Advances in Anatomy Embryology and Cell Biology Vol.106 Editors F. Beck, Lt!icester W Rild, Galveston W Kriz, Reidelberg R. Ortmann, Köln lE. Pauly, Little Rock T.R. Schiebler, Würzburg Margit Pavelka Functional Morphology of the Golgi Apparatus With 25 Figures Springer-Verlag Berlin Heidelberg NewYork London Paris Tokyo Dr. med. univ. Margit Pavelka Institut für Mikromorphologie und Elektronenmikroskopie Universität Wien Schwarzspanierstraße 17, 1090 Wien, Austria ISBN-13: 978-3-540-18062-3 e-I S BN -13: 978-3-642-72826-6 DO I: 10.1 007/978-3-642-72826-6 Library of Congress Cataloging-in-Publication Data PaveJka, Margit, 1945-. Functional morphology of the Golgi apparatus. (Advances in anatomy, embryology, and cell biology; v. 106) Bibliography: p. IncJudes index. 1. Golgi apparatus. 1. Title. H. Series. QL801.E67 vol. 106 574.4 s 87-16504 [QH603.G6] [574.8'734] This work is subject to copyright. 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Product Liability: The publisher can give no guarantee for information about drug dosage and application thereof contained in this book. In every individual case the respective user must check its accuracy by consulting other pharmaceuticalliterature. Typesetting, printing and binding: Universitätsdruckerei H. Stürtz AG, Würzburg 2121/3140-543210 Ta Ernst and Michaela Acknowledgments The author would like to express her deep gratitude to all members of the Institute of Micromorphology and Electron Microscopy of the University of Vienna for their thoughtfulness and generous help du ring the time this paper was prepared. The excellent technical assistance of Mrs. Jutta Selb mann, Mrs. Elfriede Scherzer, Mrs. Gerlinde Hartl, Mr. Helmut Oslansky, and Mr. Richard Reichhart is acknowledged with thanks. The author is also most grateful to Professor Dr. Leopold Stockinger for his generous help, critical reading of the manuscript, and continuous encouragement, and also to Professor Dr. Peter Böck for fruitful discussions and continuous stimula tion. Particular thanks are due to the author's co-worker Dr. Adolf Ellinger for many hours of stimulating discussions, for the numerous suggestions he made for improvement of the manuscript, and for his generous help with its final preparation. This work was supported by the Hochschuljubiläumsstiftung der Stadt Wien and by the Vermächtnis Josefine Hirt!. Contents 1 Introduction 1 2 Material and Methods 5 3 Architecture of the Golgi Apparatus 8 3.1 Golgi Stacks . . . . . . . . . . 8 3.1.1 Transitional Elements - Transport Vesicles 16 3.1.2 Transmost Golgi Section . . . . . . . . 17 3.1.3 Golgi Elements - "Common Secretory-Endocytic Compartments"? . . . . . . . . . . . . . . . 22 3.1.4 Endoplasmic Reticulum . . . . . . . . . 22 3.1.5 Golgi-Associated Endoplasmic Reticulum Lysosomes 23 3.1.6 Cytoplasmically Coated Membranes . 24 3.2 Golgi Architecture During Mitosis 25 3.3 Microtubules and Golgi Organization 27 3.4 Variability of Golgi Architecture 30 3.4.1 Golgi Architecture in the Course of Cell Differentiation 31 4 Cytochemistry 33 4.1 Classic Cytochemical Reactions - Enzyme Cytochemistry 33 4.1.1 Enzyme Modulations . . . . . . . . . 37 4.2 Immunocytochemistry......... 38 4.2.1 Newly Synthesized Molecules Traversing the Golgi Apparatus .... 38 4.2.1.1 Secretory Molecules 38 4.2.1.2 Lysosomal Enzymes 39 4.2.1.3 Membrane Pro teins 39 4.2.2 Enzymes Involved in Glycan Biosynthesis 40 4.2.3 Antibodies Directed Against Golgi-Associated Molecules ............. . 41 4.2.4 Lysosomal Membrane Antigens . . . . . 41 4.2.5 c-AMP-Dependent Pro tein Kinase Type 11 42 4.3 Lectin Cytochemistry 42 4.3.1 Concanavalin A 45 VII 4.3.2 Ricinus communis I Agglutinin 47 4.3.3 Helix pomatia Lectin 47 4.3.4 Pisum sativum Lectin 51 4.3.5 Lens culinaris Lectin 51 5 Uptake in the Golgi Apparatus of Internalized Moleeules .............. . 56 6 Discussion 58 6.1 Morphologie Subseetions - Functional Subcompartments ....... . 58 6.2 Morphology and Classic Cytochemistry 58 6.3 Immunocytochemistry and Lectin Cytochemistry 59 6.3.1 Exit Sites .......... . 