Stefano Lubiana Pinot Noir ferment vintage 2010

The role of yeast in winemaking is the most important element that distinguishes

wine Wine is an alcoholic drink typically made from fermented grapes. Yeast consumes the sugar in the grapes and converts it to ethanol and carbon dioxide, releasing heat in the process. Different varieties of grapes and strains of yeasts are m ...

from

fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in particu ...

juice Juice is a drink made from the extraction or pressing of the natural liquid contained in fruit and vegetables. It can also refer to liquids that are flavored with concentrate or other biological food sources, such as meat or seafood, such as ...

. In the absence of oxygen,

yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constit ...

converts the

sugars Sugar is the generic name for sweet-tasting, soluble carbohydrates, many of which are used in food. Simple sugars, also called monosaccharides, include glucose, fructose, and galactose. Compound sugars, also called disaccharides or double ...

of the fruit into alcohol and

carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is trans ...

through the process of fermentation.Jeff Cox ''"From Vines to Wines: The Complete Guide to Growing Grapes and Making Your Own Wine"'' pp. 133–36 Storey Publishing 1999 The more sugars in the grapes, the higher the potential alcohol level of the wine if the yeast are allowed to carry out fermentation to dryness.D. Bird ''"Understanding Wine Technology"'' pp. 67–73 DBQA Publishing 2005 Sometimes winemakers will stop fermentation early in order to leave some residual sugars and sweetness in the wine such as with

dessert wine Dessert wines, sometimes called pudding wines in the United Kingdom, are sweet wines typically served with dessert. There is no simple definition of a dessert wine. In the UK, a dessert wine is considered to be any sweet wine drunk with a meal ...

s. This can be achieved by dropping fermentation temperatures to the point where the yeast are inactive, sterile filtering the wine to remove the yeast or

fortification A fortification is a military construction or building designed for the defense of territories in warfare, and is also used to establish rule in a region during peacetime. The term is derived from Latin ''fortis'' ("strong") and ''facere' ...

with brandy or neutral spirits to kill off the yeast cells. If fermentation is unintentionally stopped, such as when the yeasts become exhausted of available nutrients and the wine has not yet reached dryness, this is considered a stuck fermentation.J. Robinson (ed) ''"The Oxford Companion to Wine"'' Third Edition pp. 778–80

Oxford University Press Oxford University Press (OUP) is the university press of the University of Oxford. It is the largest university press in the world, and its printing history dates back to the 1480s. Having been officially granted the legal right to print books ...

2006 The most common yeast associated with

winemaking Winemaking or vinification is the production of wine, starting with the selection of the fruit, its fermentation into alcohol, and the bottling of the finished liquid. The history of wine-making stretches over millennia. The science of wine and ...

is ''

Saccharomyces cerevisiae ''Saccharomyces cerevisiae'' () (brewer's yeast or baker's yeast) is a species of yeast (single-celled fungus microorganisms). The species has been instrumental in winemaking, baking, and brewing since ancient times. It is believed to have b ...

'' which has been favored due to its predictable and vigorous fermentation capabilities, tolerance of relatively high levels of alcohol and sulfur dioxide as well as its ability to thrive in normal wine pH between 2.8 and 4. Despite its widespread use which often includes deliberate inoculation from cultured stock, ''S. cerevisiae'' is rarely the only yeast species involved in a fermentation. Grapes brought in from harvest are usually teeming with a variety of "wild yeast" from the '' Kloeckera'' and '' Candida'' genera. These yeasts often begin the fermentation process almost as soon as the grapes are picked when the weight of the clusters in the harvest bins begin to crush the grapes, releasing the sugar-rich must.K. Fugelsang, C. Edwards ''Wine Microbiology'' Second Edition pp. 3–28 Springer Science and Business Media , New York (2010) While additions of sulfur dioxide (often added at the crusher) may limit some of the wild yeast activities, these yeasts will usually die out once the alcohol level reaches about 15% due to the

toxicity Toxicity is the degree to which a chemical substance or a particular mixture of substances can damage an organism. Toxicity can refer to the effect on a whole organism, such as an animal, bacterium, or plant, as well as the effect on a subs ...

of alcohol on the yeast cells

physiology Physiology (; ) is the scientific study of functions and mechanisms in a living system. As a sub-discipline of biology, physiology focuses on how organisms, organ systems, individual organs, cells, and biomolecules carry out the chemical ...

while the more alcohol tolerant ''

Saccharomyces ''Saccharomyces'' is a genus of fungi that includes many species of yeasts. ''Saccharomyces'' is from Greek σάκχαρον (sugar) and μύκης (fungus) and means ''sugar fungus''. Many members of this genus are considered very important in f ...

'' species take over. In addition to ''S. cerevisiae'', '' Saccharomyces bayanus'' is a species of yeast that can tolerate alcohol levels of 17–20% and is often used in fortified wine production such as

ports A port is a maritime facility comprising one or more wharves or loading areas, where ships load and discharge cargo and passengers. Although usually situated on a sea coast or estuary, ports can also be found far inland, such as H ...

and varieties such as Zinfandel and Syrah harvested at high Brix sugar levels. Another common yeast involved in wine production is ''

Brettanomyces ''Brettanomyces'' is a non-spore forming genus of yeast in the family Saccharomycetaceae, and is often colloquially referred to as "Brett". The genus name ''Dekkera'' is used interchangeably with ''Brettanomyces'', as it describes the teleomorp ...

'' whose presence in a wine may be viewed by different winemakers as either a

wine fault A wine fault or defect is an unpleasant characteristic of a wine often resulting from poor winemaking practices or storage conditions, and leading to wine spoilage. Many of the compounds that cause wine faults are already naturally present in wine ...

or in limited quantities as an added note of complexity.B. Zoecklein, K. Fugelsang, B. Gump, F. Nury ''Wine Analysis and Production'' pp. 281–90 Kluwer Academic Publishers, New York (1999)

History

For most of the

history of wine Wine has been produced for thousands of years, with evidence of ancient wine production in Georgia from BC (the earliest known traces of wine), West Azerbaijan province of Iran from BC, Armenia from BC (large-scale production), and Sicil ...

, winemakers did not know the mechanism that somehow converted sugary grape juice into alcoholic wine. They could observe the fermentation process which was often described as "boiling", "seething" or the wine being "troubled" due to release of carbon dioxide that gave the wine a frothy, bubbling appearance. This history is preserved in the

etymology Etymology ()The New Oxford Dictionary of English (1998) – p. 633 "Etymology /ˌɛtɪˈmɒlədʒi/ the study of the class in words and the way their meanings have changed throughout time". is the study of the history of the Phonological chan ...

of the word "yeast" itself which essentially means "to boil". In the mid-19th century, the French scientist Louis Pasteur was tasked by the French government to study what made some wines spoil. His work, which would later lead to Pasteur being considered one of the "Fathers of Microbiology", would uncover the connection between microscopic yeast cells and the process of the fermentation. It was Pasteur who discovered that yeast converted sugars in the must into alcohol and carbon dioxide, though the exact mechanisms of how the yeast would accomplish this task was not discovered till the 20th century with the Embden–Meyerhof–Parnas pathway.J. Robinson (ed) ''"The Oxford Companion to Wine"'' Third Edition pgs 267 & 508

2006 The yeast species commonly known as ''Saccharomyces cerevisiae'' was first identified in late 19th century enology text as ''Saccharomyces ellipsoideus'' due to the elliptical (as opposed to circular) shape of the cells. Throughout the 20th century, more than 700 different strains of ''Saccharomyces cerevisiae'' were identified. The differences between the vast majority of these strains are mostly minor, though individual winemakers will develop a preference for particular strains when making certain wines or working with particular

grape varieties This list of grape varieties includes cultivated grapes, whether used for wine, or eating as a table grape, fresh or dried (raisin, currant, sultana). For a complete list of all grape species including those unimportant to agriculture, see Viti ...

