Hanseniaspora vineae: caracterización

y su uso en la vinificación
TESIS DOCTORAL

Valentina Martı́n Russo

Programa de Posgrado de Doctorado en Biologı́a

Facultad de Ciencias, Facultat de Enologı́a
Universidad de la República - Universitat Rovira i Virgili

Montevideo – Uruguay
Noviembre de 2016
Hanseniaspora vineae: caracterización
y su uso en la vinificación
TESIS DOCTORAL

Valentina Martı́n Russo

Tesis de Doctorado presentada al Programa de

Posgrado de Doctorado en Biologı́a, Facultad de
Ciencias de la Universidad de la República, como
parte de los requisitos necesarios para la obtención
del tı́tulo de Doctor en Doctorado en Biologı́a.

Directores:
D.Sc. Prof. Francisco Carrau
D.Sc. Prof. Albert Mas

Montevideo – Uruguay
Noviembre de 2016
Martı́n Russo, Valentina
Hanseniaspora vineae: caracterización y su uso en la
vinificación / Valentina Martı́n Russo. - Montevideo:
Universidad de la República, Universitat Rovira i Virgili,
Facultad de Ciencias, Facultat de Enologı́a, 2016.
XXXII, 214 p.: il.; 29, 7cm.
Directores:
Francisco Carrau
Albert Mas
Tesis de Doctorado – Universidad de la República,
Programa de Doctorado en Biologı́a, 2016.
Referencias bibliográficas: p. 151 – 180.
1. Hanseniaspora vineae, 2. no-Saccharomyces,
3. levaduras nativas, 4. caracterización, 5. fermentación.
I. Carrau, Francisco, Mas, Albert, . II. Universidad
de la República, Programa de Posgrado de Doctorado en
Biologı́a. III. Tı́tulo.
INTEGRANTES DEL TRIBUNAL DE DEFENSA DE TESIS

D.Sc. Prof. Patrick Moyna

D.Sc. Prof. Silvana Vero

D.Sc. Prof. Inés Ponce de León

Montevideo – Uruguay
Noviembre de 2016

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 A mis padres Edison y Rossana
por su apoyo incondicional.

ix
x
Agradecimentos

Durante la elaboración de éste trabajo conté con la colaboración y el apoyo

de muchas personas. En primer lugar le agradezco a Emiliano su apoyo y com-
pañı́a incondicional. A mis padres y mis hermanos por su soporte y contención
durante el proceso de elaboración de este Tesis.
A mis tutores, los Doctores Francisco Carrau y Albert Mas quienes me guia-
ron durante el desarrollo de esta Tesis. A mis compañeros del laboratorio de
Enologı́a, Karina, Laura, los Eduardos Boido y Dellacasa, Gabriel, Bonini,
Noe, Flor, Anita, Gio y Manuel, fundamentales a lo largo de todo el proceso.
A los integrantes y responsables del grupo de investigación de Biotecnologı́a
Enológica de la Facultat de Enologı́a de la Universitat Rovira i Virgili, espe-
cialmente a Jessica, Flor y Olga.
A Gastón Ares de área Sensometrı́a y Ciencia del consumidor y a los distintos
integrantes del Departamento de Ciencia y Tecnologı́a de los Alimentos de Fa-
cultad de Quı́mica por su colaboración en las distintas etapas de elaboración
de mi trabajo.
A Carina Gaggero del Instituto de Investigaciones Biológicas Clemente Esta-
ble.
A Pablo Aguilar, Valentina Salzman y Facundo Giorelo del Instituto Pasteur
de Montevideo.
Y finalmente a las distintas instituciones que mediante su financiamiento co-
laboraron en la realización de este Tesis, PEDECIBA, ANII, CSIC y CAP.

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 “No es la especie más fuerte, ni
la mas inteligente la que
sobrevive, sino la que mejor se
adapte al cambio”

Charles Darwin

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 RESUMEN

Durante la elaboración de un vino ocurren complejas interacciones micro-

bianas entre levaduras Saccharomyces y no-Saccharomyces, y entre levaduras
y bacterias. En los viñedos las levaduras y bacterias se ubican mayoritaria-
mente sobre los granos de uva, e ingresan a la bodega en época de cosecha
como flora nativa. Esta flora está compuesta principalmente por levaduras
no-Saccharomyces y en un muy bajo porcentaje (menor al 1 %) de levaduras
Saccharomyces, levadura tı́pica de la fermentación alcohólica y la vinificación.
Entre las no-Saccharomyces las que se encuentran mas comúnmente son leva-
duras de los géneros Hanseniaspora, Candida, Pichia y Metschnikowia.
La fermentación alcohólica es un proceso durante el cual, por acción de las
levaduras, los azúcares presentes en el mosto se transforman principalmente
en etanol y dióxido de carbono. Sin embargo, al mismo tiempo se forma otro
conjunto de compuestos producto del metabolismo secundario de las levaduras,
compuestos aromáticos y otros asociados al color y a la sensación del vino
en boca, como por ejemplo, el glicerol. Por ello es importante comprender el
comportamiento de distintas levaduras durante la fermentación y su influencia
sobre estos productos secundarios que en definitiva serán el diferencial a la
hora de elaborar un vino de calidad.
Conocer dicho comportamiento puede ser también útil para intentar resol-
ver determinados problemas que puedan suscitarse, tal como pasa actualmente
con la alta concentración de azúcares en las uvas en determinadas regiones, que
genera la obtención de vinos de elevada graduación alcohólica, algo no deseado
en vinos de alta gama.
En este trabajo de tesis se realiza la caracterización de once cepas distintas
de levaduras de la especie H. vineae en cuanto a sus aptitudes enológicas. A
partir de ésta caracterización, dos de ellas fueron seleccionadas, una, por su
capacidad para la producción de fenı́lpropanoides de interés aromático en altas
concentraciones (M12 196F), fue utilizada en ensayos que tenı́an como fin com-
prender con mayor profundidad los mecanismos y vı́as de producción de esos
compuestos y los factores que la afectan. La otra para realizar fermentaciones
en mostos naturales a escala semi-industrial (T02 05F).

