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Yeasts are
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 ...
, single-celled
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s classified as members of the
fungus A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from t ...
kingdom Kingdom commonly refers to: * A monarchy ruled by a king or queen * Kingdom (biology), a category in biological taxonomy Kingdom may also refer to: Arts and media Television * ''Kingdom'' (British TV series), a 2007 British television drama s ...
. The first yeast originated hundreds of millions of years ago, and at least 1,500
species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...
are currently recognized. They are estimated to constitute 1% of all described fungal species. Yeasts are
unicellular organism A unicellular organism, also known as a single-celled organism, is an organism that consists of a single cell, unlike a multicellular organism that consists of multiple cells. Organisms fall into two general categories: prokaryotic organisms a ...
s that evolved from
multicellular A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially un ...
ancestors, with some species having the ability to develop multicellular characteristics by forming strings of connected budding cells known as pseudohyphae or false hyphae. Yeast sizes vary greatly, depending on species and environment, typically measuring 3–4  µm in
diameter In geometry, a diameter of a circle is any straight line segment that passes through the center of the circle and whose endpoints lie on the circle. It can also be defined as the longest chord of the circle. Both definitions are also valid fo ...
, although some yeasts can grow to 40 µm in size. Most yeasts reproduce asexually by mitosis, and many do so by the asymmetric division process known as
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 ...
. With their single-celled growth habit, yeasts can be contrasted with
mold A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. Not ...
s, which grow hyphae. Fungal species that can take both forms (depending on temperature or other conditions) are called
dimorphic fungi Dimorphic fungi are fungi that can exist in the form of both mold and yeast. This is usually brought about by change in temperature and the fungi are also described as thermally dimorphic fungi. An example is ''Talaromyces marneffei'', a human ...
. The yeast species ''
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 ...
'' converts
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or m ...
s to
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 ...
and alcohols through the process of fermentation. The products of this reaction have been used in baking and the production of alcoholic beverages for thousands of years. ''S. cerevisiae'' is also an important model organism in modern cell biology research, and is one of the most thoroughly studied eukaryotic microorganisms. Researchers have cultured it in order to understand the biology of the eukaryotic cell and ultimately human biology in great detail. Other species of yeasts, such as ''
Candida albicans ''Candida albicans'' is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usu ...
'', are
opportunistic pathogen An opportunistic infection is an infection caused by pathogens (bacteria, fungi, parasites or viruses) that take advantage of an opportunity not normally available. These opportunities can stem from a variety of sources, such as a weakened immune ...
s and can cause
infections An infection is the invasion of tissues by pathogens, their multiplication, and the reaction of host tissues to the infectious agent and the toxins they produce. An infectious disease, also known as a transmissible disease or communicable di ...
in humans. Yeasts have recently been used to generate electricity in
microbial fuel cell Microbial fuel cell (MFC) is a type of bioelectrochemical fuel cell system that generates electric current by diverting electrons produced from the microbial oxidation of reduced compounds (also known as fuel or electron donor) on the anode to oxid ...
s and to produce
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 ...
for the biofuel industry. Yeasts do not form a single taxonomic or
phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
grouping. The term "yeast" is often taken as a synonym for ''Saccharomyces cerevisiae'', but the phylogenetic diversity of yeasts is shown by their placement in two separate phyla: the Ascomycota and the Basidiomycota. The budding yeasts or "true yeasts" are classified in the order
Saccharomycetales Saccharomycetales belongs to the kingdom of Fungi and the division Ascomycota. It is the only order in the class Saccharomycetes. There are currently 13 families recognized as belonging to Saccharomycetales. GBIF also includes; Alloascoideacea ...
, within the phylum Ascomycota.


History

The word "yeast" comes from
Old English Old English (, ), or Anglo-Saxon, is the earliest recorded form of the English language, spoken in England and southern and eastern Scotland in the early Middle Ages. It was brought to Great Britain by Anglo-Saxon settlers in the mid-5th c ...
''gist'', ''gyst'', and from the
Indo-European The Indo-European languages are a language family native to the overwhelming majority of Europe, the Iranian plateau, and the northern Indian subcontinent. Some European languages of this family, English, French, Portuguese, Russian, Dutc ...
root ''yes-'', meaning "boil", "foam", or "bubble". Yeast microbes are probably one of the earliest domesticated organisms. Archaeologists digging in Egyptian ruins found early grinding stones and baking chambers for yeast-raised bread, as well as drawings of 4,000-year-old bakeries and breweries. Vessels studied from several archaeological sites in Israel (dating to around 5,000, 3,000 and 2,500 years ago), which were believed to have contained alcoholic beverages (beer and mead), were found to contain yeast colonies that had survived over the millennia, providing the first direct biological evidence of yeast use in early cultures. In 1680,
Dutch Dutch commonly refers to: * Something of, from, or related to the Netherlands * Dutch people () * Dutch language () Dutch may also refer to: Places * Dutch, West Virginia, a community in the United States * Pennsylvania Dutch Country People E ...
naturalist
Anton van Leeuwenhoek Antonie Philips van Leeuwenhoek ( ; ; 24 October 1632 – 26 August 1723) was a Dutch microbiologist and microscopist in the Golden Age of Dutch science and technology. A largely self-taught man in science, he is commonly known as " the ...
first
microscopically Microscopy is the technical field of using microscopes to view objects and areas of objects that cannot be seen with the naked eye (objects that are not within the resolution range of the normal eye). There are three well-known branches of micr ...
observed yeast, but at the time did not consider them to be
living organisms In biology, an organism () is any living system that functions as an individual entity. All organisms are composed of cells (cell theory). Organisms are classified by taxonomy into groups such as multicellular animals, plants, and fun ...
, but rather globular structures as researchers were doubtful whether yeasts were algae or fungi.
Theodor Schwann Theodor Schwann (; 7 December 181011 January 1882) was a German physician and physiologist. His most significant contribution to biology is considered to be the extension of cell theory to animals. Other contributions include the discovery of ...
recognized them as fungi in 1837. In 1857, French microbiologist Louis Pasteur showed that by bubbling oxygen into the yeast broth, cell growth could be increased, but fermentation was inhibited – an observation later called the " Pasteur effect". In the paper "''Mémoire sur la fermentation alcoolique,''" Pasteur proved that alcoholic fermentation was conducted by living yeasts and not by a chemical catalyst. By the late 18th century two yeast strains used in brewing had been identified: ''Saccharomyces cerevisiae'' (top-fermenting yeast) and '' S. carlsbergensis'' (bottom-fermenting yeast). ''S. cerevisiae'' has been sold commercially by the Dutch for bread-making since 1780; while, around 1800, the Germans started producing ''S. cerevisiae'' in the form of cream. In 1825, a method was developed to remove the liquid so the yeast could be prepared as solid blocks. The industrial production of yeast blocks was enhanced by the introduction of the
filter press An industrial filter press is a tool used in separation processes, specifically to separate solids and liquids. The machine stacks many filter elements and allows the filter to be easily opened to remove the filtered solids, and allows easy cleanin ...
in 1867. In 1872, Baron Max de Springer developed a manufacturing process to create granulated yeast, a technique that was used until the first World War. In the United States, naturally occurring airborne yeasts were used almost exclusively until commercial yeast was marketed at the
Centennial Exposition The Centennial International Exhibition of 1876, the first official World's Fair to be held in the United States, was held in Philadelphia, Pennsylvania, from May 10 to November 10, 1876, to celebrate the 100th anniversary of the signing of the ...
in 1876 in Philadelphia, where Charles L. Fleischmann exhibited the product and a process to use it, as well as serving the resultant baked bread. The mechanical refrigerator (first patented in the 1850s in Europe) liberated brewers and
winemakers A winemaker or vintner is a person engaged in winemaking. They are generally employed by wineries or wine companies, where their work includes: *Cooperating with viticulturists *Monitoring the maturity of grapes to ensure their quality and to dete ...
from seasonal constraints for the first time and allowed them to exit cellars and other earthen environments. For
John Molson John Molson (December 28, 1763 – January 11, 1836) was an English-born brewer and entrepreneur in colonial Quebec, which during his lifetime became Lower Canada. In addition to founding Molson Brewery, he built the first steamship and the fir ...
, who made his livelihood in
Montreal Montreal ( ; officially Montréal, ) is the second-most populous city in Canada and most populous city in the Canadian province of Quebec. Founded in 1642 as '' Ville-Marie'', or "City of Mary", it is named after Mount Royal, the triple ...
prior to the development of the fridge, the brewing season lasted from September through to May. The same seasonal restrictions formerly governed the
distiller Distillation, or classical distillation, is the process of separating the components or substances from a liquid mixture by using selective boiling and condensation, usually inside an apparatus known as a still. Dry distillation is the heatin ...
's art.