. Some of these difference include the "vigor" or speed of fermentation, temperature tolerance, the production of volatile sulfur compounds (such as hydrogen sulfide) and other compounds that may influence the

aroma An odor (American English) or odour ( Commonwealth English; see spelling differences) is caused by one or more volatilized chemical compounds that are generally found in low concentrations that humans and animals can perceive via their se ...

of the wine.

Role in winemaking

The primary role of yeast is to convert the sugars present (namely

glucose Glucose is a simple sugar with the molecular formula . Glucose is overall the most abundant monosaccharide, a subcategory of carbohydrates. Glucose is mainly made by plants and most algae during photosynthesis from water and carbon dioxide, u ...

) in the grape must into alcohol. The yeast accomplishes this by utilizing glucose through a series of metabolic pathways that, in the presence of oxygen, produces not only large amounts of energy for the cell but also many different intermediates that the cell needs to function. In the absence of oxygen ( and sometimes even in the presence of oxygenB. Zoecklein, K. Fugelsang, B. Gump, F. Nury ''Wine Analysis and Production'' pp. 97–114 Kluwer Academic Publishers, New York (1999) ), the cell will continue some metabolic functions (such as glycolysis) but will rely on other pathways such as reduction of acetaldehyde into ethanol (fermentation) to "recharge" the co-enzymes needed to keep metabolism going. It is through this process of fermentation that ethanol is released by the yeast cells as a waste product. Eventually, if the yeast cells are healthy and fermentation is allowed to run to the completion, all fermentable sugars will be used up by the yeast with only the unfermentable

pentose In chemistry, a pentose is a monosaccharide (simple sugar) with five carbon atoms. The chemical formula of many pentoses is , and their molecular weight is 150.13 g/mol.

Other compounds in wine produced by yeast

While the production of alcohol is the most noteworthy by-product of yeast metabolism from a winemaking perspective, there are a number of other products that yeast produce that can be also influence the resulting wine. This includes

glycerol Glycerol (), also called glycerine in British English and glycerin in American English, is a simple triol compound. It is a colorless, odorless, viscous liquid that is sweet-tasting and non-toxic. The glycerol backbone is found in lipids known ...

which is produced when an intermediate of the glycolysis cycle (

dihydroxyacetone Dihydroxyacetone (; DHA), also known as glycerone, is a simple saccharide (a triose) with formula . DHA is primarily used as an ingredient in sunless tanning products. It is often derived from plant sources such as sugar beets and sugar cane, an ...

) is reduced to "recharge" the NADH enzyme needed to continue other metabolic activities. This is usually produced early in the fermentation process before the mechanisms to reduce acetaldehyde into ethanol to recharge NADH becomes the cell's primary means of maintaining

redox Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a ...

balance. As glycerol contributes increased

body Body may refer to: In science * Physical body, an object in physics that represents a large amount, has mass or takes up space * Body (biology), the physical material of an organism * Body plan, the physical features shared by a group of anima ...

and a slightly sweet taste without increasing the alcohol level of the wine, some winemakers try to intentionally favor conditions that would promote glycerol production in wine. This includes selecting yeast strains that favor glycerol production (or allowing some wild yeast like ''Kloeckera'' and ''Metschnikowia'' to ferment), increased oxygen exposure and aeration as well as fermenting at higher temperatures. Glycerol production is also encouraged if most available acetaldehyde is made unavailable by binding with

bisulfite The bisulfite ion (IUPAC-recommended nomenclature: hydrogensulfite) is the ion . Salts containing the ion are also known as "sulfite lyes". Sodium bisulfite is used interchangeably with sodium metabisulfite (Na2S2O5). Sodium metabisulfite disso ...

molecules in the wine, but it would take a substantial amount of sulfur dioxide addition (far beyond legal limits) to prolong glycerol production beyond just these very nascent stages of fermentation.Dr. Yair Margalit, ''Winery Technology & Operations A Handbook for Small Wineries'' pp. 67–74 The Wine Appreciation Guild (1996) Other by-products of yeast include: * Methanol – Caused by the

demethylation Demethylation is the chemical process resulting in the removal of a methyl group (CH3) from a molecule. A common way of demethylation is the replacement of a methyl group by a hydrogen atom, resulting in a net loss of one carbon and two hydrogen at ...

pectins Ancient Greek includes the forms of the Greek language used in ancient Greece and the ancient world from around 1500 BC to 300 BC. It is often roughly divided into the following periods: Mycenaean Greek (), Dark Ages (), the Archaic peri ...

in the must by enzymes of the yeast. More commonly found in red wines than white but only in very small amounts between 20–200 mg/L. * Fusel oils – Formed by the decomposition of amino acids by the yeast. This includes

2,3-Butanediol 2,3-Butanediol is the organic compound with the formula (CH3CHOH)2. It is classified as a ''vic''-diol ( glycol). It exists as three stereoisomers, a chiral pair and the meso isomer. All are colorless liquids. Applications include precursors ...

which is formed by yeast-consuming

diacetyl Diacetyl (IUPAC systematic name: butanedione or butane-2,3-dione) is an organic compound with the chemical formula (CH3CO)2. It is a yellow liquid with an intensely buttery flavor. It is a vicinal diketone (two C=O groups, side-by-side). Diacet ...

, the compound that gives Chardonnay and other wines a "

butter Butter is a dairy product made from the fat and protein components of churned cream. It is a semi-solid emulsion at room temperature, consisting of approximately 80% butterfat. It is used at room temperature as a spread, melted as a condimen ...

y" aroma, reducing it first to acetoin and then to the more neutral-smelling 2,3-Butanediol. Many beer and winemakers who have a wine with too much "butteriness" will often "pitch" fresh yeast cultures into the no longer fermenting tank so that the yeast will consume the diacetyl and reduce the aroma. *

Succinic acid Succinic acid () is a dicarboxylic acid with the chemical formula (CH2)2(CO2H)2. The name derives from Latin ''succinum'', meaning amber. In living organisms, succinic acid takes the form of an anion, succinate, which has multiple biological ro ...

– Like glycerol, this is often formed early in fermentation. Usually found in concentrations of 500–1200 mg/L, it is a minor acid in the overall acidity of wine. * Acetic acid – Considered a main component of

volatile acidity A wine fault or defect is an unpleasant characteristic of a wine often resulting from poor winemaking practices or storage conditions, and leading to wine spoilage. Many of the compounds that cause wine faults are already naturally present in wine ...

that can make a wine taste unbalanced and overly acidic. While acetic acid is the main volatile acid produced by yeast, trace amounts of butyric, formic and propionic acids can also be formed depending on the yeast strain. Most countries have wine laws setting the legal limit of volatile acidity, usually expressed as acetic acid, to 1200–2000 mg/L. Acetic acid can also lead to the development of the wine fault

ethyl acetate Ethyl acetate ( systematically ethyl ethanoate, commonly abbreviated EtOAc, ETAC or EA) is the organic compound with the formula , simplified to . This colorless liquid has a characteristic sweet smell (similar to pear drops) and is used in glues ...

which is characterized by a "nail polish remover" smell. However, small amounts of acetic acid are actually beneficial for the yeast as they use them to synthesis lipids in the cell membrane. CatavinoEnMano

* Acetaldehyde – While most of the acetaldehyde produce gets reduced to ethanol or is bound by sulfur dioxide, concentrations between 50–100 mg/L can remain in the wine. The flor yeast strains that produce the

Spanish wine Spanish wine () includes red, white, and sparkling wines produced throughout the country. Located on the Iberian Peninsula, Spain has over 1.2 million hectares (2.9 million acres) planted in wine grapes, making it the most widely pla ...

