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Yeast
Yeast of the species Saccharomyces cerevisiae
Scientific classification
Domain:
Eukaryota
Kingdom:
Fungi
Typical divisions

Yeasts are eukaryotic micro-organisms classified in the kingdom Fungi, with about 1,500 species currently described;[1] they dominate fungal diversity in the oceans.[2] Most reproduce asexually by budding, although a few do so by binary fission. Yeasts are unicellular, although some species with yeast forms may become multicellular through the formation of a string of connected budding cells known as pseudohyphae, or false hyphae as seen in most molds.[3] Yeast size can vary greatly depending on the species, typically measuring 3–4 µm in diameter, although some yeasts can reach over 40 µm.[4]

The yeast species Saccharomyces cerevisiae has been used in baking and fermenting alcoholic beverages for thousands of years.[5] It is also extremely important as a model organism in modern cell biology research, and is one of the most thoroughly researched eukaryotic microorganisms. Researchers have used it to gather information about the biology of the eukaryotic cell and ultimately human biology.[6] Other species of yeast, such as Candida albicans, are opportunistic pathogens and can cause infections in humans. Yeasts have recently been used to generate electricity in microbial fuel cells,[7] and produce ethanol for the biofuel industry.

Yeasts do not form an exact taxonomic or phylogenetic grouping. At present it is estimated that only 1% of all yeast species have been described.[8] The term "yeast" is often taken as a synonym for S. cerevisiae,[9] but the phylogenetic diversity of yeasts is shown by their placement in both divisions Ascomycota and Basidiomycota. The budding yeasts ("true yeasts") are classified in the order Saccharomycetales.[10]

History

See also: History of wine and History of beer

The word "yeast" comes to us from Old English gist, gyst, and from the Indo-European root yes-, meaning boil, foam, or bubble.[11] Yeast microbes are probably one of the earliest domesticated organisms. People have used yeast for fermentation and baking throughout history. Archaeologists digging in Egyptian ruins found early grinding stones and baking chambers for yeasted bread, as well as drawings of 4,000-year-old bakeries and breweries.[12] In 1680 the Dutch naturalist Antonie van Leeuwenhoek first microscopically observed yeast, but at the time did not consider them to be living organisms, but rather globular structures.[13] In 1857 French microbiologist Louis Pasteur proved in the paper "Mémoire sur la fermentation alcoolique" that alcoholic fermentation was conducted by living yeasts and not by a chemical catalyst.[12][14] Pasteur showed that by bubbling oxygen into the yeast broth, cell growth could be increased, but the fermentation inhibited – an observation later called the Pasteur effect.

By the late 1700s, two yeast strains used in brewing had been identified: Saccharomyces cerevisiae, so called "high" or top yeast, and S.  carlsbergensis, "low" or bottom yeast. High yeast was sold commercially by the Dutch for bread making starting in 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.[15] The industrial production of yeast blocks was enhanced by the introduction of the filter press in 1867. In 1872, Baron Max de Springer develop a manufacturing process to create granulated yeast, a technique that was used until the first World War.[16] In the United States, naturally occurring airborne yeasts were used almost exclusively until commercial yeast was marketed at the Centennial Exposition 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.

Growth and nutrition

Yeasts are chemoorganotrophs as they use organic compounds as a source of energy and do not require sunlight to grow. Carbon is obtained mostly from hexose sugars such as glucose and fructose, or disaccharides such as sucrose and maltose. Some species can metabolize pentose sugars like ribose,[17] alcohols, and organic acids. Yeast species either require oxygen for aerobic cellular respiration (obligate aerobes), or are anaerobic but also have aerobic methods of energy production (facultative anaerobes). Unlike bacteria, there are no known yeast species that grow only anaerobically (obligate anaerobes). Yeasts grow best in a neutral or slightly acidic pH environment.

