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 Handbook of Enology
Volume 1
The Microbiology of Wine and Vinifications
2nd Edition

Handbook of Enology Volume 1 The Microbiology of Wine and Vinifications 2nd Edition P. Ribéreau-Gayon, D. Dubourdieu, B. Donèche and
A. Lonvaud  2006 John Wiley & Sons, Ltd ISBN: 0-470-01034-7
 Handbook of Enology
Volume 1
The Microbiology of Wine and Vinifications
2nd Edition

Pascal Ribéreau-Gayon
Denis Dubourdieu
Bernard Donèche
Aline Lonvaud

Faculty of Enology
Victor Segalen University of Bordeaux II, Talence, France

Original translation by

Jeffrey M. Branco, Jr.

Winemaker
M.S., Faculty of Enology, University of Bordeaux II

Revision translated by

Christine Rychlewski
Aquitaine Traduction, Bordeaux, France
Copyright  2006 John Wiley & Sons Ltd, The Atrium, Southern Gate, Chichester,
West Sussex PO19 8SQ, England

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under the terms of a licence issued by the Copyright Licensing Agency Ltd, 90 Tottenham Court Road, London W1T 4LP, UK, without the
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Library of Congress Cataloging-in-Publication Data:

Ribéreau-Gayon, Pascal.
[Traité d’oenologie. English]
Handbook of enology / Pascal Ribéreau-Gayon, Denis Dubourdieu, Bernard
Donèche ; original translation by Jeffrey M. Branco, Jr.—2nd ed. /
translation of updates for 2nd ed. [by] Christine Rychlewski.
v. cm.
Rev. ed. of: Handbook of enology / Pascal Ribéreau Gayon . . . [et al.].
c2000.
Includes bibliographical references and index.
Contents: v. 1. The microbiology of wine and vinifications
ISBN-13: 978-0-470-01034-1 (v. 1 : acid-free paper)
ISBN-10: 0-470-01034-7 (v. 1 : acid-free paper)
1. Wine and wine making—Handbooks, manuals, etc. 2. Wine and wine
making—Microbiology—Handbooks, manuals, etc. 3. Wine and wine
making—Chemistry—Handbooks, manuals, etc. I. Dubourdieu, Denis. II.
Donèche, Bernard. III. Traité d’oenologie. English. IV. Title.
TP548.T7613 2005
663
.2—dc22
2005013973

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

ISBN-13: 978-0-470-01034-1 (HB)

ISBN-10: 0-470-01034-7 (HB)

Typeset in 10/12pt Times by Laserwords Private Limited, Chennai, India

Printed and bound in Great Britain by Antony Rowe Ltd, Chippenham, Wiltshire
This book is printed on acid-free paper responsibly manufactured from sustainable forestry
in which at least two trees are planted for each one used for paper production.
Contents
Remarks Concerning the Expression of Certain Parameters of Must and Wine Composition vii
Preface to the First Edition ix
Preface to the Second Edition xiii
1 Cytology, Taxonomy and Ecology of Grape and Wine Yeasts 1
2 Biochemistry of Alcoholic Fermentation and Metabolic Pathways of Wine Yeasts 53
3 Conditions of Yeast Development 79
4 Lactic Acid Bacteria 115
5 Metabolism of Lactic Acid Bacteria 139
6 Lactic Acid Bacteria Development in Wine 161
7 Acetic Acid Bacteria 183
8 The Use of Sulfur Dioxide in Must and Wine Treatment 193
9 Products and Methods Complementing the Effect of Sulfur Dioxide 223
10 The Grape and its Maturation 241
11 Harvest and Pre-Fermentation Treatments 299
12 Red Winemaking 327
13 White Winemaking 397
14 Other Winemaking Methods 445
Index 481
Remarks Concerning the Expression
of Certain Parameters of Must
and Wine Composition
UNITS more correct expression in milliequivalents per
liter has not been embraced in France. The expres-
Metric system units of length (m), volume (l) and sion of total and volatile acidity in the equivalent
weight (g) are exclusively used. The conversion of weight of sulfuric acid has been used predomi-
metric units into Imperial units (inches, feet, gal- nantly throughout these works. In certain cases, the
lons, pounds, etc.) can be found in the following corresponding weight in tartaric acid, often used in
enological work: Principles and practices of wine- other countries, has been given.
making, R.B. Boulton, V.L. Singleton, L.F. Bisson Using the weight of the milliequivalent of the
and R.E. Kunkee, 1995, The Chapman & Hall
various acids, the below table permits the conver-
Enology Library, New York.
sion from one expression to another.
More particularly, to convert from total acidity
EXPRESSION OF TOTAL ACIDITY expressed in H2 SO4 to its expression in tartaric
AND VOLATILE ACIDITY acid, add half of the value to the original value
(4 g/l H2 SO4 → 6 g/l tartaric acid). In the other
Although EC regulations recommend the expres- direction a third of the value must be subtracted.
sion of total acidity in the equivalent weight of tar- The French also continue to express volatile
taric acid, the French custom is to give this expres- acidity in equivalent weight of sulfuric acid. More
sion in the equivalent weight of sulfuric acid. The generally, in other countries, volatile acidity is

