Ascus

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Asci of Morchella elata, Phase contrast image Morelasci.jpg
Asci of Morchella elata , Phase contrast image
There are eight ascospores in each ascus of Sordaria fimicola. Sordaria fimicola asci.jpg
There are eight ascospores in each ascus of Sordaria fimicola .

An ascus (from Ancient Greek ἀσκός (askós) 'skin bag, wineskin'; pl.: asci) [1] is the sexual spore-bearing cell produced in ascomycete fungi. Each ascus usually contains eight ascospores (or octad), 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 (e.g. Monosporascus cannonballus ), two, four, or multiples of four. In a few cases, the ascospores can bud off conidia that may fill the asci (e.g. Tympanis ) 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 (lacking a septum), 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 not used in ascospore formation) that is surrounded by the "bourrelet" (the thickened tissue near the top of the ascus). [2]

Contents

Typically, a single ascus will contain eight ascospores (or octad). The eight spores are produced by meiosis followed by a mitotic division. Two meiotic divisions turn the original diploid zygote nucleus into four haploid ones. That is, the single original diploid cell from which the whole process begins contains two complete sets of chromosomes. 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 , in a fruiting body which is visible to the naked eye, here called an ascocarp or ascoma. In other cases, such as single-celled yeasts, 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 .

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. A faint hissing sound can also be heard for species of Peziza and other cup fungi.

Asci, notably those of Neurospora crassa , have been used in laboratories for studying the process of meiosis, because the four cells produced by meiosis line up in regular order. By modifying genes 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 develop after the formation of croziers at their base. The croziers help maintain a brief dikaryon. The compatible nuclei of the dikaryon merge forming a diploid nucleus that then undergoes meiosis and ultimately internal ascospore formation. Members of the Taphrinomycotina and Saccharomycotina do not form croziers.

Classification

Asci of Hypomyces chrysospermus (they are unitunicate-inoperculate). DIC image. Hypomyces chrysospermus.jpg
Asci of Hypomyces chrysospermus (they are unitunicate-inoperculate). DIC image.

The form of the ascus, the capsule which contains the sexual spores, is important for classification of the Ascomycota. There are four basic types of ascus.

Ascus of Saccharomyces cerevisiae containing a tetrad of four spores Ascus.gif
Ascus of Saccharomyces cerevisiae containing a tetrad of four spores

Ascospores

Sordaria fimicola ascus plus ascospore Sordaria fimicola ascus plus ascospore.jpg
Sordaria fimicola ascus plus ascospore

An ascospore is a spore contained in an ascus, or that was produced inside an ascus. This kind of spore is specific to fungi classified as ascomycetes (Ascomycota).

The ascospores of Blumeria graminis are formed and released under the humid conditions. [3] After landing onto a suitable surface, unlike conidia, ascospores of Blumeria graminis showed a more variable developmental patterns. [3]

The fungi Saccharomyces 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 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. [4] The production of viable ascospores depends on successful recombinational repair during meiosis. [5] 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.

Related Research Articles

<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">Spore</span> Unit of reproduction adapted for dispersal and survival in unfavorable conditions

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, fungi and protozoa. They were thought to have appeared as early as the mid-late Ordovician period as an adaptation of early land plants.

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 most commonly bowl-shaped (apothecia) but may take on a spherical or flask-like form that has a pore opening to release spores (perithecia) or no opening (cleistothecia).

<span class="mw-page-title-main">Biological life cycle</span> Series of stages of an organism

In biology, a biological life cycle is a series of stages of the life of an organism, that begins as a zygote, often in an egg, and concludes as an adult that reproduces, producing an offspring in the form of a new zygote which then itself goes through the same series of stages, the process repeating in a cyclic fashion.

<i>Neurospora crassa</i> Species of ascomycete fungus in the family Sordariaceae

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 of French bakeries in 1843.

<i>Sordaria fimicola</i> Species of fungus

Sordaria fimicola is a species of microscopic fungus. It is commonly found in the feces of herbivores. Sordaria fimicola is often used in introductory biology and mycology labs because it is easy to grow on nutrient agar in dish cultures. The genus Sordaria, closely related to Neurospora and Podospora, is a member of the large class Sordariomycetes, or flask-fungi. The natural habitat of the three species of Sordaria that have been the principal subjects in genetic studies is dung of herbivorous animals. The species S. fimicola is common and worldwide in distribution. The species of Sordaria are similar morphologically, producing black perithecia containing asci with eight dark ascospores in a linear arrangement. These species share a number of characteristics that are advantageous for genetic studies. They all have a short life cycle, usually 7–12 days, and are easily grown in culture. Most species are self-fertile and each strain is isogenic. All kinds of mutants are easily induced and readily obtainable with particular ascospore color mutants. These visual mutants aid in tetrad analysis, especially in analysis of intragenic recombination.