Yeasts vary in what temperature range they grow best. For example, Leucosporidium frigidum grows at −2 to 20 °C (28 to 68 °F), Saccharomyces telluris at 5 to 35 °C (41 to 95 °F) and Candida slooffi at 28 to 45 °C (82 to 113 °F).[18] The cells can survive freezing under certain conditions, with viability decreasing over time.

Yeasts are generally grown in the laboratory on solid growth media or in liquid broths. Common media used for the cultivation of yeasts include; potato dextrose agar (PDA) or potato dextrose broth, Wallerstein Laboratories nutrient (WLN) agar, yeast peptone dextrose agar (YPD), and yeast mould agar or broth (YM). Homebrewers who cultivate yeast frequently use dried malt extract (DME) and agar as a solid growth medium. The antibiotic cycloheximide is sometimes added to yeast growth media to inhibit the growth of Saccharomyces 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, usually the result of exposure to air. Although harmless it can give pickled vegetables a bad flavour and so must be removed regularly during fermentation.[19]

Ecology

Yeasts are very common in the environment, but are usually isolated from sugar-rich material. Examples include naturally occurring yeasts on the skins of fruits and berries (such as grapes, apples or peaches), and exudates from plants (such as plant saps or cacti). Some yeasts are found in association with soil and insects.[20][21] The ecological function and biodiversity of yeast are relatively unknown compared to those of other microorganisms.[22] Yeasts including Candida albicans, Rhodotorula rubra, Torulopsis and Trichosporon cutaneum have been found living in between people's toes as part of their skin flora.[23] Yeasts are also present in the gut flora of mammals and some insects.[24]

An Indian study of seven bee species and 9 plant species found that 45 species from 16 genera colonise the nectaries of flowers and honey stomachs of bees. Most were members of the Candida genus; the most common species in honey stomachs was Dekkera intermedia and in flower nectaries, Candida blankii.[25] Flowers that are colonised by yeasts have less nectar to bees in them. Yeast colonising nectaries of the Stinking Hellebore have been found to raise the temperature of the flower, which may aid in attracting pollinators by increasing the evaporation of volatile organic compounds.[22][26] A black yeast has been recorded as a partner in a complex relationship between ants, their mutualistic fungus, a fungal parasite of the fungus and a bacterium that kills the parasite. The yeast have a negative effect on the bacteria that normally produce antibiotics to kill the parasite and so may affect the ants' health by allowing the parasite to spread.[27]

Reproduction

The yeast cell's life cycle:
1. Budding
2. Conjugation
3. Spore
See also: Mating of yeast

Yeasts have asexual and sexual reproductive cycles. The most common mode of vegetative growth in yeast is asexual reproduction by budding.[28] Here a small bud, or daughter cell, is formed on the parent cell. The nucleus of the parent cell splits into a daughter nucleus and migrates into the daughter cell. The bud continues to grow until it separates from the parent cell, forming a new cell.[29] Some yeasts, including Schizosaccharomyces pombe, reproduce by binary fission instead of budding.[28]

Under high stress conditions haploid cells will generally die, however under the same conditions diploid cells can undergo sporulation, entering sexual reproduction (meiosis) and producing a variety of haploid spores, which can go on to mate (conjugate), reforming the diploid.[30]

Uses

The useful physiological properties of yeast have led to their use in the field of biotechnology. 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 in bread production; brewer's yeast in beer fermentation; yeast in wine fermentation and for xylitol production.[31] So-called red rice yeast is actually a mold, Monascus purpureus. Yeasts include some of the most widely used model organisms for genetics and cell biology.

Alcoholic beverages

Alcoholic beverages are defined as beverages that contain ethanol (C2H5OH). This ethanol is almost always produced by fermentation – the metabolism of carbohydrates by certain species of yeast under anaerobic or low-oxygen conditions. Beverages such as wine, beer, or distilled spirits all use yeast at some stage of their production.

