Saccharomycotina

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Saccharomycotina
Candida albicans.jpg
Candida albicans
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Fungi
Division: Ascomycota
(unranked): Saccharomyceta
Subdivision: Saccharomycotina
O.E. Erikss. & Winka 1997 [1]
Classes
Synonyms
  • Hemiascomycotina (Brefeldt 1891) Cavalier-Smith 1998

Saccharomycotina is a subdivision (subphylum) of the division (phylum) Ascomycota in the kingdom Fungi. [2] [3] It comprises most of the ascomycete yeasts. The members of Saccharomycotina reproduce by budding and they do not produce ascocarps (fruiting bodies). [2] [4]

Contents

The subdivision includes a single class: Saccharomycetes, which again contains a single order: Saccharomycetales. [2] [3]

Notable members of Saccharomycotina are the baker's yeast Saccharomyces cerevisiae and the genus Candida that includes several human pathogens.

Etymology

The name comes from the Greek word σάκχαρον (sákkharon), meaning "sugar" and μύκης (mukēs) meaning "fungus".

History and economic importance

Historical records from ancient Egypt and China describe the processes of brewing and baking from 10,000 to 8,000 years ago, and the production of fermented beverages and foods seems to have paralleled the beginning of agriculture. [5] In the 1850s, Louis Pasteur demonstrated that yeasts are responsible for the fermentation of grape juice to wine. [6] [7]

Saccharomycotina include some of the economically most important fungi known. Members include species of industrial and agricultural importance (e.g. brewing, baking, fermentation of food products, production of citric acid, production of recombinant proteins, biofuel production, biological pest control of crops). Other species cause economic losses worldwide (plant pathogens, contaminants of foods and beverages). Yet others are animal and human pathogens. [8] [9]

Morphology

Saccharomycete yeasts usually grow as single cells. Their cellular morphology is fairly simple, although their growth form is highly adapted. Asci are naked and ascospores can have several forms. No species produce ascocarps (fruiting bodies). Saccharomycete genomes are often smaller than those of filamentous fungi. [2] [10] [11] [4]

Some species (e.g. Metschnikowia species) tend to form chains of budding cells that are termed pseudohyphae. [2] Yet other species are able to produce true septate hyphae. [4] Such species (e.g. Candida albicans ) are termed dimorphic, which means they can propagate both as budding yeasts and as filamentous hyphae.

Reproduction

Asexual reproduction

Asexual reproduction occurs mainly vegetatively by mitosis and budding. Saccharomycotina is characterized by holoblastic budding, [12] which means all layers of the parent cell wall are involved in the budding event. This leaves a scar through which no further budding occurs.

Asexual cells may vary in shape. [13] The shape of the cell may be informative in terms of detecting mode of reproduction or taxonomic placement to genera or species.

Although not commonly known, some species form endospores (e.g. Candida species). [2] These are asexual spores that are formed within their mother cell (hyphal or single cell). Strains of Candida and Metschnikowia may also form asexual resting spores called chlamydospores. [2]

Sexual reproduction

Sexual reproduction is not known for all species of Saccharomycotina, but may happen in certain species if environmental conditions favour it (e.g. deficiency in nitrogen and carbohydrate). [2] Sexual reproduction is well known in Saccharomyces cerevisiae . Here, the life cycle involves alternation between a haploid and a diploid phase. The life cycle proceeds as follows: Two cells of different mating type fuse and the nuclei undergo karyogamy. This results in a daughter cell with a diploid nucleus, functioning as an ascus, where meiosis occurs to produce haploid ascospores. When ascospores germinate, the haploid phase is established, and is maintained by further mitosis and budding. In most natural populations this phase is fairly short since ascospores fuse almost immediately after meiosis has occurred. This results in most yeast populations being diploid for most part of their life cycle. [4]

In Saccharomycotina there are two mating types present. The mating types specify peptide hormones called pheromones and corresponding receptors for each type. These pheromones organize the mating. The pheromones do not affect the same mating type or diploids, but bind to receptors of different mating type. Interaction between pheromone and receptor results in altered metabolism to allow for fusion between cells of different mating type. [4] [2]

