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The Fermentation Properties of Non-Saccharomyces Wine Yeasts and Their Interaction with Saccharomyces Cerevisiae

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The Fermentation Properties of Non-Saccharomyces Wine Yeasts and Their Interaction with Saccharomyces Cerevisiae Y s,r-( THE FERMENTATION PROPERTIES OF NON. SACCHAROMYCES YEASTS AND THEIR ¡NTERACTION WITH SACCHAROMYCES Alison Soden Depaftment of Horticulture, Viticulture and Oenology The University of Adelaide The Australian Wine Research lnstitute A thesis submitted in fulfilrhent of the requirements for the degree of Doctor of Philosophy July 1998 TABLE OF CONTENTS ABSTRACT v DECLARATION vi ACKNOWLEDGEMENTS....... vll CHAPTER 1. INTRODUCTION AND AIMS OF THIS STUDY.... I 1.1. Taxonomy of wþemaking yeasts....... 1 1.1.1. Morphology and mode of reproduction.......'... I 2 1. 1.2. Physiological features.......... ) 1. 1.3. Biochemical characteristics........ 2 1. 1.4. Genome homology .................. l. 1.5. Strain differentiation........ 3 1.2. Survey of the yeast flora of the grape......... 4 !.2.1. Factors affecting the microbial flora of grapes 4 1.3. Survey of the yeast flora of grape juice fermentation 5 1.3.1. Factors affecting yeast growth during fermentation. 5 1.4. Indigenous vs inoculated fermentation.'.....' 7 1.4.1. Mixed cultures 8 1.5. The effects of yeast on wine composition.. 9 1.5.1. Ethanol.. 10 1.5.2. Higher alcohols 10 11 t2 1.5.5. Carbonyl compounds........ """""" 15 1.5.6. Glycerol ........... 15 1.5.7. Nitrogenous compounds... 15 1.5.8. Extracellular enzymes t6 1.5.9. Volatile phenols.. t6 1.5. 10. Sulfurous compounds ....... t7 1.6. lVine spoilage by yeasts t7 1.7. Aims of this study......... t9 CHAPTER 2. THE USE OF THE PCR FOR YEAST STRAIN DIFFERENTIATION AND SPECIES IDENTIFICATION 20 2.1. Introduction 20 2.2. Mateials and methods.... 2l 2.2.1. Yeast isolates and media 2I 2.2.2. Preparation of DNA template for PCR by afreeze-boilmethod................ 23 2.2.3. Yeast DNA purification 24 2.2.4.Intron splice site primers and PCR conditions 24 2.3. Results 24 2.3.I. Design and use of intron splice site primers.. 24 2.3.2. Differentiation of unknown yeast isolates'...-......... """""'25 2.3.3. Species possess characteristic amplification fingerprints. 25 2.3.4. Species identification of indigenous wine yeasts. 29 2.3.5. Heterogeneity within the species Candida steLlata. 30 2.4. Discussion 30 2.5. Conclusion..... 33 CHAPTER 3. THE EFFECT OF FERMENTATION WITH SELECTED NON- SÀ'CChÀno¡øvCeS yEASTS UPoN WINE CoMPOSlTloN, ARoMA AND PREFERENCE 35 3.1. Introduction 351Å 3.2. Maærials and methods 36 3.2.1. Juice and winemaking .'......... 36 3.2.2. Yeast strains and fermentation treatments.... 37 3.2.3. Starter culture preparation 38 3.2.4. Chemical analysis 39 3.2.5. Preference testing 39 3.2.6. Data analysis... 40 3.3. Results 40 3.3.1. Fermentation kinetics and wine composition...'........'..""""""""'i 40 3.3.2. Wine composition................ 41, 3.3.3. Sensory analysis 44 3.3.4. Description of wine aroma. 45 3.4. Discussion ........ 46 3.5. Conclusion................. 51 CHAPTER4. THE EFFECT OF INOCULATION PROTOCOL UPON STRAIN GROWTH AND WINE COMPOSITION IN MIXED YEAST SPECIES FERMENTATION 52 4.1. Intoduction ......... 52 4.2. Matanals and methods.... 54 4.2.1. Yeast strains and inoculation protocol ............... """""""' 54 4.2.2. Chemically defined grape juice medium.... 54 4.2.3. Starter culture preparation 55 4.2.4. Fermentation and monitoring 56 4.2.5. Chemical analysis of wines 56 4.3. Results and discussion 57 4.3.1. COINOCULATION OF T'SPORA DELBRUECICIAWRI 860 AJVD S. 57 CEREVTSIAE EC 1 I I 8.................. 4,3.2. COINOCULATION OF C. STELLATA CBS 843 AND S, CEREVISIAE 8C1118 .....61 4.3.3. COINOCULATION OF C. STELLATA CBS 2649 AND ECl 1 18 66 4.3.4. SEQUENTIAL INOCULATION OF C. STELLATA CF,S 2æ9 AND S. 72 CEREVTSTAE EC 1 1 I 8.............. 4.4. Conclusion.............. 78 CHAPTER 5. THE EFFECT OF MIXED YEAST SPECIES FERMENTATION ON THE AROMA AND COMPOSITION OF CHARDONNAY WINE....... 81 5. 1. Inüoduction ......... 81 5.2. Materials and Methods . 82 5.2.1. Juice and winemaking ............... 82 5.2.2. Yeast strains 83 5.2.3. Starter culture preparation 83 5.2.4. Fermentation treatments .. 83 5.2.5. Chemical analysis.... 84 5.2.6. Quantitative descriptive analysis 85 5.2.7. Difference testing by duo-trio analysis 86 5.2.8. Data analysis.. 86 5.3. Results 87 5.3. I . Yeast growth during fermentation............... 87 5.3.2. Chemical composition...... 90 5.3.3. Sensory analysis ....... 9l 5.4. Discussion 96 5.5. Conclusion.......... 97 CHAPTER 6. CONCLUSION 99 APPENDIX 1. YEASTS ASSOCIATED WITH WINEMAKING 104 APPENDIX 2. WINE YEASTS: SEXUAL AND ASEXUAL FORMS............. 105 BIBLIOGRAPHY....... 