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 Respecto de estos estudios, los resultados fueron los siguientes:

- Se determinó el efecto negativo del agregado de fosfato diamónico (DAP)

sobre la producción, por parte de la cepa M12 196F H. vineae, de los alcoho-
les bencı́lico (BAL) y β-feniletı́lico (asociados a descriptores floral y dulce el
primero y floral, rosa y miel el segundo) y sus ésteres.
- A su vez estos compuestos fueron producidos en 1-2 órdenes de magnitud
superior por H.vineae frente a la S. cerevisiae utilizada como control bajo las
mismas condiciones de fermentación.
- Cambios en la composición de medio en cuanto a su contenido de Phe, Trp y
ρ-ABA, que mostraron un incremento en la producción de BAL, mientras que
la Tyr y el amonio disminuyeron los niveles producidos de éste alcohol.
- Cambios en los diferentes componentes nitrogenados asimilables por las le-
vaduras pueden aumentar o disminuir hasta unas 17 veces los niveles de BAL
producidos por H. vineae. Al duplicar los niveles de Phe en el medio, aumenta
significativamente la formación de BAL en H. vineae, comportamiento opuesto
al que se pudo ver en S. cerevisiae.
- El comportamiento observado en la producción de BAL sugiere la posibilidad
de que H. vineae utilice dos vı́as distintas para su producción.
- Se obtuvo un vino a escala semi-industrial de la variedad de uva blanca Ma-
cabeo, a partir de la cepa de H. vineae T02 05F en la región de Tarragona.
- Se demostró que si bien esta levadura no finaliza por sı́ misma la fermen-
tación, logra predominar en ella prácticamente hasta la mitad del proceso,
tiempo suficiente para lograr un producto diferente al obtenido a partir de S.
cerevisiae.
- En el análisis de compuestos aromáticos, se vio una mayor producción de
aquellos asociados a descriptores florales y frutados en el vino producido con
H. vineae.
- En la prueba sensorial llevada a cabo por un panel entrenado logró diferen-
ciarse notoriamente el vino inoculado con H. vineae de aquel fermentado con
S. cerevisiae (control), siendo el primero preferido por el panel, destacándose
por sus notas florales, lo cual coincide con lo obtenido en el análisis de com-
puestos aromáticos.

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 Los resultados obtenidos durante este trabajo de tesis permitieron ampliar
el conocimiento acerca del comportamiento de la especie H. vineae. Si bien
se observó diversidad entra las distintas cepas estudiadas, también se deter-
minaron caracterı́sticas compartidas entre todas las cepas analizadas, lo que
permite considerarlas como caracterı́sticas tı́picas de la especie, como son la
actividad enzimática positiva (proteasa y β-glucosidasa a pH6) y la alta pro-
ducción por parte de ésta especie de compuestos aromáticos deseables durante
la fermentacion, produciéndolos hasta en 50 órdenes más que la levadura de
fermentación convencional S. cerevisiae. Su utilización a escala semi-industrial
permitió demostrar que es viable su empleo para vinificación de vinos comer-
ciales logrando éstos diferenciarse de vinos fermentados mediante levaduras
tradicionales.

Palabras claves:
Hanseniaspora vineae, no-Saccharomyces, levaduras nativas,
caracterización, fermentación.

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Tabla de contenidos

Lista de figuras xix

Lista de tablas xxv

Lista de sı́mbolos xxvii

Lista de siglas xxvii

1 Introducción 1
1.1 Introducción general, justificación y
objetivos . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
1.1.1 Justificación del trabajo . . . . . . . . . . . . . . . . . . 4

2 Marco teórico 7
2.1 Composición quı́mica de la uva . . . . . . . . . . . . . . . . . . 7
2.2 Productos de la fermentación alcohólica . . . . . . . . . . . . . . 10
2.2.1 Compuestos aromáticos del vino . . . . . . . . . . . . . . 11
2.3 Las levaduras y su rol durante la
fermentación . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 26
2.3.1 Generalidades de las levaduras . . . . . . . . . . . . . . . 26
2.3.2 Identificación de levaduras . . . . . . . . . . . . . . . . . 32
2.3.3 Metabolismo de los azúcares . . . . . . . . . . . . . . . . 37
2.3.4 Levaduras no-Saccharomyces . . . . . . . . . . . . . . . . 41
2.3.5 Dinámica poblacional durante la fermentación . . . . . . 56