63 6.3.2 Variability of the Golgi Architecture 64 6.3.3 Mosaiclike Patterns . 65 6.3.4 "Backbone Cisternae" 68 7 Summary 69 References 70 Addendum 90 Subject Index 92 VIII 1 Introduction The complex apparatus of stacked cisternae, tubules, and vesicles, "il apparato reticulare interno " (Fig. 1), first described by Camillo Golgi in 1898 (Golgi 1898), then neglected during many years, has been rediscovered and in the last decades been proved a central crossroad in intracellular traffic. Elements of the Golgi apparatus are important stations in the routes of newly synthesized molecules, including secretory molecules, membrane constituents, and lysosomal enzymes, as weIl as internalized moleeules (Fig. 2; for recent reviews, see Bennett 1984; Dunphy and Rothman 1985; Farquhar 1985; Farquhar and Palade 1981; Goldfischer 1982; Hand and Oliver 1981; Morri~ and Ovtracht 1977; Palade 1983; Rothman 1985; Slot and Geuze 1983; Tartakoff 1980, 1983 a; V ölkl 1980; Whaley and Dauwalder 1979). Flat cisternae (saccules), interconnected by tubular-reticular elements, are arranged in parallel to form a system of stacks, the characteristic "Golgi stacks" (dictyosomes; Fig. 3), which represent morphologie subunits of the complex organelle. The stacked cisternae are the sites where multiple posttranslational modifications occur at newly synthesized molecules, such as insertion ofterminal Fig. 1 a, b. Epithelium of the rat small intestine. Affinity cytochemical labeling of the Golgi apparatus. HPA-HRP conjugates. In a, the epithelium is sectioned perpendicularly to the surface showing the characteristic supranuclear position of the Golgi apparatus in the absorp tive cells and in a go biet cell (:-); in the oblique seetion in b, the reticular, ring- or gobletlike arrangement of the Golgi elements is apparent. a x 1600; b x 1800 1 ER PM Fig. 2. Cellular transport routes involving elements of the Golgi complex. A 1, Nuclear envelope-to-Golgi route ofnewly synthesized molecules. 2, Endoplasmic reticulum-to-Golgi route of newly synthesized molecules. B, 1, Golgi intercompartmental route; traffk of newly synthesized molecules from one Golgi subcompartment to others. C, Golgi-to-final destination route of newly synthesized molecules. 1, Golgi-to-cell surface route of secretory molecules. 2, Golgi-to-plasma membrane route of plasma membrane molecules. 3, Golgi-to-lysosome route of lysosom al enzymes. D, 1, Cell surface-to-Golgi route of internalized molecules. Recycling routes. D, 1', Golgi-to-plasma membrane recycling route of plasma membrane receptors. C, 1', Plasma membrane-to-Golgi recycling route ofmolecules involved in secretory pathways. 2', Plasma membrane-to-Golgi recycling routes of molecules involved in transport of plasma membrane constituents. 3', Lysosome (or prelysosomal compartment)-to-Golgi route of receptors for lysosom al enzymes. B, 1', Golgi intercompartmental recycling route. A, 1', Golgi-to-nuclear envelope recycling route. 2', Golgi-to-endoplasmic reticulum recycling route. NE, nuclear envelope; ER, endoplasmic reticulum; PM, plasma membrane; Ly, lysosome sugars into N-glycosidically linked oligosaccharides of glycoproteins and synthe sis of O-glycosidically linked glycans (reviewed in e.g., Fleischer 1983; Hubbard and Ivatt 1981; Kornfeld and Kornfeld 1985; Northcote 1979; Schachter and Roseman 1980; Sturgess et al. 1978), formation of the lysosomal recognition marker (e.g., Pohlmann et al. 1982; Sly and Fischer 1982), sulfation (e.g., Fatem and Leblond 1985; Herzog 1985; Lee and Huttner 1985; Young 1973), and proteolytic cleavage (e.g., Orci et al. 1985b). Golgi-associated processes include steps in the synthesis and transport of proteoglycans (e.g., Kimura et al. 1984; Ratcliffe etal. 1985; Stowetal. 1985) and lipids (Keenan etal. 1974; Lipsky and Pagano 1985 a, b; Yusuf et al. 1983, 1984). Final formation and packaging of lipoprotein particles (e.g., Banerjee and Redman 1984; Christensen et al. 2
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