Sherry Sherry ( es, jerez ) is a fortified wine made from white grapes that are grown near the city of Jerez de la Frontera in Andalusia, Spain. Sherry is produced in a variety of styles made primarily from the Palomino grape, ranging from light versi ...

will produce higher amounts that contributes to the characterized "aldehydic" aromas of Sherries. In the presence of oxygen, yeast can convert some of the ethanol presence in the wine back into acetaldehyde creating

oxidized Redox (reduction–oxidation, , ) is a type of chemical reaction in which the oxidation states of substrate change. Oxidation is the loss of electrons or an increase in the oxidation state, while reduction is the gain of electrons or a ...

aromas. * Hydrogen sulfide – Often produced by yeast during fermentation because of a nitrogen deficiency in the must. This can be done by a reduction of

sulfate The sulfate or sulphate ion is a polyatomic anion with the empirical formula . Salts, acid derivatives, and peroxides of sulfate are widely used in industry. Sulfates occur widely in everyday life. Sulfates are salts of sulfuric acid and many ...

s or sulfites available in the must or by the decomposition of dead yeast cells by other yeast that releases sulfur-containing amino acids that are further broken down by the yeast. The latter often happens with wines that sit in contact with their lees for long periods of time between

racking Racking, often referred to as Soutirage or Soutirage traditionnel (meaning racking in French), also filtering or fining, is the process of moving wine or beer from one container to another using gravity rather than a pump, which can be disruptiv ...

s. In the presence of alcohol, hydrogen sulfide can react with ethanol to form ethyl mercaptans and

disulfide In biochemistry, a disulfide (or disulphide in British English) refers to a functional group with the structure . The linkage is also called an SS-bond or sometimes a disulfide bridge and is usually derived by the coupling of two thiol groups. In ...

s that contribute to off aromas and wine faults. Some commercial yeast strains, such as ''Montrachet 522'' are known to produce higher levels of hydrogen sulfides than other strains, particularly if the must has some nutrient deficiencies. * Pyruvic acid – Along with acetaldehyde, this compound can react with

anthocyanins Anthocyanins (), also called anthocyans, are water-soluble vacuolar pigments that, depending on their pH, may appear red, purple, blue, or black. In 1835, the German pharmacist Ludwig Clamor Marquart gave the name Anthokyan to a chemical com ...

extracted from contact with grape skins to create a more stable color pigment (

pyranoanthocyanin The pyranoanthocyanins are a type of pyranoflavonoids. They are chemical compounds formed in red wines by yeast during fermentation processes or during controlled oxygenation processes during the aging of wine. The different classes of pyranoant ...

) that can enhance the color of some red wines. *Various

esters In chemistry, an ester is a compound derived from an oxoacid (organic or inorganic) in which at least one hydroxyl group () is replaced by an alkoxy group (), as in the substitution reaction of a carboxylic acid and an alcohol. Glycerides ar ...

, ketones,

lactones Lactones are cyclic carboxylic esters, containing a 1-oxacycloalkan-2-one structure (), or analogues having unsaturation or heteroatoms replacing one or more carbon atoms of the ring. Lactones are formed by intramolecular esterification of the c ...

phenols In organic chemistry, phenols, sometimes called phenolics, are a class of chemical compounds consisting of one or more hydroxyl groups (— O H) bonded directly to an aromatic hydrocarbon group. The simplest is phenol, . Phenolic compounds are ...

and

acetals In organic chemistry, an acetal is a functional group with the connectivity . Here, the R groups can be organic fragments (a carbon atom, with arbitrary other atoms attached to that) or hydrogen, while the R' groups must be organic fragments no ...

Lees

When yeast cells die, they sink to the bottom of the fermentation vessel where they combine with insoluble

tartrates A tartrate is a salt or ester of the organic compound tartaric acid, a dicarboxylic acid. The formula of the tartrate dianion is O−OC-CH(OH)-CH(OH)-COO− or C4H4O62−. The main forms of tartrates used commercially are pure crystalline ta ...

, grape seeds, skin and pulp fragments to form the lees. During fermentation, the first significant racking which removes the bulk of dead yeast cells is often referred to as the ''gross lees'' as opposed to the less coarse ''fine lees'' that come as the wine continues to

settle Settle or SETTLE may refer to: Places * Settle, Kentucky, United States * Settle, North Yorkshire, a town in England ** Settle Rural District, a historical administrative district Music * Settle (band), an indie rock band from Pennsylvania * ''S ...

and age. During the time that the wine spends in contact with the lees, a number of changes can impact the wine due to both the autolysis (or self-metabolize) of the dead yeast cells as well as the reductive conditions that can develop if the lees are not aerated or stirred (a process that the French call ''bâtonnage''). The length of time that a wine spends on its lees (called ''sur lie'') will depend on the winemaking style and type of wine.J. Robinson (ed) ''"The Oxford Companion to Wine"'' Third Edition pp. 398–99

2006 The process of leaving the wine to spend some contact with the lees has a long history in winemaking, being known to the

Ancient Romans In modern historiography, ancient Rome refers to Roman civilisation from the founding of the city of Rome in the 8th century BC to the collapse of the Western Roman Empire in the 5th century AD. It encompasses the Roman Kingdom (753–509 ...

and described by

Cato the Elder Marcus Porcius Cato (; 234–149 BC), also known as Cato the Censor ( la, Censorius), the Elder and the Wise, was a Roman soldier, senator, and historian known for his conservatism and opposition to Hellenization. He was the first to write his ...

in the 2nd century BC. Today the practice is widely associated with any red wines that are barrel fermented,

Muscadet Muscadet ( , , ) is a French white wine. It is made at the western end of the Loire Valley, near the city of Nantes in the Pays de la Loire region. It is made from the Melon de Bourgogne grape, often referred to simply as ''melon''. While most ...

, sparkling wine

Champagne Champagne (, ) is a sparkling wine originated and produced in the Champagne wine region of France under the rules of the appellation, that demand specific vineyard practices, sourcing of grapes exclusively from designated places within it, ...

as well as Chardonnay produced in many wine regions across the globe. Typically when wines are left in contact with their lees, they are regularly stirred in order to release the mannoproteins,

polysaccharides Polysaccharides (), or polycarbohydrates, are the most abundant carbohydrates found in food. They are long chain polymeric carbohydrates composed of monosaccharide units bound together by glycosidic linkages. This carbohydrate can react with w ...

and other compounds that were present in the yeast cell walls and membranes. This stirring also helps avoid the development of reductive sulfur compounds like mercaptans and hydrogen sulfide that can appear if the lees layer is more than 10 cm (4 inches) thick and undisturbed for more than a week. Most of the benefits associated with lees contact deals with the influence on the wine of the mannoproteins released during the autolysis of the yeast cells. Composed primarily of

mannose Mannose is a sugar monomer of the aldohexose series of carbohydrates. It is a C-2 epimer of glucose. Mannose is important in human metabolism, especially in the glycosylation of certain proteins. Several congenital disorders of glycosylation ...

and proteins, with some glucose, mannoproteins are often bound in the cell wall of yeast with

hydrophobic In chemistry, hydrophobicity is the physical property of a molecule that is seemingly repelled from a mass of water (known as a hydrophobe). In contrast, hydrophiles are attracted to water. Hydrophobic molecules tend to be nonpolar and, t ...

aroma compounds that become volatilized as the cell wall breaks down. Not only does the release of mannoproteins impart sensory changes in the wine but they can contribute to

tartrate A tartrate is a salt or ester of the organic compound tartaric acid, a dicarboxylic acid. The formula of the tartrate dianion is O−OC-CH(OH)-CH(OH)-COO− or C4H4O62−. The main forms of tartrates used commercially are pure crystalline ta ...

and protein stability, help enhance the

and

mouthfeel Mouthfeel refers to the physical sensations in the mouth caused by food or drink, making it distinct from taste. It is a fundamental sensory attribute which, along with taste and smell, determines the overall flavor of a food item. Mouthfeel ...

of the wine as well as decrease the perception of bitterness and astringency of

tannins Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'', f ...