Beer

A mixture of diatomaceous earth and yeast after filtering beer

"Brewer's yeast" (also known as "brewing yeast") can mean any live yeast used in brewing. It can also mean yeast obtained as a by-product of brewing, dried and killed, and used as a dietary supplement for its B vitamin content.

Brewers classify yeasts as top-fermenting and bottom-fermenting. This distinction was introduced by the Dane Emil Christian Hansen. "Top-fermenting yeasts" are so called because they form a foam at the top of the wort during fermentation. They can produce higher alcohol concentrations and prefer higher temperatures, typically 16 to 24 °C (61 to 75 °F), producing fruitier, sweeter, ale-type beers. An example of a top-fermenting yeast is Saccharomyces cerevisiae, known to brewers as ale yeast. "Bottom-fermenting yeasts" are typically used to produce lager-type beers, though can also produce ale-type beers. These yeasts ferment more sugars, leaving a crisper taste, and grow well at low temperatures. An example of bottom fermenting yeast is Saccharomyces pastorianus, formerly known as Saccharomyces carlsbergensis.

For both types, yeast is fully distributed through the beer while it is fermenting, and both equally flocculate (clump together and precipitate to the bottom of the vessel) when fermentation is finished. By no means do all top-fermenting yeasts demonstrate this behaviour, but it features strongly in many English ale yeasts which may also exhibit chain forming (the failure of budded cells to break from the mother cell) which is technically different from true flocculation.

Fermenting tanks with yeast being used to brew beer

In industrial brewing, to ensure purity of strain, a "clean" sample of the yeast is stored refrigerated in a laboratory. After a certain number of fermentation cycles, a full scale propagation is produced from this laboratory sample. Typically, it is grown up in about three or four stages using sterile brewing wort and oxygen.

The most common top-fermenting brewer's yeast, Saccharomyces cerevisiae, is the same species as the common baking yeast. However, baking and brewing yeasts typically belong to different strains, cultivated to favor different characteristics: baking yeast strains are more aggressive, in order to carbonate dough in the shortest amount of time possible; brewing yeast strains act slower, but tend to produce fewer off-flavors and tolerate higher alcohol concentrations (with some strains, up to 22%).

Brettanomyces

Brettanomyces is a genus of wild yeast important in brewing lambic, a beer produced not by the deliberate addition of brewer's yeasts, but by spontaneous fermentation by wild yeasts and bacteria. Brettanomyces lambicus, B. bruxellensis and B. claussenii are native to the Senne Valley region of Belgium, where lambic beer is produced.[32]

Distilled beverages

A distilled beverage is a beverage that contains ethanol that has been purified by distillation. Carbohydrate-containing plant material is fermented by yeast, producing a dilute solution of ethanol in the process. Spirits such as whiskey and rum are prepared by distilling these dilute solutions of ethanol. Components other than ethanol are collected in the condensate, including water, esters, and other alcohols which account for the flavor of the beverage.

Fresh grapes

Wine

Main article: Fermentation (wine)

Yeast is used in winemaking where it converts the sugars present in grape juice or must into alcohol. Yeast is normally already invisibly present on the grapes. The fermentation can be done with this endogenous wild yeast;[33] however, this may give unpredictable results depending on the exact types of yeast species present. For this reason a pure yeast culture is generally added to the must, which rapidly comes to dominate the fermentation. This represses wild yeasts and ensures a reliable and predictable fermentation.[34] Most added wine yeasts are strains of Saccharomyces cerevisiae, though not all strains of the species are suitable.[34] 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.[35] Significant research has been undertaken into the development of novel wine yeast strains that produce atypical flavour profiles or increased complexity in wines.[36][37]

The growth of some yeasts such as Zygosaccharomyces and Brettanomyces in wine can result in wine faults and subsequent spoilage.[38] Brettanomyces produces an array of metabolites when growing in wine, some of which are volatile phenolic compounds. Together these compounds are often referred to as "Brettanomyces character", and are often described as antiseptic or "barnyard" type aromas. Brettanomyces is a significant contributor to wine faults within the wine industry.[39]

