Mating in fungi - Knowledge

464:, recessive mutations affecting the diploid stage of the life cycle are quite frequent in natural populations. These mutations, when homozygous in the diploid stage, often cause spores to have maturation defects or to produce barren fruiting bodies with few ascospores (sexual spores). The majority of these homozygous mutations cause abnormal meiosis (e.g. disturbed chromosome pairing or disturbed pachytene or diplotene). The number of genes affecting the diploid stage was estimated to be at least 435 (about 4% of the total number of 9,730 genes). Thus, outcrossing, promoted by the necessity for union of opposite mating types, likely provides the benefit of masking recessive mutations that would otherwise be deleterious to sexual spore formation (see 334:

necessary for expression of genes involved in sexual development. The protoperithecium consists of an ascogonium, a coiled multicellular hypha that is enclosed in a knot-like aggregation of hyphae. A branched system of slender hyphae, called the trichogyne, extends from the tip of the ascogonium projecting beyond the sheathing hyphae into the air. The sexual cycle is initiated (i.e. fertilization occurs) when a cell, usually a conidium, of opposite mating type contacts a part of the trichogyne (see Figure). Such contact can be followed by cell fusion leading to one or more nuclei from the fertilizing cell migrating down the trichogyne into the ascogonium. Since both

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30 minutes to induce germination. For normal strains, the entire sexual cycle takes 10 to 15 days. In a mature ascus containing eight ascospores, pairs of adjacent spores are identical in genetic constitution, since the last division is mitotic, and since the ascospores are contained in the ascus sac that holds them in a definite order determined by the direction of nuclear segregations during meiosis. Since the four primary products are also arranged in sequence, a first division segregation pattern of genetic markers can be distinguished from a second division segregation pattern.

228: 1012:; they sense different molecules (in this case the pheromones) outside and activate a specific pathway inside of the cell. Pheromone-receptor interaction occurs in a way that the pheromone from one individual interacts with the receptor from the partner and vice versa. The functions of these genes are to regulate reciprocal nuclear exchange, nuclear migration in both mates and ultimately clamp cell fusion. The first mating pheromone-receptor genes characterized were for 128: 108: 3158: 92: 479: 607: 80:

Mating between isogamous fungi may consist only of a transfer of a nucleus from one cell to another. Vegetative incompatibility within species often prevents a fungal isolate from mating with another isolate. Isolates of the same incompatibility group do not mate or mating does not lead to successful

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A mature perithecium may contain as many as 300 asci, each derived from identical fusion diploid nuclei. Ordinarily, in nature, when the perithecia mature the ascospores are ejected rather violently into the air. These ascospores are heat resistant and, in the lab, require heating at 60 °C for

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haploid cells have been outlined by Fincham and Day. and Wagner and Mitchell. After fusion of the cells, the further fusion of their nuclei is delayed. Instead, a nucleus from the fertilizing cell and a nucleus from the ascogonium become associated and begin to divide synchronously. The products

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reproduces by mitosis as either haploid or diploid cells. However, when starved, diploid cells undergo meiosis to form haploid spores. Mating occurs when haploid cells of opposite mating type, MATa and MATα, come into contact. Ruderfer et al. pointed out that such contacts are frequent between

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mating type strains. Both can form abundant protoperithecia, the female reproductive structure (see Figure). Protoperithecia are formed most readily in the laboratory when growth occurs on solid (agar) synthetic medium with a relatively low source of nitrogen. Nitrogen starvation appears to be

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life cycle. The haploid mycelium reproduces asexually by two processes: (1) simple proliferation of existing mycelium, and (2) formation of conidia (macro- and micro-) which can be dispersed and then germinate to produce new mycelium. In the sexual cycle, mating can only occur between individual

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mating proteins MATα2 and MATa1. In Agaricomycotina the two types of homeodomain transcription factors are termed HD1 and HD2; so the HD1 and HD2 proteins from an individual interacts with the HD2 and HD1 proteins from the other partner, respectively, generating heterodimers able to activate the

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involves activation of the same mating pathways characteristic of sex in outcrossing species, i.e. self-fertilization does not bypass required pathways for outcrossing sex but instead requires activation of these pathways within a single individual. Fusion of haploid nuclei occurs within

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Homothallic species may likely have evolved from heterothallic ancestors (Lin and Heitman 2007). In Basidiomycota homothallism is not very common and in Agaricomycotina it is estimated that only 10% of species have homothallic mating behavior. For example, one subspecies of the

542:) are unlikely to be sufficient for generally maintaining sex from one generation to the next. Instead, a short-term benefit, such as meiotic recombinational repair of DNA damages caused by stressful conditions such as starvation, may be the key to the maintenance of sex in 460:, meiosis occurring in the brief diploid stage is one of their most complex processes. The haploid multicellular vegetative stage, although physically much larger than the diploid stage, characteristically has a simple modular construction with little differentiation. In 695:. Since sexual reproduction takes place in haploid organisms, it cannot proceed until complementary genes are provided by a suitable partner through cell or hyphal fusion. The number of mating types depends on the number of genes and the number of alleles for each. 1790:

Birdsell JA, Wills C (2003). The evolutionary origin and maintenance of sexual recombination: A review of contemporary models. Evolutionary Biology Series >> Evolutionary Biology, Vol. 33 pp. 27–137. MacIntyre, Ross J.; Clegg, Michael, T (Eds.), Springer.

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is heterothallic, it appears that, in nature, mating is most often between closely related yeast cells. The relative rarity in nature of meiotic events that result from outcrossing suggests that the possible long-term benefits of outcrossing (e.g. generation of

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from a single individual are self-sterile and need to interact with another compatible individual for mating to take place. Additionally, mating compatibility in the Basidiomycota is further categorized into two types of mating systems: tetrapolar and bipolar.

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known to outcross under laboratory conditions, both mating types are not normally distributed in natural populations, with the α mating type much more commonly found (>99%), suggesting homothallism is the most prevalent mode of sexual reproduction in

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Aimi, T.; Yoshida, R.; Ishikawa, M.; Bao, D.; Kitamoto, Y. (2005). "Identification and linkage mapping of the genes for the putative homeodomain protein (hox1) and the putative pheromone receptor protein homologue (rcb1) in a bipolar basidiomycete,

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Mortimer, Robert K.; Romano, Patrizia; Suzzi, Giovanna; Polsinelli, Mario (December 1994). "Genome renewal: A new phenomenon revealed from a genetic study of 43 strains ofSaccharomyces cerevisiae derived from natural fermentation of grape musts".