Secondary fermentation

The production of

and many

sparkling wine Sparkling wine is a wine with significant levels of carbon dioxide in it, making it fizzy. While the phrase commonly refers to champagne, European Union countries legally reserve that term for products exclusively produced in the Champagne regi ...

s requires a second fermentation to occur in the bottle in order to produce the

carbonation Carbonation is the chemical reaction of carbon dioxide to give carbonates, bicarbonates, and carbonic acid. In chemistry, the term is sometimes used in place of carboxylation, which refers to the formation of carboxylic acids. In inorganic ch ...

necessary for the style. A small amount of sugared liquid is added to individual bottles, and the yeast is allowed to convert this to more alcohol and

. The lees are then ricked into the neck of the bottle, frozen, and expelled via pressure of the carbonated wine.

Types of yeasts used in winemaking

Yeast

taxonomy Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...

includes classification of yeast species depending on the presence or absence of a sexual phase. Therefore, some winemaking yeasts are classified by their asexual anamorph (or "imperfect" form) while others may be classified by their sexual

teleomorph In mycology, the terms teleomorph, anamorph, and holomorph apply to portions of the life cycles of fungi in the phyla Ascomycota and Basidiomycota: *Teleomorph: the sexual reproductive stage (morph), typically a fruiting body. *Anamorph: an asex ...

(or "perfect" form). A common example of this is ''

'' (or "Brett") that is usually referenced in wine and

viticulture Viticulture (from the Latin word for '' vine'') or winegrowing (wine growing) is the cultivation and harvesting of grapes. It is a branch of the science of horticulture. While the native territory of '' Vitis vinifera'', the common grape vine, ...

text under its asexual classification though some scientific and winemaking texts may describe specific species (such as '' Dekkera bruxellensis'') under its

sporulating In biology, a spore is a unit of sexual reproduction, sexual or asexual reproduction that may be adapted for biological dispersal, dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of ...

sexual classification of '' Dekkera''. Unless otherwise noted, this article will commonly refer to the asexual form of wine yeast. The most common yeast generally associated with winemaking is ''Saccharomyces cerevisiae'' which is also used in bread making and brewing. Other genera of yeast that can be involved in winemaking (either beneficially or as the cause of potential

wine faults A wine fault or defect is an unpleasant characteristic of a wine often resulting from poor winemaking practices or storage conditions, and leading to wine spoilage. Many of the compounds that cause wine faults are already naturally present in wine ...

) include: *''

'' (Teleomorph ''Dekkera'') *'' Candida'' (Teleomorphs for different species from several genera including '' Pichia'', '' Metschnikowia'', '' Issatchenkia'', '' Torulaspora'' and '' Kluyveromyces'') *'' Kloeckera'' (Teleomorph ''Hanseniaspora''), usually the most common "wild yeast" found in the vineyard. Some species are known as "

killer yeast A killer yeast is a yeast, such as ''Saccharomyces cerevisiae'', which is able to secrete one of a number of toxic proteins which are lethal to susceptible cells. These "killer toxins" are polypeptides that kill sensitive cells of the same or relat ...

" that produce inhibitory levels of

and acetic acid that can kill off sensitive strains of ''

'' *'' Saccharomycodes'' *'' Schizosaccharomyces'', the only wine yeast that reproduced by fission whereas most wine yeast reproduce by

budding Budding or blastogenesis is a type of asexual reproduction in which a new organism develops from an outgrowth or bud due to cell division at one particular site. For example, the small bulb-like projection coming out from the yeast cell is kno ...

. *''

Zygosaccharomyces ''Zygosaccharomyces'' is a genus of yeasts in the family Saccharomycetaceae. It was first described under the genus ''Saccharomyces'', but in 1983, it was reclassified to its current name in the work by Barnett ''et al''. The yeast has a long his ...

'', very alcohol-tolerant and can grow in wines up to 18% v/v. Additionally this yeast can survive in extremely high sugar levels (as much as 60% w/w or 60 Brix) and is very resistant to sulfur dioxide. *'' Aureobasidium'', particularly the "black yeast" species of ''

Aureobasidium pullulans ''Aureobasidium pullulans'' is a ubiquitous and generalistic black, yeast-like fungus that can be found in different environments (e.g. soil, water, air and limestone). It is well known as a naturally occurring epiphyte or endophyte of a wide ra ...

'' found in moist cellars that can contaminate

aging wine The aging of wine is potentially able to improve the quality of wine. This distinguishes wine from most other consumable goods. While wine is perishable and capable of deteriorating, complex chemical reactions involving a wine's sugars, acids an ...

in barrels.

Saccharomyces

The yeast genus ''Saccharomyces'' (sugar mold) is favored for winemaking (for both grapes as well as other

fruit wines Fruit wines are fermented alcoholic beverages made from a variety of base ingredients (other than grapes); they may also have additional flavors taken from fruits, flowers, and herbs. This definition is sometimes broadened to include any alcoholi ...

in addition also to being used in brewing and breadmaking) because of the generally reliable and positive attributes it can bring to the wine. These yeasts will usually readily

ferment Fermentation is a metabolic process that produces chemical changes in organic substrates through the action of enzymes. In biochemistry, it is narrowly defined as the extraction of energy from carbohydrates in the absence of oxygen. In food ...

, sucrose and

raffinose Raffinose is a trisaccharide composed of galactose, glucose, and fructose. It can be found in beans, cabbage, brussels sprouts, broccoli, asparagus, other vegetables, and whole grains. Raffinose can be hydrolyzed to D-galactose and sucro ...

and metabolize glucose, sucrose, raffinose, maltose and

ethanol Ethanol (abbr. EtOH; also called ethyl alcohol, grain alcohol, drinking alcohol, or simply alcohol) is an organic compound. It is an alcohol with the chemical formula . Its formula can be also written as or (an ethyl group linked to a ...

. However, ''Saccharomyces'' cannot ferment or utilize

pentose In chemistry, a pentose is a monosaccharide (simple sugar) with five carbon atoms. The chemical formula of many pentoses is , and their molecular weight is 150.13 g/mol.arabinose) which is usually present in small amount in wines as residual sugars. In addition to ''Saccharomyces cerevisiae'', other species within the genus ''Saccharomyces'' that are involved with winemaking include: *'' Saccharomyces bayanus'' *'' Saccharomyces beticus'' *'' Saccharomyces fermentati'' *'' Saccharomyces paradoxus'' *''

Saccharomyces pastorianus ''Saccharomyces pastorianus'' is a yeast used industrially for the production of lager beer, and was named in honour of Louis Pasteur by the German Max Reess in 1870. This yeast's complicated genome appears to be the result of hybridisation bet ...

'' *'' Saccharomyces uvarum''

Influences of different strains on fermentation

In 1996, ''Saccharomyces cerevisiae'' was the first single-celled,

eukaryotic Eukaryotes () are organisms whose Cell (biology), cells have a cell nucleus, nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the ...

organism to have its entire

genome In the fields of molecular biology and genetics, a genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding g ...

sequenced In genetics and biochemistry, sequencing means to determine the primary structure (sometimes incorrectly called the primary sequence) of an unbranched biopolymer. Sequencing results in a symbolic linear depiction known as a sequence which suc ...

. This sequencing helped confirm the nearly century of work by mycologists and enologists in identifying different strains of ''Saccharomyces cerevisiae'' that are used in

beer Beer is one of the oldest and the most widely consumed type of alcoholic drink in the world, and the third most popular drink overall after water and tea. It is produced by the brewing and fermentation of starches, mainly derived from ce ...

