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Effect of Low Temperature Fermentation and Nitrogen Content on Wine Yeast Metabolism

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Effect of Low Temperature Fermentation and Nitrogen Content on Wine Yeast Metabolism UNIVERSITAT ROVIRA i VIRGILI DEPARTAMENT DE BIOQUÈMICA i BIOTECNOLOGIA FACULTAT D‘ENOLOGIA EFFECT OF LOW TEMPERATURE FERMENTATION AND NITROGEN CONTENT ON WINE YEAST METABOLISM Memoria presentada per GEMMA BELTRAN i CASELLAS per optar al grau de Doctora per la Universitat Rovira i Virgili sota la direcció del Dr. José Manuel Guillamón i el Dr. Nicolas Rozès Tarragona, 2005 UNIVERSITAT ROVIRAI VIRGILI Dept.Bioquímica i Biotecnologia FACULTAT D‘ENOLOGIA C/Marcel‡lí Domingo s/n 43007 Tarragona Tel. 34-977558442 / 34-977558280 Fax. 34-977558232 E-mail: [email protected] [email protected] Els sotasignants, Dr José Manuel Guillamón Navarro, Professor Titular d‘Universitat de Nutrició i Bromatologia del Departament de Bioquímica i Biotecnologia de la Universitat Rovira i Virgili, i El Dr Nicolas Rozès, Professor Titular d‘Escola Universitària de Nutrició i Bromatologia del Departament de Bioquímica i Biotecnologia de la Universitat Rovira i Virgili FAN CONSTAR, Que el present treball, amb títol ” Effect of low temperature and nitrogen content on wine yeast metabolism‘ que presenta la Srta Gemma Beltran Casellas , per optar al Grau de Doctora per la Universitat Rovira i Virgili ha estat realitzat sota la nostra direcció, i que tots els resultats obtinguts són fruit dels experiments duts a terme per l‘esmentada doctoranda. I perquè se‘n prengui coneixement i tingui els efectes que correspongui, signem aquesta certificació. Dr. José Manuel Guillamón Dr. Nicolas Rozès Tarragona, 23 de desembre de 2004 Als meus pares A l‘Adrià AGRAÎMENTS Voldria agrair a totes les persones que d‘una manera o altra han contribuït a que aquesta tesi s‘hagi dut a terme, i que m‘han mostrat suport durant tots aquests anys. En primer lloc als meus directors de tesi, Dr. José Manuel Guillamón i Dr. Nicolas Rozès, sense els quals no hauria estat possible la realització d‘aquest treball. Als dos voldria agrair la seva dedicació i suport, així com haver-me transmès el seus coneixements però sobretot el seu entusiasme, característica indispensable en qualsevol investigador i que intentaré guardar sempre amb mi. Al Dr. Albert Mas, per confiar en mi des d‘un primer moment, i oferir-me la possibilitat d‘incorporar-me al seu grup de treball, i d‘aquesta manera permetre‘m descobrir el món dels llevats i la fermentació alcohòlica, tant desconegut per a mi en un principi, però que poc a poc ha arribat a captivar-me. A tots membres del tribunal de la tesi, per acceptar formar-ne part del mateix, així com a Jean Luc Parrou i Paolo Guidicci, per acceptar fer l‘informe d‘avaluació de la tesi. És realment un honor i un plaer per a mi. Moltes gràcies. A tots els professors i persones del Departament de Bioquímica i Biotecnologia, així com als de la Facultat d‘Enologia que m‘han donat suport i m‘han ajudat en que ha fet falta durant tots aquest anys. A les secretaries del departament, Mireia i Cristina, per la seva eficiència i disposició, en especial aquests últims mesos, i també a la Montse Poblet, Isabel Baiges i Santiago Moreno, que m‘han aconsellat i ajudat a obtenir el que he necessitat per la realització dels experiments. A tots els meus companys i persones que he tingut a prop durant aquest temps. En especial voldria agrair la seva amistat a la Xus i la Maite (mis angeles) i com no, a la Sumpta, companyes i amigues, que han estat sempre al meu costat, tant dins com fora del laboratori, gràcies per ser-hi, pels savis consells i per tants bons moments, que espero continuin durant molt de temps. A la resta de companys del laboratori, amb els que he compartit poiata, tertúlies al pigsa, cine, i molts altres moments, gràcies a tots, tant als d‘abans (la vieja guardia), ja doctors: Miriam, Josep, Ramon, Cristina, Pep, Judit,... que hem van fer sentir molt ben acollida des d‘un primer moment i em van transmetre la seva experiència com a investigadors, com els d‘ara: Núria, Rosanna, Mireia, Angel, Marian, Zoel, Roser, Carmen, els analítics Marc, Pili i Montse, i tots els que han estat de pas, per fer del petit convent del Ramon y Cajal nº70 quelcom més que un simple lloc de treball. A Emilia Matallana per acollir-me en el seu laboratori del IATA a València durant una breu estada i permetre‘m realitzar i aprendre la tècnica de Northerns-blot. Gràcies en especial a la Geno, la meva mestra durant 2 setmanes, amb qui va ser un plaer treballar i conversar, i a la resta de companys, Agustín, Marcel‡lí, Roberto, Puri, Elena i Aurora, que, tot i que per poc temps, em van fer sentir com a casa. Je voudrais remercier à Jean Maire François pour m‘avoir accueilli dans son labo et me permettre réaliser et apprendre l‘analyse de puces de DNA, aussi lui remercier sa confiance en moi et m‘avoir donné l‘opportunité de travailler dans son équipe pendant une année. C‘était pour moi un vrai plaisir travailler avec Jean Luc, un modèle à suivre au niveau scientifique et aussi personnel, je voudrais lui remercier sa patience et son bon encadrement, et m‘avoir faire connaître la tréhalose chez les levures. Au personnel