Baking

Main article: Baker's yeast

Yeast, most commonly Saccharomyces cerevisiae, is used in baking as a leavening agent, where it converts the fermentable sugars present in dough into the gas carbon dioxide. 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, or sugar in a bread dough accelerates the growth of yeasts. Salt , hot water and fats such as butter slow yeast growth [citation needed]. Most yeasts used in baking are of the same species common in alcoholic fermentation. Additionally, Saccharomyces exiguus (also known as S. minor) is a wild yeast found on plants, fruits, and grains that is occasionally used for baking. Sugar and vinegar are the best conditions for yeast to ferment. In bread making the yeast initially respires aerobically, producing carbon dioxide and water. When the oxygen is depleted anaerobic respiration begins, producing ethanol as a waste product; however, this evaporates during baking.[40]

A block of fresh yeast.

It is not known when yeast was first used to bake bread. The first records that show this use came from Ancient Egypt.[41] Researchers speculate that 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.

Active dried yeast, a granulated form in which yeast is commercially sold.

Today there are several retailers of baker's yeast; one of the best-known in North America is Fleischmann’s Yeast, which was developed in 1868. During World War II Fleischmann's developed a granulated active dry yeast, which did not require refrigeration and had a longer shelf life than fresh yeast. The company created yeast that would rise twice as fast, reducing baking time. Baker's yeast is also sold as a fresh yeast compressed into a square "cake". This form perishes quickly, and must therefore be used soon after production. A weak solution of water and sugar can be used to determine if 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 using a sourdough starter, flour and water are added instead of sugar; this is referred to as proofing the sponge.

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. Some bread doughs are knocked back after one rising and left to rise again. A longer rising time gives a better flavour, but the yeast can fail to raise the bread in the final stages if it is left for too long initially. The dough is then shaped into loaves, left to rise until it is the correct size, and then baked. Dried yeast is usually specified for use in a bread machine, however a (wet) sourdough starter can also work.

Bioremediation

Some yeasts can find potential application in the field of bioremediation. One such yeast, Yarrowia lipolytica, is known to degrade palm oil mill effluent,[42] TNT (an explosive material),[43] and other hydrocarbons such as alkanes, fatty acids, fats and oils.[44] It can also tolerate high concentrations of salt and heavy metals,[45] and is being investigated for its potential as a heavy metal biosorbent.[46]

Industrial ethanol production

The ability of yeast to convert sugar into ethanol has been harnessed by the biotechnology industry to produce ethanol fuel. The process starts by milling a feedstock, such as sugar cane, field corn, or cheap cereal grains, and then adding dilute sulfuric acid, or fungal alpha amylase enzymes, to break down the starches into complex sugars. A gluco amylase 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 concentration.[47]

Saccharomyces yeasts have been genetically engineered to ferment xylose, one of the major fermentable sugars present in cellulosic biomasses, such as agriculture residues, paper wastes, and wood chips.[48][49] Such a development means that ethanol can be efficiently produced from more inexpensive feedstocks, making cellulosic ethanol fuel a more competitively priced alternative to gasoline fuels.[50]

Non-alcoholic beverages

A Kombucha culture fermenting in a jar

Root beer and other sweet carbonated beverages can be produced using the same methods as beer, except that fermentation is stopped sooner, producing carbon dioxide, but only trace amounts of alcohol, and a significant amount of sugar is left in the drink. Yeast in symbiosis with acetic acid bacteria is used in the preparation of Kombucha, a fermented sweetened tea. Species of yeast found in the tea can vary, and may include: Brettanomyces bruxellensis, Candida stellata, Schizosaccharomyces pombe, Torulaspora delbrueckii and Zygosaccharomyces bailii.[51] Kombucha is a popular beverage among Eastern Europe and some former soviet republics, under the name Kvas. Kefir and kumis are made by fermenting milk with yeast and bacteria.[52]

Nutritional supplements

See also: Tibicos

Yeast is used in nutritional supplements popular with vegans and the health conscious, where it is often referred to as "nutritional yeast". It is a deactivated yeast, usually Saccharomyces cerevisiae. It is an excellent source of protein and vitamins, especially the B-complex vitamins, whose functions are related to metabolism as well as other minerals and cofactors required for growth. It is also naturally low in fat and sodium. Some brands of nutritional yeast, though not all, are fortified with vitamin B12, which is produced separately from bacteria. Nutritional yeast, though it has a similar appearance to brewer's yeast, is very different and has a very different taste.

