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Vilela, A. Wine Aroma and Flavor Compounds from Fermentative Origin. Encyclopedia. Available online: https://encyclopedia.pub/entry/49062 (accessed on 19 May 2024).
Vilela A. Wine Aroma and Flavor Compounds from Fermentative Origin. Encyclopedia. Available at: https://encyclopedia.pub/entry/49062. Accessed May 19, 2024.
Vilela, Alice. "Wine Aroma and Flavor Compounds from Fermentative Origin" Encyclopedia, https://encyclopedia.pub/entry/49062 (accessed May 19, 2024).
Vilela, A. (2023, September 12). Wine Aroma and Flavor Compounds from Fermentative Origin. In Encyclopedia. https://encyclopedia.pub/entry/49062
Vilela, Alice. "Wine Aroma and Flavor Compounds from Fermentative Origin." Encyclopedia. Web. 12 September, 2023.
Wine Aroma and Flavor Compounds from Fermentative Origin
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This entry is adapted from the peer-reviewed paper 10.3390/fermentation6010022

Wine sensory experience includes flavor, aroma, color, and (for some) even acoustic traits, which impact consumer acceptance. The quality of the wine can be negatively affected by the presence of off-flavors and aromas, dubious colors, or sediments present in the bottle or glass after pouring (coloring matter that precipitates or calcium bitartrate crystals). Flavor profiles of wines result from many variations in vineyard and winery production, including grape selection, winemaker’s knowledge and technique, and tools used to produce wines with a specific flavor. 

wine yeasts lactic acid bacteria co-inoculation sequence inoculation flavor compounds

1. The Human Senses in Wine Evaluation

Five senses are involved in perceiving wine sensory quality: sight, taste, hearing, touch, and smell. Color perception results from the stimulus of the retina by light (wavelengths 380 to 760 nm). In wine, color and appearance are the first attributes by which quality is assessed. According to Spence [1], color is the most critical product-intrinsic indicator consumers use when searching, purchasing, and consuming food or a libation. Color, clarity, and hue affect the perception of other attributes, such as flavor, due to their association with color. For example, a yellow/green beverage is expected to have a lemon flavor and an acidic taste.
Taste is a chemical sense and happens when taste stimuli come into contact with the taste receptors located on the tongue, called taste buds. Humans can distinguish six basic tastes: sweet, sour, salty, bitter, umami, and fatty [2][3]. Between 20 and 30 levels of intensity can be distinguished for each taste, and each taste quality represents different nutritional or physiological requirements or a potential dietary risk [4].
Sound (waves that strike the eardrum, causing it to vibrate [5]) is also essential when judging a wine. For instance, when we hear a champagne cork popping, it is a sign that the wine has an enjoyable gas.
Texture in wine can be defined as the total sum of kinesthetic sensations derived from oral manipulation. It encompasses mouthfeel, masticatory properties, residual properties, and visual and auditory properties [6].
Aroma and flavor are chemical senses stimulated by the chemical properties of odor molecules, which must reach the olfactory bulb to interact with olfactory cells in the olfactory mucosa [7]; therefore, to smell, molecules must be airborne (i.e., volatile). The sensory term “flavor” is a mingled experience based on human judgment, built on personal differences in perception thresholds.
In conclusion, as reported by Swiegers et al. [8], all of the senses play a crucial role in wine/flavor development—color, aroma, mouthfeel, sound, and taste. Altogether, these sensory perceptions are very complex. Wine contains many flavor and aroma-active compounds. Terpenes, methoxypyrazines, esters, ethanol, and other alcohols and aldehydes impart distinct flavors and aromas (floral, pepper, fruit, woody, and vinylic flavors, among others) to wine [8][9][10]. The taste of wine can be described as sweet, sour, salty, umami, bitter, and, to a lesser extent, fat [11]. These properties result from sugars, polyols, salts, polyphenols, flavonoid compounds, amino acids, and fatty acids. Compounds such as glycerol, polysaccharides, and mannoproteins contribute to the viscosity and mouthfeel of wines [12]; grape anthocyanins contribute to the color [13], and ethanol, by sheer mass, also carries other alcohols along, promoting a mouth-warming effect [14].

