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An ascus (; ) is the sexual
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
-bearing
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
produced in
ascomycete 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 defi ...
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
. Each ascus usually contains eight ascospores (or octad), produced by
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
followed, in most species, by a
mitotic In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintai ...
cell division. However, asci in some genera or species can occur in numbers of one (e.g. ''
Monosporascus cannonballus ''Monosporascus cannonballus'' is a species of fungus in the Sordariales The order Sordariales is one of the most diverse taxonomic groups within the Sordariomycetes (subdivision Pezizomycotina, division Ascomycota). Species in the order Sor ...
''), two, four, or multiples of four. In a few cases, the ascospores can bud off conidia that may fill the asci (e.g. ''
Tympanis ''Tympanis'' is a genus of fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are class ...
'') with hundreds of conidia, or the ascospores may fragment, e.g. some ''
Cordyceps ''Cordyceps'' is a genus of ascomycete fungi (sac fungi) that includes about 600 species. Most ''Cordyceps'' species are endoparasitoids, parasitic mainly on insects and other arthropods (they are thus entomopathogenic fungi); a few are parasiti ...
'', also filling the asci with smaller cells. Ascospores are nonmotile, usually single celled, but not infrequently may be coenocytic (lacking a
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interatri ...
), 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 (the portion of
cytoplasm In cell biology, the cytoplasm is all of the material within a eukaryotic cell, enclosed by the cell membrane, except for the cell nucleus. The material inside the nucleus and contained within the nuclear membrane is termed the nucleoplasm. The ...
not used in ascospore formation) that is surrounded by the "bourrelet" (the thickened tissue near the top of the ascus). Typically, a single ascus will contain eight ascospores (or octad). The eight spores are produced by
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
followed by a
mitotic In cell biology, mitosis () is a part of the cell cycle in which replicated chromosomes are separated into two new nuclei. Cell division by mitosis gives rise to genetically identical cells in which the total number of chromosomes is maintai ...
division. Two meiotic divisions turn the original
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
zygote A zygote (, ) is a eukaryotic cell formed by a fertilization event between two gametes. The zygote's genome is a combination of the DNA in each gamete, and contains all of the genetic information of a new individual organism. In multicellula ...
nucleus into four
haploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
ones. That is, the single original diploid cell from which the whole process begins contains two complete sets of
chromosome A chromosome is a long DNA molecule with part or all of the genetic material of an organism. In most chromosomes the very long thin DNA fibers are coated with packaging proteins; in eukaryotic cells the most important of these proteins are ...
s. In preparation for meiosis, all the DNA of both sets is duplicated, to make a total of four sets. The nucleus that contains the four sets divides twice, separating into four new nuclei – each of which has one complete set of chromosomes. Following this process, each of the four new nuclei duplicates its DNA and undergoes a division by mitosis. As a result, the ascus will contain four pairs of spores. Then the ascospores are released from the ascus. In many cases the asci are formed in a regular layer, the ''
hymenium The hymenium is the tissue layer on the hymenophore of a fungal fruiting body where the cells develop into basidia or asci, which produce spores. In some species all of the cells of the hymenium develop into basidia or asci, while in others some ...
'', in a fruiting body which is visible to the naked eye, here called an ''
ascocarp An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are m ...
'' or ''ascoma''. In other cases, such as single-celled