Nutrition and growth

Yeasts are chemoorganotrophs, as they use
organic compound In chemistry, organic compounds are generally any chemical compounds that contain carbon-hydrogen or carbon-carbon bonds. Due to carbon's ability to catenate (form chains with other carbon atoms), millions of organic compounds are known. T ...
s as a source of energy and do not require sunlight to grow. Carbon is obtained mostly from
hexose In chemistry, a hexose is a monosaccharide (simple sugar) with six carbon atoms. The chemical formula for all hexoses is C6H12O6, and their molecular weight is 180.156 g/mol. Hexoses exist in two forms, open-chain or cyclic, that easily convert ...
sugars, such as
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 ...
and fructose, or disaccharides such as sucrose and maltose. Some species can metabolize
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.organic acids. Yeast species either require oxygen for aerobic
cellular respiration Cellular respiration is the process by which biological fuels are oxidised in the presence of an inorganic electron acceptor such as oxygen to produce large amounts of energy, to drive the bulk production of ATP. Cellular respiration may be des ...
(
obligate aerobe An obligate aerobe is an organism that requires oxygen to grow. Through cellular respiration, these organisms use oxygen to metabolise substances, like sugars or fats, to obtain energy. In this type of respiration, oxygen serves as the terminal ...
s) or are anaerobic, but also have aerobic methods of energy production (
facultative anaerobe 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 '' Staphylococc ...
s). Unlike
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
, no known yeast species grow only anaerobically (
obligate anaerobe Obligate anaerobes are microorganisms killed by normal atmospheric concentrations of oxygen (20.95% O2). Oxygen tolerance varies between species, with some species capable of surviving in up to 8% oxygen, while others lose viability in environme ...
s). Most yeasts grow best in a neutral or slightly acidic pH environment. Yeasts vary in regard to the temperature range in which they grow best. For example, '' Leucosporidium frigidum'' grows at , ''
Saccharomyces telluris ''Saccharomyces telluris'' is a species of 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 ...
'' at , and '' Candida slooffi'' at . The cells can survive freezing under certain conditions, with viability decreasing over time. In general, yeasts are grown in the laboratory on solid
growth media A growth medium or culture medium is a solid, liquid, or semi-solid designed to support the growth of a population of microorganisms or cells via the process of cell proliferation or small plants like the moss ''Physcomitrella patens''. Different ...
or in liquid
broth Broth, also known as bouillon (), is a savory liquid made of water in which meat, fish or vegetables have been simmered for a short period of time. It can be eaten alone, but it is most commonly used to prepare other dishes, such as soups, ...
s. Common media used for the cultivation of yeasts include
potato dextrose agar Potato dextrose agar (BAM Media M127
from the U.S.
or
potato dextrose broth Potato dextrose agar (BAM Media M127
from the U.S.
, Wallerstein Laboratories nutrient agar, yeast peptone dextrose agar, and yeast mould agar or broth. Home brewers who cultivate yeast frequently use dried
malt extract Malt is germinated cereal grain that has been dried in a process known as "malting". The grain is made to germinate by soaking in water and is then halted from germinating further by drying with hot air. Malted grain is used to make beer, whi ...
and agar as a solid growth medium. The fungicide cycloheximide is sometimes added to yeast growth media to inhibit the growth of ''
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 ...
'' yeasts and select for wild/indigenous yeast species. This will change the yeast process. The appearance of a white, thready yeast, commonly known as kahm yeast, is often a byproduct of the lactofermentation (or pickling) of certain vegetables. It is usually the result of exposure to air. Although harmless, it can give pickled vegetables a bad flavor and must be removed regularly during fermentation.


