Metabolites produced and/or physical properties enhanced in wines fermented with non-conventional yeasts as single or co-fermentations compared with pure fermentation with
Abstract
Non-Saccharomyces yeasts play a substantial role in the early stages of wine fermentation. With the increase in alcohol concentration, indigenous or commercial strains of Saccharomyces cerevisiae take over and complete the transformation of the grape must sugars into ethanol, CO2, and other secondary metabolites. The presence of non-Saccharomyces during the fermentation has an impact on the wine composition, and consequently, their contribution during the fermentation process cannot be ignored. The new challenges to enhance the appeal and value of wine elaborated by traditional technology are being achieved by selecting and using autochthonous non-Saccharomyces and Saccharomyces strains that may enhance regional identity of wines. Greater understanding of yeast biochemistry and physiology is enabling the selection and development of yeast strains that have defined specific influences on process efficiency and wine quality. The aim of this chapter was to show the different aspects of non-Saccharomyces species that may play a positive incidence in the biotechnological process to conduct to wine elaboration.
Keywords
- non-Saccharomyces yeast
- wine production
- aroma
- enzymes
- mixed culture
1. Introduction
Since 1866 when Louis Pasteur first elucidated the bioconversion of grape juice into wine, this complex biochemical process and the role of the yeast therein have been studied continuously.
Yeast and bacterial microbial dynamics have been studied since the 1970s [4]. Using classical methods, microbial count and diversity is determined by employing synthetic culture media containing agar. The biodiversity in the complex ecosystems is impossible to characterize with precision using classical microbiological culture-dependent methods. When using enrichment methods and growth on culture medium, the microbiota naturally present in the sample are open to undergo important changes because of the ability of the certain species take control of the system and overcome other microbial components [5]. Due to this, populations less numerous or sensitive to the stress conditions are hard to find again and identify. Therefore, the use of culture-dependant methods could cause a misidentification of the microbial ecology of complex ecosystems [6].
Since the end of the 1990s, molecular techniques have helped provide a good overview of microbial ecology. These methods, generally named culture-independent methods, are used for the identification of microorganisms directly in the system through the study of their DNA and RNA without the need for isolation and cultivation. There are several advantages of the direct characterization of wine microbial DNA against to culture-dependent methods. Firstly, not all microbial populations are able to grow in enrichment media due to injury, lack of appropriate nutrients, or presence of viable but not culturable states. Secondly, the direct analyses allow saving time in comparison with the enrichment methods. This advantage could enable winemakers to use microbial detection data during the fermentation process, being able to anticipate possible spoilage problems in the wine. Furthermore, the DNA-based studies allow processing a larger numbers of samples than plating methods [7].
Nevertheless, in the past few years, successful culture-independent methods such as denaturing gradient gel electrophoresis (DGGE), real-time PCR, fluorescence in situ hybridization (FISH), or Fourier transform infrared spectroscopy (FT-IR) have been described. The PCR-DGGE method was first developed for the study of the microbial ecology in the environmental samples but soon found application in food microbiology [8]. This method is based on the separation of same length DNA fragments, but of different sequences. PCR-DGGE method has reported detection limits between 102 CFU/mL in pure cultures and 104 CFU/mL in wine or must samples [9].
In recent years, scientists have used real-time quantitative PCR (QPCR) to detect and quantity microorganisms in different alimentary environments [10]. The advantages of QPCR are the low detection level, often as low as one cell per mL, the speed by which assays are performed, and the ability to quantify yeasts present following alcoholic fermentation.
Fluorescence in situ hybridization (FISH) is a very promising technique for wine ecology studies for its simplicity and rapidity as well as the ability to observe the cell morphology by a microscope and the high sensitivity obtained using a flow cytometer [11].
Fourier transform infrared (FT-IR) spectroscopy is used to identify isolates according to the different components of the cell [12]. The relative success of this method is directly dependent on the complexity within a reference spectral library, identification results on genus and species level. Due to high automation and cost efficiency, this high-throughput method gives much deeper insights into functional yeast diversity during wine production.
Fortunately, DNA-based approaches have largely helped to clarify modern taxonomy. DNA sequence analysis is now widely used in the identification and classification of yeasts and other fungi therefore helping to reassign species within genus level because of the new species that are now discovered [13]. In food and beverage industry, these name changes influence our ability to notify the identity of spoilage microorganisms and to be in accordance with the regulations governing the presence of certain microorganisms in this industry. Current taxonomies recognize 149 yeast genera comprising nearly 1500 species [14]. Of these, more than 40 species have been isolated from grape must [15].
