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Yeasts in Natural and Artificial Habitats Springer-Verlag Berlin Heidelberg GmbH J.F.T. Spencer D.M. Spencert (Eds.) Yeasts in Natural and Artificial Habitats With 59 Figures and 28 Tables Springer John F.T. Spencer Dorothy M. Spencert PROIMI Avenida Belgrano y Pasaje Caseros 4000 S.M. de Tucuman Argentina ISBN 978-3-642-08160-6 Library of Congress Cataloging-in-Publication Data. Spencer, John F.T. Yeasts in natural and artificial habitats/).F.T. Spencer, D.M. Spencer. p. em. Includes bibliographical references and index. ISBN 978-3-642-08160-6 ISBN 978-3-662-03370-8 (eBook) DOI 10.1007/978-3-662-03370-8 I. Yeast fungi. I. Spencer, Dorothy M. II. Title. QK617.S.S65 1997 579.5'63-dc21 96-50359 This work is subject to copyright. All rights are reserved, whether the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations, recitation, broad casting, reproduction on microfilm or in any other way, and storage in data banks. Duplication of this publication or parts thereof is permitted only under the provisions of the German Copyright Law of September 9, 1965, in its current version, and permission for use must always be obtained from Springer-Verlag Berlin Heidelberg GmbH. Violations are liable for prosecution under the German Copyright Law. ©Springer-Verlag Berlin Heidelberg 1997 Originally published by Springer-Verlag Berlin Heidelberg New York in 1997 The use of general descriptive names, registered names, trademarks, etc. in this publication does not imply, even in the absence of a specific statement, that such names are exempt from the relevant protective laws and regulations and therefore free for general use. Cover design: Design & Production, Heidelberg Typesetting: Best-set Typesetter Ltd., Hong Kong SPIN: 10047436 31/3137/SPS-5 4 3 2 I 0-Printed on acid-free paper Dedication This book is dedicated to our many friends throughout the world who share our enthusiasm for the yeasts. It is dedicated in particular to my coauthor and devoted wife, Dorothy Spencer, who, besides her part in the writing of the book, did a great deal of the work of correcting my numerous typographical and similar errors, and who died of cancer before she could have the satisfaction of seeing that her labors were not in vain. J.F.T. Spencer Contents A Guide to the World of the Yeasts J.F.T. Spencer and D.M. Spencer ........................ . 1 Historical Introduction: Yeasts and Man in the Past J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 2 2 Taxonomy: The Names of the Yeasts J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 11 3 Ecology: Where Yeasts Live J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 33 4 Ecology: The Bad Guys: Pathogens of Humans and Other Animals J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 59 5 Yeasts as Living Objects: Yeast Nutrition J.F.T. Spencer, D.M. Spencer, and L.I.C. de Figueroa 68 6 Outside and Inside: The Morphology and Cytology of the Yeast Cell J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 80 7 Membranes M. Hofer.......................................... 95 8 The Yeasts: Sex and Nonsex. Life Cycles, Sporulation and Genetics J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 133 9 Chemical Warfare Among the Yeasts: the "Killer" Phenomenon, Genetics and Applications V. Vondrejs and Z. Palkova . . . . . . . . . . . . . . . . . . . . . . . . . . 153 VIII Contents 10 Inside the Inside: Part I: Yeasts and Molecular Biology, A Recipe for Alphabet Soup. Chromosome Structure, Replication, Transcription, and Translation J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 171 11 Inside the Inside: Part II: The Regulators. Cell Specialization: a Regulatory Hierarchy ].F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 204 12 Yeasts and the Life of Man: Part I: Helpers and Hinderers. "Traditional" Yeast-Based Industries; Spoilage Yeasts J.F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 226 13 Yeasts and the Life of Man: Part II: Genetics and Molecular Biology of Industrial Yeasts and Processes ].F.T. Spencer and D.M. Spencer . . . . . . . . . . . . . . . . . . . . . . 243 14 Genetics and Molecular Biology of Methylotrophic Yeasts E. Berardi . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 264 15 Yeasts in Food Fermentations and Therapeutics G. Vijayalakshmi, B.K. Lonsane, and M.M. Krishnaiah 295 16 Yeasts and Energy. The Production of Fuel-Grade Ethanol B.K. Lonsane, G. Vijayalakshmi, and M.M. Krishnaiah . . . 315 Notes Added in Proof.................................. 353 Subject Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 357 Species Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37 3 List of Contributors Berardi, Enrico, Dipartimento di Biologie Agrarie ed Ambientale, Universita degli Studi di Ancona, 60131 Ancona, Italia de Figueroa, L.I.C., PROIMI, Avenida Belgrano y Pasaje Caseros, 4000 S.M. de Tucuman, Argentina HOfer, Milan, Botanisches Institut, Rheinische Friedrich-Wilhelms Universitlit, Kirschallee 1, 53115 Bonn, Germany Krishnaiah, M.M., Central Food Technological Research Institute, Microbiology & Fermentations, Mysore 570013, India Lonsane, B.K., Central Food Technological Research Institute, Microbiology & Fermentations, Mysore 570013, India Palkova, Z., Dept. of Genetics and Microbiology, Faculty of Science, Charles University, Vinicna 5, 128 44 Prague, Czech Republic Spencer\ Dorothy M., PROIMI, Avenida Belgrano y Pasaje Caseros, 4000 S.M. de Tucuman, Argentina Spencer, John F.T., PROIMI, Avenida Belgrano y Pasaje Caseros, 4000 S.M. de Tucuman, Argentina