Ecology

Yeasts are very common in the environment, and are often isolated from sugar-rich materials. Examples include naturally occurring yeasts on the skins of fruits and berries (such as grapes, apples, or
peach The peach (''Prunus persica'') is a deciduous tree first domesticated and cultivated in Zhejiang province of Eastern China. It bears edible juicy fruits with various characteristics, most called peaches and others (the glossy-skinned, non-f ...
es), and exudates from plants (such as plant saps or cacti). Some yeasts are found in association with soil and insects. Yeasts from the soil and from the skins of fruits and berries have been shown to dominate fungal succession during fruit decay. The ecological function and
biodiversity Biodiversity or biological diversity is the variety and variability of life on Earth. Biodiversity is a measure of variation at the genetic (''genetic variability''), species (''species diversity''), and ecosystem (''ecosystem diversity'') l ...
of yeasts are relatively unknown compared to those of other
microorganism A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
s. Yeasts, including ''
Candida albicans ''Candida albicans'' is an opportunistic pathogenic yeast that is a common member of the human gut flora. It can also survive outside the human body. It is detected in the gastrointestinal tract and mouth in 40–60% of healthy adults. It is usu ...
'', ''
Rhodotorula ''Rhodotorula'' is a genus of pigmented yeasts, part of the division Basidiomycota. It is readily identifiable by distinctive orange/red colonies when grown on Sabouraud's dextrose agar (SDA). This distinctive color is the result of pigme ...
rubra'', '' Torulopsis'' and ''
Trichosporon cutaneum ''Trichosporon'' is a genus of anamorphic fungi in the family Trichosporonaceae. All species of ''Trichosporon'' are yeasts with no known teleomorphs (sexual states). Most are typically isolated from soil, but several species occur as a natura ...
'', have been found living in between people's toes as part of their skin flora. Yeasts are also present in the gut flora of mammals and some insects and even deep-sea environments host an array of yeasts. An Indian study of seven bee species and nine plant species found 45 species from 16 genera colonize the
nectaries Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
of flowers and honey stomachs of bees. Most were members of the genus '' Candida''; the most common species in honey stomachs was '' Dekkera intermedia'' and in flower nectaries, '' Candida blankii''. Yeast colonising nectaries of the
stinking hellebore ''Helleborus foetidus'', known variously as stinking hellebore , dungwort, setterwort and bear's foot, is a species of flowering plant in the buttercup family Ranunculaceae, native to the mountainous regions of Central and Southern Europe and Asi ...
have been found to raise the temperature of the flower, which may aid in attracting pollinators by increasing the evaporation of
volatile organic compound Volatile organic compounds (VOCs) are organic compounds that have a high vapour pressure at room temperature. High vapor pressure correlates with a low boiling point, which relates to the number of the sample's molecules in the surrounding air, a ...
s. A
black yeast “Black yeasts”, sometimes also black fungi, dematiaceous fungi, microcolonial fungi or meristematic fungi is a diverse group of slow-growing microfungi which reproduce mostly asexually ( fungi imperfecti). Only few genera reproduce by budding ...
has been recorded as a partner in a complex relationship between
ant Ants are eusocial insects of the family Formicidae and, along with the related wasps and bees, belong to the order Hymenoptera. Ants evolved from vespoid wasp ancestors in the Cretaceous period. More than 13,800 of an estimated total of ...
s, their mutualistic fungus, a fungal
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
of the fungus and a bacterium that kills the parasite. The yeast has a negative effect on the bacteria that normally produce antibiotics to kill the parasite, so may affect the ants' health by allowing the parasite to spread. Certain strains of some species of yeasts produce proteins called yeast killer toxins that allow them to eliminate competing strains. (See main article on
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 ...
.) This can cause problems for winemaking but could potentially also be used to advantage by using killer toxin-producing strains to make the wine. Yeast killer toxins may also have medical applications in treating yeast infections (see "Pathogenic yeasts" section below). Marine yeasts, defined as the yeasts that are isolated from marine environments, are able to grow better on a medium prepared using seawater rather than freshwater. The first marine yeasts were isolated by Bernhard Fischer in 1894 from the Atlantic Ocean, and those were identified as ''Torula'' sp. and ''Mycoderma'' sp. Following this discovery, various other marine yeasts have been isolated from around the world from different sources, including seawater, seaweeds, marine fish and mammals. Among these isolates, some marine yeasts originated from terrestrial habitats (grouped as facultative marine yeast), which were brought to and survived in marine environments. The other marine yeasts were grouped as obligate or indigenous marine yeasts, which are confined to marine habitats. However, no sufficient evidence has been found to explain the indispensability of seawater for obligate marine yeasts. This article contains quotations from this source, which is available under a Creative Commons Attribution license. It has been reported that marine yeasts are able to produce many bioactive substances, such as amino acids, glucans, glutathione, toxins, enzymes, phytase, and vitamins with potential applications in the food, pharmaceutical, cosmetic, and chemical industries as well as for marine culture and environmental protection. Marine yeast was successfully used to produce bioethanol using seawater-based media which will potentially reduce the
water footprint A water footprint shows the extent of water use in relation to consumption by people. The water footprint of an individual, community, or business is defined as the total volume of fresh water used to produce the goods and services consumed by t ...
of bioethanol.


Reproduction

Yeasts, like all fungi, may have asexual and
sexual Sex is the biological distinction of an organism between male and female. Sex or SEX may also refer to: Biology and behaviour *Animal sexual behaviour **Copulation (zoology) **Human sexual activity **Non-penetrative sex, or sexual outercourse ** ...
reproductive cycles. The most common mode of vegetative growth in yeast is asexual reproduction 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 ...
, where a small bud (also known as a bleb or daughter cell) is formed on the parent cell. The
nucleus Nucleus ( : nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom * Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucl ...
of the parent cell splits into a daughter nucleus and migrates into the daughter cell. The bud then continues to grow until it separates from the parent cell, forming a new cell. The daughter cell produced during the budding process is generally smaller than the mother cell. Some yeasts, including ''
Schizosaccharomyces pombe ''Schizosaccharomyces pombe'', also called "fission yeast", is a species of yeast used in traditional brewing and as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically measur ...
'', reproduce by fission instead of budding, and thereby creating two identically sized daughter cells. In general, under high-stress conditions such as nutrient starvation, haploid cells will die; under the same conditions, however, diploid cells can undergo sporulation, entering sexual reproduction (
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately r ...
) and producing a variety of haploid spores, which can go on to mate (conjugate), reforming the diploid. The haploid fission yeast ''Schizosaccharomyces pombe'' is a facultative sexual microorganism that can undergo mating when nutrients are limiting. Exposure of ''S. pombe'' to hydrogen peroxide, an agent that causes oxidative stress leading to oxidative DNA damage, strongly induces mating and the formation of meiotic spores. The budding yeast ''Saccharomyces cerevisiae'' reproduces by mitosis as diploid cells when nutrients are abundant, but when starved, this yeast undergoes meiosis to form haploid spores. Haploid cells may then reproduce asexually by mitosis. Katz Ezov et al. presented evidence that in natural ''S. cerevisiae'' populations clonal reproduction and selfing (in the form of intratetrad mating) predominate. In nature, mating of haploid cells to form diploid cells is most often between members of the same clonal population and
out-crossing Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a usefu ...
is uncommon. Analysis of the ancestry of natural ''S. cerevisiae'' strains led to the conclusion that out-crossing occurs only about once every 50,000 cell divisions. These observations suggest that the possible long-term benefits of outcrossing (e.g. generation of diversity) are likely to be insufficient for generally maintaining sex from one generation to the next. Rather, a short-term benefit, such as recombinational repair during meiosis, may be the key to the maintenance of sex in ''S. cerevisiae''. Some pucciniomycete yeasts, in particular species of ''
Sporidiobolus ''Sporidiobolus'' is a genus of Basidiomycota found in the family Sporidiobolaceae. It contains 8 species. It is one of the few genera within Basidiomycota that can reproduce asexually in yeast-form. Species *''Sporidiobolus johnsonii'' *''Spor ...
'' and ''
Sporobolomyces ''Sporobolomyces'' is a fungal genus of uncertain familial placement in the order Sporidiobolales. Species in the genus are anamorphic yeasts. Teleomorphic forms are included in the genus ''Sporidiobolus''. Generic synonyms of ''Sporobolomyce ...
'', produce aerially dispersed, asexual ballistoconidia.