Distribution and ecology

Saccharomycete yeasts are found in nearly all regions of the world, including hot deserts, polar areas, in freshwater, in salt water, and in the atmosphere. [2] Their growth is mainly saprotrophic, but some members are important pathogens of plants and animals, including humans. They are often found in specialized habitats, e.g. small volumes of organic carbon rich liquid (e.g. flower nectar). [4]

Examples of ecological modes in Saccharomycotina:

Although yeasts are commonly isolated from soil, few are believed to have soil as a primary habitat. [2]

Accurate identification of species is important for understanding yeast ecology, something that is now possible with the increased use of DNA-based methods. Before molecular methods were available, identification was mainly based on morphology, something that resulted in misclassifications and further prevented reliable results of ecological research.

Taxonomy

Saccharomycotina is a subdivision (subphylum) of the division (phylum) Ascomycota. It is a sister group to Pezizomycotina. [2] [3]

Yeasts were traditionally classified as a separate group of the fungal kingdom, but in recent years[ when? ] DNA-based methods have changed the understanding of phylogenetic relationships among fungi. Yeasts are considered to be a polyphyletic group, [2] [11] consisting of members of Basidiomycota, Taphrinomycotina, as well as Saccharomycotina. This realization has led to major changes in the phylogeny and taxonomy of Saccharomycotina. [2]

In addition, the recent[ when? ] changes in the International Code of Nomenclature for algae, fungi and plants [17] [18] have had a major impact on the classification of fungi, including Saccharomycotina. The changes imply that a fungus can only bear one correct name, i.e. separate names for anamorphs and teleomorphs are no longer allowed. This involves major changes in Saccharomycotina taxonomy, as many species are currently described from both anamorphic and teleomorphic stages. [18] The genus Candida is an example of a genus that is undergoing large-scale revisions.

Molecular identification methods are important tools for discovery of new species and subsequently give better understanding of biodiversity in this group. Much of the future classification of Saccharomycotina will rest on phylogenetic analysis of DNA sequences rather than on the morphological and developmental characters.[ citation needed ]

Phylogeny

Phylogeny by Groenewald et al. 2023 [19]

Saccharomycotina
Lipomycetes
Lipomycetales

Lipomycetaceae Novák & Zsolt 1961

Trigonopsidomycetes
Trigonopsidales

Trigonopsidaceae Lachance & Kurtzman 2013

Dipodascomycetes
Alloascoideomycetes
Alloascoideales

Alloascoideaceae Kurtzman & Robnett 2013

Sporopachydermiomycetes
Sporopachydermiales

Sporopachydermiaceae Groenewald et al. 2023

Pichiomycetes
Alaninales

Pachysolenaceae Groenewald et al. 2023

Pichiales

Pichiaceae Zender 1925

Serinales

Cephaloascaceae Batra 1973

Metschnikowiaceae Kamienski 1899 ex Doweld 2013

Debaryomycetaceae Kurtzman & Suzuki 2010

Saccharomycetes
Ascoideales

Ascoideaceae Engl. 1892

Saccharomycopsidaceae Arx & Van der Walt 1987

Phaffomycetales

Wickerhamomycetaceae Kurtzman, Robnett & Bas.-Powers 2008

Phaffomycetaceae Yamada et al. 1999

Saccharomycodales

Saccharomycodaceae Kudrjanzev 1960

Saccharomycetales

Endomycetaceae Schröter 1893

Saccharomycetaceae Winter 1881

See also

Related Research Articles

<span class="mw-page-title-main">Yeast</span> Informal group of fungi

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

<span class="mw-page-title-main">Basidiomycota</span> Division of fungi

Basidiomycota is one of two large divisions that, together with the Ascomycota, constitute the subkingdom Dikarya within the kingdom Fungi. Members are known as basidiomycetes. More specifically, Basidiomycota includes these groups: agarics, puffballs, stinkhorns, bracket fungi, other polypores, jelly fungi, boletes, chanterelles, earth stars, smuts, bunts, rusts, mirror yeasts, and Cryptococcus, the human pathogenic yeast.