Desired Expression
Known Expression meq/l g/l g/l g/l
H2 SO4 tartaric acid acetic acid
meq/l 1.00 0.049 0.075 0.060
g/l H2 SO4 20.40 1.00 1.53 1.22
g/l tartaric acid 13.33 0.65 1.00
g/l acetic acid 16.67 0.82 1.00
Multiplier to pass from one expression of total or volatile acidity to another
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viii Remarks Concerning the Expression of Certain Parameters of Must and Wine Composition

expressed in acetic acid. It is rarely expressed and then divided by 17 to obtain a fairly good
in milliequivalents per liter. The below table also approximation of the TAP.
allows simple conversion from one expression to
another. In any case, the determination of the Brix or TAP
The expression in acetic acid is approximately of a must is approximate. First of all, it is not
20% higher than in sulfuric acid. always possible to obtain a representative grape
or must sample for analysis. Secondly, although
physical, densimetric or refractometric measure-
EVALUATING THE SUGAR ments are extremely precise and rigorously express
CONCENTRATION OF MUSTS the sugar concentration of a sugar and water mix-
ture, these measurements are affected by other sub-
This measurement is important for tracking grape
stances released into the sample from the grape
maturation, fermentation kinetic and if necessary
and other sources. Furthermore, the concentrations
determining the eventual need for chaptalization.
of these substances are different for every grape
This measurement is always determined by
or grape must sample. Finally, the conversion rate
physical, densimetric or refractometric analysis.
of sugar into alcohol (approximately 17 to 18 g/l)
The expression of the results can be given accord-
varies and depends on fermentation conditions and
ing to several scales: some are rarely used, i.e.
yeast properties. The widespread use of selected
degree Baumé and degree Oechsle. Presently, two yeast strains has lowered the sugar conversion rate.
systems exist (Section 10.4.3):

1. The potential alcohol content (titre alcoomét- Measurements Using Visible

raque potential or TAP, in French) of musts and Ultraviolet Spectrometry
can be read directly on equipment, which is The measurement of optic density, absorbance, is
graduated using a scale corresponding to 17.5 widely used to determine wine color (Volume 2,
or 17 g/l of sugar for 1% volume of alcohol. Section 6.4.5) and total phenolic compounds con-
Today, the EC recommends using 16.83 g/l as centration (Volume 2, Section 6.4.1). In these
the conversion factor. The ‘mustimeter’ is a works, the optic density is noted as OD, OD 420
hydrometer containing two graduated scales: (yellow), OD 520 (red), OD 620 (blue) or OD 280
one expresses density and the other gives a (absorption in ultraviolet spectrum) to indicate the
direct reading of the TAP. Different methods optic density at the indicated wavelengths.
varying in precision exist to calculate the TAP Wine color intensity is expressed as:
from a density reading. These methods take var-
ious elements of must composition into account CI = OD 420 + OD 520 + OD 620,
(Boulton et al., 1995).
Or is sometimes expressed in a more simplified
2. Degree Brix expresses the percentage of sugar form: CI = OD 420 + OD 520.
in weight. By multiplying degree Brix by 10, Tint is expressed as:
the weight of sugar in 1 kg, or slightly less
OD 420
than 1 liter, of must is obtained. A conversion T =
table between degree Brix and TAP exists in OD 520
Section 10.4.3 of this book. 17 degrees Brix The total phenolic compound concentration is
correspond to an approximate TAP of 10% and expressed by OD 280.
20 degrees Brix correspond to a TAP of about The analysis methods are described in Chapter 6
12%. Within the alcohol range most relevant to of Handbook of Enology Volume 2, The Chemistry
enology, degree Brix can be multiplied by 10 of Wine.
Preface to the First Edition
Wine has probably inspired more research and Wine Analysis J. Ribéreau-Gayon and E. Peynaud
publications than any other beverage or food. In (1947 and 1958); Treatise on Enology (2 Volumes)
fact, through their passion for wine, great scientists J. Ribéreau-Gayon and E. Peynaud (1960 and
have not only contributed to the development of 1961); Wine and Winemaking E. Peynaud (1971
practical enology but have also made discoveries and 1981); Wine Science and Technology (4 volu-
in the general field of science. mes) J. Ribéreau-Gayon, E. Peynaud, P. Ribéreau-
A forerunner of modern enology, Louis Pasteur Gayon and P. Sudraud (1975–1982).
developed simplified contagious infection mod- For an understanding of current advances in
els for humans and animals based on his obser- enology, the authors propose this book Handbook
vations of wine spoilage. The following quote of Enology Volume 1: The Microbiology of Wine
clearly expresses his theory in his own words: and Vinifications and the second volume of the
‘when profound alterations of beer and wine are Handbook of Enology Volume 2: The Chemistry of
observed because these liquids have given refuge Wine: Stabilization and Treatments.
to microscopic organisms, introduced invisibly and Although written by researchers, the two vol-
accidentally into the medium where they then
umes are not specifically addressed to this group.
proliferate, how can one not be obsessed by the
Young researchers may, however, find these books
thought that a similar phenomenon can and must
useful to help situate their research within a par-
sometimes occur in humans and animals.’
ticular field of enology. Today, the complexity of
Since the 19th century, our understanding of
modern enology does not permit a sole researcher
wine, wine composition and wine transformations
has greatly evolved in function of advances in rel- to explore the entire field.
evant scientific fields i.e. chemistry, biochemistry, These volumes are also of use to students and
microbiology. Each applied development has lead professionals. Theoretical interpretations as well
to better control of winemaking and aging con- as solutions are presented to resolve the problems
ditions and of course wine quality. In order to encountered most often at wineries. The authors
continue this approach, researchers and winemak- have adapted these solutions to many different sit-
ers must strive to remain up to date with the latest uations and winemaking methods. In order to make
scientific and technical developments in enology. the best use of the information contained in these
For a long time, the Bordeaux school of enology works, enologists should have a broad understand-
was largely responsible for the communication of ing of general scientific knowledge. For example,
progress in enology through the publication of the understanding and application of molecular
numerous works (Béranger Publications and later biology and genetic engineering have become
Dunod Publications): indispensable in the field of wine microbiology.
Wine Analysis U. Gayon and J. Laborde (1912); Similarly, structural and quantitative physiochem-
Treatise on Enology J. Ribéreau-Gayon (1949); ical analysis methods such as chromatography,
x Preface to the First Edition