Uses

The useful physiological properties of yeast have led to their use in the field of
biotechnology Biotechnology is the integration of natural sciences and engineering sciences in order to achieve the application of organisms, cells, parts thereof and molecular analogues for products and services. The term ''biotechnology'' was first used ...
. Fermentation of sugars by yeast is the oldest and largest application of this technology. Many types of yeasts are used for making many foods:
baker's yeast Baker's yeast is the common name for the strains of yeast commonly used in baking bread and other bakery products, serving as a leavening agent which causes the bread to rise (expand and become lighter and softer) by converting the fermentabl ...
in bread production, brewer's yeast in
beer fermentation Brewing is the production of beer by steeping a starch source (commonly cereal grains, the most popular of which is barley) in water and fermenting the resulting sweet liquid with yeast. It may be done in a brewery by a commercial brewer ...
, and yeast in wine fermentation and for
xylitol Xylitol is a chemical compound with the formula , or HO(CH2)(CHOH)3(CH2)OH; specifically, one particular stereoisomer with that structural formula. It is a colorless or white crystalline solid that is freely soluble in water. It can be classifi ...
production. So-called red rice yeast is actually a
mold A mold () or mould () is one of the structures certain fungi can form. The dust-like, colored appearance of molds is due to the formation of spores containing fungal secondary metabolites. The spores are the dispersal units of the fungi. Not ...
, ''
Monascus purpureus ''Monascus purpureus'' (syn. ''M. albidus'', ''M. anka'', ''M. araneosus'', ''M. major'', ''M. rubiginosus'', and ''M. vini''; , lit. "red yeast") is a species of mold that is purplish-red in color. It is also known by the names ang-khak rice mo ...
''. Yeasts include some of the most widely used model organisms for
genetics Genetics is the study of genes, genetic variation, and heredity in organisms.Hartl D, Jones E (2005) It is an important branch in biology because heredity is vital to organisms' evolution. Gregor Mendel, a Moravian Augustinian friar wor ...
and cell biology.


Alcoholic beverages

Alcoholic beverages are defined as
beverages A drink or beverage is a liquid intended for human consumption. In addition to their basic function of satisfying thirst, drinks play important roles in human culture. Common types of drinks include plain drinking water, milk, juice, smoothies ...
that contain
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 ...
(C2H5OH). This ethanol is almost always produced by fermentation – the
metabolism Metabolism (, from el, μεταβολή ''metabolē'', "change") is the set of life-sustaining chemical reactions in organisms. The three main functions of metabolism are: the conversion of the energy in food to energy available to run c ...
of
carbohydrate In organic chemistry, a carbohydrate () is a biomolecule consisting of carbon (C), hydrogen (H) and oxygen (O) atoms, usually with a hydrogen–oxygen atom ratio of 2:1 (as in water) and thus with the empirical formula (where ''m'' may or m ...
s by certain species of yeasts under anaerobic or low-oxygen conditions. Beverages such as mead, wine, beer, or
distilled spirits Liquor (or a spirit) is an alcoholic drink produced by distillation of grains, fruits, vegetables, or sugar, that have already gone through alcoholic fermentation. Other terms for liquor include: spirit drink, distilled beverage or hard l ...
all use yeast at some stage of their production. A distilled beverage is a beverage containing ethanol that has been purified by
distillation Distillation, or classical distillation, is the process of separating the components or substances from a liquid mixture by using selective boiling and condensation, usually inside an apparatus known as a still. Dry distillation is the heat ...
. Carbohydrate-containing plant material is fermented by yeast, producing a dilute solution of ethanol in the process. Spirits such as
whiskey Whisky or whiskey is a type of distilled alcoholic beverage made from fermented grain mash. Various grains (which may be malted) are used for different varieties, including barley, corn, rye, and wheat. Whisky is typically aged in wooden ...
and
rum Rum is a liquor made by fermenting and then distilling sugarcane molasses or sugarcane juice. The distillate, a clear liquid, is usually aged in oak barrels. Rum is produced in nearly every sugar-producing region of the world, such as the Ph ...
are prepared by distilling these dilute solutions of ethanol. Components other than ethanol are collected in the condensate, including water,
ester 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 a ...
s, and other alcohols, which (in addition to that provided by the oak in which it may be aged) account for the flavour of the beverage.