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.

<span class="mw-page-title-main">Heterokaryon</span>

A heterokaryon is a multinucleate cell that contains genetically different nuclei. Heterokaryotic and heterokaryosis are derived terms. This is a special type of syncytium. This can occur naturally, such as in the mycelium of fungi during sexual reproduction, or artificially as formed by the experimental fusion of two genetically different cells, as e.g., in hybridoma technology.

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

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.

<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.

<span class="mw-page-title-main">Erysiphales</span> Order of fungi

Erysiphales are an order of ascomycete fungi. The order contains one family, Erysiphaceae. Many of them cause plant diseases called powdery mildew.

This is a glossary of some of the terms used in phytopathology.

Sporogenesis is the production of spores in biology. The term is also used to refer to the process of reproduction via spores. Reproductive spores were found to be formed in eukaryotic organisms, such as plants, algae and fungi, during their normal reproductive life cycle. Dormant spores are formed, for example by certain fungi and algae, primarily in response to unfavorable growing conditions. Most eukaryotic spores are haploid and form through cell division, though some types are diploid sor dikaryons and form through cell fusion.we can also say this type of reproduction as single pollination

<span class="mw-page-title-main">Crozier (mycology)</span> Anatomic feature in ascomycete fungi

A crozier is an anatomical feature of many fungi in the phylum Ascomycota that forms at the base of asci and looks like a hook-topped shepherd’s staff or stylized religious crosier. Croziers resemble and function similarly to clamp connections on the dikaryotic hyphae of Basidiomycota.

<i>Xylaria polymorpha</i> Species of fungus

Xylaria polymorpha, commonly known as dead man's fingers, is a cosmopolitan saprobic fungus. It is characterized by its elongated upright, clavate, or strap-like stromata poking up through the ground, much like fingers.

<span class="mw-page-title-main">Tetrad (meiosis)</span> Product of meiosis in spore-producing organisms

The tetrad is the four spores produced after meiosis of a yeast or other Ascomycota, Chlamydomonas or other alga, or a plant. After parent haploids mate, they produce diploids. Under appropriate environmental conditions, diploids sporulate and undergo meiosis. The meiotic products, spores, remain packaged in the parental cell body to produce the tetrad.

<span class="mw-page-title-main">Brachymeiosis</span>

Brachymeiosis was a hypothesized irregularity in the sexual reproduction of ascomycete fungi, a variant of meiosis following an "extra" karyogamy step. The hypothesized process would have transformed four diploid nuclei into eight haploid ones. The current scientific consensus is that brachymeiosis does not occur in any fungi.

Ascodesmis nigricans is a coprophilous fungus that could be isolated from the dung of various animals. It was firstly introduced by Philippe Édouard Léon Van Tieghem, a French botanist, and was the type species of the genus Ascodesmis. It is an uncommon species but its development of the fruit body has been the subject of much laboratory study due to the easy nature of its cultivation. Ascodesmis nigricans is not pathogenic to human, animals or plants. This species has world-wide distribution.

<span class="mw-page-title-main">Glossary of mycology</span>

This glossary of mycology is a list of definitions of terms and concepts relevant to mycology, the study of fungi. Terms in common with other fields, if repeated here, generally focus on their mycology-specific meaning. Related terms can be found in glossary of biology and glossary of botany, among others. List of Latin and Greek words commonly used in systematic names and Botanical Latin may also be relevant, although some prefixes and suffixes very common in mycology are repeated here for clarity.

References

  1. Henry Liddell; Robert Scott. "ἀσκός". A Greek-English Lexicon . Retrieved 27 February 2018 via Perseus Project.
  2. Hawksworth DL. (2013). Ascomycete Systematics: Problems and Perspectives in the Nineties. Springer. p. 116. ISBN   978-1-4757-9290-4.
  3. 1 2 Zhu, Mo; Riederer, Markus; Hildebrandt, Ulrich (2017). "Very-long-chain aldehydes induce appressorium formation in ascospores of the wheat powdery mildew fungus Blumeria graminis". Fungal Biology. 121 (8): 716–728. doi:10.1016/j.funbio.2017.05.003. PMID   28705398.
  4. Bernstein C, Johns V (April 1989). "Sexual reproduction as a response to H2O2 damage in Schizosaccharomyces pombe". J. Bacteriol. 171 (4): 1893–7. doi:10.1128/jb.171.4.1893-1897.1989. PMC   209837 . PMID   2703462.
  5. Guo H, King MC (2013). "A quality control mechanism linking meiotic success to release of ascospores". PLOS ONE. 8 (12): e82758. Bibcode:2013PLoSO...882758G. doi: 10.1371/journal.pone.0082758 . PMC   3846778 . PMID   24312672.