2. Main Wine Aroma and Flavor Compounds from the Fermentative Origin

Yeast and bacteria are vital to the development of wine flavor. Many biosynthetic pathways in wine yeast and malolactic bacteria are responsible for forming wine aroma and flavor. However, we cannot discard the other factors influencing the wine's chemical composition, such as viticultural practices, grape-must composition, pH, fermentation temperature, and technological aspects associated with the vinification process [15]. So, depending on their origin, wine aroma, and flavor compounds can be named varietal aromas (originating from the grapes), fermentative aromas (created during alcoholic and malolactic fermentations), and aging aromas (developed during the reductive or oxidative wine-aging that depends on storage conditions) [16].
Most of the wine aroma and flavor compounds are produced or released during wine fermentation due to microbial activities of Saccharomyces and non-Saccharomyces yeast genera (Brettanomyces, Candida, Debaryomyces, Hanseniaspora, Hansenula, Kloeckera, Kluyveromyces, Lachancea, Metschnikowia, Pichia, Saccharomycodes, Schizosaccharomyces, Torulaspora, and Zygosaccharomyces). Both in spontaneous and inoculated wine fermentations, non-Saccharomyces are essential in the early stages of the fermentation, before Saccharomyces become dominant in the culture, and contribute meaningfully to the global aroma profile of wines by producing flavor-active compounds [17][18].
Aroma compounds have been directly linked to specific varietal flavors and wine aromas [19][20]. Most of these compounds are present in grapes and fermented wine at low concentrations. These aroma compounds are found in grapes in the form of non-odorant precursors that, due to the metabolic activity of Saccharomyces and non-Saccharomyces yeast during fermentation, are transformed to aromas and flavor that are of great relevance in the sensory perception of wines [20] (Table 1).
Table 1. Main odorants contribute to the varietal aromas of some monovarietal wines.

Compounds

Main Cultivars

Odour Descriptor

Ref.

Geraniol

Muscat, Gewurztraminer

Citrus, floral, geranium

[21]

Linalool

Muscat, Gewurztraminer

Floral, lavender

[21]

Nerol

Muscat

Floral

[21]

Tetrahydro-4-methyl-2-(2-methyl-1-propenyl)-2,5-cis-2h-pyran (cis-rose oxide)

Gewurztraminer

Geranium oil

[22]

3,6-Dimethyl-3a,4,5,7a-Tetrahydro-3h-1-Benzofuran-2-One

Gewurztraminer

Coconut, woody, sweet

[23]

3-Isobutyl-2-Methoxypyrazines

Sauvignon blanc

Bell pepper

[24][25]

4-Methyl-4-Mercaptopentan-2-One

Sauvignon blanc

Black currant

[24][25]

3-Mercapto-1-Hexanol (R Isomer)

Sauvignon blanc

Grapefruit, citrus peel

[24][25]

1,1,6-Trimethyl-1,2-Dihydronaphthalene

Riesling

Kerosene

[26]

3-Mercapto-1-Hexanol (S Isomer)

Semillon

Passion fruit

[27]