, bread and

. Today there are several hundred different strains of ''S. cerevisiae'' identified. Not all of the strains are suitable for winemaking and even among the strains that are, there is debate among winemakers and scientists about the actual magnitude of differences between the various strains and their potential impact on the wine. Even among strains that have demonstrated distinctive difference when compared among young wines, these differences seem to fade and become less distinctive as the wines age. Some distinct difference among various strains include the production of certain "off-flavor" and aromas that may be temporary (but producing a "stinky fermentation") or could stay with the wine and either have to be dealt with through other winemaking means (such as the presence of volatile sulfur compounds like hydrogen sulfide) or leave a faulty wine. Another difference includes the "vigor" or speed of fermentation (which can also be influenced by other factors beyond yeast selection) with some yeast strains having the tendency to do "fast ferments" while others may take longer to get going. Another less measurable difference that are subject to more debate and questions of winemakers preference is the influence of strain selection on the

varietal A varietal wine is a wine made primarily from a single named grape variety, and which typically displays the name of that variety on the wine label.The American Heritage Dictionary of the English Language, Fourth Edition, 2000.winepros.com.au. ...

flavors of certainly grape varieties such as Sauvignon blanc and

Sémillon Sémillon is a golden-skinned grape used to make dry and sweet white wines, mostly in France and Australia. Its thin skin and susceptibility to botrytis make it dominate the sweet wine region Sauternes AOC and Barsac AOC. History The Sémil ...

. It is believed that these wines can be influenced by

thiols In organic chemistry, a thiol (; ), or thiol derivative, is any organosulfur compound of the form , where R represents an alkyl or other organic substituent. The functional group itself is referred to as either a thiol group or a sulfhydryl grou ...

produced by the

hydrolysis Hydrolysis (; ) is any chemical reaction in which a molecule of water breaks one or more chemical bonds. The term is used broadly for substitution, elimination, and solvation reactions in which water is the nucleophile. Biological hydrolys ...

of certain cysteine-linked compounds by enzymes that are more prevalent in particular strains. Other aromatic varieties such as

Gewürztraminer Gewürztraminer () is an aromatic wine grape variety, used in white wines, and performs best in cooler climates. In English, it is sometimes referred to colloquially as Gewürz (; although this is never the case in German, because "Gewürz" me ...

Riesling Riesling (, ; ) is a white grape variety that originated in the Rhine region. Riesling is an aromatic grape variety displaying flowery, almost perfumed, aromas as well as high acidity. It is used to make dry, semi-sweet, sweet, and sparkling wh ...

and

Muscat Muscat ( ar, مَسْقَط, ) is the capital and most populated city in Oman. It is the seat of the Governorate of Muscat. According to the National Centre for Statistics and Information (NCSI), the total population of Muscat Governorate was ...

may also be influenced by yeast strains containing high levels of

glycosidases Glycoside hydrolases (also called glycosidases or glycosyl hydrolases) catalyze Catalysis () is the process of increasing the rate of a chemical reaction by adding a substance known as a catalyst (). Catalysts are not consumed in the rea ...

enzymes that can modify monoterpenes. Similarly, though potentially to a much smaller extent, other varieties could be influenced by hydrolytic enzymes working on

aliphatics In organic chemistry, hydrocarbons ( compounds composed solely of carbon and hydrogen) are divided into two classes: aromatic compounds and aliphatic compounds (; G. ''aleiphar'', fat, oil). Aliphatic compounds can be saturated, like hexane, o ...

, norisoprenoids, and

benzene Benzene is an organic chemical compound with the molecular formula C6H6. The benzene molecule is composed of six carbon atoms joined in a planar ring with one hydrogen atom attached to each. Because it contains only carbon and hydrogen atoms ...

derivatives such as

polyphenols Polyphenols () are a large family of naturally occurring organic compounds characterized by multiples of phenol units. They are abundant in plants and structurally diverse. Polyphenols include flavonoids, tannic acid, and ellagitannin, some of ...

in the must. In

sparkling wine production Sparkling wine production is the method of winemaking used to produce sparkling wine. The oldest known production of sparkling wine took place in 1531 with the ''ancestral method''. Pressure and terminology In popular parlance and also in the ...

some winemakers select strains (such as one known as ''Épernay'' named after the town in the

Champagne wine region The wine region within the historical province of Champagne in the northeast of France is best known for the production of champagne, the sparkling white wine that bears the region's name. EU law and the laws of most countries reserve the term " ...

France France (), officially the French Republic ( ), is a country primarily located in Western Europe. It also comprises of overseas regions and territories in the Americas and the Atlantic, Pacific and Indian Oceans. Its metropolitan area ...

and ''California Champagne'', also known as

UC-Davis The University of California, Davis (UC Davis, UCD, or Davis) is a public land-grant research university near Davis, California. Named a Public Ivy, it is the northernmost of the ten campuses of the University of California system. The institu ...

strain 505) that are known to

flocculate Flocculation, in the field of chemistry, is a process by which colloidal particles come out of suspension to sediment under the form of floc or flake, either spontaneously or due to the addition of a clarifying agent. The action differs from p ...

well, allowing the dead yeast cells to be removed easily by riddling and

disgorgement Disgorgement is defined by ''Black's Law Dictionary'' as "the act of giving up something (such as profits illegally obtained) on demand or by legal compulsion." Overview Disgorgement is a remedy or penalty used in US securities law. For exampl ...

. In

production, the surface film of yeast known as flor used to make the distinctive style of fino and manzanilla sherries comes from different strains of ''Saccharomyces cerevisiae'', though the commercial flor yeast available for inoculation is often from different species of ''Saccharomyces'', '' Saccharomyces beticus'', '' Saccharomyces fermentati'' and '' Saccharomyces bayanus''.

Wild yeasts and natural fermentation

In winemaking, the term "wild yeast" has multiple meanings. In its most basic context, it refers to yeast that has not been introduced to the must by intentional inoculation of a cultured strain. Instead, these "wild yeasts" often come into contact with the must through their presence on harvest equipment, transport bins, the surface winemaking equipment and as part of the natural

flora Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous (ecology), indigenous) native plant, native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' ...

of a winery. Very often these are strains of ''Saccharomyces cerevisiae'' that have taken residence in these places over the years, sometimes being previously introduced by inoculation of prior vintages. In this context, these wild yeasts are often referred to as ambient, indigenous or natural yeast as opposed to inoculated, selected or cultured yeast. Wineries that often solely rely on these "in-house" strains will sometimes market their wines as being the product of wild or ''natural fermentations''. The (c. 304) '' Nanfang Caomu Zhuang'' has the earliest description of winemaking using " herb ferment" (''cǎoqū'' 草麴) wild yeast with rice and various herbs, including the poisonous ''

Gelsemium elegans ''Gelsemium elegans'', commonly known as heartbreak grass, is a poisonous plant of the family Gelsemiaceae found in China and other Asian countries. It contains toxic alkaloids such as gelsemine, gelsenicine, gelsevirine and koumine. Crumbled ...

'' (''yěgé'' 冶葛).Li Hui-Lin (1979), ''Nan-fang ts'ao-mu chuang: a fourth century flora of Southeast Asia'', The Chinese University Press, p. 59. Another use of the term "wild yeast" refers to the non-''Saccharomyces'' genera of yeasts that are present in the vineyard, on the surface of

grapevines ''Vitis'' (grapevine) is a genus of 79 accepted species of vining plants in the flowering plant family Vitaceae. The genus is made up of species predominantly from the Northern Hemisphere. It is economically important as the source of grapes, ...

and of the grapes themselves. Anywhere from 160 to 100,000 colony forming units of wild yeasts per berry could exist in a typical vineyard. These yeasts can be carried by air currents, birds and insects through the vineyard and even into the winery (such as by

fruit flies Fruit fly may refer to: Organisms * Drosophilidae, a family of small flies, including: ** ''Drosophila'', the genus of small fruit flies and vinegar flies ** ''Drosophila melanogaster'' or common fruit fly ** '' Drosophila suzukii'' or Asian frui ...

). The most common wild yeasts found in the vineyard are from the genera '' Kloeckera'', '' Candida'' and '' Pichia'' with the species ''

Kloeckera apiculata ''Hanseniaspora guilliermondii'' is a species of yeast in the family Saccharomycetaceae. In its anamorph form, it is called ''Kloeckera apis''. Taxonomy The initial sample of the species was isolated by South African pathologist Adrianus P ...