NMR and mass spectrometry must now be treatments’ is completed when the wine is bottled.
mastered in order to explore wine chemistry. Aging specifically refers to the transformation of
The goal of these two works was not to create bottled wine.
an exhaustive bibliography of each subject. The This distinction of two phases is certainly the
authors strove to choose only the most relevant and result of commercial practices. Traditionally in
significant publications to their particular field of France, a vine grower farmed the vineyard and
research. A large number of references to French transformed grapes into an unfinished wine. The
enological research has been included in these wine merchant transferred the bulk wine to his cel-
works in order to make this information available lars, finished the wine and marketed the product,
to a larger non-French-speaking audience. preferentially before bottling. Even though most
In addition, the authors have tried to convey wines are now bottled at the winery, these long-
a French and more particularly a Bordeaux per- standing practices have maintained a distinction
spective of enology and the art of winemaking. between ‘wine grower enology’ and ‘wine mer-
The objective of this perspective is to maximize chant enology’. In countries with a more recent
the potential quality of grape crops based on the viticultural history, generally English speaking, the
specific natural conditions that constitute their ‘ter- vine grower is responsible for winemaking and
roir’. The role of enology is to express the char- wine sales. For this reason, the Anglo-Saxon tradi-
acteristics of the grape specific not only to variety tion speaks of winemaking, which covers all oper-
and vineyard practices but also maturation condi- ations from harvest reception to bottling.
tions, which are dictated by soil and climate. In these works, the distinction between ‘vinifi-
It would, however, be an error to think that the cation’ and ‘stabilization and treatments’ has been
world’s greatest wines are exclusively a result of maintained, since the first phase primarily concerns
tradition, established by exceptional natural con- microbiology and the second chemistry. In this
ditions, and that only the most ordinary wines, manner, the individual operations could be linked
produced in giant processing facilities, can ben- to their particular sciences. There are of course lim-
efit from scientific and technological progress. its to this approach. Chemical phenomena occur
Certainly, these facilities do benefit the most from during vinification; the stabilization of wines dur-
high performance installations and automation of ing storage includes the prevention of microbial
operations. Yet, history has unequivocally shown contamination.
that the most important enological developments Consequently, the description of the different
in wine quality (for example, malolactic fermenta- steps of enology does not always obey logic as
tion) have been discovered in ultra premium wines. precise as the titles of these works may lead
The corresponding techniques were then applied to to believe. For example, microbial contamination
less prestigious products. during aging and storage are covered in Vol-
High performance technology is indispensable ume 1. The antiseptic properties of SO2 incited the
for the production of great wines, since a lack description of its use in the same volume. This line
of control of winemaking parameters can easily of reasoning lead to the description of the antioxi-
compromise their quality, which would be less of dant related chemical properties of this compound
a problem with lower quality wines. in the same chapter as well as an explanation of
The word ‘vinification’ has been used in this adjuvants to sulfur dioxide: sorbic acid (antisep-
work and is part of the technical language of tic) and ascorbic acid (antioxidant). In addition,
the French tradition of winemaking. Vinification the on lees aging of white wines and the result-
describes the first phase of winemaking. It com- ing chemical transformations cannot be separated
prises all technical aspects from grape maturity from vinification and are therefore also covered
and harvest to the end of alcoholic and some- in Volume 1. Finally, our understanding of pheno-
times malolactic fermentation. The second phase lic compounds in red wine is based on complex
of winemaking ‘winematuration, stabilization and chemistry. All aspects related to the nature of the
Preface to the First Edition xi