Vijayalakshmi, G., Central Food Technological Research Institute, Microbiology & Fermentations, Mysore 570013, India Vondrejs, V., Dept. of Genetics and Microbiology, Faculty of Science, Charles University, Vinicna 5, 128 44 Prague, Czech Republic INTRODUCTION A Guide to the World of the Yeasts J.F.T. Spencer and D.M. Spencert As the well-known authority on yeasts, the late Professor Rose, frequently pointed out, it is impossible for one person to present, in a single volume, the details of the life, composiotion, habitats, relationships, and actual and potential uses to man kind of the 500 (at last count) known species of yeasts. This book confirms the truth of this statement. However, our aim is actually more modest than that, and this book is an attempt to introduce the general reader, and possibly some inter ested specialists, to the lives of the yeasts in their natural and more artificial habitats, their use by human beings, and to give some idea of the wonderfully complex activities within the yeast cell, the characteristics of the metabolism and molecular biology of yeasts, and the applications of these characteristics to life in the present-dayworld ofhuman existence. The book proceeds from a brief chapter on what is and is not known of the origins and early history of the yeasts, through a description of their classification, relationships, habitats and general life style, their external morphology and internal structures and mechanisms within their cells, the regulatory mechanisms controlling processes such as signal transmis sion, mating, cell fusion, and many others. To prove the earlier contention that it is impossible for one person to describe adequately everything about every yeasts, there are special sections dealing with yeast membranes, killer factor, and in the part of the book dealing with the impact of yeasts on the human condition, we have included sections on fundamental and applied aspects of the methylotrophic yeasts, yeasts in fermented foods, and use of yeasts in production of fuel alcohol. As the originators and authors of the book, we are extremely grateful to these specialists for their valuable contributions to the volume. In conclusion, we would like to thank our friends, once again, who have made substantial contributions to this task, in the hope that they will enjoy seeing the book in print and reading it. We thank them for their patience and understanding of the task we have had as authors ourselves, and as editors of their work. We would like also to express our thanks to Dr. Faustino Sifieriz and all our friends at PROIMI for their advice and encouragement during our work on the book. ).F.T. Spencer/D.M. Spencer (eds) Yeasts in Natural and Artificial Habitats ©Springer-Verlag Berlin Heidelberg 1997 CHAPTER 1 Historical Introduction: Yeasts and Man in the Past J.F.T. Spencer and D.M. Spencer 1 From Earliest Times to the Beginning of the Modern Era Fossil yeasts are unknown. Fungal spores have been found in fossiliferous materi als, and yeasts may have been associated with them, but they have not been recognized. The sight of a yeast ascus containing ascospores, or of the zigzag configuration of the arthrospores of a fossilized Trichosporon species, would en able a paleobotanist to place the yeasts more accurately in time. There may be a long wait for such a discovery, though ancient diatoms, algae, and other micro organisms have been seen, and eventually a yeast ascus may be found, resting peacefully in its amber tomb (Poinar et al. 1993). Yeasts have been known, indirectly, "only" for a few thousand years. The first peoples to eat leavened bread and drink beverages such as wines were either the Egyptians or one of the other civiizations of the Fertile Crescent. They knew the uses of yeasts but not their identity. Beer may have been discovered at about the same time, possibly by some farmer who was soaking a pot of grain in water, to make it more palatable to his domestic animals. It is known that the Egyptians knew bread, beer, and wine. The knowledge was probably shared by the other peoples of the region, and by those of the land with whose inhabitants they traded -India. The use of yeasts in breadmaking may have arisen through some of the early methods of cooking raw flour. For instance, in making salt-rising bread, the flour and a weak brine are mixed and kept warm, probably at about 30 °C, until the volume of the mixture approximately doubles, and this is then mixed with more flour and warm water and allowed to rise again. The normal practice in all breadmaking, when commercial yeast or spent yeast from breweries is not avail able, is to save a portion of the uncooked dough to start the next batch. The better batches of starter would probably be propagated as long as possible, and would have consisted predominantly of yeasts. In this way, the characteristic odor of yeast starters would have been recognized, very early, and would have been fa vored by the women who did the real work of baking; they would in all probability have traded the "good" starters, which would have been largely yeasts. Unleavened bread is known from flat loaves dating from at least 6000 B.C. in Egypt. If a dough mix of this sort were left to stand for any reason, fermented spontaneously, and was then baked, the result would be naturally leavened bread. From there to deliberately leavened loaves is only a short step. The odor of rising J.F.T. Spencer/D.M. Spencer (eds) Yeasts in Natural and Artificial Habitats ©Springer-Verlag Berlin Heidelberg 1997
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