Yeasts are predominant in the ancient and complex process of winemaking. In spontaneous fermentations, there is a progressive growth pattern of indigenous yeasts, with the final stages invariably being dominated by the alcohol-tolerant strains of Saccharomyces cerevisiae. This species is universally known as the ‘wine yeast’ and is widely preferred for initiating wine fermentations. The primary role of wine yeast is to catalyze the rapid, complete and efficient conversion of grape sugars to ethanol, carbon dioxide and other minor, but important, metabolites without the development of off-flavours. However, due to the demanding nature of modern winemaking practices and sophisticated wine markets, there is an ever-growing quest for specialized wine yeast strains possessing a wide range of optimized, improved or novel oenological properties. This review highlights the wealth of untapped indigenous yeasts with oenological potential, the complexity of wine yeasts’ genetic features and the genetic techniques often used in strain development. The current status of genetically improved wine yeasts and potential targets for further strain development are outlined. In light of the limited knowledge of industrial wine yeasts’ complex genomes and the daunting challenges to comply with strict statutory regulations and consumer demands regarding the future use of genetically modified strains, this review cautions against unrealistic expectations over the short term. However, the staggering potential advantages of improved wine yeasts to both the winemaker and consumer in the third millennium are pointed out. Copyright # 2000 John Wiley & Sons, Ltd.

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Tailoring wine yeast for the new millennium: novel approaches to the ancient art of winemaking

Wine is a highly complex mixture of compounds which largely define its appearance, aroma, flavour and mouth-feel properties. The compounds responsible for those attributes have been derived in turn from three major sources, viz. grapes, microbes and, when used, wood (most commonly, oak). The grape-derived compounds provide varietal distinction in addition to giving wine its basic structure. Thus, the floral monoterpenes largely define Muscat-related wines and the fruity volatile thiols define Sauvignon-related wines; the grape acids and tannins, together with alcohol, contribute the palate and mouth-feel properties. Yeast fermentation of sugars not only produces ethanol and carbon dioxide but a range of minor but sensorially important volatile metabolites which gives wine its vinous character. These volatile metabolites, which comprise esters, higher alcohols, carbonyls, volatile fatty acids and sulfur compounds, are derived from sugar and amino acid metabolism. The malolactic fermentation, when needed, not only provides deacidification, but can enhance the flavour profile.

The aroma and flavour profile of wine is the result of an almost infinite number of variations in production, whether in the vineyard or the winery. In addition to the obvious, such as the grapes selected, the winemaker employs a variety of techniques and tools to produce wines with specific flavour profiles. One of these tools is the choice of microorganism to conduct fermentation. During alcoholic fermentation, the wine yeast Saccharomyces cerevisiae brings forth the major changes between grape must and wine: modifying aroma, flavour, mouth-feel, colour and chemical complexity. The wine bacterium Oenococcus oeni adds its contribution to wines that undergo malolactic fermentation. Thus flavour-active yeasts and bacterial strains can produce desirable sensory results by helping to extract compounds from the solids in grape must, by modifying grape-derived molecules and by producing flavour-active metabolites. This article reviews some of the most important flavour compounds found in wine, and their microbiological origin.

Yeast and bacterial modulation of wine aroma and flavour

The most mysterious aspect of wine is the endless variety of flavours that stem from a complex, completely non-linear system of interactions among many hundreds of compounds. In its widest sense, wine flavour refers to the overall impression of both aroma and taste components. Aroma is usually associated with odorous, volatile compounds; the bouquet of wine refers to the more complex flavour compounds which evolve as a result of fermentation, elevage and ageing. With the exception of terpenes in the aromatic grape varieties and alkoxypyrazines in the herbaceous cultivars, perceived flavour is the result of absolute amounts and specific ratios of many of these interactive compounds, rather than being attributable to a single "impact" compound. Without underestimating the complexity of these interactive effects or negating the definitive role played by the accumulated secondary grape metabolites in the varietal character of wine, this review will focus mainly on the contribution of yeast fermentation to the sensorial quality of the final product. Yeast and fermentation conditions are claimed to be the most important factors influencing the flavours in wine. Both spontaneous and inoculated wine fermentations are affected by the diversity of yeasts associated with the vineyard and winery. During the primary alcoholic fermentation of sugar, the wine yeast, Saccharomyces cerevisiae, together with other indigenous non-Saccharomyces species, produce ethanol, carbon dioxide and a number of by-products. Of these yeast-derived metabolites, the alcohols, acetates and C4-C8 1tfatty acid ethyl esters are found in the highest concentration in wine. While the volatile metabolites contribute to the fermentation bouquet ubiquitous to all young wines, the production levels of these by-products are variable and yeast strain specific. Therefore, this article also highlights the importance of untapping the hidden wealth of indigenous yeast species present on grapes, and the selection and genetic development of yeast starter culture strains with improved flavour profiles. In the future, some winemakers may prefer to use mixtures of indigenous yeast species and tailored S. cerevisiae strains as starter cultures to reflect the biodiversity and stylistic distinctiveness of a given region. This will help winemakers to fullfil the consumer's demand for individual wines with intact local character and to ensure the survival of wine's most enthralling aspect - its endless variety.