'' being the most dominant species by far. ''Saccharomyces cerevisiae'', itself, is actually quite rarely found in the vineyard or on the surface freshly harvested wine grapes unless the winery frequently reintroduced winery waste (such as lees and

pomace Pomace ( ), or marc (; from French ''marc'' ), is the solid remains of grapes, olives, or other fruit after pressing for juice or oil. It contains the skins, pulp, seeds, and stems of the fruit. Grape pomace has traditionally been used to pro ...

) into the vineyard. Unlike the "ambient" ''Saccharomyces'' wild yeast, these genera of wild yeasts have very low tolerance to both alcohol and sulfur dioxide. They are capable of starting a fermentation and often begin this process as early as the harvest bin when clusters of grapes get slightly crushed under their own weight. Some winemakers will try to "knock out" these yeasts with doses of sulfur dioxide, most often at the crusher before the grapes are pressed or allowed to macerate with skin contact. Other winemakers may allow the wild yeasts to continue fermenting until they succumb to the toxicity of the alcohol they produce which is often between 3–5% alcohol by volume and then letting either inoculated or "ambient" ''Saccharomyces'' strains finish the fermentation. Pomace in the vineyard after pressing

The use of both "ambient" and non-''Saccharomyces'' wild yeasts carries both potential benefits and risk. Some winemakers feel that the use of resident/indigenous yeast helps contribute to the unique expression of ''

terroir (, ; from ''terre'', "land") is a French term used to describe the environmental factors that affect a crop's phenotype, including unique environment contexts, farming practices and a crop's specific growth habitat. Collectively, these contex ...

'' in the wine. In wine regions such as

Bordeaux Bordeaux ( , ; Gascon oc, Bordèu ; eu, Bordele; it, Bordò; es, Burdeos) is a port city on the river Garonne in the Gironde department, Southwestern France. It is the capital of the Nouvelle-Aquitaine region, as well as the prefect ...

, classified and highly regarded estates will often tout the quality of their resident "chateau" strains. To this extent, wineries will often take the leftover

and lees from winemaking and return them to the vineyard to be used as compost in order to encourage the sustained presence of favorable strains. But compared to inoculated yeast, these ambient yeasts hold the risk of having a more unpredictable fermentation. Not only could this unpredictability include the presence of off-flavors/aromas and higher

but also the potential for a stuck fermentation if the indigenous yeast strains are not vigorous enough to fully convert all the sugars. It is virtually inevitable that non-''Saccharomyces'' wild yeast will have a role in beginning the fermentation of virtually every wine but for the wineries that choose to allow these yeasts to continue fermenting versus minimizing their influence do so with the intent of enhancing complexity through bio-diversity. While these non-''Saccharomyces'' ferment glucose and fructose into alcohol, they also have the potential to create other intermediates that could influence the aroma and flavor profile of the wine. Some of these intermediates could be positive, such as phenylethanol, which can impart a

rose A rose is either a woody perennial flowering plant of the genus ''Rosa'' (), in the family Rosaceae (), or the flower it bears. There are over three hundred species and tens of thousands of cultivars. They form a group of plants that can be ...

-like aroma. However, as with ambient yeasts, the products of these yeasts can be very unpredictable – especially in terms of the types of flavors and aromas that these yeasts can produce.

Inoculated yeast

When winemakers select a cultured yeast strain, it is largely done because the winemaker wants a predictable fermentation taken to completion by a strain that has a track record of dependability. Among the particular considerations that are often important to winemakers is a yeast's tendency to: *Quickly begin fermentation, out-competing other "wild yeasts" for nutrients in the must *Completely utilize all fermentable sugars with a predictable sugar-to-alcohol conversion rate *Have an alcohol-tolerance up to 15% or even higher depending on the winemaking style *Have a high sulfur dioxide tolerance but low production of sulfur compounds such as hydrogen sulfide or dimethyl sulfide *Produce a minimum amount of residual pyruvate, acetic acid and acetaldehyde *Produce minimum foaming during fermentation which may create difficulties for cap management during maceration or cause bungs to pop out during barrel fermentation. *Have high levels of

flocculation Flocculation, in the field of chemistry, is a process by which colloidal particles come out of suspension to sediment under the form of floc or flake, either spontaneously or due to the addition of a clarifying agent. The action differs from ...

and lees compaction that makes

, fining and filtering of the wine easier. Inoculated (or ''pure cultured'') yeasts are strains of ''Saccharomyces cerevisiae'' that have been identified and plated from wineries across the world (including notable producers from well-known wine regions such as

, Burgundy,

Napa Valley Napa Valley is an American Viticultural Area (AVA) located in Napa County in California's Wine Country. It was established by the Bureau of Alcohol, Tobacco and Firearms (ATF) on January 27, 1981. Napa Valley is considered one of the premier ...

and the

Barossa Valley The Barossa Valley (Barossa German: ''Barossa Tal'') is a valley in South Australia located northeast of Adelaide city centre. The valley is formed by the North Para River. It is notable as a major wine-producing region and tourist destina ...

). These strains are tested in laboratories to determine a strain's vigor, sulfur dioxide and alcohol tolerance, production levels of acetic acid and sulfur compounds, ability to re-ferment (positive for sparkling wine but a negative attribute for sweet late-harvest wines), development of surface film on the wine (positive for some Sherry styles but a negative attribute for many other wines), enhancement of a wine's color or certain varietal characteristics by enzymes in the yeast cells and other metabolic products produced by the yeast, foaming and flocculation tendencies, yeasticidal properties (a trait known as "

Killer yeast A killer yeast is a yeast, such as ''Saccharomyces cerevisiae'', which is able to secrete one of a number of toxic proteins which are lethal to susceptible cells. These "killer toxins" are polypeptides that kill sensitive cells of the same or relat ...

") and tolerance for nutritional deficiencies in a must that may lead to a stuck fermentation.

Re-hydrating freeze dried yeast cultures

Adding Agria wine to rehydrated yeast to prepare culture inoculum

Pure culture yeasts that are grown in a lab are often

freeze dried Freeze drying, also known as lyophilization or cryodesiccation, is a low temperature dehydration process that involves freezing the product and lowering pressure, removing the ice by sublimation. This is in contrast to dehydration by most conve ...

and packaged for commercial use. Prior to their addition into must, these yeasts need to be re-hydrated in "starter cultures" that must be carefully monitored (particularly in regards to temperature) to ensure that the yeast cells are not killed off by

cold shock Cold shock response is a series of neurogenic cardio-respiratory responses caused by sudden immersion in cold water. In cold water immersions, such as by falling through thin ice, cold shock response is perhaps the most common cause of death. Als ...

. Ideally winemakers want to add enough inoculum to have a viable cell population density of 5 million cells per milliliter. The exact amount of freeze-dried culture varies by manufacturer and strain of yeast but it is often around 1 gram per gallon (or 25 grams per 100 liters). Wines that could have potentially problematic fermentation (such as high sugar level late harvest or botryized wines) may have more yeast added. Similarly, re-hydration procedures will also vary depending on the manufacturer and winery. Yeast is often inoculated in a volume of water or grape must that is 5–10 times the weight of the dry yeast. This liquid is often brought to temperature of 40 °C (104 °F) prior to the introduction of the yeast (though some yeast strains may need temperatures below 38 °C (100 °F)) to allow the cells to disperse easily rather than clump and sink to the bottom of the container. The heat activation also allows the cells to quickly reestablish their membrane barrier before soluble cytoplasmic components escape the cell. Re-hydration at lower temperatures can greatly reduce the viability of the yeast with up to 60% cell death if the yeast is re-hydrated at 15 °C (60 °F). The culture is then stirred and aerated to incorporate oxygen into the culture which the yeast uses in the synthesis of needed survival factors. The temperature of the starter culture is then slowly reduced, often by the graduated addition of must to get within 5–10 °C (9–18 °F) of the must that the culture will be added to. This is done to avoid the sudden cold shock that the yeast cells may experience if the starter culture was added directly to the must itself which can kill up to 60% of the culture. Additionally, surviving cells exposed to cold shock tend to see an increase in hydrogen sulfide production.