Beer

Brewing yeasts may be classed as "top-cropping" (or "top-fermenting") and "bottom-cropping" (or "bottom-fermenting"). Top-cropping yeasts are so called because they form a foam at the top of the
wort Wort () is the liquid extracted from the mashing process during the brewing of beer or whisky. Wort contains the sugars, the most important being maltose and maltotriose, that will be fermented by the brewing yeast to produce alcohol. Wort als ...
during fermentation. An example of a top-cropping yeast 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 ...
'', sometimes called an "ale yeast". Bottom-cropping yeasts are typically used to produce
lager Lager () is beer which has been brewed and conditioned at low temperature. Lagers can be pale, amber, or dark. Pale lager is the most widely consumed and commercially available style of beer. The term "lager" comes from the German for "storag ...
-type beers, though they can also produce ale-type beers. These yeasts ferment well at low temperatures. An example of bottom-cropping yeast is ''
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 ...
'', formerly known as ''S. carlsbergensis''. Decades ago, taxonomists reclassified ''S. carlsbergensis'' (uvarum) as a member of ''S. cerevisiae'', noting that the only distinct difference between the two is metabolic. Lager strains of ''S. cerevisiae'' secrete an enzyme called melibiase, allowing them to hydrolyse
melibiose Melibiose is a reducing disaccharide formed by an α-1,6 linkage between galactose and glucose (D-Gal-α(1→6)-D-Glc). It differs from lactose in the chirality of the carbon where the galactose ring is closed and that the galactose is linked to a ...
, a
disaccharide A disaccharide (also called a double sugar or ''biose'') is the sugar formed when two monosaccharides are joined by glycosidic linkage. Like monosaccharides, disaccharides are simple sugars soluble in water. Three common examples are sucrose, la ...
, into more fermentable monosaccharides. Top- and bottom-cropping and cold- and warm-fermenting distinctions are largely generalizations used by laypersons to communicate to the general public. The most common top-cropping brewer's yeast, ''S. cerevisiae'', is the same species as the common baking yeast. Brewer's yeast is also very rich in essential minerals and the
B vitamin B vitamins are a class of water-soluble vitamins that play important roles in cell metabolism and synthesis of red blood cells. Though these vitamins share similar names (B1, B2, B3, etc.), they are chemically distinct compounds that often coexist ...
s (except B12), a feature exploited in food products made from leftover ( by-product) yeast from brewing. However, baking and brewing yeasts typically belong to different strains, cultivated to favour different characteristics: baking yeast strains are more aggressive, to carbonate
dough Dough is a thick, malleable, sometimes elastic paste made from grains or from leguminous or chestnut crops. Dough is typically made by mixing flour with a small amount of water or other liquid and sometimes includes yeast or other leavenin ...
in the shortest amount of time possible; brewing yeast strains act more slowly but tend to produce fewer off-flavours and tolerate higher alcohol concentrations (with some strains, up to 22%). '' Dekkera/Brettanomyces'' is a genus of yeast known for its important role in the production of ' lambic' and specialty
sour ale Sour beer, also known as Sours, is beer which has an intentionally acidic, tart, or sour taste. Traditional sour beer styles include Belgian lambics, gueuze and Flanders red ale, and German gose and Berliner Weisse. Brewing Unlike modern brewin ...
s, along with the secondary conditioning of a particular Belgian
Trappist beer Trappist beer is brewed by Trappist monks. Thirteen Trappist monasteries—six in Belgium, two in the Netherlands, and one each in Austria, Italy, England, France, and Spain—currently produce beer, but the ''Authentic Trappist Product'' labe ...
. The taxonomy of the genus ''Brettanomyces'' has been debated since its early discovery and has seen many reclassifications over the years. Early classification was based on a few species that reproduced asexually (anamorph form) through multipolar budding. Shortly after, the formation of ascospores was observed and the genus ''Dekkera'', which reproduces sexually (teleomorph form), was introduced as part of the taxonomy. The current taxonomy includes five species within the genera of ''Dekkera/Brettanomyces''. Those are the anamorphs ''
Brettanomyces bruxellensis ''Brettanomyces bruxellensis'' (the anamorph of ''Dekkera bruxellensis'') is a yeast associated with the Zenne, Senne valley near Brussels, Belgium. Despite its Latin species name, ''B. bruxellensis'' is found all over the globe. In the wild, it ...
'', '' Brettanomyces anomalus'', ''
Brettanomyces custersianus ''Brettanomyces'' is a spore, 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 tele ...
'', '' Brettanomyces naardenensis'', and '' Brettanomyces nanus'', with teleomorphs existing for the first two species, '' Dekkera bruxellensis'' and '' Dekkera anomala''. The distinction between ''Dekkera'' and ''Brettanomyces'' is arguable, with Oelofse et al. (2008) citing Loureiro and Malfeito-Ferreira from 2006 when they affirmed that current molecular DNA detection techniques have uncovered no variance between the anamorph and teleomorph states. Over the past decade, ''Brettanomyces'' spp. have seen an increasing use in the craft-brewing sector of the industry, with a handful of breweries having produced beers that were primarily fermented with pure cultures of ''Brettanomyces'' spp. This has occurred out of experimentation, as very little information exists regarding pure culture fermentative capabilities and the aromatic compounds produced by various strains. ''Dekkera''/''Brettanomyces'' spp. have been the subjects of numerous studies conducted over the past century, although a majority of the recent research has focused on enhancing the knowledge of the wine industry. Recent research on eight ''Brettanomyces'' strains available in the brewing industry focused on strain-specific fermentations and identified the major compounds produced during pure culture anaerobic fermentation in wort.


Wine

Yeast is used in
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 ...
, where it converts the sugars present (
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 ...
and fructose) in
grape juice Grape juice is obtained from crushing and blending grapes into a liquid. In the wine industry, grape juice that contains 7–23 percent of pulp, skins, stems and seeds is often referred to as '' must''. The sugars in grape juice allow it to be ...
( must) into ethanol. Yeast is normally already present on grape skins. Fermentation can be done with this endogenous "wild yeast", but this procedure gives unpredictable results, which depend upon the exact types of yeast species present. For this reason, a pure yeast culture is usually added to the must; this yeast quickly dominates the fermentation. The wild yeasts are repressed, which ensures a reliable and predictable fermentation. Most added wine yeasts are strains of ''S. cerevisiae'', though not all strains of the species are suitable. Different ''S. cerevisiae'' yeast strains have differing physiological and fermentative properties, therefore the actual strain of yeast selected can have a direct impact on the finished wine. Significant research has been undertaken into the development of novel wine yeast strains that produce atypical flavour profiles or increased complexity in wines. The growth of some yeasts, such as ''
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 ...
'' and ''
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 ...
'', in wine can result in
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 ...
s and subsequent spoilage. ''Brettanomyces'' produces an array of metabolites when growing in wine, some of which are volatile
phenol Phenol (also called carbolic acid) is an aromatic organic compound with the molecular formula . It is a white crystalline solid that is volatile. The molecule consists of a phenyl group () bonded to a hydroxy group (). Mildly acidic, it ...
ic compounds. Together, these compounds are often referred to as "''Brettanomyces'' character", and are often described as "
antiseptic An antiseptic (from Greek ἀντί ''anti'', "against" and σηπτικός ''sēptikos'', "putrefactive") is an antimicrobial substance or compound that is applied to living tissue/skin to reduce the possibility of infection, sepsis, or putre ...
" or "barnyard" type aromas. ''Brettanomyces'' is a significant contributor to wine faults within the wine industry. Researchers from the
University of British Columbia The University of British Columbia (UBC) is a public research university with campuses near Vancouver and in Kelowna, British Columbia. Established in 1908, it is British Columbia's oldest university. The university ranks among the top thre ...
, Canada, have found a new strain of yeast that has reduced
amine In chemistry, amines (, ) are compounds and functional groups that contain a basic nitrogen atom with a lone pair. Amines are formally derivatives of ammonia (), wherein one or more hydrogen Hydrogen is the chemical element wi ...
s. The amines in red wine and Chardonnay produce off-flavors and cause headaches and hypertension in some people. About 30% of people are sensitive to biogenic amines, such as
histamine Histamine is an organic nitrogenous compound involved in local immune responses, as well as regulating physiological functions in the gut and acting as a neurotransmitter for the brain, spinal cord, and uterus. Since histamine was discovered ...
s.


