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WINE MICROBIOLOGY the Chapman & Hall Enology Library WINE MICROBIOLOGY The Chapman & Hall Enology Library Principles and Practices of Winemaking by Roger B. Boulton, Vernon L. Singleton, Linda F. Bisson, and Ralph E. Kunkee Wine Microbiology by Kenneth C. Fugelsang Winery Utilities Planning, Design and Operation by David R. Storm Wine Analysis and Production by Bruce W. Zoecklein, Kenneth C. Fugelsang, Barry H. Gump, and Fred S. Nury Forthcoming Titles Winemaking From Grape Growing to Marketplace by Richard P. Vine, Bruce Bordelon, Ellen M. Harkness, Theresa Browning and Cheri Wagner WINE MICROBIOLOGY Kenneth C. Fugelsang California State University at Fresno Springer-Science+Business Media, B.V Cover design: Sald Sayrafiezadeh, emDASH inc. Art Direction: Andrea Meyer Copyright © 1997 by Springer Science+Business Media Dordrecht Originally published by Chapman & HaU in 1997 Softcover reprint of the hardcover 1st edition 1997 ISBN 978-1-4757-6972-2 ISBN 978-1-4757-6970-8 (eBook) DOI 10.1007/978-1-4757-6970-8 AU rights reserved. No part of this book covered by the copyright hereon may be reproduced or used in any form or by any means-graphic, electronic, or mechanical, including photocopying, recording, taping, or information storage and retrieval systems-without the written permission of the publisher. 2 3 4 5 6 7 8 9 10 XXX 01 00 99 98 97 Library of Congress Cataloging-in-Publication Data Fugelsang, K. C. Wine microbiology / Kenneth C. Fugelsang. p. em. -- (The Chapman & Hali enology library) Jncludes bibliographical referenees and index. ISBN 978-1-4757-6972-2 1. Wine and wine making--Microbiology. L Title. II. Series. QR151.F676 1996 663' .203--de20 96-9521 CIP British Library Cataloguing in Publication Data available CONTENTS Preface Xlll Acknowledgments XVI Introduction XIX Chapter 1: The Lactic Acid Bacteria 3 1.1 INTRODUCTION 3 1.1.1 Heterofermentative and Homofermentative Metabolism 4 1.1.2 Ecology and Population Dynamics of LAB 7 Presence of LAB in the Vineyard 7 Growth of Native LAB in Must, Juice, and Wine 10 1.2 UTILIZATION OF COMMERCIAL "STARTERS" FOR MLF 11 1.2.1 Starter Culture Preparation 12 Rehydration, Growth, and Expansion Media Composition 12 Nitrogen requirements 13 Acid Adjustment 14 Sterilization 14 1.2.2 Pure and Coculture Methods 15 1.2.3 Monitoring Population Changes 16 1.2.4 Timing of LAB Addition 17 1.2.5 Use of Ongoing MLF or ML-lees as Inoculum 18 v vi Contents 1.3 GROWTH OF LAB IN WINE (THE MALOLACTIC FERMENTATION) 18 1.3.1 Biochemistry and Physiology 19 1.3.2 Changes in Wine 20 1.3.3 Environmental Conditions and Winemaking Decisions 21 Impacting MLF 21 pH 21 Alcohol Levels 22 Nutritional Status 22 Cellar Temperature 23 Sulfur Dioxide 23 Microbial Antaganism 25 Yeasts 26 Lactic Acid Bacteria 27 Bacteriophage 28 1.4 LAB SPOILAGE 29 1.4.1 Acetic Acid Formation 29 1.4.2 Refermentation 29 1.4.3 Geranium Tone 31 1.4.4 Mousiness 32 1.4.5 Diacetyl (Citric Acid Utilization) 33 1.4.6 Tartaric Acid Utilization 34 1.4.7 Mannitol Formation, Ropiness, and Polysaccaride Formation 36 1.4.8 Metabolites of Glycerol (Acrolein) 37 1.5 LABORATORY PROCEDURES (IDENTIFICATION OF BACTERIAL ISOLATES) 37 1.5.1 Gram stain 37 1.5.2 Catalase