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Yeast and its Importance to Wine Aroma - A Review

Probiotics are live microorganisms that provide health benefits to the host when ingested in adequate amounts. The strains most frequently used as probiotics include lactic acid bacteria and bifidobacteria. Probiotics have demonstrated significant potential as therapeutic options for a variety of diseases, but the mechanisms responsible for these effects have not been fully elucidated yet. Several important mechanisms underlying the antagonistic effects of probiotics on various microorganisms include the following: modification of the gut microbiota, competitive adherence to the mucosa and epithelium, strengthening of the gut epithelial barrier and modulation of the immune system to convey an advantage to the host. Accumulating evidence demonstrates that probiotics communicate with the host by pattern recognition receptors, such as toll-like receptors and nucleotide-binding oligomerization domain-containing protein-like receptors, which modulate key signaling pathways, such as nuclear factor-ĸB and mitogen-activated protein kinase, to enhance or suppress activation and influence downstream pathways. This recognition is crucial for eliciting measured antimicrobial responses with minimal inflammatory tissue damage. A clear understanding of these mechanisms will allow for appropriate probiotic strain selection for specific applications and may uncover novel probiotic functions. The goal of this systematic review was to explore probiotic modes of action focusing on how gut microbes influence the host.

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Probiotic Mechanisms of Action

This review discusses the impacts of nitrogen addition in the vineyard and winery, and establishes the effects that nitrogen has on grape berry and wine composition and the sensory attributes of wine. Nitrogen is the most abundant soil-derived macronutrient in a grapevine, and plays a major role in many of the biological functions and processes of both grapevine and fermentative microorganisms. Manipulation of grapevine nitrogen nutrition has the potential to influence quality components in the grape and, ultimately, the wine. In addition, fermentation kinetics and formation of flavour-active metabolites are also affected by the nitrogen status of the must, which can be further manipulated by addition of nitrogen in the winery. The only consistent effect of nitrogen application in the vineyard on grape berry quality components is an increase in the concentration of the major nitrogenous compounds, such as total nitrogen, total amino acids, arginine, proline and ammonium, and consequently yeast-assimilable nitrogen (YAN). Both the form and amount of YAN have significant implications for wine quality. Low must YAN leads to low yeast populations and poor fermentation vigour, increased risk of sluggish/stuck/slow fermentations, increased production of undesirable thiols (e.g. hydrogen sulfide) and higher alcohols, and low production of esters and long chain volatile fatty acids. High must YAN leads to increased biomass and higher maximum heat output due to greater fermentation vigour, and increased formation of ethyl acetate, acetic acid and volatile acidity. Increased concentrations of haze-causing proteins, urea and ethyl carbamate and biogenic amines are also associated with high YAN musts. The risk of microbial instability, potential taint from Botrytis-infected fruit and possibly atypical ageing character is also increased. Intermediate must YAN favours the best balance between desirable and undesirable chemical and sensory wine attributes. ‘Macro tuning’, of berry nitrogen status can be achieved in the vineyard, given genetic constraints, but the final ‘micro tuning’ can be more readily achieved in the winery by the use of nitrogen supplements, such as diammonium phosphate (DAP) and the choice of fermentation conditions. This point highlights the need to monitor nitrogen not only in the vineyard but also in the must immediately before fermentation, so that appropriate additions can be made when required. Overall, optimisation of vineyard and fermentation nitrogen can contribute to quality factors in wine and hence affect its value. However, a better understanding of the effect of nitrogen on grape secondary metabolites and different types of nitrogen sources on yeast flavour metabolism and wine sensory properties is still required.

Implications of nitrogen nutrition for grapes, fermentation and wine

Nine Saccharomyces cerevisiae and four Saccharomyces bayanus strains used in the production of Trebbiano wine were examined. The aim of this study is to evaluate the different abilities of various yeast strains in wine-making. The possibility of yeast discrimination on the basis of their volatile production is another possibility. Wine chemical analyses showed statistically significant differences depending on the yeast strain used. Some compounds such as 2-phenylethanol, 2-phenethyl acetate, ethyl lactate, 3-ethoxypropanol, and, to a lesser extent, diethyl succinate and propionic acid characterized examined Saccharomyces bayanus yeasts. Moreover, these strains did not produce any undesirable compounds, such as acetic acid and sulfur anhydride. For these reasons and because they synthesized malic acid, they could be more suitable for white wine production. The other yeasts showed great differences, which are difficult to correlate with the strain. However, some strains had peculiar characteristics, such as an uncommonly high concentration of n-propanol and 3-ethoxypropanol.

Yeast influence on volatile composition of wines.

A hundred strains of Saccharomyces cerevisiae were examined for the ability to produce higher alcohols. In the strains tested the production of higher alcohols was found to be an individual strain ...

A biometric study of higher alcohol production in Saccharomyces cerevisiae.

Saccharomyces sensu stricto: systematics, genetic diversity and evolution.

A natural subgroup (that we refer to as Saccharomyces uvarum) was identified, within the heterogeneous species Saccharomyces bayanus. The typical electrophoretic karyotype, interfertility of hybrids between strains, distinctive sugar fermentation pattern, and uniform fermentation characteristics in must, indicated that this subgroup was not only highly homogeneous, but also clearly distinguishable from other species within the Saccharomyces sensu stricto group. Investigation of the S. bayanus type strain and other strains that have been classified as S. bayanus, confirmed the apparent lack of homogeneity and, in some cases, supported the hypothesis that they are natural hybrids.

Saccharomyces uvarum, a distinct group within Saccharomyces sensu stricto

Saccharomyces uvarum is proposed as a proper species within the complex Saccharomyces sensu stricto. Molecular characteristics including the similarity of the restriction profile of the non-transcribed spacer 2 (NTS2) and of the D1/D2 sequences of the rDNA, as well as other genotypic and phenotypic characteristics confirm that this group of strains is highly homogeneous and distinguishable from other species of the Saccharomyces sensu stricto group.

Carlo Zambonelli

Papers

Yeast influence on volatile composition of wines.

A biometric study of higher alcohol production in Saccharomyces cerevisiae.

Saccharomyces sensu stricto: systematics, genetic diversity and evolution.

Saccharomyces uvarum, a distinct group within Saccharomyces sensu stricto

Saccharomyces uvarum, a proper species within Saccharomyces sensu stricto