Baking

Yeast, the most common one being ''S. cerevisiae'', is used in baking as a
leavening agent In cooking, a leavening agent () or raising agent, also called a leaven () or leavener, is any one of a number of substances used in doughs and batters that cause a foaming action (gas bubbles) that lightens and softens the mixture. An altern ...
, where it converts the food/fermentable sugars present in dough into the gas
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 ...
. This causes the dough to expand or rise as gas forms pockets or bubbles. When the dough is baked, the yeast dies and the air pockets "set", giving the baked product a soft and spongy texture. The use of potatoes, water from potato boiling,
eggs Humans and human ancestors have scavenged and eaten animal eggs for millions of years. Humans in Southeast Asia had domesticated chickens and harvested their eggs for food by 1,500 BCE. The most widely consumed eggs are those of fowl, especial ...
, or sugar in a bread dough accelerates the growth of yeasts. Most yeasts used in baking are of the same species common in alcoholic fermentation. In addition, ''Saccharomyces exiguus'' (also known as ''S. minor''), a wild yeast found on plants, fruits, and grains, is occasionally used for baking. In breadmaking, the yeast initially respires aerobically, producing carbon dioxide and water. When the oxygen is depleted, fermentation begins, producing ethanol as a waste product; however, this evaporates during baking. It is not known when yeast was first used to bake bread. The first records that show this use came from Ancient Egypt. Researchers speculate a mixture of flour meal and water was left longer than usual on a warm day and the yeasts that occur in natural contaminants of the flour caused it to ferment before baking. The resulting bread would have been lighter and tastier than the normal flat, hard cake. Today, there are several retailers of baker's yeast; one of the earlier developments in North America is Fleischmann's Yeast, in 1868. During World War II, Fleischmann's developed a
granulated Granulation is the process of forming grains or granules from a powdery or solid substance, producing a granular material. It is applied in several technological processes in the chemical and pharmaceutical industries. Typically, granulation invo ...
active dry yeast which did not require refrigeration, had a longer
shelf life Shelf life is the length of time that a commodity may be stored without becoming unfit for use, consumption, or sale. In other words, it might refer to whether a commodity should no longer be on a pantry shelf (unfit for use), or no longer on a ...
than fresh yeast, and rose twice as fast. Baker's yeast is also sold as a fresh yeast compressed into a square "cake". This form perishes quickly, so must be used soon after production. A weak solution of water and sugar can be used to determine whether yeast is expired. In the solution, active yeast will foam and bubble as it ferments the sugar into ethanol and carbon dioxide. Some recipes refer to this as proofing the yeast, as it "proves" (tests) the viability of the yeast before the other ingredients are added. When a sourdough starter is used, flour and water are added instead of sugar; this is referred to as proofing the
sponge Sponges, the members of the phylum Porifera (; meaning 'pore bearer'), are a basal animal clade as a sister of the diploblasts. They are multicellular organisms that have bodies full of pores and channels allowing water to circulate throug ...
. When yeast is used for making bread, it is mixed with flour, salt, and warm water or milk. The dough is kneaded until it is smooth, and then left to rise, sometimes until it has doubled in size. The dough is then shaped into loaves. Some bread doughs are knocked back after one rising and left to rise again (this is called dough proofing) and then baked. A longer rising time gives a better flavor, but the yeast can fail to raise the bread in the final stages if it is left for too long initially.


Bioremediation

Some yeasts can find potential application in the field of
bioremediation Bioremediation broadly refers to any process wherein a biological system (typically bacteria, microalgae, fungi, and plants), living or dead, is employed for removing environmental pollutants from air, water, soil, flue gasses, industrial effluent ...
. One such yeast, ''
Yarrowia lipolytica ''Yarrowia'' is a fungal genus in the family Dipodascaceae. For a while the genus was monotypic, containing the single species ''Yarrowia lipolytica'', a yeast that can use unusual carbon sources, such as hydrocarbons. This has made it of interes ...
'', is known to degrade palm oil mill effluent,
TNT Trinitrotoluene (), more commonly known as TNT, more specifically 2,4,6-trinitrotoluene, and by its preferred IUPAC name 2-methyl-1,3,5-trinitrobenzene, is a chemical compound with the formula C6H2(NO2)3CH3. TNT is occasionally used as a reagen ...
(an explosive material), and other
hydrocarbon In organic chemistry, a hydrocarbon is an organic compound consisting entirely of hydrogen and carbon. Hydrocarbons are examples of group 14 hydrides. Hydrocarbons are generally colourless and hydrophobic, and their odors are usually weak or ...
s, such as alkanes,
fatty acid In chemistry, particularly in biochemistry, a fatty acid is a carboxylic acid with an aliphatic chain, which is either saturated or unsaturated. Most naturally occurring fatty acids have an unbranched chain of an even number of carbon atoms, ...
s, fats and oils. It can also tolerate high concentrations of salt and heavy metals, and is being investigated for its potential as a heavy metal biosorbent. ''Saccharomyces cerevisiae'' has potential to bioremediate toxic pollutants like
arsenic Arsenic is a chemical element with the symbol As and atomic number 33. Arsenic occurs in many minerals, usually in combination with sulfur and metals, but also as a pure elemental crystal. Arsenic is a metalloid. It has various allotropes, ...
from industrial effluent. Bronze statues are known to be degraded by certain species of yeast. Different yeasts from Brazilian gold mines
bioaccumulate Bioaccumulation is the gradual accumulation of substances, such as pesticides or other chemicals, in an organism. Bioaccumulation occurs when an organism absorbs a substance at a rate faster than that at which the substance is lost or eliminated ...
free and complexed silver ions.


Industrial ethanol production

The ability of yeast to convert sugar into ethanol has been harnessed by the biotechnology industry to produce
ethanol fuel Ethanol fuel is ethyl alcohol, the same type of alcohol found in alcoholic beverages, used as fuel. It is most often used as a motor fuel, mainly as a biofuel additive for gasoline. The first production car running entirely on ethanol was t ...
. The process starts by milling a feedstock, such as
sugar cane Sugarcane or sugar cane is a species of (often hybrid) tall, perennial grass (in the genus '' Saccharum'', tribe Andropogoneae) that is used for sugar production. The plants are 2–6 m (6–20 ft) tall with stout, jointed, fibrous stalk ...
,
field corn Field corn, also known as cow corn, is a North American term for maize (''Zea mays'') grown for livestock fodder (silage), ethanol, cereal, and processed food products. The principal field corn varieties are dent corn, flint corn, flour corn (also ...
, or other
cereal grain A cereal is any grass cultivated for the edible components of its grain (botanically, a type of fruit called a caryopsis), composed of the endosperm, germ, and bran. Cereal grain crops are grown in greater quantities and provide more food ...
s, and then adding dilute sulfuric acid, or fungal alpha
amylase An amylase () is an enzyme that catalyses the hydrolysis of starch (Latin ') into sugars. Amylase is present in the saliva of humans and some other mammals, where it begins the chemical process of digestion. Foods that contain large amounts of ...
enzymes, to break down the starches into complex sugars. A glucoamylase is then added to break the complex sugars down into simple sugars. After this, yeasts are added to convert the simple sugars to ethanol, which is then distilled off to obtain ethanol up to 96% in purity. ''Saccharomyces'' yeasts have been
genetically engineered Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including t ...
to ferment
xylose Xylose ( grc, ξύλον, , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional g ...
, one of the major fermentable sugars present in cellulosic biomasses, such as agriculture residues, paper wastes, and wood chips. Such a development means ethanol can be efficiently produced from more inexpensive feedstocks, making
cellulosic ethanol Cellulosic ethanol is ethanol (ethyl alcohol) produced from cellulose (the stringy fiber of a plant) rather than from the plant's seeds or fruit. It can be produced from grasses, wood, algae, or other plants. It is generally discussed for use as a ...
fuel a more competitively priced alternative to gasoline fuels.