Nutritional yeast has a nutty, cheesy, creamy flavor which makes it popular as an ingredient in cheese substitutes. It is often used by vegans in place of Parmesan cheese. Another popular use is as a topping for popcorn. It can also be used in mashed and fried potatoes, as well as putting it into scrambled eggs. It comes in the form of flakes, or as a yellow powder similar in texture to cornmeal, and can be found in the bulk aisle of most natural food stores. In Australia it is sometimes sold as "savory yeast flakes". Though "nutritional yeast" usually refers to commercial products, inadequately fed prisoners have used "home-grown" yeast to prevent vitamin deficiency.[53]

Probiotics

Some probiotic supplements use the yeast Saccharomyces boulardii to maintain and restore the natural flora in the large and small gastrointestinal tract. S. boulardii has been shown to reduce the symptoms of acute diarrhea in children,[54][55] prevent reinfection of Clostridium difficile,[56] reduce bowel movements in diarrhea predominant IBS patients,[57] and reduce the incidence of antibiotic,[58] traveler's,[59] and HIV/AIDS[60] associated diarrheas.

Aquarium hobby

Yeast is often used by aquarium hobbyists to generate carbon dioxide (CO2) to fertilize plants in planted aquariums. A homemade setup is widely used as a cheap and simple alternative to pressurized CO2 systems. While not as effective as these, the homemade setup is considerably cheaper for less demanding hobbyists.

There are several recipes for homemade CO2, but they are variations of the basic recipe: Baking yeast is inserted in a plastic bottle together with sugar, baking soda and water. This produces CO2 for about 2 or 3 weeks. The CO2 is injected in the aquarium via a narrow hose and released through a CO2 diffuser that helps dissolve the gas in the water. The CO2 is used by plants in the photosynthesis process.

CO2 injection is very important to plant growth in planted aquariums.[61]

Science

Diagram showing a yeast cell

Several yeasts, particularly Saccharomyces cerevisiae, have been widely used in genetics and cell biology. This is largely because S. cerevisiae is a simple eukaryotic cell, serving as a model for all eukaryotes including humans for the study of fundamental cellular processes such as the cell cycle, DNA replication, recombination, cell division and metabolism. Also yeasts are easily manipulated and cultured in the lab which has allowed for the development of powerful standard techniques, such as Yeast two-hybrid, Synthetic genetic array analysis and tetrad analysis. Many proteins important in human biology were first discovered by studying their homologs in yeast; these proteins include cell cycle proteins, 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.[62] At the time it was the most complex organism to have its full genome sequenced and took 7 years and the involvement of more than 100 laboratories to accomplish.[63] The second yeast species to have its genome sequenced was Schizosaccharomyces pombe, which was completed in 2002.[64][65] It was the sixth eukaryotic genome sequenced and consists of 13.8 million base pairs.

Yeast extract

Main article: Yeast extract
Marmite and Vegemite have a distinctive dark colour
Vegemite and Marmite, products made from yeast extract

Yeast extract is the common name for various forms of processed yeast products that are used as food additives or flavours. They are often used in the same way that monosodium glutamate (MSG) is used, and like MSG, often contain free glutamic acid. The general method for making yeast extract for food products such as Vegemite and Marmite on a commercial scale is to add salt to a suspension of yeast making the solution hypertonic, which leads to the cells shrivelling up. This triggers autolysis, where the yeast's 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 which would give poor texture) are separated. Yeast autolysates are used in Vegemite and Promite (Australia); Marmite, Bovril and Oxo (the United Kingdom, Republic of Ireland and South Africa); and Cenovis (Switzerland).