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of cells directly produced by a single meiosis, and these cells can mate with each other. The second reason is that haploid cells of one mating type, upon cell division, often produce cells of the opposite mating type with which they may mate.

546:. Alternatively, recessive deleterious mutations accumulate during the diploid expansion phase, and are purged during selfing: this purging has been termed "genome renewal" and provides an advantage of sex that does not depend on outcrossing. 50:

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.

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As the above events are occurring, the mycelial sheath that had enveloped the ascogonium develops as the wall of the perithecium becomes impregnated with melanin, and blackens. The mature perithecium has a flask-shaped structure.

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analysis simple, since recessive traits will show up in the offspring. Analysis of genetic recombination is facilitated by the ordered arrangement of the products of meiosis within a sac-like structure called an

795:, determine the "specificity" or sexual identity of the individual harboring them. Only individuals with different mating types are compatible with each other and therefore able to start the mating event. 2610:

Lin, X.; Heitman, J. (2007). "Mechanisms of Homothallism in Fungi and Transitions between Heterothallism and Homothallism". In J. Heitman; J. W. Kronstad; J. W. Taylor; L. A. Casselton (eds.).

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hyphae (containing separate haploid nuclei from both initial parents). Dikaryotic hyphae, under the appropriate environmental conditions will give rise to the fruiting body which contains the

174:. The zygosporangium is a unique structure to the Zygomycota and is easily recognizable in microscopy due to its characteristic dark color and spiky shape. The nuclei join in a process called 2034:

James, Timothy (2007). "Analysis of mating type locus organization and synteny in mushroom fungi: Beyond model species". In Heitman, J.; Kronstad, J.W.; Taylor, J.W.; Casselton, L.A. (eds.).

806:– specialized cells in which sexual recombination via karyogamy and meiosis occurs. This dikaryotic condition in Basidiomycota is often maintained by a specialized hyphal structure called a 2625:

Kües, U; Navarro-González, M. (2010). "Mating-type orthologous genes in the primarily homothallic Moniliophthora perniciosa, the causal agent of Witches' Broom Disease in cacao".

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pair of haploid nuclei within the crozier divide synchronously. Next, septa form to divide the crozier into three cells. The central cell in the curve of the hook contains one

2271:"Evolution of the bipolar mating system of the mushroom Coprinellus disseminatus from its tetrapolar ancestors involves loss of mating-type-specific pheromone receptor function" 201:

develops one of two complementary organs, a "female" ascogonium or a "male" antheridium. These organs resemble gametangia except that they contain only nuclei. A bridge, the

409:. The diploid nucleus has 14 chromosomes formed from the two fused haploid nuclei that had 7 chromosomes each. Formation of the diploid nucleus is immediately followed by 456:

can only occur between strains of different mating type suggests that some degree of outcrossing is favored by natural selection. In haploid multicellular fungi, such as

182:, which grows into a mature diploid zygomycete. A diploid zygomycete can then undergo meiosis to create spores, which disperse and germinate. The following generations of 529:

populations clonal reproduction and a type of “self-fertilization” (in the form of intratetrad mating) predominate. Ruderfer et al. analyzed the ancestry of natural

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of the opposite mating type through the trichogyne. Fusion of the nuclei of opposite mating types occurs within the protoperithecium to form a zygote (2N) nucleus.

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forms around the merged tips, enclosing nuclei from both isolates. A second pair of septa forms two adjacent cells, one on each side. These adjacent cells, called

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strains have the same sexual structures, neither strain can be regarded as exclusively male or female. However, as a recipient, the protoperithecium of both the

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mating types, approach close to 100% chance of encountering a compatible partner in nature, due to the huge number of mating types generated by these systems.

202: 751:(the mushroom-forming fungi) about 90% of the species are heterothallic. The tetrapolar type of mating system is ruled by two unlinked mating loci termed 313:. Sexual fruiting bodies (perithecia) can only be formed when two cells of different mating type come together (see Figure). Like other Ascomycetes, 1027:

mating locus contains genes that code for two types of homeodomain transcription factor proteins, usually tightly linked, that are homologues to the

366:) migrate into numerous ascogenous hyphae, which then begin to grow out of the ascogonium. Each of these ascogenous hyphae bends to form a hook (or 565:

is the most common fungal pathogen in humans. It causes both debilitating mucosal infections and potentially life-threatening systemic infections.

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nucleus in each ascus. Meiosis is an essential part of the life cycle of all sexually reproducing organisms, and in its main features, meiosis in

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species that exhibit a sexual cycle the overwhelming majority in nature are homothallic (self-fertilizing). Selfing in the homothallic fungus

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and diploid forms. This contrasts with most multicellular eukaryotes such as mammals, where the adults are usually diploid and produce haploid

2103:"The b alleles of U. maydis, whose combinations program pathogenic development, code for polypeptides containing a homeodomain-related motif" 237:

strains of different mating type, A and a. Fertilization occurs by the passage of nuclei of conidia or mycelium of one mating type into the

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nuclei fuse with each other to form a diploid nucleus (see Figure). This nucleus is the only diploid nucleus in the entire life cycle of

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that can grow to form a further crozier that can then form its own ascus-initial cell. This process can then be repeated multiple times.

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Ruderfer DM, Pratt SC, Seidel HS, Kruglyak L (September 2006). "Population genomic analysis of outcrossing and recombination in yeast".

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or hyphal fusion. When a receptor on one haploid detects a pheromone from a complementary mating type, it approaches the source through

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needs to switch from white to opaque cells. The latter are more efficient in mating and referred to as the mating competent cells of

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reproductive structures termed cleistothecia, in which the diploid zygote undergoes meiotic divisions to yield haploid ascospores.

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mating types, an individual has an approximately 50% chance to encounter a compatible mate in nature. However, species such as

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closely related yeast cells for two reasons. The first is that cells of opposite mating type are present together in the same

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is homothallic, although other subspecies have maintained their ability to outcross. Also, a variety of the edible mushroom

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mating has focused on several model species with different behaviour. Not all fungi reproduce sexually and many that do are

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strains can be thought of as the female structure, and the fertilizing conidium can be thought of as the male participant.

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nucleus (see Figure). This binuclear cell initiates ascus formation and is called an “ascus-initial” cell. Next the two

2054:"The pheromone cell signaling components of the Ustilago a mating-type loci determine intercompatibility between species" 1604:

Raju NB, Leslie JF (October 1992). "Cytology of recessive sexual-phase mutants from wild strains of Neurospora crassa".