yeast Yeasts are eukaryotic, single-celled microorganisms classified as members of the fungus kingdom. The first yeast originated hundreds of millions of years ago, and at least 1,500 species are currently recognized. They are estimated to constitut ...
s, no such structures are found. In rare cases asci of some genera can regularly develop inside older discharged asci one after another, e.g. ''
Dipodascus ''Dipodascus'' is a genus of fungi in the family Dipodascaceae The Dipodascaceae are a family of yeasts in the order Saccharomycetales. According to the 2007 Outline of Ascomycota, the family contains four genera; however, the placement of ''Sp ...
''. Asci normally release their spores by bursting at the tip, but they may also digest themselves, passively releasing the ascospores either in a liquid or as a dry powder. Discharging asci usually have a specially differentiated tip, either a pore or an operculum. In some hymenium forming genera, when one ascus bursts, it can trigger the bursting of many other asci in the ascocarp resulting in a massive discharge visible as a cloud of spores – the phenomenon called "puffing". This is an example of
positive feedback Positive feedback (exacerbating feedback, self-reinforcing feedback) is a process that occurs in a feedback loop which exacerbates the effects of a small disturbance. That is, the effects of a perturbation on a system include an increase in the ...
. A faint hissing sound can also be heard for species of ''
Peziza ''Peziza'' is a large genus of saprophytic cup fungi that grow on the ground, rotting wood, or dung. Most members of this genus are of unknown edibility and are difficult to identify as separate species without use of microscopy. The polyphylet ...
'' and other
cup fungi The Pezizaceae (commonly referred to as cup fungi) are a family of fungi in the Ascomycota which produce mushrooms that tend to grow in the shape of a "cup". Spores are formed on the inner surface of the fruit body (mushroom). The cup shape ty ...
. Asci, notably those of ''
Neurospora crassa ''Neurospora crassa'' is a type of red bread mold of the phylum Ascomycota. The genus name, meaning "nerve spore" in Greek, refers to the characteristic striations on the spores. The first published account of this fungus was from an infestation ...
'', have been used in laboratories for studying the process of meiosis, because the four
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
s produced by meiosis line up in regular order. By modifying
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
s coding for spore color and nutritional requirements, the biologist can study crossing over and other phenomena. The formation of asci and their use in genetic analysis are described in detail in ''Neurospora crassa''. Asci of most
Pezizomycotina Pezizomycotina make up most of the Ascomycota fungi and include most lichenized fungi too. Pezizomycotina contains the filamentous ascomycetes and is a subdivision of the Ascomycota (fungi that form their spores in a sac-like ''ascus''). It is mo ...
develop after the formation of croziers at their base. The croziers help maintain a brief
dikaryon The dikaryon is a nuclear feature which is unique to certain fungi. (The green alga ''Derbesia'' had been long considered an exception, until the heterokaryotic hypothesis was challenged by later studies.) Compatible cell-types can fuse cytoplasms ( ...
. The compatible nuclei of the dikaryon merge forming a
diploid Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectively ...
nucleus that then undergoes
meiosis Meiosis (; , since it is a reductional division) is a special type of cell division of germ cells in sexually-reproducing organisms that produces the gametes, such as sperm or egg cells. It involves two rounds of division that ultimately resu ...
and ultimately internal ascospore formation. Members of the
Taphrinomycotina The Taphrinomycotina are one of three subdivisions constituting the Ascomycota (fungi that form their spores in a sac-like ascus) and is more or less synonymous with the slightly older invalid name Archiascomycetes (sometimes spelled Archaeascom ...
and
Saccharomycotina Saccharomycotina is a subdivision (subphylum) of the division (phylum) Ascomycota in the kingdom Fungi. It comprises most of the ascomycete yeasts. The members of Saccharomycotina reproduce by budding and they do not produce ascocarps (fruitin ...
do not form croziers.