Test 38 1.5.3 General Growth Medium for LAB 39 1.5.4 Heterofermentative Versus Homofermentative Utilization of Sugar 40 1.5.5 Mannitol Salt Formation 41 1.5.6 Ammonia Production from Arginine 42 1.5.7 Monitoring MLF (Paper Chromatography) 44 Chapter 2: Acetic Acid Bacteria 48 2.1 INTRODUCTION 48 2.1.2 Growth on Carbohydrates 49 2.1.3 Growth of Ethanol 51 2.1.4 Ecology of Acetic Acid Bacteria 52 Vineyard 53 Primary Processing and Fermentation 54 Post-Fermentation 55 2.1.5 Control of Acetic Acid Bacteria 57 2.1.6 Reduction in Volatile Acidity 58 Contents vii 2.1.7 Sensory Implications 59 Acetic Acid 59 Ethyl Acetate 59 Acetaldehyde 60 Acetoin 60 Dihydroxyacetone 60 2.2 OTHER WINE-ASSOCIATED BACTERIA 60 2.2.1 Bacillus 61 2.2.2 Other Bacterial Involvement 61 2.3 LABORATORY PROCEDURES FOR IDENTIFICATION OF ACETIC ACID BACTERIA 62 2.3.1 Gram Staining 63 2.3.2 Catalase Test 63 2.3.3 Ethanol Oxidation: Separation of Acetobacter from Gluconobacter 64 Carr Medium 64 Calcium Carbonate-Ethanol Medium (Frateur's Medium) 65 Calcium Lactate Yeast Extract Agar 65 Glucose-Yeast Extract-Carbonate Medium 66 2.3.4 Glycerol Medium (Test for Ketogenesis) 66 Chapter 3: Yeasts and Molds 68 3.1 YEAST CLASSIFICATION AND ECOLOGY 68 3.1.1 Yeasts in the Vineyard 69 3.1.2 Yeasts in the Winery 70 3.2 NATIVE FLORA AND FERMENTATION 71 3.3 SELECTED NATIVE YEASTS 72 3.3.1 Brettanomyces/Dekkera 72 Taxonomy 73 Morphology 73 Distribution and Ecology 74 Monitoring 77 Sensory Properties 79 3.3.2 Zygosaccharomyces 80 Taxonomy 80 Morphological Properties 81 Habitat 82 3.3.3 Hansenula anomala 83 3.3.4 Kloeckera apiculata (Hanseniaspora uvarum) 84 3.3.5 Pichia 84 3.3.6 Candida 85 3.3.7 Other Spoilage Yeasts 85 3.4 YEAST IDENTIFICATION 86 3.4.1 Cell Morphology 86 3.4.2 Colony Appearance (Color) 87 viii Contents 3.4.3 Asexual (Vegetative) Reproduction 87 Multilateral Budding 88 Bipolar Budding 89 "Fission" 89 Pseudomycelium Formation 89 3.4.4 Sexual Reproduction (Ascospore Formation) 89 3.4.5 Oxidative Requirements for Nitrogen and Carbon 90 3.4.6 Taxonomic Key 91 3.5 LABORATORY PROCEDURES (FOR IDENTIFICATION OF YEASTS) 96 3.5.1 Isolation 96 3.5.2 Demonstration of Ascospores 98 3.5.3 Carbon and Nitrogen Assimilation Tests 101 3.5.4 Fermentation Broths 105 3.5.5 Diagnostic and "Differential" Media 106 WL-Nutritional and WL-Differential Media 107 Brettanomyces/ Dekkera-Selective Medium 107 Zygosaccharomyces-Selective Medium 108 Slide Culture 109 3.6 MOLDS OF IMPORTANCE IN WINEMAKING 110 3.6.1 Identification 111 Mold Life Cycle III 3.6.2 Botrytis cinerea (Gray Mold) 112 3.6.3 Other Molds 113 Penicillum sp. (Blue-Green Molds) 114 Aspergillus (Black Mold) 114 Mucor and Rhizopus (Pin Molds) 114 3.7 LABORATORY PROCEDURES (FOR OBSERVATION AND IDENTIFICATION OF MOLDS) 114 Chapter 4: Prefermentation Processing 117 4.1 INTRODUCTION 117 4.2 HARVEST AND TRANSPORT 118 4.3 FRUIT QUALI1Y ASSESSMENT 118 4.4 PREFERMENTATION PROCESSING 119 4.4.1 Sulfur Dioxide 120 4.4.2 Initial Fermentable Sugar Levels 121 4.4.3 Hydrogen Ion Concentration (pH) 121 4.4.4 Suspended Solids 122 4.4.5 Redox Potential (Oxygen) 123 4.4.6 Available Nitrogen 124 4.4.7 Inert Gassing 127 4.5 JUICE (MUTE) STORAGE 127 4.6 PROCESSING MODIFICATIONS FOR MICROBIALLY DETERIORATED FRUIT 129 4.6.1 