Cell Biology Monographs

Transcription

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2 Cell Biology Monographs Edited by M. Alfert, Berkeley, Calif. w. Beermann, Tilbingen L. Goldstein, Lexington, Ky. K. R. Porter, Boulder, Colo. Advisory Board J. Brachet, Bruxelles D. Branton, Cambridge, Mass. H. G. Callan, St. Andrews E. C. Cocking, Nottingham W. W. Franke, Heidelberg N. Kamiya, Okazaki G. Rudkin, Philadelphia, Pa. G. F. Springer, Evanston, Ill. L. Stockinger, Wien B. F. Trump, Baltimore, Md. Volume 13 Springer-Verlag Wien New York

3 The Cytoskeleton An Introductory Survey M. Schliwa Springer-Verlag Wien New York

4 Prof. Dr. MANFRED SCHLIWA Department of Zoology University of California Berkeley, California, U.S.A. This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically those of translation, reprinting, re-use of illustrations, broadcasting, reproduction by photocopying machine or similar means, and storage in data banks by Springer-VerlaglWien Softcover reprint oflhe hardcover 1st edition 1986 With 88 Figures ISSN ISBN-13: e-isbn-13: DOl: /

5 Fur Usch

6 Preface The term cytoskeleton has become firmly established in today's scientific vocabulary. Indeed, it is difficult to believe that only ten years ago, it was virtually non-existent. Since then, the modern field of research on the structural organization of the cytoplasm has turned into one of the most productive and rapidly expanding research areas in Cell Biology today. Considerable progress has been made towards the identification of the various structural components of the cytoskeleton and their interactions with one another and with membranes. The first attempts to understand, in molecular terms, complex cellular processes such as shape changes, locomotion, division, and organelle movements have been made. And it is now apparent that the cytoskeleton has impact on other biological processes such as the control of gene expression, protein synthesis, cell cycle regulation, and development. This monograph outlines the basic properties of the major components of the polymeric filament networks and their interactions and associations. Wherever possible, emphasis is placed on more recent references. Any attempt to cover a research field this complex in an introductory monograph is, by necessity, fragmentary, and oversights or omissions are inevitable. I wish to apologize in advance to all those colleagues who feel that their work is not adequately represented. This book would not have possible without the help of Ursula Euteneuer and Susan Spath, whose numerous thoughtful suggestions and comments on the final draft essentially made the text readable. To them I am deeply indebted. I also thank Paul Licht, Mona Radice, and Paul Tibbetts for their help in the struggle with the idiosyncrasies ofthe bibliography program, and the Department of Zoology at Berkeley for providing access to the departmental computer to prepare the list of references. I am grateful to the members of my laboratory for their moral support and indulgence during the months of my preoccupation with this undertaking, and hope that their patience is partly rewarded by the final result. Finally, I wish to thank the many colleagues who kindly supplied illustrative material. They are individually acknowledged in the figure legends. Berkeley, October 1985 MANFRED SCHLIWA

7 Prologue.... Contents Part A. Components I. Cytoplasmic Contractile Proteins II Historical Aspects Actin Structure Actin Assembly in vitro Inhibitors Cytochalasins Phalloidin Proteins Associated with Cytoplasmic Actin Depolymerizing and Polymerization-Inhibiting Proteins Cross-Linking Proteins End-Binding Proteins Some General Considerations on Actin Networks 104. Myosin Structure Myosin Filament Formation 1.5. Actin-Myosin Interaction Regulation of Actin-Myosin Interaction Tropomyosin, Troponin, and Alpha-Actinin in Smooth and Nonmuscle Cells Myosin and Its Interaction with Actin: Outlook Aspects of Contractile Protein Distribution and Function Microtubules II.3. II.4. Historical Aspects.. Microtubule Structure Tubulin Interactions with Other Molecules IIA.1. Ions IIA.2. Nucleotides.... IIA.3. Microtubule-Associated Proteins IIAA. Drugs I Colchicine and Its Analogues 1I Vinblastine and Vincristine II Nocodazole Taxo!... II Other Microtubule Poisons

8 x Contents Mechanism of Microtubule Polymerization Microtubule Polarity Nucleation Sites and Organizing Centers Structure Demonstration of Nucleating Activity Biochemistry of MTOCs Microtubule Machineries: Cilia and Flagella Structure Function Ciliary and Cytoplasmic Dyneins Distribution and Function of Microtubules in Cells: A Summary. 81 III. Intermediate Filaments Historical Aspects Classification of Intermediate Filaments: Molecular Composition and Tissue Distribution Vimentin Desmin 87 III.2.3. Cytokeratins Neurofilaments Glial Filaments Cells Without Intermediate Filaments? Intermediate Filaments as Markers For Tumor Characterization Structural, Physical, and Chemical Similarities Between Intermediate Filament Proteins Co-Existence and Co-Assembly of Different Intermediate Filament Subunits III.5. Intermediate Filament-Associated Proteins Phosphorylation of Intermediate Filament Proteins Regulation of Intermediate Filament Assembly in the Cell Aspects of the Cellular Distribution and Function of Intermediate Filaments IV. Dynamic Aspects of Filament Assembly 115 V. Other Filament Types V.1. Spasmin-Like Filaments... V.2. Connectin/Titin-Like Filaments V.3. 3 nm Linkers Part B. Interactions VI. Some General Aspects of the Three-Dimensional Organization of Cytoskeletal Components VII. Interactions Between the Major Filamentous Components of the Cytoskeleton. 145 VII.1. Some Methods for the Detection of Filament Interactions. 146 VII.2. Self-Associations of the Major Cytoskeletal Filament Types. 148 VII.2.1. Actin Filaments

9 Contents VII.2.2. Microtubules.. VII Intermediate Filaments.. VII.3. Interactions Among Different Filament Components VII.3.I. Actin Filaments and Microtubules... VII.3.2. Actin Filaments and Intermediate Filaments VII.3.3. Microtubules and Intermediate Filaments VII.4. Filament Interactions: A Perspective. VIII. The Cytoskeleton and the Cytoplasmic Matrix XI IX. Membrane-Cytoskeleton Interactions IX.I. Introduction IX.2. Paradigm Erythrocyte. 180 IX.2.I. Spectrin 180 IX.2.2. Ankyrin 182 IX.2.3. Band IX.2.4. Actin. 184 IX.2.5. Band IX.2.6. Construction of the Subplasmalemmal Network. 185 IX.3. Proteins Related to the Erythrocyte Skeleton in Other Cell Types. 188 IX.4. Association of Membrane Proteins with the Cytoskeleton 196 IX.5. Association of Cytoskeletal Fibers with the Membranes 203 IX.6. Some Aspects of Organelle-Cytoskeleton Relationships 208 IX.7. Specialized Filament-Membrane Assemblies IX.7.I. Adherens Junctions and Adhesion Plaques IX.7.2. Desmosomes IX.7.3. The Brush Border of Intestinal Epithelial Cells 227 IX.7.3.I. Morphology and Ultrastructure IX Biochemical Anatomy IX A Note on Brush Border "Motility" 234 IX.8. Transmembrane Interactions of the Cytoskeleton with the Extracellular Matrix. 236 References.. Subject Index