<span class="mw-page-title-main">Ascomycota</span> Division or phylum of fungi

Ascomycota is a phylum of the kingdom Fungi that, together with the Basidiomycota, forms the subkingdom Dikarya. Its members are commonly known as the sac fungi or ascomycetes. It is the largest phylum of Fungi, with over 64,000 species. The defining feature of this fungal group is the "ascus", a microscopic sexual structure in which nonmotile spores, called ascospores, are formed. However, some species of Ascomycota are asexual and thus do not form asci or ascospores. Familiar examples of sac fungi include morels, truffles, brewers' and bakers' yeast, dead man's fingers, and cup fungi. The fungal symbionts in the majority of lichens such as Cladonia belong to the Ascomycota.

<span class="mw-page-title-main">Ascus</span> Spore-bearing cell in ascomycete fungi

An ascus is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores, produced by meiosis followed, in most species, by a mitotic cell division. However, asci in some genera or species can occur in numbers of one, two, four, or multiples of four. In a few cases, the ascospores can bud off conidia that may fill the asci with hundreds of conidia, or the ascospores may fragment, e.g. some Cordyceps, also filling the asci with smaller cells. Ascospores are nonmotile, usually single celled, but not infrequently may be coenocytic, and in some cases coenocytic in multiple planes. Mitotic divisions within the developing spores populate each resulting cell in septate ascospores with nuclei. The term ocular chamber, or oculus, refers to the epiplasm that is surrounded by the "bourrelet".

<i>Candida albicans</i> Species of fungus

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 usually a commensal organism, but it can become pathogenic in immunocompromised individuals under a variety of conditions. It is one of the few species of the genus Candida that cause the human infection candidiasis, which results from an overgrowth of the fungus. Candidiasis is, for example, often observed in HIV-infected patients. C. albicans is the most common fungal species isolated from biofilms either formed on (permanent) implanted medical devices or on human tissue. C. albicans, C. tropicalis, C. parapsilosis, and C. glabrata are together responsible for 50–90% of all cases of candidiasis in humans. A mortality rate of 40% has been reported for patients with systemic candidiasis due to C. albicans. By one estimate, invasive candidiasis contracted in a hospital causes 2,800 to 11,200 deaths yearly in the US. Nevertheless, these numbers may not truly reflect the true extent of damage this organism causes, given new studies indicating that C. albicans can cross the blood–brain barrier in mice.

Heterothallic species have sexes that reside in different individuals. The term is applied particularly to distinguish heterothallic fungi, which require two compatible partners to produce sexual spores, from homothallic ones, which are capable of sexual reproduction from a single organism.

<i>Saccharomyces</i> Genus of fungi

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 food production where they are known as brewer's yeast, baker's yeast and sourdough starter among others. They are unicellular and saprotrophic fungi. One example is Saccharomyces cerevisiae, which is used in making bread, wine, and beer, and for human and animal health. Other members of this genus include the wild yeast Saccharomyces paradoxus that is the closest relative to S. cerevisiae, Saccharomyces bayanus, used in making wine, and Saccharomyces cerevisiaevar. boulardii, used in medicine.

<span class="mw-page-title-main">Mating of yeast</span> Biological process of yeast

The mating of yeast, also known as yeast sexual reproduction, is a fundamental biological process that promotes genetic diversity and adaptation in yeast species. Yeast species, such as Saccharomyces cerevisiae, are single-celled eukaryotes that can exist as either haploid cells, which contain a single set of chromosomes, or diploid cells, which contain two sets of chromosomes. Haploid yeast cells come in two mating types, a and α, each producing specific pheromones to identify and interact with the opposite type, thus displaying simple sexual differentiation. A yeast cell's mating type is determined by a specific genetic locus known as MAT, which governs its mating behaviour. Haploid yeast can switch mating types through a form of genetic recombination, allowing them to change mating type as often as every cell cycle. When two haploid cells of opposite mating types encounter each other, they undergo a complex signaling process that leads to cell fusion and the formation of a diploid cell. Diploid cells can reproduce asexually, but under nutrient-limiting conditions, they undergo meiosis to produce new haploid spores.

<i>Torulaspora delbrueckii</i> Species of fungus

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 10662, etc.. The type strain of T. delbrueckii CBS 1146 T was sequenced in 2009, and is composed of 8 chromosomes in addition to a mitochondrial genome.