Classification

The form of the ascus, the capsule which contains the sexual spores, is important for classification of the
Ascomycota 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 def ...
. There are four basic types of ascus. *A unitunicate-operculate ascus has a "lid", the Operculum, which breaks open when the spores are mature and allows the spores to escape. Unitunicate-operculate asci only occur in those ascocarps which have apothecia, for instance the morels. 'Unitunicate' means 'single-walled'. *Instead of an operculum, a unitunicate-inoperculate ascus has an elastic ring that functions like a pressure valve. Once mature the elastic ring briefly expands and lets the spores shoot out. This type appears both in
apothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mo ...
and in
perithecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mos ...
; an example is the illustrated ''Hypomyces chrysospermus''. *A bitunicate ascus is enclosed in a double wall. This consists of a thin, brittle outer shell and a thick elastic inner wall. When the spores are mature, the shell splits open so that the inner wall can take up water. As a consequence this begins to extend with its spores until it protrudes above the rest of the ascocarp so that the spores can escape into free air without being obstructed by the bulk of the fruiting body. Bitunicate asci occur only in
pseudothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mos ...
and are found only in the classes
Dothideomycetes Dothideomycetes is the largest and most diverse class of ascomycete fungi. It comprises 11 orders 90 families, 1300 genera and over 19,000 known species. Traditionally, most of its members were included in the loculoascomycetes, which is not par ...
and Chaetothyriomycetes (which were formerly united in the old class Loculoascomycetes). Examples: ''Venturia inaequalis'' (
apple scab Apple scab is a common disease of plants in the rose family ( Rosaceae) that is caused by the ascomycete fungus ''Venturia inaequalis''. While this disease affects several plant genera, including ''Sorbus, Cotoneaster,'' and '' Pyrus'', it is m ...
) and ''Guignardia aesculi'' (Brown Leaf Mold of Horse Chestnut). *Prototunicate asci are mostly spherical in shape and have no mechanism for forcible dispersal. The mature ascus wall dissolves allowing the spores to escape, or it is broken open by other influences, such as animals. Asci of this type can be found both in
perithecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are mos ...
and in
cleistothecia An ascocarp, or ascoma (), is the fruiting body ( sporocarp) of an ascomycete phylum fungus. It consists of very tightly interwoven hyphae and millions of embedded asci, each of which typically contains four to eight ascospores. Ascocarps are ...
, for instance with
Dutch elm disease Dutch elm disease (DED) is caused by a member of the sac fungi (Ascomycota) affecting elm trees, and is spread by elm bark beetles. Although believed to be originally native to Asia, the disease was accidentally introduced into Americas, America ...
(''Ophiostoma''). This is something of a catch-all term for cases which do not fit into the other three ascus types, and they probably belong to several independent groups which evolved separately from unitunicate asci.


Ascospores

An ascospore is a
spore In biology, a spore is a unit of sexual or asexual reproduction that may be adapted for dispersal and for survival, often for extended periods of time, in unfavourable conditions. Spores form part of the life cycles of many plants, algae, f ...
contained in an ascus, or that was produced inside an ascus. This kind of spore is specific to
fungi A fungus ( : fungi or funguses) is any member of the group of eukaryotic organisms that includes microorganisms such as yeasts and molds, as well as the more familiar mushrooms. These organisms are classified as a kingdom, separately from ...
classified as
ascomycete 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 defi ...
s (
Ascomycota 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 def ...
). The ascospores of ''
Blumeria graminis ''Blumeria graminis'' (commonly called barley powdery mildew or corn mildew) is a fungus that causes powdery mildew on grasses, including cereals. It is the only species in the genus ''Blumeria''. It has also been called ''Erysiphe graminis'' and ...
'' are formed and released under the humid conditions. After landing onto a suitable surface, unlike conidia, ascospores of ''Blumeria graminis'' showed a more variable developmental patterns. The fungi ''
Saccharomyces ''Saccharomyces'' is a genus of fungi that includes many species of yeasts. ''Saccharomyces'' is from Greek σάκχαρον (sugar) and μύκης (fungus) and means ''sugar fungus''. Many members of this genus are considered very important in f ...
'' produces ascospores when grown on V-8 medium, acetate ascospore agar, or Gorodkowa medium. These ascospores are globose and located in asci. Each ascus contains one to four ascospores. The asci do not rupture at maturity. Ascospores are stained with Kinyoun stain and ascospore stain. When stained with Gram stain, ascospores are gram-negative while vegetative cells are gram-positive. The fission yeast ''
Schizosaccharomyces pombe ''Schizosaccharomyces pombe'', also called "fission yeast", is a species of yeast used in traditional brewing and as a model organism in molecular and cell biology. It is a unicellular eukaryote, whose cells are rod-shaped. Cells typically meas ...
'' is a single-celled haploid organism that reproduces asexually by mitosis and fission. However, exposure to the DNA damaging agent hydrogen peroxide induces pair-wise mating of haploid cells of opposite mating type to form transient diploid cells that then undergo meiosis to form asci, each with four ascospores. The production of viable ascospores depends on successful recombinational repair during meiosis. When this repair is defective a quality control mechanism prevents germination of damaged ascospores. These findings suggest that mating followed by meiosis is an adaptation for repairing DNA damage in the parental haploid cells in order to allow production of viable progeny ascospores.


References


External links


IMA Mycological Glossary: Ascus


{{Fungus structure Mycology Fungal morphology and anatomy