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with conserved residues and the pheromone receptors belong to the G protein-coupled family of receptors located in the

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has maintained an elaborate, but largely hidden, mating apparatus. Johnson suggested that mating strategies may allow

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Westergaard, Mogens; Mitchell, Herschel K. (1947). "Neurospora V. A Synthetic Medium Favoring Sexual Reproduction".

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lives mainly in tropical and sub-tropical regions. It often can be found growing on dead plant matter after fires.

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Both mating loci have become physically linked such that they now act as a single locus; this has occurred in the

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Schulz, B.; Banuett, F.; Dahl, M.; Schlesinger, R.; Schäfer, W.; Martin, T.; Herskowitz, I.; Kahmann, R. (1990).

1736:"Heterothallism in Saccharomyces cerevisiae isolates from nature: effect of HO locus on the mode of reproduction" 798:

A successful mating interaction begins with nuclear exchange and nuclear migration resulting in the formation of

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Brown, A. J.; Casselton, L. A. (2001). "Mating in mushrooms: increasing the chances but prolonging the affair".

3057: 1857:"The consequences of rare sexual reproduction by means of selfing in an otherwise clonally reproducing species" 1557:"Recessive mutations from natural populations of Neurospora crassa that are expressed in the sexual diplophase" 632: 3114: 3192: 3129: 2929: 2912: 1734:

Katz Ezov T, Chang SL, Frenkel Z, Segrè AV, Bahalul M, Murray AW, Leu JY, Korol A, Kashi Y (January 2010).

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strains and concluded that outcrossing occurs only about once every 50,000 cell divisions. Thus, although

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Lengeler, K.B.; Fox, D.S.; Fraser, J.A.; Forrester, K.; Dietrich, F.S.; Heitman, J.; et al. (2002).

954: 2320:"Linkage of mating-type loci distinguishes bipolar from tetrapolar mating in basidiomycetous smut fungi" 1278:

Turgeon, B. Gillian (1998). "Application of Mating Type Gene Technology to Problems in Fungal Biology".

948:. Virulence success in these two pathogens is highly associated with mating and their mating type locus. 730:, when mating occurs within a single individual, or in other words each individual is self-fertile; and 3067: 2907: 465: 2897: 1947:

Bennett, R.J.; Johnson, A.D. (2005). "Mating in Candida Albicans and the Search for a Sexual Cycle".

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A picture of the mating type mechanism has begun to emerge from studies of particular fungi such as

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Among the 250 known species of aspergilli, about 36% have an identified sexual state Among those

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mating locus is predominantly diallelic, which reduces the occurrence of outcrossing within these

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Wagner RP, Mitchell HK. (1964). Genetics and Metabolism. John Wiley and Sons, Inc., New York

2714:"Mating type and the genetic basis of self-fertility in the model fungus Aspergillus nidulans" 2246:

Casselton, L.A.; Kües, U. (2007). "The origin of multiple mating types in the model mushrooms

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In contrast, bipolar mating systems are ruled by a single allelic mating locus, termed either

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forms, that provides a passage for nuclei to travel from the antheridium to the ascogonium. A

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Fincham J RS, Day PR (1963). Fungal Genetics. Blackwell Scientific Publications, Oxford, UK.

831: 69:; thus, for many members of the fungal kingdom, the terms "male" and "female" do not apply. 3074: 2725: 2331: 1960: 1747: 1109: 298: 35: 8: 2971: 2939: 825: 290: 39: 2729: 2712:

Paoletti M, Seymour FA, Alcocer MJ, Kaur N, Calvo AM, Archer DB, Dyer PS (August 2007).

2335: 1751: 1229:"An overview of the function and maintenance of sexual reproduction in dikaryotic fungi" 1036:

transcriptional regulated pathway, which involves formation of clamp cells, coordinated

3084: 3062: 3047: 2650: 2513: 2460: 2431: 2295: 2270: 2220: 2195: 2168: 2143: 2078: 2053: 2013: 1929: 1881: 1856: 1834: 1768: 1735: 1716: 1581: 1556: 1497: 1472: 1453: 1449: 1413: 1388: 1339: 1314: 1255: 1228: 1204: 1171: 1133: 768: 2673:"Sexual development and cryptic sexuality in fungi: insights from Aspergillus species" 2547: 2407: 2378: 2000: 1983: 1665: 1640: 3187: 3089: 2875: 2804: 2743: 2694: 2689: 2672: 2642: 2592: 2587: 2570: 2551: 2505: 2465: 2412: 2359: 2354: 2319: 2300: 2225: 2173: 2124: 2119: 2102: 2083: 2005: 1964: 1921: 1886: 1826: 1792: 1773: 1759: 1708: 1670: 1621: 1586: 1502: 1418: 1367: 1344: 1295: 1260: 1209: 1191: 1139: 1063: 1001: 925:

locus lost functionality in determining mating type, as has occurred in the mushroom

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species. Most species can reproduce both sexually and asexually, alternating between

2654: 2517: 2196:"Orchestration of sexual reproduction and virulence by the fungal mating-type locus" 2017: 1838: 390:

cells on either side of the first ascus-forming cell fuse with each other to form a

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Proceedings of the National Academy of Sciences of the United States of America

901:. In Agaricomycotina, bipolar organisms mostly have multiple alleles for their 259: 227: 167: 74: 2738: 2713: 2501: 2211: 1872: 3176: 3094: 3025: 2991: 2902: 2868: 1404: 1245: 1195: 1073: 1041: 1009: 935: 731: 719: 214: 134: 81:

offspring. High variation has been reported including same-chemotype mating,

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to survive in the hostile environment of a mammalian host. In order to mate

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hypha grows towards a compatible mate and they both form a bridge, called a

3161: 3013: 2890: 2885: 2831: 2747: 2698: 2646: 2638: 2555: 2509: 2469: 2416: 2304: 2268: 2229: 2177: 2009: 1968: 1925: 1890: 1852: 1777: 1712: 1422: 1348: 1299: 1264: 1213: 1187: 1145: 1005: 703: 286: 70: 2596: 2571:"The a mating type locus of U. maydis specifies cell signaling components" 2363: 2128: 2087: 1830: 1822: 1674: 1625: 1590: 1506: 687:. Sexual reproduction thereby depends on pheromones produced from variant 2254:". In Heitman, J.; Kronstad, J.W.; Taylor, J.W.; Casselton, L.A. (eds.). 1128: 851: 727: 387: 58: 2451: 2159: 1904:

Johnson A (November 2003). "The biology of mating in Candida albicans".

