes also dictates that environmental changes in temperature, ionic strength, pressure, binding of material to membrane "receptors" and many other stimuli can induce drastic changes in the physical state of the membrane, leading to phase changes and separations whereby molecules are excluded away from or sequestered into specific membrane regions or domains. Finally, trans-membrane interaction of components within the plasma membrane with membrane-associated structures in the cytoplasm offers a potential mechanism for the transmission of "in formation" across the membrane in either direction via various combinations of cooperative, allosteric and transductive coupling mechanisms. These and other possible regulatory devices may well allow different cell types to react individually to the same stimuli, thereby creating opportunities for a large repertoire of cell-specific responses and membrane specialization based on a relatively restricted and conservative framework of membrane structure. Membrane dynamics embraces a broad range of physical, chemical and biological processes. Comprehensive discussion of this subject demands con sideration of the basic physico-chemical properties of the different molecules and macromolecules found in membranes, analysis of their functional in terrelationships within membranes, characterization of the diverse factors that influence membrane organization in both physiological and pathological states and, finally, identification of relationships between the control of plasma membrane organization and changes in cell surface properties and cell be havior. Insight into these various aspects of membrane dynamics has come from work in many disciplines, from detailed observations at the molecular level as well as phenomenological descriptions of the behavior of intact cells, and from the application of numerous techniques, some old but improved, and some entirely new. This volume, the third in the Cell Surface Reviews series, contains fourteen chapters which reflect the dramatic growth of information of the dynamic nature of membrane organization. No attempt has been made to cover all aspects of this vast subject. However, the reviews in this volume offer a broad perspective of current concepts of plasma membrane organization and the range of experimental strategies and techniques used to investigate this im portant structure. Discussion of the importance of the biosynthesis, assembly and turnover of plasma membrane components in cell surface dynamics has been largely excluded from this volume since the next volume in this series will xiii be devoted entirely to this subject. Together, volumes 3 and 4 of Cell Surface Reviews provide an extensive and up-to-date discussion of what we feel to be the more important aspects of cell surface dynamics. We thank the contributors for their authoritative and comprehensive chap ters. We are also grateful to Judy Kaiser, Adele Brodginski, Shirley Guagliardi and Molly Terhaar for their assistance in preparing the edited manuscripts. George Poste Buffalo, New York August, 1976 Garth L. Nicolson Irvine, California August, 1976
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