106 v ABSTRACT This thesis is concerned with the contribution of some non-SacchaÍomyces yeasts, particularly strains of Candida, to grape juice fermentation. An emphasis was placed upon the investigation of fermentation by non-Saccåarcmyces strains in mixed culture with a winemaking strain of Saccharomyces cerevisiae. A polymerase chain reaction (PCR) method using intron splice site specific primers was developed that permitted rapid identification of a number of species associated with winemaking, and also differentiation of isolates of the same species at the strain level. For preliminary assessment of their fermentation characteristics and aroma profiles, a number of yeasts were used as pure cultures for winemaking. Differences in fermentative capacity, the effect upon wine composition, aroma description and preference were identified between S. cerevisiae and the non-Saccharomyce^s yeasts. Strains within a species also had notably different effects upon fermentation. As few of these yeasts were able to completely ferment grape juice, acceptable isolates were also assessed for their growth and metabolic activities during mixed culture fermentation of a chemically defined grape juice-like medium, in conjunction with a commercial winemaking strain of Saccharomyces cerevisiae. The effect on growth kinetics and wine composition were found to depend upon the relative growth rate and fermentative capacity of the two strains involved. Coinoculation and sequential inoculation strategies were identified that could promote the growth and metabolic activity of a weakly fermentative strain and similarly suppress the strongly fermentative S. cerevisiae strain. Formal sensory analysis was used to describe the aroma of wines made by monocultures of C' ste|latacBs 2649 and S. cerevisiae ECl118, and by coinoculation and sequential inoculation of both of these yeasts. The inoculation protocol was found to have a significant effect upon utilisation pattern, the wine composition the fermentation kinetics - particularly the sugar and the wine aroma. Sequential inoculation, where the fermentation was initiated with C. stellata and completed with S. cerevisiae, produced a wine that had constitutional and aroma characteristics of both strains, but with significant differences in intensity. This work demonstrates the potential use of selected non-Saccåarcmyces yeasts in commercial winemaking. v1 DEGLARATION This thesis contains no material which has been accepted for the award of any other degree or diploma in any university or tertiary institution and, to the best of my knowledge and belief contains no material previously published or written by another person, except where due reference has been made in the text. I give consent to this thesis, when deposited in the University Library, being available for loan and photocopying. vll ACKNOWLEDGEMENTS The receipt of an APRA/Industry postgraduate award in conjunction with Lallemand Pty Ltd Australia is gratefully acknowledged. I would like to thank: My supervisors Dr Paul Henschke, Dr Miguel de Barros Lopes, Dr Vladimir Jiranek and (formerly) Prof. Terry Lee for their guidance during this work and the preparation of this thesis. Dr Miguel de Barros Lopes and Ms Anna Martens for their essential roles in the collaborative development of a PCR method for yeast identification. Dr Leigh Francis and Mr Phil Spillman for guidance on the conduct of sensory analysis and for advice on statistics. Mr Holger Gockowiak and Mr Jeff Eglinton for guidance and advice on just about everything microbiological. Ms Mariola Kwiatkowski and Dr Leigh Francis for programming the Cobas Fara autoanalyser for glucose and fructose analysis. Mr Alan Pollnitz for setting up a GC/IVIS assay for wine volatiles, for running the samples, and for tuition on chromatogram analysis. Dr Paul Monk and Lallemand Pty Ltd for the provision of yeast isolates, wine bottles and cross-flow filtration services Mr Jeff Cohn, Ms Marty Bannister and staff of Vinquiry, California for yeast isolation. Petaluma Ltd for providing grape juice. Dr Peter Langridge for the use of laboratory facilities during the PCR work. The AWRI Analytical Service for the use of their facilities, and in particular Mr Adrian Coulter for HPLC analysis. The staff and postgraduate students of the AWRI who participated as sensory judges, for their considerable efforts and enthusiasm. SìTY QF 1 Ghapter 1 l¿lr INTRODUCTION AND AIMS OF THIS STUDY An understanding of the microbiology of wine production originated with the pioneering . studies of Louis Pasteur over 130 years ago (see Amerine et aL.l98O), which showed that yeasts were responsible for the fermentation of grape juice into wine. Since Pasteur's time, the microbiology of winemaking has been studied extensively, revealing it to be a complex interaction of yeasts and bacteria. During fermentation, yeasts utilise the constituents of grape juice, principally converting sugars into ethancl and carbon dioxide, and generating an array of metabolic products which determine the chemical composition and sensory attributes
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