Nutritional needs of wine yeast

In order to successfully complete a fermentation with minimum to no negative attributes being added to the wine, yeast needs to have the full assortment of its nutritional needs met. These include not only an available energy source (carbon in the form of sugars such as glucose) and

yeast assimilable nitrogen Yeast assimilable nitrogen or YAN is the combination of free amino nitrogen (FAN), ammonia (NH3) and ammonium (NH4+) that is available for a yeast, e.g. the wine yeast ''Saccharomyces cerevisiae'', to use during fermentation. Outside of the f ...

(

ammonia Ammonia is an inorganic compound of nitrogen and hydrogen with the formula . A stable binary hydride, and the simplest pnictogen hydride, ammonia is a colourless gas with a distinct pungent smell. Biologically, it is a common nitrogenous wa ...

and amino acids or YAN) but also

mineral In geology and mineralogy, a mineral or mineral species is, broadly speaking, a solid chemical compound with a fairly well-defined chemical composition and a specific crystal structure that occurs naturally in pure form.John P. Rafferty, ed. (2 ...

s (such as

magnesium Magnesium is a chemical element with the symbol Mg and atomic number 12. It is a shiny gray metal having a low density, low melting point and high chemical reactivity. Like the other alkaline earth metals (group 2 of the periodic ta ...

) and

vitamin A vitamin is an organic molecule (or a set of molecules closely related chemically, i.e. vitamers) that is an essential micronutrient that an organism needs in small quantities for the proper functioning of its metabolism. Essential nutrie ...

s (such as

thiamin Thiamine, also known as thiamin and vitamin B1, is a vitamin, an essential micronutrient, that cannot be made in the body. It is found in food and commercially synthesized to be a dietary supplement or medication. Phosphorylated forms of t ...

and riboflavin) that serve as important growth and survival factors. Among the other nutritional needs of wine yeast: *

Phosphate In chemistry, a phosphate is an anion, salt, functional group or ester derived from a phosphoric acid. It most commonly means orthophosphate, a derivative of orthophosphoric acid . The phosphate or orthophosphate ion is derived from phosph ...

– used for the production of

nucleic acids Nucleic acids are biopolymers, macromolecules, essential to all known forms of life. They are composed of nucleotides, which are the monomers made of three components: a 5-carbon sugar, a phosphate group and a nitrogenous base. The two main cl ...

phospholipids Phospholipids, are a class of lipids whose molecule has a hydrophilic "head" containing a phosphate group and two hydrophobic "tails" derived from fatty acids, joined by an alcohol residue (usually a glycerol molecule). Marine phospholipids t ...

(an important component of the cell membrane) and ATP (

Adenosine triphosphate Adenosine triphosphate (ATP) is an organic compound that provides energy to drive many processes in living cells, such as muscle contraction, nerve impulse propagation, condensate dissolution, and chemical synthesis. Found in all known forms o ...

which the cell uses for transferring energy for metabolism). *

Potassium Potassium is the chemical element with the symbol K (from Neo-Latin ''kalium'') and atomic number19. Potassium is a silvery-white metal that is soft enough to be cut with a knife with little force. Potassium metal reacts rapidly with atmosph ...

– important for the uptake and utilization of phosphate * Biotin – involved in the synthesis of proteins,

fatty acids In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated and unsaturated compounds#Organic chemistry, saturated or unsaturated. Most naturally occurring fatty acids have an B ...

and nucleic acids. *

Pantothenic acid Pantothenic acid, also called vitamin B5 is a water-soluble B vitamin and therefore an essential nutrient. All animals require pantothenic acid in order to synthesize coenzyme A (CoA) – essential for fatty acid metabolism – as well as to, ...

– involved in the metabolism of sugars and

lipid Lipids are a broad group of naturally-occurring molecules which includes fats, waxes, sterols, fat-soluble vitamins (such as vitamins A, D, E and K), monoglycerides, diglycerides, phospholipids, and others. The functions of lipids includ ...

s. A deficiency of this vitamin could lead into increase hydrogen sulfide production with off-aromas in the resulting wine. * Nicotinic acid – involved in the synthesis of Nicotinamide adenine dinucleotide (NAD+), a co-enzyme that is important in maintaining the

balance of the cell as well as in the process of

ethanol fermentation Ethanol fermentation, also called alcoholic fermentation, is a biological process which converts sugars such as glucose, fructose, and sucrose into cellular energy, producing ethanol and carbon dioxide as by-products. Because yeasts perform this ...

itself. * Inositol – involved with the

secondary messenger Second messengers are intracellular signaling molecules released by the cell in response to exposure to extracellular signaling molecules—the first messengers. (Intercellular signals, a non-local form or cell signaling, encompassing both first m ...

molecules that facilitate

cell division Cell division is the process by which a parent cell divides into two daughter cells. Cell division usually occurs as part of a larger cell cycle in which the cell grows and replicates its chromosome(s) before dividing. In eukaryotes, there ar ...

. *Trace amounts of

calcium Calcium is a chemical element with the symbol Ca and atomic number 20. As an alkaline earth metal, calcium is a reactive metal that forms a dark oxide-nitride layer when exposed to air. Its physical and chemical properties are most similar t ...

chlorine Chlorine is a chemical element with the symbol Cl and atomic number 17. The second-lightest of the halogens, it appears between fluorine and bromine in the periodic table and its properties are mostly intermediate between them. Chlorine i ...

copper Copper is a chemical element with the symbol Cu (from la, cuprum) and atomic number 29. It is a soft, malleable, and ductile metal with very high thermal and electrical conductivity. A freshly exposed surface of pure copper has a pinkis ...

iron Iron () is a chemical element with Symbol (chemistry), symbol Fe (from la, Wikt:ferrum, ferrum) and atomic number 26. It is a metal that belongs to the first transition series and group 8 element, group 8 of the periodic table. It is, Abundanc ...

manganese Manganese is a chemical element with the symbol Mn and atomic number 25. It is a hard, brittle, silvery metal, often found in minerals in combination with iron. Manganese is a transition metal with a multifaceted array of industrial alloy use ...

and

zinc Zinc is a chemical element with the symbol Zn and atomic number 30. Zinc is a slightly brittle metal at room temperature and has a shiny-greyish appearance when oxidation is removed. It is the first element in group 12 (IIB) of the periodi ...

for healthy cell function. Red wine grape pomace

Many of these nutrients are available in the must and skins of the grapes themselves but sometimes are supplemented by winemakers with additions such as

diammonium phosphate Diammonium phosphate (DAP; IUPAC name diammonium hydrogen phosphate; chemical formula (NH4)2(HPO4) is one of a series of water-soluble ammonium phosphate salts that can be produced when ammonia reacts with phosphoric acid. Solid diammonium phosp ...

(DAP), freeze-dried micro-nutrients (such as ''Go-Ferm'' and ''Ferm-K'') and even the remnant of dead or extracted yeast cells such that the fermenting yeast can break down to mine for available nitrogen and nutrients. One historical winemaking tradition that is still practiced in some

Italian wine Italian wine is produced in every region of Italy. Italy is the world's largest producer of wine, with an area of under vineyard cultivation, and contributing a 2013–2017 annual average of 48.3 million hl of wine. In 2018 Italy accounted fo ...

regions is the ''

ripasso Valpolicella (, , ) is a viticultural zone of the province of Verona, Italy, east of Lake Garda. The hilly agricultural and marble-quarrying region of small holdings north of the Adige is famous for wine production. Valpolicella ranks just after ...

