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Handbook of Enology Volume 1 Handbook of Enology VOLUME 1 THE MICROBIOLOGY OF WINE AND VINIFICATIONS Third Edition Pascal Ribéreau‐Gayon, Denis Dubourdieu, Bernard B. Donèche and Aline A. Lonvaud Research Unit in Enology, Institute of Vine and Wine Sciences, University of Bordeaux, Villenave d’Ornon, France Coordinator: Philippe Darriet Research Unit in Enology, Institute of Vine and Wine Sciences, University of Bordeaux, Villenave d’Ornon, France Translator: John Towey With assistance from Juliet Towey Vista Translations LLC, Portland, OR, USA This edition first published 2021 © 2021 John Wiley & Sons Ltd Edition History 1e Wiley 2000 2e Wiley 2005 All rights reserved. No part of this publication may be reproduced, stored in a retrieval system, or transmitted, in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, except as permitted by law. Advice on how to obtain permission to reuse material from this title is available at http://www.wiley.com/go/permissions. The right of Pascal Ribéreau‐Gayon, Denis Dubourdieu, Bernard B. Donèche and Aline A. Lonvaud to be identified as the authors of this work has been asserted in accordance with law. Originally published in France as Traité d’oenologie Volume 1 – Microbiologie du Vin, Vinifications By Pascal Ribéreau‐Gayon, Denis Dubourdieu, Bernard B. Donèche, Aline A. Lonvaud © Dunod Editeur, 2017, Malakoff Registered Offices John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, USA John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK Editorial Office The Atrium, Southern Gate, Chichester, West Sussex, PO19 8SQ, UK For details of our global editorial offices, customer services, and more information about Wiley products visit us at www.wiley.com. Wiley also publishes its books in a variety of electronic formats and by print‐on‐demand. Some content that appears in standard print versions of this book may not be available in other formats. Limit of Liability/Disclaimer of Warranty In view of ongoing research, equipment modifications, changes in governmental regulations, and the constant flow of information relating to the use of experimental reagents, equipment, and devices, the reader is urged to review and evaluate the information provided in the package insert or instructions for each chemical, piece of equipment, reagent, or device for, among other things, any changes in the instructions or indication of usage and for added warnings and precautions. While the publisher and authors have used their best efforts in preparing this work, they make no representations or warranties with respect to the accuracy or completeness of the contents of this work and specifically disclaim all warranties, including without limitation any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives, written sales materials or promotional statements for this work. The fact that an organization, website, or product is referred to in this work as a citation and/or potential source of further information does not mean that the publisher and authors endorse the information or services the organization, website, or product may provide or recommendations it may make. This work is sold with the understanding that the publisher is not engaged in rendering professional services. The advice and strategies contained herein may not be suitable for your situation. You should consult with a specialist where appropriate. Further, readers should be aware that websites listed in this work may have changed or disappeared between when this work was written and when it is read. Neither the publisher nor authors shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. Library of Congress Cataloging‐in‐Publication Data Names: Ribéreau-Gayon, Pascal, author. | Dubourdieu, Denis, author. | Donèche, Bernard B., author. | Lonvaud, Aline A., author. | Glories, Yves, author. | Maujean, Alain, author. | Towey, John, translator. Title: Handbook of enology. Other titles: Traite´ d’oenologie. English Description: Third edition / coordinator, Philippe Darriet ; translator John Towey. | Hoboken, NJ : Wiley, 2020-2021. | Includes bibliographical references and indexes. | Contents: volume 1. The microbiology of wine and vinifications / Pascal Ribéreau-Gayon, Denis Dubourdieu, Bernard B. Donèche and Aline A. Lonvaud – volume 2. The chemistry of wine stabilization and treatments / Pascal Ribéreau-Gayon, Denis Dubourdieu, Yves Glories and Alain Maujean. Identifiers: LCCN 2020033294 (print) | LCCN 2020033295 (ebook) | ISBN 9781119584681 (v. 1 ; hardback) | ISBN 9781119587767 (v. 2 ; hardback) | ISBN 9781119584629 (v. 1 ; epub) | ISBN 9781119588498 (v. 2 ; epub) | ISBN 9781119584698 (v. 1 ; adobe pdf) | ISBN 9781119588443 (v. 2 ; adobe pdf) Subjects: LCSH: Wine and wine making–Handbooks, manuals, etc. | Wine and wine making–Microbiology–Handbooks, manuals, etc. | Wine and wine making–Chemistry–Handbooks, manuals, etc. Classification: LCC TP548 .T7613 2020 (print) | LCC TP548 (ebook) | DDC 663/.2–dc23 LC record available at https://lccn.loc.gov/2020033294 LC ebook record available at https://lccn.loc.gov/2020033295 Cover Design: Wiley Cover Image: © Digoarpi/Shutterstock Set in 10/12pt StixTwo Text by SPi Global, Pondicherry, India 10 9 8 7 6 5 4 3 2 1 CONTENTS Foreword xi Preface to the Second Edition xiii Preface to the First Edition xv Remarks Concerning the Expression of Certain Parameters of Must and Wine Composition xix Part I Microbiology of Wine 1 1 Yeasts 3 1.1 . Introduction 3 1.2 . The Cell Wall 5 1.3 . The Plasma Membrane 9 1.4 . The Cytoplasm and Its Organelles 15 1.5 . The Nucleus 18 1.6 . Reproduction and the Yeast Biological Cycle 19 1.7 . The Killer Phenomenon 23 1.8 . Classifi cation of Yeast Species 27 1.9 . Identifi cation of Wine Yeast Strains 49 1.10. Ecology of Grape and Wine Yeasts 57 References 67 2 Yeast Metabolism 73 2.1 . Introduction 73 2.2 . Sugar Degradation Pathways 74 2.3 . Regulation of Sugar-Utilizing Metabolic Pathways 82 2.4 . Metabolism of Nitrogen Compounds 93 References 101 vi Contents 3 Conditions of Yeast Development 103 3.1. Introduction 103 3.2. Monitoring and Controlling Fermentations 104 3.3. Yeast Growth Cycle and Fermentation Kinetics 108 3.4. Nutrition Requirements 110 3.5. Fermentation Activators 115 3.6. Inhibition of Fermentation 122 3.7. Physicochemical Factors Affecting Yeast Growth and Fermentation Kinetics 126 3.8. Stuck Fermentations 135 References 142 4 Lactic Acid Bacteria 145 4.1. The Different Components of the Bacteria Cell 145 4.2. Taxonomy of Lactic Acid Bacteria 154 4.3. Identification of Lactic Acid Bacteria 156 4.4. The Oenococcus oeni Species 170 References 171 5 Metabolism of Lactic Acid Bacteria 175 5.1. Generalities: A Review 175 5.2. Metabolism of Sugars: Lactic Acid Fermentation 177 5.3. Metabolism of the Principal Organic Acids of Wine 182 5.4. Other Transformations Likely to Occur in Winemaking 189 5.5. Effect of the Metabolism of Lactic Acid Bacteria on Wine Composition and Quality 195 References 197 6 Lactic Acid Bacteria Development in Wine 201 6.1. Lactic Acid Bacteria Nutrition in Wine 201 6.2. Physicochemical Factors of Bacterial Growth 204 6.3. Evolution of Lactic Acid Bacteria Microflora: Influence on Wine Composition 210 6.4. Microbial Interactions During Winemaking 218 6.5. The Importance of Bacteriophages 224 References 226 Contents vii 7 Acetic Acid Bacteria 229 7.1. Principal Characteristics and Cytology 229 7.2. Classification and Identification 230 7.3. Principal Physiological Characteristics 232 7.4. Metabolism of Acetic Acid Bacteria 233 7.5. Acetic Acid Bacteria Development in Grape Musts 237 7.6. Evolution of Acetic Acid Bacteria During Winemaking and Wine Aging and the Impact on Wine Quality 238 References 240 8 The Use of Sulfur Dioxide in Must and Wine Treatment 243 8.1. Introduction 243 8.2. Physiological Effects 246 8.3. Chemistry of Sulfur Dioxide 248 8.4. Molecules Binding Sulfur Dioxide 252 8.5. Practical Consequences: The State of Sulfur Dioxide in Wines 261 8.6. Antimicrobial Properties of Sulfur Dioxide 262 8.7. The Role of Sulfur Dioxide in Winemaking 265 8.8. The Use of Sulfur Dioxide in the Winery 270 References 277 9 Products and Methods Complementing the Effect of Sulfur Dioxide 279 9.1. Introduction 279 9.2. Sorbic Acid 280 9.3. Octanoic and Decanoic Acids (Saturated Short-Chain Fatty Acids) 283 9.4. Dimethyl Dicarbonate (DMDC) 285 9.5. Lysozyme 286 9.6. Destruction of Yeasts by Heat (Pasteurization) 289 9.7. Ascorbic Acid 293 9.8. The Use of Inert Gases 297 References 301 viii Contents Part II Vinification 303 Reflections on Global Taste and Typicity of Wines 305 10 The Grape and Its Maturation 309 10.1. Introduction 309 10.2. Description and Composition of the Mature Grape 310 10.3. Changes in the Grape During Maturation 322 10.4. Definition of Ripeness—Concept of Vintage 343 10.5. Impact of Various Other Factors on Maturation and Grape Composition at Ripeness 366 10.6. Botrytis cinerea 375 References 391 11 Harvest and Pre-fermentation Treatments 399 11.1. Introduction 399 11.2. Improving Grape Quality by Overripening 400 11.3. Harvest Date and Operations 401 11.4. Acidity Adjustments of the Harvested Grapes 