Nonalcoholic beverages

A number of sweet carbonated beverages can be produced by the same methods as beer, except the fermentation is stopped sooner, producing carbon dioxide, but only trace amounts of alcohol, leaving a significant amount of residual sugar in the drink. *
Root beer Root beer is a sweet North American soft drink traditionally made using the root bark of the sassafras tree '' Sassafras albidum'' or the vine of '' Smilax ornata'' (known as sarsaparilla, also used to make a soft drink, sarsaparilla) as the ...
, originally made by Native Americans, commercialized in the United States by
Charles Elmer Hires Charles Elmer Hires (August 19, 1851 – July 31, 1937) was an American pharmacist and an early promoter of commercially prepared root beer. He founded the Charles E. Hires Co., which manufactured and distributed Hires Root Beer. Biography Early ...
and especially popular during
Prohibition Prohibition is the act or practice of forbidding something by law; more particularly the term refers to the banning of the manufacture, storage (whether in barrels or in bottles), transportation, sale, possession, and consumption of alcohol ...
* Kvass, a
fermented drink This is a list of fermented foods, which are foods produced or preserved by the action of microorganisms. In this context, fermentation typically refers to the fermentation of sugar to alcohol using yeast, but other fermentation processes involv ...
made from rye, popular in Eastern Europe. It has a recognizable, but low alcoholic content. *
Kombucha Kombucha (also tea mushroom, tea fungus, or Manchurian mushroom when referring to the culture; Latin name ''Medusomyces gisevii'') is a fermented, lightly effervescent, sweetened black tea drink commonly consumed for its purported health b ...
, a fermented sweetened tea. Yeast in symbiosis with
acetic acid bacteria Acetic acid bacteria (AAB) are a group of Gram-negative bacteria which oxidize sugars or ethanol and produce acetic acid during fermentation. The acetic acid bacteria consist of 10 genera in the family Acetobacteraceae. Several species of acetic ...
is used in its preparation. Species of yeasts found in the tea can vary, and may include: ''
Brettanomyces bruxellensis ''Brettanomyces bruxellensis'' (the anamorph of ''Dekkera bruxellensis'') is a yeast associated with the Zenne, Senne valley near Brussels, Belgium. Despite its Latin species name, ''B. bruxellensis'' is found all over the globe. In the wild, it ...
'', '' Candida stellata'', ''
Schizosaccharomyces pombe ''Schizosaccharomyces pombe'', also called "fission yeast", is a species of yeast used in traditional brewing and as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically measur ...
'', ''
Torulaspora delbrueckii ''Torulaspora delbrueckii'' is a ubiquitous yeast species with both wild and anthropic habitats. The type strain of ''T. delbrueckii'' is CBS 1146T, equivalent to CLIB 230 or ATCC 10662etc. The type strain of ''T. delbrueckii'' CBS 1146 T was seq ...
'' and ''
Zygosaccharomyces bailii ''Zygosaccharomyces bailii'' is a species in the genus ''Zygosaccharomyces''. It was initially described as ''Saccharomyces bailii'' by Lindner in 1895, but in 1983 it was reclassified as ''Zygosaccharomyces bailii'' in the work by Barnett et al ...
''. Also popular in Eastern Europe and some
former Soviet republics The post-Soviet states, also known as the former Soviet Union (FSU), the former Soviet Republics and in Russia as the near abroad (russian: links=no, ближнее зарубежье, blizhneye zarubezhye), are the 15 sovereign states that wer ...
under the name ''chajnyj grib'' (russian: Чайный гриб), which means "tea mushroom". *
Kefir Kefir ( ; also spelled as kephir or kefier; ; ; ) is a fermented milk drink similar to a thin yogurt or ayran that is made from kefir grains, a specific type of mesophilic symbiotic culture. The drink originated in the North Caucasus, in p ...
and kumis are made by fermenting milk with yeast and bacteria. *
Mauby Mauby (in Trinidad and Tobago, Saint Lucia, Jamaica, St. Vincent and The Grenadines, Grenada, Guyana, Bermuda, Barbados, Antigua and Barbuda and Anguilla), also known as ''maví (or mabí)'' in the Dominican Republic and Puerto Rico, ''mabi ...
( es, mabí), made by fermenting sugar with the wild yeasts naturally present on the bark of the ''Colubrina elliptica'' tree, popular in the Caribbean


Foods and

Yeast is used as an ingredient in foods for its umami flavor, in much of the same way that monosodium glutamate (MSG) is used and, like MSG, often contain free glutamic acid. Examples include: * Yeast extract, made from the intracellular contents of yeast and used as food additives or Flavoring, flavours. The general method for making yeast extract for food products such as Vegemite and Marmite on a commercial scale is ''heat autolysis'', i.e. to add salt to a suspension of yeast, making the solution hypertonic, which leads to the cells' shrivelling up. This triggers autolysis (biology), autolysis, wherein the yeast's digestion, digestive enzymes break their own proteins down into simpler compounds, a process of self-destruction. The dying yeast cells are then heated to complete their breakdown, after which the husks (yeast with thick cell walls that would give poor texture) are removed. Yeast autolysates are used in Vegemite and Promite (Australia); Marmite (the United Kingdom); the unrelated Marmite (New Zealand), Marmite (New Zealand); Vitam-R (Germany); and Cenovis (Switzerland). * Nutritional yeast, which is whole dried, deactivated yeast cells, usually ''S. cerevisiae''. Usually in the form of yellow flake or powder, its nutty and umami flavor makes it a vegan substitute for cheese powder. Another popular use is as a topping for popcorn. It can also be used in mashed and fried potatoes, as well as in scrambled eggs. It comes in the form of flakes, or as a yellow powder similar in texture to cornmeal. In Australia, it is sometimes sold as "savoury yeast flakes". Both types of yeast foods above are rich in Vitamin B, B-complex vitamins (besides vitamin B12 unless fortified), making them an attractive nutritional supplement to vegans. The same vitamins are also found in some yeast-fermented products mentioned above, such as kvass. Nutritional yeast in particular is naturally low in fat and sodium and a source of protein and vitamins as well as other minerals and cofactor (biochemistry), cofactors required for growth. Many brands of nutritional yeast and yeast extract spreads, though not all, are fortified with Cyanocobalamin, vitamin B12, which is produced separately by
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
. In 1920, the Fleischmann's Yeast, Fleischmann Yeast Company began to promote yeast cakes in a "Yeast for Health" campaign. They initially emphasized yeast as a source of vitamins, good for skin and digestion. Their later advertising claimed a much broader range of health benefits, and was censured as misleading by the Federal Trade Commission. The fad for yeast cakes lasted until the late 1930s.


Probiotics

Some probiotic supplements use the yeast ''Saccharomyces boulardii, S. boulardii'' to maintain and restore the natural flora in the gastrointestinal tract. ''S. boulardii'' has been shown to reduce the symptoms of acute diarrhea, reduce the chance of infection by ''Clostridium difficile (bacteria), Clostridium difficile'' (often identified simply as C. difficile or C. diff), reduce bowel movements in diarrhea-predominant irritable bowel syndrome, IBS patients, and reduce the incidence of antibiotic-associated diarrhea, antibiotic-, travelers' diarrhea, traveler's-, and HIV/AIDS-associated diarrheas.


