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The Effect of Sulfur Dioxide Addition and Alternative THE EFFECT OF SULFUR DIOXIDE ADDITION AND ALTERNATIVE FERMENTATION TECHNIQUES ON THE MICROBIAL COMMUNITIES AND SENSORY PROFILES OF WINE By Sydney Christian Morgan A THESIS SUBMITTED IN PARTIAL FULFILLMENT OF THE REQUIREMENTS FOR THE DEGREE OF DOCTOR OF PHILOSOPHY in THE COLLEGE OF GRADUATE STUDIES (Biology) THE UNIVERSITY OF BRITISH COLUMBIA (Okanagan) August 2019 © Sydney Christian Morgan, 2019 The following individuals certify that they have read, and recommend to the College of Graduate Studies for acceptance, a dissertation entitled: THE EFFECT OF SULFUR DIOXIDE ADDITION AND ALTERNATIVE FERMENTATION TECHNIQUES ON THE MICROBIAL COMMUNITIES AND SENSORY PROFILES OF WINE submitted by Sydney Christian Morgan in partial fulfillment of the requirements of the degree of Doctor of Philosophy. Dr. Daniel Durall, Department of Biology, University of British Columbia, Okanagan Supervisor Dr. Michael Deyholos, Department of Biology, University of British Columbia, Okanagan Supervisory Committee Member Dr. John Klironomos, Department of Biology, University of British Columbia, Okanagan Supervisory Committee Member Dr. Richard Plunkett, Department of Biology, University of British Columbia, Okanagan Supervisory Committee Member Dr. David Scott, Department of Earth and Environmental Sciences, University of British Columbia, Okanagan University Examiner Dr. Thomas Henick-Kling, School of Food Science, Washington State University External Examiner ii Abstract Modern winemaking often involves the addition of sulfur dioxide (SO2), to remove potential spoilage microbes from the grape juice, and the addition of commercial wine yeasts, to ensure a successful and timely completion of alcoholic fermentation. However, consumer demand is shifting towards low-SO2 wines, and wines fermented by a collection of indigenous yeasts and bacteria. Many winemakers wish to produce these wines for their customers, but there is a current lack of research into the relative risks and rewards of these winemaking strategies. We investigated these topics, and found that both SO2 addition and inoculation technique can significantly influence the fungal and bacterial communities and the sensory profiles of commercially-produced wines. Fermentations conducted without SO2 were found to contain more vineyard-derived microorganisms and produced more fruity characters, while the fermentations conducted with SO2 produced wines with less-desirable characteristics. Pied de cuve inoculation had a limited effect on the wine microbial communities and was inhibited by the addition of SO2; when it persisted, it resulted in slower fermentations, and produced wines with more sweetness and body as compared to wines that did not receive the pied de cuve inoculum. At one commercial winery, we identified a highly diverse indigenous population of Saccharomyces uvarum, a yeast that is related to the main winemaking yeast, Saccharomyces cerevisiae. S. uvarum was able to compete with a commercial S. cerevisiae strain in controlled fermentations, and produced wines with a unique composition of aroma-active volatile compounds, showing its potential to be used as an indigenous yeast inoculum for locally-made wines. In summary, we iii showed that both SO2 addition and fermentation technique (uninoculated or pied de cuve inoculation) can alter the microbial communities and sensory profiles of commercial and laboratory-made wines, and reported an indigenous yeast population in the Okanagan Valley winemaking region. iv Lay summary Yeasts and bacteria make wine by converting grape sugars to ethanol, and also produce hundreds of compounds that contribute to the aroma and flavour of wine. Today, most winemakers add commercial yeasts and bacteria to their grape must to ensure a successful fermentation and a consistent, predictable flavour profile. However, some winemakers are looking for minimal intervention approaches to winemaking, which can include adding fewer chemicals such as sulfur dioxide (SO2), and letting native vineyard and winery microbes conduct fermentation. We found that wines made with different levels of SO2 contained different communities of microbes. We also found that low-SO2 wines contain more vineyard-derived yeasts and bacteria, and produce wines with more fruity aromas and flavours. In summary, we found that these minimal-intervention techniques can create wines with unique sensory profiles, and can be used successfully by winemakers to produce quality wines with more regional character. v Preface A version of Chapter 2 has been published: Morgan, S.C., Scholl, C.M., Benson, N.L., Stone, M.L., Durall, D.M., 2017. Sulfur dioxide addition at crush alters Saccharomyces cerevisiae strain composition in spontaneous fermentations at two Canadian wineries. Int. J. Food. Microbiol. 