408 11.5. Increasing Sugar Concentrations 412 11.6. Enzymatic Transformations of the Grape After Harvest 418 11.7. Use of Commercial Enzymes in Winemaking 427 References 429 12 Red Winemaking 431 12.1. Generalities 431 12.2. Mechanical Processing of the Harvested Grapes 433 12.3. Tank Filling 440 12.4. Controlling Alcoholic Fermentation 446 12.5. Maceration 452 12.6. Draining Off the Skins and Pressing 469 12.7. Malolactic Fermentation 480 12.8. Automated Red Winemaking Methods 494 12.9. Carbonic Maceration 499 References 511 Contents ix 13 White Winemaking 513 13.1. Distinctive Characteristics of White Winemaking 513 13.2. White Grape Quality and Picking Criteria 517 13.3. Juice Extraction 526 13.4. Protecting Juice from Oxidation 538 13.5. Clarification 543 13.6. Juice Treatments and the Advisability of Bentonite Treatments 552 13.7. Fermentation Operations 552 13.8. Making Dry White Wines in Barrels 557 13.9. Controlling Reduction Off-Aromas During White Wine Aging 564 References 568 14 Other Winemaking Methods 571 14.1. Rosé Wines 571 14.2. Botrytized Sweet Wines (Sauternes and Tokaji) 577 14.3. Champagne and Sparkling Wines 588 14.4. Fortified Wines 602 14.5. Flor Wines 608 References 612 Index 615 FOREWORD Over 70 years ago, in 1949, under the and wine, a greater understanding of direction of Professor Jean Ribéreau‐Gayon, chemical and microbiological parameters, the fi rst edition of T raité d’oenologie ( Hand- with the aim of ensuring satisfactory fer- book of Enology ) appeared. That publication mentations and predicting the evolution of aimed to gather scientifi c and technical wines, and better mastery of wine stabili- knowledge in the main fi elds of a science zation processes. As a result, the purpose still in its early stages, originating from the of this publication is to guide readers in works of Louis Pasteur. Later on, the book their thought processes with a view to pre- was regularly republished by integrating the serving and optimizing the identity and most recent experimental results, especially taste of wine and its aging potential. In those conducted at the Bordeaux School other words, this publication was meant, as of Enology. In 1997, the fourth edition, pub- the late Professor Denis Dubourdieu wrote, lished under the direction of Professor to help “. . .obtain original products that Pascal Ribéreau‐Gayon and Professors are suffi ciently complex, fi ne, and appreci- Denis Dubourdieu, Yves Glories, Aline A. ated by modern consumers. . .,” in a con- Lonvaud, Bernard B. Donèche, and Alain text where grape ripeness characteristics Maujean, was completely reshaped into are evolving and greater attention is being two volumes. placed on chemical additives. The current book, which is the seventh The outline of this edition of the Hand- French edition (third in English) of the book of Enology does not diff er from the Handbook of Enology , updates and enriches previous ones. Some of the chapters have the knowledge found in the two‐volume been signifi cantly reworked in light of edition that fi rst appeared 20 years ago. recent research, while others have been Subsequent editions were printed in 2004 changed only slightly or not at all. This is a and then in 2012. As Émile Peynaud said in collective work; it is the fruit of eff orts by Le Vin et les Jours , “. . .enology is at the ser- the researchers and professors of the Enol- vice of wine, it is a science in movement, ogy Research Unit at the I nstitut des Sciences advancing both in its research and applica- de la Vigne et du Vin (ISVV), who provided tions. . .” As an applied science, it is irri- their expertise to update this Volume 1: gated by knowledge from the fundamental sciences (chemistry, biochemistry, micro- • Patricia Ballestra, Associate Professor, biology, bioengineering, psychophysics, IUT Périgueux, Université de Bor- cognitive psychology, etc.) and nourished deaux, researcher in the Enology by empirical observations. The approach Research Unit, ISVV, Université de used in the H andbook of Enology is thus Bordeaux (Volume 1, Chapter  5 ), still the same. It is about providing practi- regarding spoilage microorganisms tioners, winemakers, technicians, and related to mousiness. enology students with solid knowledge and • Philippe Darriet, Director of the Enol- the most recent research results. This ogy Research Unit, Université de Bor- knowledge can be used to contribute to a deaux (Volume 1, Chapters 1 0 and 1 3 ) , better defi nition of the quality of grapes regarding the biochemistry of aromas xi

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