Yeast Cell Envelopes: Biochemistry, Biophysics, and Ultrastructure Volun1e I Editor Wilfred Niels Arnold, Ph.D. Professor Department of Biochemistry University of Kansas Medical Center Kansas City, Kansas Boca Raton London New York CRC Press is an imprint of the Taylor & Francis Group, an informa business First published 1981 by CRC Press Taylor & Francis Group 6000 Broken Sound Parkway NW, Suite 300 Boca Raton, FL 33487-2742 Reissued 2018 by CRC Press © 1981 by CRC Press, Inc. CRC Press is an imprint of Taylor & Francis Group, an Informa business No claim to original U.S. Government works This book contains information obtained from authentic and highly regarded sources. Reasonable efforts have been made to publish reliable data and information, but the author and publisher cannot assume responsibility for the validity of all materials or the consequences of their use. 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CCC is a not-for-profit organization that provides licenses and registration for a variety of users. For organizations that have been granted a photocopy license by the CCC, a separate system of payment has been arranged. Trademark Notice: Product or corporate names may be trademarks or registered trademarks, and are used only for identification and explanation without intent to infringe. Library of Congress Cataloging in Publication Data Main entry under title: Yeast cell envelopes: biochemistry, biophysics, and ultrastructure. (Uniscience series) Includes bibliographies and indexes. 1. Yeast fungi. 2. Fungi—Cytology. I. Arnold, Wilfred Niels, 1936-. II. Title: Cell en- velopes. [DNLM: 1. Yeasts—Cytology. 2. Cell membrane. QK 617.5 Y39] QK617.5.Y4 589.2’33 81-415 ISBN 0-8493-5965-1 (v. 1) AACRl ISBN 0-8493-5966-X (v. 2) A Library of Congress record exists under LC control number: 81000415 Publisher’s Note The publisher has gone to great lengths to ensure the quality of this reprint but points out that some imperfections in the original copies may be apparent. Disclaimer The publisher has made every effort to trace copyright holders and welcomes correspondence from those they have been unable to contact. ISBN 13: 978-1-315-89869-8 (hbk) ISBN 13: 978-1-351-07779-8 (ebk) Visit the Taylor & Francis Web site at http://www.taylorandfrancis.com and the CRC Press Web site at http://www.crcpress.com PREFACE Yeasts hold important associations with man in at least three arenas. As industrial agents their positions in the fermentative and baking industries are household knowledge. A few yeast or yeast-like species are pathogenic. And, as experimental eukaryotic cells, yeasts continue their long history as useful subjects for studies on metabolism, genetics, and molecular biology. A comprehensive review of the yeast cell envelope has not appeared previously and we trust that this attempt will be timely. The title of this volume was chosen to reflect the three major areas of contribution to our current understanding of the cell envelope, but we have not attempted to group chapters into subdivisions. That would be somewhat arbitrary at best. In fact, the contributing authors were recruited for their interdisciplinary work as well as their special expertise. The approach is to describe phenomena, to review the literature, and to illuminate outstanding problems. We have also attempted to generate working hypotheses which may stimulate further studies. That some of these ideas be of germinal value is of more concern to us than that all of the hypotheses should stand the test of further experimentation. Brenda Johnson has given special assistance in the assemblage of this volume. I also wish to acknowledge my former teachers and mentors Drs. G. Langdon, J. Middleton, J. Bald, and J. Thompson, as well as two colleagues, E. Juni and I. Goldstein, for encouragement during my formative years in biological chemistry. W. N. Arnold Westwood Hills, Kansas July 1980 THE EDITOR Wilfred N. Arnold, Ph.D., is Professor of Biochemistry at the University of Kansas Medical Center, Kansas City, Kansas. Dr. Arnold received a B.S. from the University of Queensland, Australia in 1956, an M.A. from the University of California, Los Angeles in 1958, and a Ph.D. from Cornell University, Ithaca, N.Y. in 1962. He has held postdoctoral fellowships at the Waite Institute, University of Adelaide, Australia and at the University of Wisconsin Medical School, Madison. Before coming to Kansas in 1971, he held research positions at the Commonwealth Scientific and Industrial Research Organization (CSIRO) in Australia and the University of California at Riverside; and an Assistant Professorship at Wayne State University Medical School, Detroit, Mich. Dr. Arnold is a member of the American Society of Biological Chemists, American Chemical Society, American Association for the Advancement of Science, American Society for Microbiology, Society for Complex Carbohydrates, and the honorary society, Sigma Xi. Dr. Arnold has authored 40 publications in biochemistry. His current research interests are centered on the yeast cell envelope with particular emphasis on questions at the interface of biochemistry and ultrastructure of pathogenic and nonpathogenic species of yeast. This research has been sponsored by N.I.H., Research Corporation, and Sigma Xi. Dr. Arnold has taught biochemistry to undergraduate, graduate, and medical students and has authored and directed three educational films. CONTRIBUTORS JohnS. D. Bacon, Sc.D., F.R.S.E. Oldfich Necas, M.D., D.Sc. Honorary Research Associate Professor and Head University of Aberdeen and Department of Biology Head Medical Faculty Carbohydrate Biochemistry J. E. Purkyne University Department Brno, Czechoslovakia Rowett Research Institute Bucksburn, Aberdeen Armando J. Parodi, Ph.D. Scotland, United Kingdom Visiting Scientist Department of Microbiology Robert G. Garrison, Ph.D. The Wellcome Research Laboratories Research Microbiologist Research Triangle Park, North Veterans Administration Medical Carolina Center Kansas City, Missouri and Martin L. Slater, Ph.D. Associate Professor Executive Secretary Department of Microbiology Microbial Physiology Study Section University of Kansas Medical Center Division of Research Grants Kansas City, Kansas National Institutes of Health Bethesda, Maryland Jaroslav Horak, Ph.D. Senior Worker Department of Cell Physiology Eva Streiblova, Ph.D. Institute of Microbiology Senior Scientific Worker Czechoslovak Academy of Sciences Department of Cell Physiology Videnska, Prague Institute of Microbiology Czechoslovakia Czechoslovak Academy of Sciences Videnska, Prague Arnost Kotyk, D.Sc. Czechoslovakia Head Department of Cell Physiology Institute of Microbiology Shigeo Suzuki, Ph.D. Czechoslovak Academy of Sciences Professor and The Second Department of Hygienic Professor Chemistry Charles University Tohoku College of Pharmacy Videnska, Prague Komatsushima, Sendai Czechoslovakia Japan Augustin Svoboda, M.D., Ph.D. Assistant Professor Department of Biology Medical Faculty J. E. Purkyne University Brno, Czechoslovakia TABLE OF CONTENTS Volume I Chapter 1 Introduction ............................................................................................. 