2. Use of non-Saccharomyces yeasts in the wine production
Non-
The initial belief that all non-
During fermentation, and more evident in spontaneous fermentations, which lack the initial high-density inoculum of
Despite the sustained presence of certain non-
Species | Metabolites and/or physical properties | References |
---|---|---|
Acetic acid Dodecanoic acid |
[9, 35, 42, 43] | |
Ethyl octanoate | ||
Glycerol | ||
Higher alcohols | ||
2-Methyl propanoic acid | ||
Succinic acid | ||
Terpenoids | ||
Esters Medium-chain fatty acids |
[44] | |
Terpenoids | ||
Acetate and ethyl esters | [9, 47] | |
Sulfur compounds | ||
Higher alcohols | ||
Medium-chain fatty acids | ||
2-Phenylethyl acetate | [48] | |
Acetate and ethyl esters | [47] | |
Acetone | ||
Heavy sulfur compounds | ||
Ethyl acetate | [49, 50] | |
Ethyl lactate | ||
Acetate and ethyl esters Higher alcohols |
[37, 50, 51] | |
Color | [41] | |
Methanol | ||
Glycerol Lactic acid 2-Phenylethanol |
[17, 38] | |
Esters | [35, 46] | |
Color | [53] | |
Thiols | [52] | |
Acetaldehyde | [29, 49] | |
2,3-Butanediol | ||
Ethyl esters | ||
Higher alcohols | ||
Polysaccharides | ||
Maloalcoholic deacidification | [39, 40] | |
Pyruvic acid | ||
Color | ||
Propanol | ||
Succinic acid | [35–37, 66] | |
Linalool Polysaccharides |
||
Polysaccharides | [49] | |
Polysaccharides | [50] |
The non-
3. Influence of non-Saccharomyces yeasts in mixed fermentations
In recent years, re-evaluation of the role of non-
The use of controlled mixed fermentations of non-
The influence of multistarter fermentation practices on final wine composition and on growth and death rates of the
Recent studies using strains of
Other studies have been carried out with the aim of de-acidifying the grape must or wine through malic acid degradation using mixed fermentations of
The use of
Another non-
Yeasts belonging to the
Benito et al. [53] have studied the hydroxycinnamate decarboxylase (HCDC) activity of
4. Highlights produced by non-Saccharomyces yeasts
4.1. Enzymes with oenological interest
Over the last several decades, the utilization of enzymes has become more important in winemaking. Enzymatic treatments of grapes, musts, and wines are nowadays useful for multiple positive aims, reduction of times maceration, clarification and filtration, increase in free and press juice yields, improvements in color and aroma extraction as well as wine stability [54]. The enzymes are proteins usually produced by bacteria or by filamentous fungi [55]. These proteins are very valuable tools for the winemakers; they now strengthen the use of endogenous enzymes over commercial exogenous enzymes. The production of extracellular hydrolytic enzymes by indigenous yeast could be notable and a better understanding to their benefit of wine production is required. Moreover, wine yeast has a decisive role in the production of commercial enzymes to be used in the wine elaboration process [56]. The principal wine yeast,
On the contrary than
Terpenoids, fatty acid esters, higher alcohols, glycerol, acetaldehyde, acetic acid, and succinic acid are some metabolic products generated from non-
Proteolytic and pectinolytic (polygalacturonase) are other extracellular enzymatic activities produced by non-
Non-
The role of pectinases in winemaking has been evaluated by Canal-Llaubères [67]. Some of the applications in mash treatment are to improve juice extraction, clarification process, filterability, and also color extraction. The use of pectolytic enzymes for maceration may also accelerate the extraction of phenolic compounds, reducing the maceration time needed for high quality of wine [68]. The addition of fungal pectinase preparations is a normal practice in wine industry even though pectin esterase and polygalacturonase enzymatic activities increase during grape ripening and are produced by non-
The accumulation of esters in wine is known to be a result of the balance between the yeast’s ester-synthesizing enzymes and hydrolysis reactions involving esterases (responsible for cleavage and in some cases, formation of ester bonds). The production of extracellular esterases in
The lipids proceeding from the grape or from autolytic activity of yeasts can be degraded by lipases. After this enzymatic reaction, free fatty acids would be released into the juice or wine, which can lead to changes in wine quality. While properties of lipoxygenase and peroxide-cleaving enzymes from grapes have been well established [73], the knowledge about lipase enzyme production by non-
4.2. Use of lower ethanol efficiency yeasts
In recent decades, the increasing alcohol level in wine is one of the most important challenges facing in the enological industry. The problem is related in part to global warming, which results in modifications of fruit maturation patterns, as well as a lack of balance between sugar accumulation and the phenolic ripeness of berries [77]. Fermentations with higher initial sugar content combined with high final ethanol concentration may have impact on microbiological, technological, sensorial, and financial aspects of winemaking. Higher sugar level delivers shifts in alcohol, altering flavors and mouthfeel. Musts with higher sugar concentrations cause a stress response in yeast leading to an increased formation of fermentation co-products, such as acetic acid. Also, this increasing sugar content leads to delay harvest period so as to insure appropriate aromatic and phenolic maturity. On the commercial side, excess ethanol can get worse sensory quality of wine, discourage consumers, because of the health effects associated with the excessive alcohol consumption or become a drawback in the global market, due to regulations and taxes associated with the alcohol content of beverages. All these reasons stimulate the creation of strategies directed to reduce alcohol level in wine.