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Heterothallism is the most common mating system in Basidiomycota and in

726:. In general for fungi there are two main types of sexual reproduction: 1917: 1540: 1524: 1457: 997: 905:

mating locus; however, in Ustilaginomycotina and Pucciniomycotina, the

810:. The formation of clamp connections is regulated by both mating loci. 799: 707: 362:

of these nuclear divisions (still in pairs of unlike mating type, i.e.

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Philosophical Transactions of the Royal Society B: Biological Sciences

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because it is easy to grow and has a haploid life cycle: this makes

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Sex in Fungi: Molecular Determination and Evolutionary Implications

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Sex in Fungi: Molecular determination and evolutionary implications

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Sex in Fungi: Molecular Determination and Evolutionary Implications

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is a diploid fungus that grows both as a yeast and as a filament.

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Depending on the species, sexual reproduction takes place through

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have the most complex systems of sexual reproduction known among

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in nature. Finally, the fungus causing witches' broom in cacao,

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Slonczewski, Joan L.; Foster, John W.; Zinser, Erik R. (2020).

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in 1958. The results of these experiments led directly to the "

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to gametophyte mating and biparental transfer of mitochondria.

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are a diverse group of organisms that employ a huge variety of

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Combination of genetic material between compatible mating types

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Nieuwenhuis, Bart P. S.; James, Timothy Y. (19 October 2016).

1807: 723: 514: 302: 171: 325:. There is no evident morphological difference between the 2809: 1690: 1387:

Horowitz NH, Berg P, Singer M, et al. (January 2004).

913:. Bipolarity likely arose via one of two potential routes: 692: 317:

has two mating types that, in this case, are symbolized by

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James, T.Y.; Srivilai, P.; Kües, U.; Vilgalys, R. (2006).

2047: 2045: 1733: 1641:"Life cycle of the budding yeast Saccharomyces cerevisiae" 2788: 2241: 2239: 2144:"Basidiomycete mating type genes and pheromone signaling" 775:), both of which can be multiallelic. The combination of 421:

mating type. One further mitotic division leads to four

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provide structural support. The central cell, called the

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species only isolates of opposite mating types can mate.

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Typical mating fusion of two compatible monokaryons in

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James, T.Y.; Lee, M.; van Diepen, L.T.A. (2011).

2236: 444: 2533: 2529: 2527: 2189: 2187: 2029: 2027: 1088:, has a primarily homothallic biology despite having 967: 525:

Katz Ezov et al. presented evidence that in natural

1435: 952:Other bipolar species include the white rot fungus 73:species are able to mate with themselves, while in 2524: 2184: 2024: 1386: 1982:Miller, Mathew G.; Johnson, Alexander D. (2002). 3174: 2670: 2141: 2051: 1470: 1312: 1227:Wallen, R. Margaret; Perlin, Michael H. (2018). 1169: 850:It is believed that multi-allelic systems favor 101:form progametangia with suspensors during mating 1473:"Sexual development genes of Neurospora crassa" 813:Examples of tetrapolar organisms are the smuts 504:, brewer's and baker's yeast, is in the phylum 397:After formation of the ascus-initial cell, the 2609: 2258:. Washington, DC: ASM Press. pp. 283–300. 2038:. Washington, DC: ASM Press. pp. 317–331. 1946: 854:in Basidiomycota. For example, in the case of 2773: 2383:: A step in the evolution of sex chromosomes" 1981: 1315:"Dancing genomes: fungal nuclear positioning" 2705: 2193: 1727: 1638: 1226: 186:can undergo asexual or sexual reproduction. 2614:. Washington DC: ASM Press. pp. 35–57. 2569:Bölker, M.; Urban, M.; Kahmann, R. (1992). 1851: 1686: 1684: 1632: 1471:Nelson MA, Metzenberg RL (September 1992). 1389:"A centennial: George W. Beadle, 1903–1989" 996:locus there are linked genes that code for 635:. Unsourced material may be challenged and 471: 2780: 2766: 1603: 1554: 713: 293:in the experiments for which they won the 250:is a type of red bread mold of the phylum 2737: 2688: 2586: 2459: 2406: 2353: 2343: 2294: 2219: 2167: 2118: 2077: 1999: 1903: 1880: 1767: 1664: 1580: 1550: 1548: 1496: 1412: 1338: 1254: 1244: 1203: 655:Learn how and when to remove this message 353:The subsequent steps following fusion of 1784: 1681: 477: 226: 2666: 2664: 1897: 1277: 671:. The mating type genes are located in 589:since meiosis is still not observed in 189: 141: 3175: 1961:10.1146/annurev.micro.59.030804.121310 1545: 121:and antheridium with trichogyne bridge 2761: 2671:Dyer PS, O'Gorman CM (January 2012). 2318:Bakkeren, G.; Kronstad, J.W. (1994). 2052:Bakkeren, G.; Kronstad, J.W. (1996). 2033: 295:Nobel Prize in Physiology or Medicine 2661: 1855:; Lyttle, David N. (December 2011). 1555:Leslie JF, Raju NB (December 1985). 743:Tetrapolar and bipolar mating system 633:adding citations to reliable sources 600: 433:seems typical of meiosis generally. 220: 154:, by joining at the hyphal tips via 2876:Heterogametic sex / Homogametic sex 2142:Raudaskoski, M.; Kothe, E. (2010). 1313:Gladfelter, A.; Berman, J. (2009). 549: 13: 1450:10.1002/j.1537-2197.1947.tb13032.x 14: 3204: 2194:Hsueh, Y.P.; Heitman, J. (2008). 1364:Microbiology: An Evolving Science 57:is a complex process governed by 3157: 3156: 2950:Evolution of sexual reproduction 2690:10.1111/j.1574-6976.2011.00308.x 1760:10.1111/j.1365-294X.2009.04436.x 1069:A. bisporus var. eurotetrasporus 858:, which bears more than 25 605: 275:). In its natural environment, 126: 106: 90: 2618: 2603: 2562: 2476: 2423: 2370: 2311: 2262: 2200:Current Opinion in Microbiology 2135: 2094: 1975: 1940: 1845: 1801: 1597: 1529: 1280:Annual Review of Phytopathology 1172:"The frequency of sex in fungi" 1047: 882:, which has more than 339 870:, which has more than 240 209:grows from the ascogonium, and 3058:Sexual reproduction in animals 1861:Theoretical Population Biology 1657:10.1128/MMBR.52.4.536-553.1988 1639:Herskowitz I (December 1988). 1513: 1464: 1429: 1380: 1366:(5th ed.). W. W. Norton. 1355: 1306: 1292:10.1146/annurev.phyto.36.1.115 1271: 1220: 1163: 1096:mating type-like genes in its 596: 1: 2627:Journal of Basic Microbiology 2548:10.1016/s0168-9525(01)02343-5 2001:10.1016/S0092-8674(02)00837-1 1949:Annual Review of Microbiology 1156: 482:The yeast cell's life cycle: 2930:Sex as a biological variable 2913:Simultaneous hermaphroditism 2588:10.1016/0092-8674(92)90182-c 2120:10.1016/0092-8674(90)90744-y 710:movement if it is a gamete. 517:, the sac that contains the 7: 2434:Phanerochaete chrysosporium 2399:10.1128/ec.1.5.704-718.2002 2287:10.1534/genetics.105.051128 2070:10.1093/genetics/143.4.1601 1906:Nature Reviews Microbiology 1319:Nature Reviews Microbiology 1117: 1004:. The pheromones are short 955:Phanerochaete chrysosporium 718:Some of the species within 417:mating type and two of the 301:" hypothesis that specific 194:As it approaches a mate, a 10: 3209: 3068:Penile-vaginal intercourse 2908:Sequential hermaphroditism 1573:10.1093/genetics/111.4.759 1489:10.1093/genetics/132.1.149 1438:American Journal of Botany 1151:Fungus § Reproduction 942:and in the human pathogen 466:Complementation (genetics) 445:Benefit of mating type in 170:, is destined to become a 20: 3152: 3105: 2938: 2898:Testis-determining factor 2795: 2739:10.1016/j.cub.2007.07.012 2677:FEMS Microbiology Reviews 2502:10.1007/s00294-005-0012-7 2212:10.1016/j.mib.2008.09.014 1873:10.1016/j.tpb.2011.08.004 1233:Frontiers in Microbiology 1146:Dioecy § In mycology 1085:Moniliophthora perniciosa 886:mating types and 64 23:Sexual selection in fungi 2839:Sex-determination system 1405:10.1534/genetics.166.1.1 1246:10.3389/fmicb.2018.00503 1029:Saccharomyces cerevisiae 958:and the edible mushroom 928:Coprinellus disseminatus 878:mating types, each, and 759:(in Agaricomycotina) or 501:Saccharomyces cerevisiae 473:Saccharomyces cerevisiae 309:. This concept launched 2787: 2381:Cryptococcus neoformans 2345:10.1073/pnas.91.15.7085 1645:Microbiological Reviews 945:Cryptococcus neoformans 714:Mating in Basidiomycota 32:reproductive strategies 2822:Sexual differentiation 2639:10.1002/jobm.201000013 2379:"Mating-type locus of 1188:10.1098/rstb.2015.0540 1059:Sistotrema brinkmannii 496: 242: 38:to almost exclusively 2864:Temperature-dependent 2252:Schizophyllum commune 1823:10.1002/yea.320101203 917:During evolution the 832:Schizophyllum commune 481: 370:) at its tip and the 230: 34:, ranging from fully 1325:(875–886): 875–886. 