1 W. N. Arnold Chapter 2 Atlas of Cell Morphology ........................................................................... 5 R. G. Garrison and W. N. Arnold Chapter 3 Physical Aspects of the Yeast Cell Envelope ................................................ 25 W. N. Arnold Chapter 4 Transport Processes in the Plasma Membrane .............................................. 49 A. Kotyk and J. Horak Chapter 5 Nature and Disposition of Polysaccharides within the Cell Envelope ................ 65 J. S. D. Bacon Chapter 6 Antigenic Determinants ............................................................................ 85 S. Suzuki Chapter 7 Lipids .................................................................................................... 97 W. N. Arnold Subject Index ........................................................................................ 115 Taxonomic Index ................................................................................... 123 TABLE OF CONTENTS Volume II Chapter 1 Enzymes .................................................................................................. 1 W. N. Arnold Chapter 2 Biosynthetic Mechanisms for Cell Envelope Polysaccharides ........................... 47 A. J. Parodi Chapter 3 Molecular Biology of Budding ................................................................... 65 M. L. Slater Chapter 4 Fission ................................................................................................... 79 E. Streiblova Chapter 5 Protoplasts ............................................................................................. 93 W. N. Arnold Chapter 6 Morphogenesis in Protoplasts ................................................................... 105 0. Neeas and A. Svoboda Chapter 7 Autolysis .............................................................................................. 129 W. N. Arnold Chapter 8 Vegetative Ultrastructure ......................................................................... 139 R. G. Garrison Subject Index ........................................................................................ 161 Taxonomic Index ................................................................................... 169 1 Chapter 1 INTRODUCTION W. N. Arnold A working definition of the class of organisms collectively known as "yeasts" is not easily postulated. They are unicellular, eukaryotic microorganisms that typically reproduce by budding. So much is true for the majority of yeasts but one soon runs into exceptions; there are species with a propensity for mycelium formation, others with a fission type of cell division, and still another genus that bears conidia on sterigmata. As discussed by Ladder' (Table 1) the term "yeasts" developed historically and really includes a heterogeneous group of microorganisms. Phaff et al. ,Z have reviewed the etymology of "yeast", and equivalent terms in other languages, and point out the consistent relationship to fermentation. However, they also conclude that the name embraces a heterogeneous collection of fungi. The number of defined yeast species is now somewhat in excess of 500. In addition there are a number of fungi with "yeast-like" cells as part of their cell cycle. At the other extreme some biochemists incorrectly use the term yeast as synonymous with Saccharomyces cerevisiae. One of the themes that will emerge in the following chapters is that a rich variety of additional species awaits further biological investigation and that in some circumstances the more exotic species may offer advantages for experimental study. This book is devoted to the yeast cell envelope. The accumulated primary literature on various aspects of this subject now merits this degree of specialization. More general references on yeast include volumes edited by Cook,3 and by Rose and Harrison.4 The text by Phaff et al., 2 includes an introduction to yeast biology. The cell envelope consists of the plasma membrane, the periplasmic space, cell wall, and (for some species) a slime layer. A schematic representation is given in Figure 1, primarily to establish relative location of components rather than actual dimensions. The cell envelope is thus bounded on the inside by cytoplasm, and on the outside by the medium. The definitions that follow were developed with the assistance of contributors and other colleagues and have been applied consistently. In the majority of cases I have chosen not to belabor synonymy. Also, inappropriate terms or incorrect applications of terms in current usage have been avoided. One outstanding example of the latter from the older literature is the generic term "membrane" which was sometimes used to describe any membrane of the protoplasm (including the plasma membrane), the cell wall, or even the cell envelope. In the current setting this is, of course, intolerably confusing. The plasma membrane is the boundary of the cell protoplasm and is microscopically discernible as a bilayer. The plasma membrane is the primary site for solute regulation between the cytoplasm and the medium. Plasma membrane, plasmalemma, ectoplast, protoplasmic membrane, and cytoplasmic membrane are used synonymously in the literature. Semantic arguments against the last two terms have been registered; plasma membrane is in vogue although plasmalemma is frequently encountered. Ectoplast is valid but has not been popularly embraced. Cytoplasm plus plasma membrane is defined as protoplasm. The protoplasm of a cell gives rise to a protoplast, i.e., an unnaturally produced entity derived from a cell by denuding the cell of cell wall (and slime layer).