Pathogenic yeasts

A photomicrograph of Candida albicans showing hyphal outgrowth and other morphological characteristics.

Some species of yeast are opportunistic pathogens where they can cause infection in people with compromised immune systems.

Cryptococcus neoformans is a significant pathogen of immunocompromised people causing the disease termed cryptococcosis. This disease occurs in about 7–9% of AIDS patients in the USA, and a slightly smaller percentage (3–6%) in western Europe.[66] The cells of the 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 Candida genus are another group of opportunistic pathogens which causes oral and vaginal infections in humans, known as candidiasis. Candida is commonly found as a commensal yeast in the mucus membranes of humans and other warm-blooded animals. However, sometimes these same strains can become pathogenic. Here the yeast cells sprout a hyphal outgrowth, which locally penetrates the mucosal membrane, causing irritation and shedding of the tissues.[66] The pathogenic yeasts of candidiasis in probable descending order of virulence for humans are: C. albicans, C. tropicalis, C. stellatoidea, C. glabrata, C. krusei, C. parapsilosis, C. guilliermondii, C. viswanathii, C. lusitaniae and Rhodotorula mucilaginosa.[67] Candida glabrata is the second most common Candida pathogen after C. albicans, causing infections of the urogenital tract, and of the bloodstream (candidemia).[68]

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.[69] During their growth, yeasts metabolize some food components and produce metabolic end products. This causes the physical, chemical, and sensory properties of a food to change, and the food is spoiled.[70] The growth of yeast within food products is often seen on their surface, as in cheeses or meats, or by the fermentation of sugars in beverages, such as juices, and semi-liquid products, such as syrups and jams.[69] The yeast of the Zygosaccharomyces genus have had a long history as a spoilage yeast within the food industry. This is mainly due to the fact that these species can grow in the presence of high sucrose, ethanol, acetic acid, sorbic acid, benzoic acid, and sulfur dioxide concentrations,[71] representing some of the commonly used food preservation methods. Methylene blue is used to test for the presence of live yeast cells.

See also

Template:Fungiportal

Look up yeast in Wiktionary, the free dictionary.
Wikimedia Commons has media related to Yeasts.

Footnotes

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  19. ^ Kaufmann, Klaus (2002). Making Sauerkraut and Pickled Vegetables at Home: Creative Recipes for Lactic Fermented Food to Improve Your Health. Google books: Book Publishing Company. {{cite book}}: Unknown parameter |coauthors= ignored (|author= suggested) (help)
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  26. ^ Barley, Shanta (10 February 2010). "Stinky flower is kept warm by yeast partner". New Scientist. Retrieved 10 February 2010.
  27. ^ Little, Ainslie. "Black yeast symbionts compromise the efficiency of antibiotic defenses in fungus-growing ants". Ecology. 89 (5): 1216–1222. ISSN 10.1890/07-0815.1 doi: 10.1890/07-0815.1. {{cite journal}}: Check |issn= value (help); Unknown parameter |coauthors= ignored (|author= suggested) (help)
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Cited texts

  • Alexopoulos CJ, Mims CW, Blackwell M. (1996). Introductory Mycology. New York: Wiley. ISBN 0-471-52229-5.{{cite book}}: CS1 maint: multiple names: authors list (link)
  • Kirk PM, Cannon PF, Minter DW, Stalpers JA. (2008). Dictionary of the Fungi. 10th ed. Wallingford: CABI. ISBN 0-85199-826-7.{{cite book}}: CS1 maint: multiple names: authors list (link)
  • Moore-Landecker E. (1996). Fundamentals of the Fungi. Englewood Cliffs, New Jersey: Prentice Hall. ISBN 0-13-376864-3.

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