Researchers, engineers, and oenologists are working together to develop approaches to limit ethanol content of wines, targeting almost all the steps in the production cycle [78], including among other examples, grapevine clonal selection, vineyard management, winemaking practices adapted to unripe grapes [79], use of lower ethanol efficiency yeast strains [80] or metabolic inhibitors [81] and partial dealcoholization by physical means [82]. While some of these technologies are still in need of fundamental research, others are in several stages of regulatory support and implementation by the industry.
The development of low-alcohol yeasts is a current challenge in wine industry. During the last years, researchers have been investigating
Given the vast potential for diverse wine relevant phenotypes among non-
Respiration and fermentation are two pathways for ATP production from glucose used by yeasts. Both pathways start with glycolysis, the major process for sugar degradation where the breakage of one glucose molecule results in the production of two molecules of pyruvate and ATP. In fermentation, pyruvate is finally transformed into ethanol by pyruvate decarboxylase (
Yeasts can be classified depending on the way they regulate their respiro-fermentative metabolism. Crabtree-positive yeasts could ferment under aerobic conditions only if sugar concentration is above certain thresholds. The prime example of Crabtree-positive species is
It is thought that redox balance in the metabolism of sugars generates the production of metabolic by-products as acetic acid, ethanol, and glycerol. In recent years, it is trying to take the control of metabolic systems in order to redirect carbon flux towards desirable compounds release, for example, glycerol overproduction. An added benefit of this approach is that enhanced glycerol concentrations can have a favorable influence on wine by enhancing its sweetness, smoothness, and overall body [90].
Several yeast strains, including
4.3. Bioprotection by non-Saccharomyces yeasts
Vinification process is composed by different and delicate steps as growing, harvesting, fermentation, and aging and storage in the winery. Unsuitable precautions or poor practice during any of these steps can lead to growth of wine spoilage organisms and consequent production losses. The major microorganisms involved in wine spoilage are acetic acid bacteria from genera
Traditionally, sulfur dioxide (as potassium metabisulphite), sorbic acid, fumaric acid, and dimethyl dicarbonate (DMDC) are used for preservation of different wines in various countries. Due to these drawbacks and growing consumer bias against chemical preservatives, research efforts are directed towards use of different physical methods and exploitation of natural antimicrobial compounds obtained from plants, animals, and microorganisms for wine preservation. Many studies have demonstrated the potential of natural products such as hydroxycinnamates and organic acids [94], chitosan [95], nisin [96], lysozyme [97], antimicrobial peptides [98], killer toxins [99], natamycin [100], β-glucanases [101], bovine lactoferrin-derived peptides [102], carvacrol and thymol [103], and vitamin K5 [104] for the control of wine spoilage yeasts and bacteria.
One of the biological mechanisms for the regulation of population dynamics in several microbial ecosystems is the production of toxins capable of kill or inhibit other microorganisms, taxonomically related or not to the producing strains. The toxins synthesized by yeasts, known as killer factor, are proteins or glycoproteins whose action is mediated by specific receptors in the cell wall of the sensitive microorganism. The killer character, first reported on the decade of the 1960s in a
The killer toxins secreted by the yeast species
5. Concluding remarks
Strain selection is of key importance, as not all strains within a species will necessarily show the same desirable characteristics [110]. The accepted list of desirable characteristics as pertaining to the wine yeast
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