'' method of adding the leftover

from the pressing of other wines into a newly fermenting batch of wine as an additional food source for the yeast. ''Saccharomyces cerevisiae'' can assimilate nitrogen from both inorganic (ammonia and ammonium) and organic forms (amino acids, particularly arginine). As yeast cells die, enzymes within the cells begin autolyzing by breaking down the cell, including the amino acids. This autolysis of the cell provides an available nitrogen source for the still-fermenting and viable yeast cells. However, this autolysis can also release sulfur-link compounds (such as the breakdown of amino acid cysteine) which can combine with other molecules and react with alcohol to create volatile

that can contribute to a "stinky fermentation" or later development into various wine faults.

The role of oxygen

Yeasts are

facultative anaerobes A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...

meaning that they can exist in both the presence and absence of oxygen. While fermentation is traditionally thought of as an

anaerobic Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to: * Anaerobic adhesive, a bonding a ...

process done in the absence of oxygen, early exposure of the yeast to oxygen can be a vital component in the successful completion of that fermentation. This is because oxygen is important in the synthesis of cell "survival factors" such as ergosterol and

lanosterol Lanosterol is a tetracyclic triterpenoid and is the compound from which all animal and fungal steroids are derived. By contrast plant steroids are produced via cycloartenol. Role in biosynthesis of other steroids Elaboration of lanosterol under en ...

. These sterols are important in maintaining the

selective permeability Semipermeable membrane is a type of biological or synthetic, polymeric membrane that will allow certain molecules or ions to pass through it by osmosis. The rate of passage depends on the pressure, concentration, and temperature of the molecule ...

of the yeast cell membrane which becomes critical as the yeast becomes exposed to increasing osmotic pressure and levels of alcohol in the wine. As a waste product of its own metabolism, alcohol is actually very toxic to yeast cells. Yeast with weak survival factors and lacking sterols may succumb to these conditions before fermenting a wine to complete dryness, leaving a stuck fermentation. Cultured yeasts that are freeze-dried and available for inoculation of wine must are deliberately grown in commercial labs in high oxygen/low sugar conditions that favor the development of these survival factors. One of the reasons that some winemakers prefer using inoculated yeast is the predictability of fermentation due to the high level of survival factors that cultured yeast are assured of having without necessarily needing to expose the wine to additional levels of oxygen. Winemakers using "ambient" yeasts that are resident in their winery may not have this same assurance of survival factors and may need to compensate with other winemaking techniques. Wild non-''Saccharomyces'' yeasts often need a much greater exposure to oxygen in order to build up survival factors which is why many of these yeasts are often found living oxidatively as "film yeast" on the surface of wines in tanks or barrels.

Wine faults related to yeast

Either directly or indirectly, wine yeast can be a culprit behind a wide variety of

. These can include the presence of " off flavors" and aromas that can be the by-product of some "wild yeast" fermentation such as those by species within the genera of ''Kloeckera'' and ''Candida''. Even the common wine yeast ''Saccharomyces cerevisiae'' can be behind some wine faults with some strains of the yeast known to produce higher than ideal levels of acetic acid, acetaldehyde and volatile sulfur compounds such as

. Also any yeast can have a low tolerance to nutritional deficiencies, temperature fluctuation or extremes and excessive or low sugar levels that may lead to a stuck fermentation. In the presence of

oxygen Oxygen is the chemical element with the symbol O and atomic number 8. It is a member of the chalcogen group in the periodic table, a highly reactive nonmetal, and an oxidizing agent that readily forms oxides with most elements as ...

several species of ''Candida'' and ''Pichia'' can create a film surface on top of the wine in the tank of barrel. Allowed to go unchecked, these yeasts can rapidly deplete the available free sulfur compounds that keeps a wine protected from oxidation and other microbial attack. The presence of these yeasts is often identified by elevated levels of

, particularly acetic acid. Some strains of ''Pichia'' will metabolize acetic acid (as well as

and isoamyl acetate that may also be produced) with the side-effect of substantially decreasing the titratable acidity and shifting the pH of wine upwards to levels that make the wine prone to attack by other spoilage microbes. Commonly called "film yeast", these yeasts are distinguished from the flor sherry yeast that are usually welcomed by winemakers in producing the delicate fino-style wines. Growth of many unfavorable wild yeasts is generally slowed at lower cellar temperatures, so many winemakers who wish to inhibit the activities of these yeasts before the more favorable ''Saccharomyces'' yeast kick in, will often chill their must, such as the practice of "cold soaking" the must during a pre-fermentation maceration at temperatures between 4–15 °C (39–50 °F). Though some species, such as ''Brettanomyces'', will not be inhibited and may even thrive during an extended period of cold soaking.

Brettanomyces

The wine yeast ''Brettanomyces'' (or "Brett") produces very distinctive aroma compounds,

4-Ethylphenol Ethylphenol (4-EP) is an organic compound with the formula C2H5C6H4OH. It is one of three isomeric ethylphenols. A white solid, it occurs as an impurity in xylenols and as such is used in the production of some commercial phenolic resins. It is ...

(4-EP) and 4-Ethylguaiacol (4-EG), that can have a wine being described as smelling like a "barnyard", "wet saddle" or "band-aid". To some winemakers and with some wine styles (such as Pinot noir from Burgundy), a limited amount of these compounds could be considered a positive attribute that adds to the complexity of wine. To other winemakers and with other wine styles (such as

from the Mosel), the presence of any Brett will be considered a fault.M. Baldy ''The University Wine Course'' p. 80 The Wine Appreciation Guild Third Edition 2009

Fruit flies Fruit fly may refer to: Organisms * Drosophilidae, a family of small flies, including: ** ''Drosophila'', the genus of small fruit flies and vinegar flies ** ''Drosophila melanogaster'' or common fruit fly ** '' Drosophila suzukii'' or Asian frui ...

are common

vector Vector most often refers to: *Euclidean vector, a quantity with a magnitude and a direction *Vector (epidemiology), an agent that carries and transmits an infectious pathogen into another living organism Vector may also refer to: Mathematic ...

in the transfer of ''Brettanomyces'' between tanks and even nearby wineries. As a fermentation yeast, ''Brettanomyces'' can usually ferment a wine up to 10–11% alcohol levels before they die out. Sometimes ''Brettanomyces'' already present in a wine that has been inoculated with ''Saccharomyces cerevisiae'' will out compete the ''Saccharomyces'' strain for nutrients and even inhibit it due to the high levels of acetic acid,

decanoic acid Capric acid, also known as decanoic acid or decylic acid, is a saturated fatty acid, medium-chain fatty acid (MCFA), and carboxylic acid. Its formula is CH3(CH2)8COOH. Salts and esters of decanoic acid are called caprates or decanoates. The te ...

and

octanoic acid Caprylic acid (), also known under the systematic name octanoic acid or C8 Acid, is a saturated fatty acid, medium-chain fatty acid (MCFA). It has the structural formula , and is a colorless oily liquid that is minimally soluble in water with a s ...

that many strains of ''Brettanomyces'' can produce. Once Brett is in a winery, it is very difficult to control even with strict hygiene and the discarding of barrels and equipment that has previously come into contact with "Bretty" wine. This is because many species of ''Brettanomyces'' can use a wide variety of carbon sources in wine and grape must, including

, for metabolism. Additionally, Brett can produce a wide range of by-products that could influence the wine beyond just the 4-EP and 4-EG compounds previously discussed. Many of these compounds, such as the "footprints" of the 4-EP and 4-EG, will still remain in the wine even after yeast cells die and are removed by racking and sterile filtration.

References

External links

*P. Romano, C. Fiore, M. Paraggio, M. Caruso, A. Capece '
Function of yeast species and strains in wine flavour
'' International Journal of Food Microbiology 86 (2003) 169–180 {{Authority control Food microbiology Saccharomycetes Winemaking