244, 96–102. DOI:10.1016/j.ijfoodmicro.2016.12.025. A version of Chapter 3 has been published: Morgan, S.C., Tantikachornkiat, M., Scholl, C.M., Benson, N.L., Cliff, M.A., Durall, D.M., 2019. The effect of sulfur dioxide addition at crush on the fungal and bacterial communities and the sensory attributes of Pinot gris wines. Int. J. Food Microbiol. 290, 1–14. DOI:10.1016/j.ijfoodmicro.2018.09.020. A version of Chapter 4 has been published: Morgan, S.C., Haggerty, J.J., Johnston, B., Jiranek, V., Durall, D.M., 2019. Response to sulfur dioxide addition by two commercial Saccharomyces cerevisiae strains. Fermentation. 5, 1-20. DOI:10.3390/fermentation5030069. A version of Chapter 5 has been published: Morgan, S.C., McCarthy, G.C., Watters, B.S., Tantikachornkiat, M., Zigg, I., Cliff, M.A., Durall, D.M., 2019. Effect of sulfite addition and pied de cuve inoculation on the microbial communities and sensory profiles of Chardonnay wines: dominance of indigenous Saccharomyces uvarum at a commercial winery. FEMS Yeast Res. 19, foz049. DOI: 10.1093/femsyr/foz049. The winery experiments described in Chapters 2, 3, and 5 were designed in consultation with my supervisor, Dr. Daniel Durall, and realized with assistance from vi the winemakers Darryl Brooker, Taylor Whelan, Alexandra Haselich, Grant Stanley, and Matthew Fortuna. Sample collection was performed by myself, Natasha Benson (honours student), Brittany Watters (honours student), and Chrystal Scholl (laboratory technician). Chemical analysis was performed by myself, Natasha Benson, and Morgan Stone (laboratory technician). S. cerevisiae and S. uvarum strain typing was performed by myself, Natasha Benson, Ieva Zigg (work-study student), Garrett McCarthy (graduate student) and Chrystal Scholl. Library preparation for Illumina sequencing was performed by myself and Mansak Tantikachornkiat, a fellow graduate student. Illumina sequencing was performed by Dan New and fellow technicians at IBEST at the University of Idaho. Illumina sequencing results were analyzed by myself, with assistance from Mansak Tantikachornkiat. Technical assistance and guidance was provided by Marissa Neuner (graduate student) and Stacey Sakakibara (visiting researcher). Sensory evaluations of wine were performed in collaboration with the Summerland Research and Development Centre (part of Agriculture and Agri-Food Canada), under the supervision of Dr. Margaret Cliff and with assistance from Kareen Stanich (sensory laboratory technician). I performed all statistical analysis and data visualization, and wrote all three manuscripts for publication. The laboratory experiments described in Chapters 4 and 6 were conducted at the University of Adelaide in Adelaide, Australia, and were designed in consultation with Dr. Daniel Durall, as well as Dr. Vladimir Jiranek and Dr. Tommaso Watson from the University of Adelaide. I performed all experiments, including data collection and analysis, statistical analysis and data visualization, and the writing of vii both manuscripts. Chemical analysis of the wine was performed under the invaluable guidance of Dr. Jade Haggerty. I also received training, guidance, and technical assistance from the following members of the Wine Microbiology Group at the University of Adelaide: Nicholas Van Holst, Dr. Joanna Sundstrom, Louise Bartle, Dr. Ee Lin Tek, Dr. Krista Sumby, Liang Chen, Tom Lang, Chen Liang, and Dr. Jennifer Gardner. viii Table of contents Abstract ..................................................................................................................... iii Lay summary ............................................................................................................. v Preface ...................................................................................................................... vi Table of contents ....................................................................................................... ix List of tables ............................................................................................................ xiv List of figures ........................................................................................................... xix Acknowledgements ................................................................................................ xxiv Dedication ............................................................................................................. xxvii 1. Introduction ............................................................................................................ 1 1.1 History of winemaking ...................................................................................... 1 1.2 Yeasts involved in winemaking ........................................................................
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