3 Metodologı́a 63
3.1 Caracterización de levaduras . . . . . . . . . . . . . . . . . . . . 63
3.1.1 Aislamiento de microorganismos . . . . . . . . . . . . . . 63
3.1.2 Identificación molecular . . . . . . . . . . . . . . . . . . 64

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 3.1.3 Caracterización de actividad enzimática
extracelular . . . . . . . . . . . . . . . . . . . . . . . . . 65
3.1.4 Microvinificaciones . . . . . . . . . . . . . . . . . . . . . 67
3.1.5 Producción de glicerol . . . . . . . . . . . . . . . . . . . 69
3.1.6 Resistencia al SO2 . . . . . . . . . . . . . . . . . . . . . 69
3.1.7 Determinación de polisacáridos . . . . . . . . . . . . . . 69
3.1.8 Extracción e identificación de compuestos aromáticos . . 71
3.1.9 Evaluación sensorial . . . . . . . . . . . . . . . . . . . . 74
3.1.10 Determinación del tamaño del genoma y ploidı́a de H.
vineae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
3.2 Efecto de compuestos nitrogenados en la producción de bence-
noides y fenı́lpropanoides . . . . . . . . . . . . . . . . . . . . . . 77
3.2.1 Fermentaciones a escala de laboratorio . . . . . . . . . . 77
3.2.2 Extracción y análisis de compuestos aromáticos . . . . . 78
3.3 Vinificaciones a escala semi-industrial . . . . . . . . . . . . . . . 79
3.3.1 Cepas de levaduras . . . . . . . . . . . . . . . . . . . . . 79
3.3.2 Condiciones de fermentación . . . . . . . . . . . . . . . . 79
3.3.3 Seguimiento de las fermentaciones . . . . . . . . . . . . . 80
3.3.4 Identificación de levaduras . . . . . . . . . . . . . . . . . 80
3.3.5 Análisis quı́micos en WineScan . . . . . . . . . . . . . . 80
3.3.6 Análisis sensorial . . . . . . . . . . . . . . . . . . . . . . 80
3.3.7 Extracción y análisis de compuestos aromáticos . . . . . 81

4 Resultados y discusión 83
4.1 Caracterización de las levaduras . . . . . . . . . . . . . . . . . . 83
4.1.1 Identificación molecular . . . . . . . . . . . . . . . . . . 84
4.1.2 Caracterización enzimática . . . . . . . . . . . . . . . . . 85
4.1.3 Capacidad fermentativa . . . . . . . . . . . . . . . . . . 89
4.1.4 Producción de glicerol . . . . . . . . . . . . . . . . . . . 92
4.1.5 Resistencia al SO2 . . . . . . . . . . . . . . . . . . . . . 93
4.1.6 Liberación de polisacáridos durante la
fermentación alcohólica . . . . . . . . . . . . . . . . . . . 95
4.1.7 Composición aromática de las fermentaciones . . . . . . 97
4.1.8 Evaluación sensorial . . . . . . . . . . . . . . . . . . . . 110
4.1.9 Determinación del tamaño del genoma y ploidı́a de H.
vineae . . . . . . . . . . . . . . . . . . . . . . . . . . . . 111

xxx
 4.1.10 Conclusiones . . . . . . . . . . . . . . . . . . . . . . . . 113
4.2 Efecto del nitrógeno sobre la composición de fenı́lpropanoides . . 116
4.2.1 Diferencias en la producción de fenı́lpropanoides entre S.
cerevisiae y H. vineae . . . . . . . . . . . . . . . . . . . 116
4.2.2 Efecto del nitrógeno inorgánico sobre la sı́ntesis de
fenı́lpropanoides . . . . . . . . . . . . . . . . . . . . . . . 119
4.2.3 Efecto de los aminoácidos sobre los compuestos bence-
noides . . . . . . . . . . . . . . . . . . . . . . . . . . . . 121
4.2.4 Propuestas de vı́as metabólicas para la formación de
fenı́lpropanoides . . . . . . . . . . . . . . . . . . . . . . . 125
4.2.5 Conclusiones . . . . . . . . . . . . . . . . . . . . . . . . 130
4.3 Vinificaciones a nivel semi-industrial . . . . . . . . . . . . . . . 131
4.3.1 Seguimiento de las fermentaciones . . . . . . . . . . . . . 131
4.3.2 Determinaciones analı́ticas . . . . . . . . . . . . . . . . . 138
4.3.3 Composición de compuestos volátiles . . . . . . . . . . . 139
4.3.4 Análisis sensorial . . . . . . . . . . . . . . . . . . . . . . 142
4.3.5 Conclusiones . . . . . . . . . . . . . . . . . . . . . . . . 143

5 Consideraciones finales 145

Referencias bibliográficas 151

Glosario 180

Apéndices 181
Apéndice 1 Artı́culos publicados en el marco de éste trabajo de Tesis183

Anexos 213

xxxi
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