de la plate-forme, Véronique, Adilia, Sergueï et Delphine, qui m‘ont aidé avec l‘analyse et interprétation des puces, merci pour votre aide et patience. Je voudrais aussi remercier au reste de copains du labo: Marie-Ange, Matthieu, Olivier, Marlene, Helène C., Helène M., Lidwine, Celine, Fadi, et tout l‘équipe JMF en général, pour son accueil, aide et conseils quand il a fallu. Y una mención especial a la parte hispana de Toulouse (Filipa, Angelica, Carles, Juan Carlos, Maria, Maite y Xus) por su apoyo y sus risas, y con los que la estancia en Francia fue muy agradable. Y como no, a Sandra, Juan y Santi, amigos ya para siempre, por hacernos sentir como una gran familia y demostrarnos que aún existe gente con el corazón tan grande como ellos. Je voudrais bien evidentment remercier à Tristan, pour une très bonne concurrance scientiphique, pour son soutien et aide, même en la distance. Gràcies a tots els que d‘alguna manera m‘heu ajudat aquest últims mesos en la escriptura i realització de la tesi. Al Xavi, per ajudar-me amb el disseny i elaboració de la portada, així com pel seu suport i amistat durant tot aquest temps; a la Cristina, Tristan i els companys del servei lingüístic per ajudar-me i aconsellar-me en bona part de les correccions lingüístiques de la tesi; a la Xus, la Maite, el Josep, la Rosanna per ajudar-me a resoldre aspectes i dubtes tant tècnics com científics, a la Susanna per la seva paciència, consells i aliments i a la Sumpta, per escoltar-me i ajudar-me a desconnectar i desestressar-me quan m‘ha fet falta. Als meus amics de sempre, per ser-hi i estar sempre al meu costat. I per acabar voldria agrair de manera molt especial als meus pares, àvia i germans, Susanna, Mari i Xavier, així com a la resta de família, per haver-me donat suport incondicional en tot moment i haver acceptat i entès les meves inquietuds. Al petit Adrià, que sols la seva presència transmet alegria i il‡lusió, i m‘ha ajudat a superar de manera més amena aquests difícils mesos d‘escriptura. Gràcies a tots i molta sort. INDEX OBJECTIVES & OUTLINE OF THESIS__________________________________1 INTRODUCTION_____________________________________________________7 CHAPTER 1_________________________________________________________63 —Effect of fermentation temperature and culture media on the lipid composition and volatile compounds in wine“ CHAPTER 2_________________________________________________________87 —Integration of transcriptomic and metabolomic analyses for understanding the global responses of low temperature winemaking fermentations.“ CHAPTER 3________________________________________________________123 —Nitrogen Catabolite Repression in Saccharomyces cerevisiae during wine fermentation.“ CHAPTER 4________________________________________________________139 —Influence of the timing of nitrogen additions during wine fermentations on the fermentation kinetics and nitrogen consumption.“ CHAPTER 5________________________________________________________161 —Effect of low temperature fermentation on yeast nitrogen metabolism.“ CONCLUSIONS AND PERSPECTIVES________________________________173 ANNEXES__________________________________________________________187 Annex1: Materials and Methods 189 Annex2: Supplementary data of yeast functional genomics at low temperature 221 Annex3: —Fermentaciones a bajas temperaturas“ 235 OBJECTIVES & OUTLINE OF THESIS Objectives OBJECTIVES The winemaking industry is facing continuous challenges in the present global market. Those involving the quality and reproducibility of the product, which depend heavily on yeast metabolism, are of particular interest. Finding ways to improve fermentation performance, wine flavour and other sensory qualities are becoming important objectives for wine producers. When I joined the research group into Enological Biotechnology at the Department of Biochemistry and Biotechnology (Faculty of Enology, Rovira i Virgili University, Tarragona, SPAIN), the research objectives focused on the characterization of yeast metabolism in low temperature fermentations. Low temperature fermentations have interesting applications in the wine industry, especially in the production of wines with more pronounced aromatic profile. Our research group had already collaborated with the cellar Torres S.A (Spain) and the yeast producers Lallemand S.A. (Toulouse, France) to further our knowledge of low temperature fermentations, particularly in terms of changes in yeast metabolism and improvements in fermentation performance. This work was supported by grants from the Spanish government (FEDER 2FD97- 1854, AGL2000-0205-P4-0) and the France-Spanish Picasso Integrated Project (Acciones Integradas HF2001-0015). In modern winemaking, consumer olfactory profiling is becoming increasingly important as a guide for production decisions. Today the wine industry manages to control the fermentation temperature effectively, and a low temperature is becoming more common for white and rosé fermentation. Wines produced at low temperatures (10-15ºC) are known to develop certain characteristics of taste and aroma. A first hypothesis was that these improved sensorial properties were generally due to a greater retention of volatile substances at low temperature.
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