1110:Aspergillus nidulans 823:, and the mushrooms 629:improve this section 299:one gene, one enzyme 190:Mating in Ascomycota 142:Mating in Zygomycota 3193:Sexual reproduction 3080:Hormonal motivation 3053:Fungal reproduction 2972:Reproductive system 2730:2007CBio...17.1384P 2452:10.1128/ec.00212-10 2336:1994PNAS...91.7085B 2248:Coprinopsis cinerea 2160:10.1128/ec.00319-09 1752:2010MolEc..19..121K 1331:10.1038/nrmicro2249 1002:pheromone receptors 826:Coprinopsis cinerea 291:George Wells Beadle 3085:Human reproduction 3063:Sexual intercourse 3048:Plant reproduction 2536:Trends in Genetics 1918:10.1038/nrmicro752 1182:(1706): 20150540. 1134:Mating-type region 791:) alleles, termed 769:Ustilaginomycotina 679:for production of 497: 305:code for specific 243: 3170: 3169: 3090:Lordosis behavior 2805:Sexual dimorphism 2330:(15): 7085–7089. 1817:(12): 1543–1552. 1740:Molecular Ecology 1373:978-0-393-42865-0 1140:Neurospora crassa 1064:Agaricus bisporus 665: 664: 657: 540:genetic diversity 311:molecular biology 247:Neurospora crassa 233:Neurospora crassa 222:Neurospora crassa 3200: 3160: 3159: 3120:Animal sexuality 3043:Sexual selection 2782: 2775: 2768: 2759: 2758: 2752: 2751: 2741: 2709: 2703: 2702: 2692: 2668: 2659: 2658: 2622: 2616: 2615: 2607: 2601: 2600: 2590: 2566: 2560: 2559: 2531: 2522: 2521: 2490:Current Genetics 2480: 2474: 2473: 2463: 2427: 2421: 2420: 2410: 2374: 2368: 2367: 2357: 2347: 2315: 2309: 2308: 2298: 2281:(3): 1877–1891. 2266: 2260: 2259: 2243: 2234: 2233: 2223: 2191: 2182: 2181: 2171: 2139: 2133: 2132: 2122: 2098: 2092: 2091: 2081: 2064:(4): 1601–1613. 2049: 2040: 2039: 2031: 2022: 2021: 2003: 1979: 1973: 1972: 1944: 1938: 1937: 1901: 1895: 1894: 1884: 1849: 1843: 1842: 1805: 1799: 1788: 1782: 1781: 1771: 1731: 1725: 1724: 1688: 1679: 1678: 1668: 1636: 1630: 1629: 1601: 1595: 1594: 1584: 1552: 1543: 1533: 1527: 1517: 1511: 1510: 1500: 1468: 1462: 1461: 1433: 1427: 1426: 1416: 1384: 1378: 1377: 1359: 1353: 1352: 1342: 1310: 1304: 1303: 1275: 1269: 1268: 1258: 1248: 1224: 1218: 1217: 1207: 1167: 1038:nuclear division 924: 920: 908: 904: 900: 896: 889: 885: 877: 873: 865: 862:but only 2 861: 844:Laccaria bicolor 838:Pleurotus djamor 808:clamp connection 790: 786: 782: 778: 773:Pucciniomycotina 766: 762: 758: 754: 660: 653: 649: 646: 640: 609: 601: 587:parasexual cycle 558:Candida albicans 551:Candida albicans 130: 110: 94: 3208: 3207: 3203: 3202: 3201: 3199: 3198: 3197: 3173: 3172: 3171: 3166: 3148: 3135:Differentiation 3125:Human sexuality 3115:Plant sexuality 3101: 2997:Spermatogenesis 2941: 2934: 2797: 2791: 2786: 2756: 2755: 2718:Current Biology 2710: 2706: 2669: 2662: 2623: 2619: 2608: 2604: 2567: 2563: 2532: 2525: 2486:Pholiota nameko 2481: 2477: 2440:Eukaryotic Cell 2428: 2424: 2387:Eukaryotic Cell 2375: 2371: 2316: 2312: 2267: 2263: 2244: 2237: 2192: 2185: 2148:Eukaryotic Cell 2140: 2136: 2099: 2095: 2050: 2043: 2032: 2025: 1980: 1976: 1945: 1941: 1902: 1898: 1850: 1846: 1806: 1802: 1789: 1785: 1732: 1728: 1693:Nature Genetics 1689: 1682: 1637: 1633: 1618:10.1139/g92-124 1602: 1598: 1553: 1546: 1534: 1530: 1518: 1514: 1469: 1465: 1444:(10): 573–577. 1434: 1430: 1385: 1381: 1374: 1360: 1356: 1311: 1307: 1276: 1272: 1225: 1221: 1168: 1164: 1159: 1124:Mating of yeast 1120: 1055:ectomycorrhizal 1050: 986: 961:Pholiota nameko 938:plant pathogen 922: 918: 906: 902: 898: 894: 887: 883: 880:S. commune 875: 871: 868:C. cinerea 863: 859: 816:Ustilago maydis 788: 784: 780: 776: 764: 760: 756: 752: 749:Agaricomycotina 745: 716: 661: 650: 644: 641: 626: 610: 599: 554: 495: 476: 452:That mating in 450: 392:binucleate cell 239:protoperithecia 225: 192: 144: 137: 131: 122: 111: 102: 95: 55:Mating in fungi 25: 19: 12: 11: 5: 3206: 3196: 3195: 3190: 3185: 3168: 3167: 3165: 3164: 3153: 3150: 3149: 3147: 3146: 3145: 3144: 3143: 3142: 3137: 3132: 3117: 3111: 3109: 3103: 3102: 3100: 3099: 3098: 3097: 3092: 3087: 3082: 3077: 3072: 3071: 3070: 3055: 3050: 3045: 3040: 3039: 3038: 3033: 3023: 3022: 3021: 3016: 3006: 3005: 3004: 2999: 2989: 2984: 2979: 2974: 2969: 2964: 2963: 2962: 2957: 2946: 2944: 2936: 2935: 2933: 2932: 2927: 2922: 2917: 2916: 2915: 2910: 2900: 2895: 2894: 2893: 2888: 2881:Sex chromosome 2878: 2873: 2872: 2871: 2866: 2861: 2856: 2851: 2846: 2836: 2835: 2834: 2829: 2819: 2818: 2817: 2812: 2801: 2799: 2793: 2792: 2785: 2784: 2777: 2770: 2762: 2754: 2753: 2724:(16): 1384–9. 2704: 2660: 2633:(5): 442–451. 2617: 2602: 2581:(3): 441–450. 