corresponding substances, their properties and their cases i.e. phenolic compounds, whose different
evolution during grape maturation, vinification and families are often defined by analytical criteria.
aging are therefore covered in Volume 2. The authors thank the following people who
These works only discuss the principles of have contributed to the creation of this work:
equipment used for various enological operations J.F. Casas Lucas, Chapter 14, Sherry; A. Brugi-
and their effect on product quality. For example, rard, Chapter 14, Sweet wines; J.N. de Almeida,
temperature control systems, destemmers, crushers Chapter 14, Port wines; A. Maujean, Chapter 14,
and presses as well as filters, inverse osmosis Champagne; C. Poupot for the preparation of
machines and ion exchangers are not described in material in Chapters 1, 2 and 13; Miss F. Luye-
detail. Bottling is not addressed at all. An in-depth Tanet for her help with typing.
description of enological equipment would merit a They also thank Madame B. Masclef in particu-
detailed work dedicated to the subject. lar for her important part in the typing, preparation
Wine tasting, another essential role of the and revision of the final manuscript.
winemaker, is not addressed in these works.
Many related publications are, however, readily Pascal Ribéreau-Gayon
available. Finally, wine analysis is an essential tool Bordeaux
that a winemaker should master. It is, however, not
covered in these works except in a few particular
Preface to the Second Edition
The two-volume Enology Handbook was pub- individual terroirs. Scientific winemaking has not
lished simultaneously in Spanish, French, and Ital- resulted in standardization or leveling of quality.
ian in 1999 and has been reprinted several times. On the contrary, by making it possible to correct
The Handbook has apparently been popular with defects and eliminate technical imperfections, it
students as an educational reference book, as well has revealed the specific qualities of the grapes
as with winemakers, as a source of practical solu- harvested in different vineyards, directly related to
tions to their specific technical problems and sci- the variety and terroir, more than ever before.
entific explanations of the phenomena involved. Interest in wine in recent decades has gone
It was felt appropriate at this stage to prepare beyond considerations of mere quality and taken
an updated, reviewed, corrected version, including on a truly cultural dimension. This has led some
the latest enological knowledge, to reflect the many people to promote the use of a variety of tech-
new research findings in this very active field. The niques that do not necessarily represent significant
outline and design of both volumes remain the progress in winemaking. Some of these are sim-
same. Some chapters have changed relatively little ply modified forms of processes that have been
as the authors decided there had not been any sig- known for many years. Others do not have a suf-
nificant new developments, while others have been ficiently reliable scientific interpretation, nor are
modified much more extensively, either to clarify their applications clearly defined. In this Hand-
and improve the text, or, more usually, to include book, we have only included rigorously tested
new research findings and their practical applica- techniques, clearly specifying the optimum con-
tions. Entirely new sections have been inserted in ditions for their utilization.
some chapters. As in the previous edition, we deliberately
We have made every effort to maintain the same omitted three significant aspects of enology: wine
approach as we did in the first edition, reflecting analysis, tasting, and winery engineering. In view
the ethos of enology research in Bordeaux. We use of their importance, these topics will each be
indisputable scientific evidence in microbiology, covered in separate publications.
The authors would like to take the opportunity
biochemistry, and chemistry to explain the details
of the publication of this new edition of Volume 1
of mechanisms involved in grape ripening, fermen-
to thank all those who have contributed to updating
tations and other winemaking operations, aging,
this work:
and stabilization. The aim is to help winemakers
achieve greater control over the various stages in
— Marina Bely for her work on fermentation
winemaking and choose the solution best suited
kinetics (Section 3.4) and the production of
to each situation. Quite remarkably, this scientific
volatile acidity (Sections 2.3.4 and 14.2.5)
approach, most intensively applied in making the
finest wines, has resulted in an enhanced capac- — Isabelle Masneuf for her investigation of the
ity to bring out the full quality and character of yeasts’ nitrogen supply (Section 3.4.2)
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