Aquarium hobby

Yeast is often used by aquarium hobbyists to generate carbon dioxide (CO2) to nourish plants in Aquascaping, planted aquaria. CO2 levels from yeast are more difficult to regulate than those from pressurized CO2 systems. However, the low cost of yeast makes it a widely used alternative.


Scientific research

Several yeasts, in particular ''S. cerevisiae'' and ''S. pombe'', have been widely used in genetics and cell biology, largely because they are simple eukaryote, eukaryotic cells, serving as a model for all eukaryotes, including humans, for the study of fundamental cellular processes such as the cell cycle, DNA replication, genetic recombination, recombination, cell division, and metabolism. Also, yeasts are easily manipulated and cultured in the laboratory, which has allowed for the development of powerful standard techniques, such as yeast two-hybrid, synthetic genetic array analysis, and tetrad (genetics), tetrad analysis. Many proteins important in human biology were first discovered by studying their homology (biology), homologues in yeast; these proteins include cell cycle proteins, Cell signaling, signaling proteins, and protein-processing enzymes. On 24 April 1996, ''S. cerevisiae'' was announced to be the first eukaryote to have its genome, consisting of 12 million base pairs, fully sequenced as part of the Genome Project. At the time, it was the most complex organism to have its full genome sequenced, and the work seven years and the involvement of more than 100 laboratories to accomplish. The second yeast species to have its genome sequenced was ''Schizosaccharomyces pombe'', which was completed in 2002. It was the sixth eukaryotic genome sequenced and consists of 13.8 million base pairs. As of 2014, over 50 yeast species have had their genomes sequenced and published. Genomic and functional gene annotation of the two major yeast models can be accessed via their respective model organism databases: SGD and PomBase.


Genetically engineered biofactories

Various yeast species have been genetically engineered to efficiently produce various drugs, a technique called metabolic engineering. ''S. cerevisiae'' is easy to genetically engineer; its physiology, metabolism and genetics are well known, and it is amenable for use in harsh industrial conditions. A wide variety of chemical in different classes can be produced by engineered yeast, including phenols, phenolics, isoprenoids, alkaloids, and polyketides. About 20% of biopharmaceuticals are produced in ''S. cerevisiae'', including insulin, vaccines for hepatitis, and human serum albumin.


Pathogenic yeasts

Some species of yeast are
opportunistic pathogen An opportunistic infection is an infection caused by pathogens (bacteria, fungi, parasites or viruses) that take advantage of an opportunity not normally available. These opportunities can stem from a variety of sources, such as a weakened immune ...
s that can cause infection in people with compromised immune systems. ''Cryptococcus neoformans'' and ''Cryptococcus gattii'' are significant pathogens of immunocompromised people. They are the species primarily responsible for cryptococcosis, a fungal infection that occurs in about one million HIV/AIDS patients, causing over 600,000 deaths annually. The cells of these yeast are surrounded by a rigid polysaccharide capsule, which helps to prevent them from being recognised and engulfed by white blood cells in the human body. Yeasts of the genus '' Candida'', another group of opportunistic pathogens, cause Oral candidiasis, oral and vaginal infections in humans, known as candidiasis. ''Candida'' is commonly found as a commensal yeast in the mucous membranes of humans and other warm-blooded animals. However, sometimes these same strains can become pathogenic. The yeast cells sprout a hyphal outgrowth, which locally penetrates the mucous membrane, mucosal membrane, causing irritation and shedding of the tissues. A book from the 1980s listed the pathogenic yeasts of candidiasis in probable descending order of virulence for humans as: ''Candida albicans, C. albicans'', ''Candida tropicalis, C. tropicalis'', ''Candida stellatoidea, C. stellatoidea'', ''Candida glabrata, C. glabrata'', ''Candida krusei, C. krusei'', ''Candida parapsilosis, C. parapsilosis'', ''Candida guilliermondii, C. guilliermondii'', ''Candida viswanathii, C. viswanathii'', ''Candida lusitaniae, C. lusitaniae'', and ''Rhodotorula, Rhodotorula mucilaginosa''. ''Candida glabrata'' is the second most common ''Candida'' pathogen after ''C. albicans'', causing infections of the urogenital tract, and of the bloodstream (candidemia). ''Candida auris, C. auris'' has been more recently identified.


Food spoilage

Yeasts are able to grow in foods with a low pH (5.0 or lower) and in the presence of sugars, organic acids, and other easily metabolized carbon sources. During their growth, yeasts metabolize some food components and produce metabolic end products. This causes the physical, chemical, and sensible properties of a food to change, and the food is spoiled. The growth of yeast within food products is often seen on their surfaces, as in cheeses or meats, or by the fermentation of sugars in beverages, such as juices, and semiliquid products, such as syrups and jams. The yeast of the genus ''
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 ...
'' have had a long history as spoilage yeasts within the food industry. This is mainly because these species can grow in the presence of high sucrose, ethanol, acetic acid, sorbic acid, benzoic acid, and sulfur dioxide concentrations, representing some of the commonly used food preservation methods. Methylene blue is used to test for the presence of live yeast cells. In oenology, the major spoilage yeast is ''
Brettanomyces bruxellensis ''Brettanomyces bruxellensis'' (the anamorph of ''Dekkera bruxellensis'') is a yeast associated with the Zenne, Senne valley near Brussels, Belgium. Despite its Latin species name, ''B. bruxellensis'' is found all over the globe. In the wild, it ...
''. '' Candida blankii'' has been detected in Iberian ham and meat.


Symbiosis

An Indian study of seven bee species and nine plant species found 45 yeast species from 16 genera colonise the
nectaries Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualist ...
of flowers and honey stomachs of bees. Most were members of the genus '' Candida''; the most common species in honey bee stomachs was '' Dekkera intermedia'', while the most common species colonising flower nectaries was '' Candida blankii''. Although the mechanics are not fully understood, it was found that ''A. indica'' flowers more if '' Candida blankii'' is present. In another example, ''Spathaspora passalidarum'', found in the digestive tract of scarab beetles, aids the digestion of plant cells by fermenting
xylose Xylose ( grc, ξύλον, , "wood") is a sugar first isolated from wood, and named for it. Xylose is classified as a monosaccharide of the aldopentose type, which means that it contains five carbon atoms and includes an aldehyde functional g ...
.


See also

* Bioaerosol * Ethanol fermentation * Evolution of aerobic fermentation * ''Kazachstania yasuniensis'' – a yeast isolated in 2015 * Mycosis (fungal infection in animals) * Start point (yeast) * WHI3 * Plasmid#Yeast plasmids, Yeast plasmids * Zymology


References


Further reading

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External links


''Saccharomyces'' genome database

Yeast growth and the cell cycle

Yeast virtual library
{{Authority control Yeasts, Abortifacients Brewing ingredients Fermentation in food processing Food additives Fungus common names Leavening agents Medicinal fungi