2561: 2542:(7): 393–400. 2523: 2496:(3): 184–194. 2475: 2446:(2): 249–261. 2422: 2393:(5): 704–718. 2369: 2310: 2261: 2235: 2206:(6): 517–524. 2183: 2154:(6): 847–859. 2134: 2113:(2): 295–306. 2093: 2041: 2023: 1994:(3): 293–302. 1974: 1939: 1896: 1867:(4): 317–322. 1844: 1800: 1797:978-0306472619 1783: 1726: 1705:10.1038/ng1859 1699:(9): 1077–81. 1680: 1631: 1596: 1544: 1528: 1512: 1463: 1428: 1379: 1372: 1354: 1305: 1270: 1219: 1161: 1160: 1158: 1155: 1154: 1153: 1148: 1143: 1136: 1131: 1126: 1119: 1116: 1057:Basidiomycete 1049: 1046: 985: 966: 950: 949: 932: 856:U. maydis 744: 741: 732:heterothallism 715: 712: 683:and pheromone 663: 662: 613: 611: 604: 598: 595: 553: 548: 494: 493: 490: 487: 483: 475: 470: 449: 443: 260:model organism 224: 219: 213:occurs in the 191: 188: 168:zygosporangium 143: 140: 139: 138: 132: 125: 123: 112: 105: 103: 96: 89: 61:. Research on 17: 9: 6: 4: 3: 2: 3205: 3194: 3191: 3189: 3186: 3184: 3181: 3180: 3178: 3163: 3155: 3154: 3151: 3141: 3138: 3136: 3133: 3131: 3128: 3127: 3126: 3123: 3122: 3121: 3118: 3116: 3113: 3112: 3110: 3108: 3104: 3096: 3095:Pelvic thrust 3093: 3091: 3088: 3086: 3083: 3081: 3078: 3076: 3073: 3069: 3066: 3065: 3064: 3061: 3060: 3059: 3056: 3054: 3051: 3049: 3046: 3044: 3041: 3037: 3034: 3032: 3029: 3028: 3027: 3026:Fertilization 3024: 3020: 3017: 3015: 3012: 3011: 3010: 3007: 3003: 3000: 2998: 2995: 2994: 2993: 2992:Gametogenesis 2990: 2988: 2985: 2983: 2980: 2978: 2975: 2973: 2970: 2968: 2965: 2961: 2958: 2956: 2953: 2952: 2951: 2948: 2947: 2945: 2943: 2937: 2931: 2928: 2926: 2925:parasexuality 2923: 2921: 2918: 2914: 2911: 2909: 2906: 2905: 2904: 2903:Hermaphrodite 2901: 2899: 2896: 2892: 2889: 2887: 2884: 2883: 2882: 2879: 2877: 2874: 2870: 2869:Haplodiploidy 2867: 2865: 2862: 2860: 2857: 2855: 2852: 2850: 2847: 2845: 2842: 2841: 2840: 2837: 2833: 2830: 2828: 2825: 2824: 2823: 2820: 2816: 2813: 2811: 2808: 2807: 2806: 2803: 2802: 2800: 2794: 2790: 2783: 2778: 2776: 2771: 2769: 2764: 2763: 2760: 2749: 2745: 2740: 2735: 2731: 2727: 2723: 2719: 2715: 2708: 2700: 2696: 2691: 2686: 2683:(1): 165–92. 2682: 2678: 2674: 2667: 2665: 2656: 2652: 2648: 2644: 2640: 2636: 2632: 2628: 2621: 2613: 2606: 2598: 2594: 2589: 2584: 2580: 2576: 2572: 2565: 2557: 2553: 2549: 2545: 2541: 2537: 2530: 2528: 2519: 2515: 2511: 2507: 2503: 2499: 2495: 2491: 2487: 2479: 2471: 2467: 2462: 2457: 2453: 2449: 2445: 2441: 2437: 2435: 2426: 2418: 2414: 2409: 2404: 2400: 2396: 2392: 2388: 2384: 2382: 2373: 2365: 2361: 2356: 2351: 2346: 2341: 2337: 2333: 2329: 2325: 2321: 2314: 2306: 2302: 2297: 2292: 2288: 2284: 2280: 2276: 2272: 2265: 2257: 2253: 2249: 2242: 2240: 2231: 2227: 2222: 2217: 2213: 2209: 2205: 2201: 2197: 2190: 2188: 2179: 2175: 2170: 2165: 2161: 2157: 2153: 2149: 2145: 2138: 2130: 2126: 2121: 2116: 2112: 2108: 2104: 2097: 2089: 2085: 2080: 2075: 2071: 2067: 2063: 2059: 2055: 2048: 2046: 2037: 2030: 2028: 2019: 2015: 2011: 2007: 2002: 1997: 1993: 1989: 1985: 1978: 1970: 1966: 1962: 1958: 1954: 1950: 1943: 1935: 1931: 1927: 1923: 1919: 1915: 1912:(2): 106–16. 1911: 1907: 1900: 1892: 1888: 1883: 1878: 1874: 1870: 1866: 1862: 1858: 1854: 1853:Masel, Joanna 1848: 1840: 1836: 1832: 1828: 1824: 1820: 1816: 1812: 1804: 1798: 1794: 1787: 1779: 1775: 1770: 1765: 1761: 1757: 1753: 1749: 1746:(1): 121–31. 1745: 1741: 1737: 1730: 1722: 1718: 1714: 1710: 1706: 1702: 1698: 1694: 1687: 1685: 1676: 1672: 1667: 1662: 1658: 1654: 1651:(4): 536–53. 1650: 1646: 1642: 1635: 1627: 1623: 1619: 1615: 1612:(5): 815–26. 1611: 1607: 1600: 1592: 1588: 1583: 1578: 1574: 1570: 1567:(4): 759–77. 1566: 1562: 1558: 1551: 1549: 1542: 1538: 1532: 1526: 1522: 1516: 1508: 1504: 1499: 1494: 1490: 1486: 1483:(1): 149–62. 1482: 1478: 1474: 1467: 1459: 1455: 1451: 1447: 1443: 1439: 1432: 1424: 1420: 1415: 1410: 1406: 1402: 1398: 1394: 1390: 1383: 1375: 1369: 1365: 1358: 1350: 1346: 1341: 1336: 1332: 1328: 1324: 1320: 1316: 1309: 1301: 1297: 1293: 1289: 1285: 1281: 1274: 1266: 1262: 1257: 1252: 1247: 1242: 1238: 1234: 1230: 1223: 1215: 1211: 1206: 1201: 1197: 1193: 1189: 1185: 1181: 1177: 1173: 1166: 1162: 1152: 1149: 1147: 1144: 1142: 1141: 1137: 1135: 1132: 1130: 1127: 1125: 1122: 1121: 1115: 1112: 1111: 1106: 1101: 1099: 1095: 1091: 1087: 1086: 1081: 1080:C. neoformans 1076: 1075: 1074:C. neoformans 1070: 1066: 1065: 1060: 1056: 1045: 1043: 1039: 1035: 1030: 1026: 1022: 1017: 1015: 1011: 1010:cell membrane 1007: 1003: 999: 995: 991: 983: 979: 975: 971: 965: 963: 962: 957: 956: 947: 946: 941: 937: 933: 930: 929: 916: 915: 914: 912: 891: 881: 869: 857: 853: 848: 846: 845: 840: 839: 834: 833: 828: 827: 822: 821:U. longissima 818: 817: 811: 809: 805: 801: 796: 794: 774: 770: 750: 740: 737: 733: 729: 725: 721: 720:Basidiomycota 711: 709: 705: 701: 696: 694: 690: 686: 682: 678: 674: 670: 669:S. cerevisiae 659: 656: 648: 638: 634: 630: 624: 623: 619: 614:This section 612: 608: 603: 602: 594: 592: 588: 584: 580: 576: 572: 568: 564: 560: 559: 552: 547: 545: 544:S. cerevisiae 541: 536: 535:S. cerevisiae 532: 531:S. cerevisiae 528: 527:S. cerevisiae 523: 520: 516: 511: 510:S. cerevisiae 507: 503: 502: 491: 488: 485: 484: 480: 474: 469: 467: 463: 459: 455: 448: 442: 438: 