Rotundone

Shiraz

Black pepper

[28]
During alcoholic fermentation, some yeast, mainly non-Saccharomyces yeasts, can release β-glucosidases that hydrolyze the glycosidic bonds of the odorless, non-volatile glycoside-linked forms of monoterpenes (geraniol, linalool, nerol, among others), releasing the odor compounds to the wine [29]. Volatile thiols that give Sauvignon blanc wines their characteristic aroma (bell pepper, black currant, grapefruit, and citrus peel) are not present in grape juice but occur in grape must as odorless, non-volatile, cysteine-bound conjugates. During fermentation, the wine yeasts are responsible for cleaving the thiol from the precursor [30].
However, the significant aromas and flavor compounds from the fermentative origin are ethanol, higher alcohols or fusel alcohols, and esters. The biosynthetic pathways responsible for forming higher alcohols, the Ehrlich pathway, or the enzymes accountable for forming esters, have been studied in wine yeasts [31].
Higher alcohols are derived from amino acid catabolism via a pathway first described by Ehrlich [32] and later revised by Neubauer and Fromherz in 1911 [33]. Amino acids assimilated by the Ehrlich pathway (valine, leucine, isoleucine, methionine, and phenylalanine), present in grapes must be metabolized by yeasts sequentially throughout the fermentation. Figure 1 shows the metabolism of phenylalanine with the production of 2-phenylethanol and, after oxidation of phenylacetaldehyde, the formation of phenylacetate. Both compounds possess a pleasant rose-like aroma/flavor.
Figure 1. Schematic representation of the Ehrlich pathway for the catabolism of the aromatic amino acid phenylalanine, leading to the formation of 2-phenylethanol [34]. This biosynthetic pathway consists of three steps (reactions 1, 2, and 3): first, amino acids are deaminated to the corresponding α-ketoacids in transaminase reactions. In the second step, α-ketoacids are decarboxylated and converted to their corresponding aldehydes (five decarboxylases are involved in this process); in the third step, alcohol dehydrogenases (Adh1p to Adh6p and Sfa1p) catalyze the reduction of aldehydes to their corresponding higher alcohols [35].
Studies have shown that profiles and concentrations of higher alcohols produced vary by yeast species, even when the fermentation conditions are similar, which indicates that the mechanisms that regulate the Ehrlich pathway are diverse in non-Saccharomyces yeasts compared to Saccharomyces [16][36]. So, Ehrlich pathway mechanisms should be explored in detail in non-Saccharomyces yeasts as they form essential and flavorful wine aromas [36].
The most critical esters are synthesized by yeasts during alcoholic fermentation as a detoxification mechanism since they are less toxic than their correspondent alcohol or acidic precursors. Moreover, their synthesis serves as a mechanism for the regeneration of free CoA from its conjugates [16][37].
Esters (Figure 2) that contribute to wine aroma, derived from fermentation, belong to two categories: the acetate esters of higher alcohols and the ethyl esters of medium-chain fatty acids (MCFA). Acetate esters are formed inside the yeast cell, and in S. cerevisiae, the reaction is metabolized by two alcohol acetyltransferases, AATase I and AATase II (encoded by genes ATF1 and ATF2 [35][38]). Eat1p is responsible for producing acetate and propanoate esters [39][40]. Most medium-chain fatty acid ethyl ester biosynthesis during fermentation is catalyzed by two enzymes, Eht1p and Eeb1p [38][41].
Figure 2. Schematic representation of the essential wine esters: ethyl acetate (glue-like aroma), isoamyl acetate (banana aroma), 2-phenyl ethyl acetate (roses and honey aromas), isobutyl acetate (sweet-fruits aromas), ethyl caproate and ethyl caprylate with a sour-apple aroma [38].
Volatile fatty acids also contribute to the flavor and aroma of the wine. During yeast fermentation, long-chain fatty acids (LCFAs) are also formed via the fatty-acid synthesis pathway from acetyl-CoA in concentrations varying from ng/L to g/L [42]. Medium-chain fatty acids (MCFAs (C6 to C12)) are produced primarily by yeasts as intermediates in the biosynthesis of LCFAs that are prematurely released from the fatty acid synthase complex. These acids (Table 2) directly contribute to the flavor of wine or serve as substrates that participate in the formation of ethyl acetates [43]. Most have unpleasant aromas (see Table 2), so their formation should be minimized.
Table 2. Yeasts produce primary medium-chain fatty acids (MCFAs (C6 to C12)) during alcoholic fermentation.

Fatty Acid

Associated Aroma

Odor Threshold (μg/L) 1

Butanoic Acid

Rancid butter or baby vomit aroma

173

Hexanoic Acid

Sour, fatty, sweat, cheese

420

Octanoic Acid

Fatty, waxy, rancid oily, vegetable, cheesy

500

Decanoic Acid

Unpleasant rancid, sour, rich, citrus

1000

2-Methylpropanoic Acid

Acidic sour, cheese, dairy, buttery, rancid

2300

2- and 3-Methylbutyric Acid

Sour, stinky feet, sweaty, cheese, tropical

33
1 Measured in model wine, water/ethanol (90 + 10, w/w) [44].
Yeasts can also form sulfur-containing compounds during alcoholic fermentation. They are usually perceived as off-flavors. The sulfur-containing compounds can be derived from the grape and the metabolic activities of yeast and bacteria. They can also occur due to the chemical reactions during the wine aging and storage and environmental contamination [45]. They can be formed by enzymatic mechanisms as the products of metabolic and fermentative pathways whose substrates are both amino acids and some sulfur-containing pesticides. When wine microorganisms metabolize these thiols, the sulfur compounds formed are considered off-flavors [46], which convey negative notes such as cabbage, garlic, onion, rotten eggs, rubber, and sulfur to wines [47]. However, some volatile thiols may confer enjoyable aromatic notes at trace levels, such as 4-mercapto-4-methylpentan-2-one (4MMP), 3-mercaptohexan-1-ol (3MH), already mentioned in Table 1, and 3-mercaptohexyl acetate (3MHA), necessary for the characterization of the typical Sauvignon Blanc wine aroma [24][25][48].
Finally, the carbonyl compounds are another important family of aromatic compounds present in wines. This group may include acetaldehyde, acrolein, ethyl carbamate, formaldehyde, and furfural [49]. Several factors may contribute to the presence of carbonyl compounds in wines, including the fermentation of over-ripe grapes and increasing the maceration time, probably due to increased concentration of precursors like amino acids and glucose in the must [50]. Due to their carbonyl group, carbonyl compounds present a high reactivity with the nucleophile’s cellular constituents [51] and may cause cell damage. So, these compounds are toxic, and their formation should be avoided.

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