<span class="mw-page-title-main">Saccharomycetes</span> Class of fungi

Saccharomycetes belongs to the Ascomycota division of the kingdom Fungi. It is the only class in the subdivision Saccharomycotina, the budding yeasts. Saccharomycetes contains a single order, Saccharomycetales.

<span class="mw-page-title-main">Mating in fungi</span> Combination of genetic material between compatible mating types

Fungi are a diverse group of organisms that employ a huge variety of reproductive strategies, ranging from fully asexual to almost exclusively sexual species. Most species can reproduce both sexually and asexually, alternating between haploid and diploid forms. This contrasts with most multicellular eukaryotes such as mammals, where the adults are usually diploid and produce haploid gametes which combine to form the next generation. In fungi, both haploid and diploid forms can reproduce – haploid individuals can undergo asexual reproduction while diploid forms can produce gametes that combine to give rise to the next generation.

<i>Nakaseomyces glabratus</i> Species of fungus

Nakaseomyces glabratus is a species of haploid yeast of the genus Nakaseomyces, previously known as Candida glabrata. Despite the fact that no sexual life cycle has been documented for this species, N. glabratus strains of both mating types are commonly found. N. glabrata is generally a commensal of human mucosal tissues, but in today's era of wider human immunodeficiency from various causes, N. glabratus is often the second or third most common cause of candidiasis as an opportunistic pathogen. Infections caused by N. glabratus can affect the urogenital tract or even cause systemic infections by entrance of the fungal cells in the bloodstream (Candidemia), especially prevalent in immunocompromised patients.

<span class="mw-page-title-main">Taphrinomycotina</span> Subdivision of fungi

The Taphrinomycotina are one of three subdivisions constituting the Ascomycota and is more or less synonymous with the slightly older invalid name Archiascomycetes. Recent molecular studies suggest that the group is monophyletic and basal to the rest of the Ascomycota.

<i>Kluyveromyces marxianus</i> Species of fungus

Kluyveromyces marxianus in ascomycetous yeast and member of the genus, Kluyveromyces. It is the sexual stage of Atelosaccharomyces pseudotropicalis also known as Candida kefyr. This species has a homothallic mating system and is often isolated from dairy products.

The mating-type locus is a specialized region in the genomes of some yeast and other fungi, usually organized into heterochromatin and possessing unique histone methylation patterns. The genes in this region regulate the mating type of the organism and therefore determine key events in its life cycle, such as whether it will reproduce sexually or asexually. In fission yeast such as S. pombe, the formation and maintenance of the heterochromatin organization is regulated by RNA-induced transcriptional silencing, a form of RNA interference responsible for genomic maintenance in many organisms. Mating type regions have also been well studied in budding yeast S. cerevisiae and in the fungus Neurospora crassa.

Pathogenic fungi are fungi that cause disease in humans or other organisms. Although fungi are eukaryotic, many pathogenic fungi are microorganisms. Approximately 300 fungi are known to be pathogenic to humans; their study is called "medical mycology". Fungal infections are estimated to kill more people than either tuberculosis or malaria—about two million people per year.

Brettanomyces claussenii is a wild yeast of the genus Brettanomyces which has a negative Pasteur effect. It and Brettanomyces anomalus share identical mtDNA. In the wild, it is found on the skins of fruit. It has been shown to be useful for wine and beer fermentation as well as ethanol production.

Sexual selection has been observed in fungi as a part of their reproduction, although they also often reproduce asexually. In the basidiomycetes, the sex ratio is biased towards males, implying sexual selection there. Male–male competition to fertilize occurs in fungi including yeasts. Pheromone signaling is used by female gametes and by conidia, implying male choice in these cases. Female–female competition may also occur, indicated by the much faster evolution of female-biased genes in fungi.

<i>Candida tropicalis</i> Species of fungus

Candida tropicalis is a species of yeast in the genus Candida. It is a common pathogen in neutropenic hosts, in whom it may spread through the bloodstream to peripheral organs. For invasive disease, treatments include amphotericin B, echinocandins, or extended-spectrum triazole antifungals.

Candida catenulata is a yeast-form fungus in the phylum Ascomycota. It is distributed globally and commonly found on the skin of humans and animals, in soil, and in dairy products.

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