434: 432: 428: 424: 420: 416: 412: 408: 404: 400: 395: 393: 389: 385: 381: 377: 373: 369: 365: 360: 356: 351: 349: 345: 341: 337: 332: 328: 324: 320: 316: 312: 308: 304: 300: 296: 292: 288: 284: 280: 278: 274: 270: 265: 261: 258:is used as a 257: 253: 249: 248: 240: 235: 234: 229: 223: 218: 216: 215:fruiting body 212: 208: 204: 200: 197: 187: 185: 181: 177: 173: 169: 165: 161: 157: 153: 152:progametangia 149: 136: 135:Basidiomycota 129: 124: 120: 116: 113:Fungi within 109: 104: 100: 97:Fungi within 93: 88: 87: 86: 84: 78: 76: 75:heterothallic 72: 68: 64: 60: 56: 52: 49: 45: 41: 37: 33: 29: 24: 16: 3052: 3014:spermatozoon 2942:reproduction 2891:Y chromosome 2886:X chromosome 2832:Virilization 2827:Feminization 2721: 2717: 2707: 2680: 2676: 2630: 2626: 2620: 2611: 2605: 2578: 2574: 2564: 2539: 2535: 2493: 2489: 2485: 2478: 2443: 2439: 2433: 2425: 2390: 2386: 2380: 2372: 2327: 2323: 2313: 2278: 2274: 2264: 2255: 2251: 2247: 2203: 2199: 2151: 2147: 2137: 2110: 2106: 2096: 2061: 2057: 2035: 1991: 1987: 1977: 1952: 1948: 1942: 1909: 1905: 1899: 1864: 1860: 1847: 1814: 1810: 1803: 1786: 1743: 1739: 1729: 1696: 1692: 1648: 1644: 1634: 1609: 1605: 1599: 1564: 1560: 1531: 1515: 1480: 1476: 1466: 1441: 1437: 1431: 1396: 1392: 1382: 1363: 1357: 1322: 1318: 1308: 1283: 1279: 1273: 1236: 1232: 1222: 1179: 1175: 1165: 1138: 1108: 1104: 1102: 1093: 1089: 1083: 1079: 1072: 1068: 1062: 1058: 1051: 1048:Homothallism 1033: 1028: 1024: 1020: 1018: 1013: 1006:polypeptides 993: 989: 987: 981: 977: 973: 969: 959: 953: 951: 943: 939: 926: 892: 879: 867: 855: 849: 842: 836: 830: 824: 820: 814: 812: 797: 792: 746: 728:homothallism 717: 697: 691:of the same 668: 666: 651: 642: 627:Please help 615: 590: 585:is termed a 582: 581:. Mating in 578: 574: 570: 566: 562: 556: 555: 550: 543: 534: 530: 526: 524: 509: 505: 499: 498: 472: 461: 457: 453: 451: 446: 439: 435: 430: 426: 422: 418: 414: 406: 402: 398: 396: 383: 379: 375: 371: 367: 363: 358: 354: 352: 347: 343: 339: 335: 330: 326: 322: 318: 314: 287:Edward Tatum 285:was used by 282: 281: 276: 272: 268: 255: 245: 244: 231: 221: 193: 158:. A pair of 145: 79: 59:mating types 54: 53: 26: 15: 1399:(1): 1–10. 1286:: 115–137. 1129:Mating type 1105:Aspergillus 984:mating loci 852:outcrossing 793:mating type 708:chemotactic 704:chemotropic 675:and encode 597:Mating type 591:C. albicans 583:C. albicans 579:C. albicans 575:C. albicans 571:C. albicans 567:C. albicans 563:C. albicans 489:Conjugation 388:uninucleate 71:Homothallic 3177:Categories 3075:Copulation 2796:Biological 1955:: 233–55. 1541:B00BXTC5BO 1525:B000W851KO 1157:References 998:pheromones 800:dikaryotic 706:growth or 681:pheromones 506:Ascomycota 283:Neurospora 252:Ascomycota 203:trichogyne 199:sac fungus 178:to form a 164:suspensors 156:plasmogamy 148:zygomycete 119:ascogonium 115:Ascomycota 99:Zygomycota 83:sporophyte 21:See also: 3130:Mechanics 3107:Sexuality 3002:Oogenesis 2977:Sex organ 2967:Germ cell 2955:Anisogamy 1196:0962-8436 1042:septation 1014:U. maydis 940:U. hordei 685:receptors 645:June 2021 616:does not 462:N. crassa 458:N. crassa 454:N. crassa 447:N. crassa 431:N. crassa 425:and four 407:N. crassa 315:N. crassa 277:N. crassa 256:N. crassa 211:karyogamy 176:karyogamy 67:isogamous 3188:Mycology 3162:Category 3140:Activity 3036:Internal 3031:External 2920:Intersex 2748:17669651 2699:22091779 2655:36453261 2647:20586074 2556:11418220 2518:25670803 2510:16096791 2470:21131435 2417:12455690 2305:16461425 2275:Genetics 2230:18935978 2178:20190072 2058:Genetics 2018:13859192 2010:12176317 1969:15910278 1926:15035040 1891:21888925 1839:11989104 1778:20002587 1713:16892060 1561:Genetics 1477:Genetics 1423:15020400 1393:Genetics 1349:19898490 1300:15012495 1265:29619017 1214:27619703 1118:See also 673:homeobox 382:and one 307:proteins 207:dikaryon 184:mycelium 2987:Meiosis 2960:Isogamy 2726:Bibcode 2597:1310895 2461:3067404 2364:7913746 2332:Bibcode 2296:1456265 2221:2622727 2169:2901643 2129:1967554 2088:8844149 2079:1207424 1934:1826178 1882:3218209 1831:7725789 1769:3892377 1748:Bibcode 1675:3070323 1626:1427061 1591:2933298 1582:1202670 1507:1356883 1498:1205113 1458:2437339 1414:1470705 1340:2794368 1256:5871698 1239:: 503. 1205:5031624 988:In the 911:species 804:basidia 734:, when 700:gametes 689:alleles 677:enzymes 637:removed 622:sources 486:Budding 411:meiosis 368:crozier 264:genetic 196:haploid 48:gametes 44:haploid 36:asexual 3183:Mating 3009:Gamete 2982:Mating 2940:Sexual 2815:Female 2746: 2697: 2653: 2645: 2595: 2554: 2516: 2508: 2468: 2458: 2415: 2408:126754 2405: 2362: 2352: 2303: 2293: 2228: 2218: 2176: 2166: 2127: 2086: 2076: 2016: 2008: 1967: 1932: 1924: 1889: 1879: 1837: 1829: 1795: 1776: 1766: 1721:783720 1719: 1711: 1673: 1666:373162 1663: 1624: 1606:Genome 1589: 1579: 1539: 1523: 1505: 1495: 1456: 1421: 1411: 1370: 1347: 1337: 1298: 1263: 1253: 1212: 1202: 1194: 1098:genome 736:hyphae 519:tetrad 180:zygote 63:fungal 40:sexual 2798:terms 2651:S2CID 2514:S2CID 2355:44343 2014:S2CID 1930:S2CID 1835:S2CID 1811:Yeast 1717:S2CID 1454:JSTOR 724:fungi 515:ascus 492:Spore 303:genes 271:(pl. 269:ascus 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