Processing Botrytis-Infected Fruit 129 Contents ix Chapter 5: Fermentation and Post-Fermentation Processing 132 5.1 INTRODUCTION 132 5.2 YEAST STARTERS (PREPARATION AND PROPAGATION) 133 5.3 FERMENTATION TEMPERATURE 136 5.4 NATIVE FLORA AND FERMENTATION 136 5.5 STUCK FERMENTATION 137 5.5.1 Temperature and Alcohol-Related Problems 138 5.5.2 Nitrogen Supplementation 139 5.5.3 Revitalizing Fermentations 140 5.6 POST-FERMENTATION GROWTH 141 Chapter 6: Bottling 143 6.1 INTRODUCTION 143 6.2 FILTRATION 144 6.3 PRESERVATIVES AND STERILANTS 146 6.3.1 Sulfur Dioxide 147 6.3.2 Sorbic Acid 148 6.3.3 Fumaric Acid 151 6.3.4 Benzoic Acid 151 6.3.5 Dimethyldicarbonate 152 6.3.6 OTHER ANTIMICROBIAL AGENTS 153 Lysozyme 153 Nisin 154 Carbon Monoxide 154 6.4 LABORATORY PROCEDURES 155 6.4.1 Bottling Line Sampling 155 6.4.2 Sampling Low-Density Populations 155 Chapter 7: Wmery Sanitation 159 7.1 INTRODUCTION 159 7.2 PRELIMINARY CLEANING 160 7.3 WATER QUALI1Y 160 7.4 DETERGENTS 161 7.4.1 Alkalies 161 7.4.2 Sequestering Agents 162 7.4.3 Surfactants 162 7.4.4 Acids 162 7.5 SANITIZERS 162 7.5.1 Chlorine-Based Sanitizers 164 7.5.2 Iodine 165 7.5.3 Quaternary Ammonium Compounds 165 7.5.4 Detergent-Sanitizer Formulations 165 7.5.5 Sulfur Dioxide 166 7.5.6 Physical Sterilants 166 x Contents 7.6 SANITATION MONITORING 166 7.6.1 Swab Tests 167 7.6.2 Direct-Contact Tests 167 Appendix A: Basic Microscopy and Laboratory Setup 169 Al MICROSCOPE 169 Magnification 170 Resolution 170 Contrast 170 A2 CENTRIFUGE/FILTERS 171 A3 AUTOCLAVE 172 A.4 INCUBATOR 172 A.5 WATERBATHS 173 A.6 DISPOSABLE VERSUS REUSABLE PIPETfES 173 A.7 MEDIA 174 A.8 MISCELLANEOUS SUPPLIES 174 A.9 ALTERNATIVES 175 Appendix B: Media Preparation and Transfer Techniques 176 B.1 INTRODUCTION 176 B.2 PHYSICAL/CHEMICAL REQUIREMENTS FOR MEDIA 177 B.2.1 Utilizable Source of Carbon and Nitrogen 178 B.2.2 Oxygen Requirements 178 B.2.3 Hydrogen Ion Concentration (pH) 179 B.2.4 Moisture and Water Activity (Aw) 179 B.2.5 Incubation Temperature 180 B.3 STERILIZATION OF LABORATORY MEDIA AND SUPPLIES 180 B.3.1 Steam Sterilization 181 B.3.2 Boiling Water 182 B.3.3 Dry Heat 182 B.3.4 Sterile Filtration 183 B.3.5 Chemical Sterilization 183 Ethylene Oxide 183 Dimethyldicarbonate 183 Ethanol 184 B.4 MEDIA PREPARATION 184 B.4.1 Fermentation Broths 185 B.4.2 Solidified Media 186 Plates 186 Slants (Slopes) 187 Agar Deeps 188 B.5 STERILE TRANSFER TECHNIQUES 188 B.5.1 Transfers from Plate to Plate ("Streaking for Isolation") 188 B.5.2 Transfer from Slant to Slant 189 B.5.3 Transfer Between Agar "Deeps" 190 B.5.4 Transfer from Plates to Liquids 190 B.5.5 Liquid-to-Liquid and Liquid-to-Solid Transfers (Using Pipettes) 191 Contents xi Appendix C: Estimation of Population Density 193 C.1 INTRODUCTION 193 C.2 PREPARATION OF DILUTIONS 194 C.3 POPULATION MEASUREMENTS (PLATING OR INDIRECT METHODS) 197 C.3.1 Pour-Plate Technique 197 C.3.2 Spread-Plate Technique 198 C.3.3 Membrane Filtration 199 C.4 MICROSCOPIC (DIRECT) EXAMINATION USING VIABILI1Y STAINS 201 C.4.1 Methylene blue 201 C.4.2 Ponceau-S 202 C.4.3 Walford's stain 202 C.4.4 Viable Cell Counting 203 C.4.5 Epifluorescence Methods 205 C.4.6 Bioluminescence 206 C.5 RECENT TECHNOLOGY 207 C.5.1
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