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Before we go on to explore the many orders of ascomycetes, we must take a closer look at the ascus itself. All asci are not the same. There are four flavours. (1) Unitunicate-operculate asci Unitunicate asci have a single wall. Some have a built-in lid or operculum (stained blue - left) - at maturity this pops open around a built-in line of weakness (below, left) so that the spores can be ejected. Unitunicate- operculate asci are found only in apothecial ascomata. .

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The photomicrograph on the khan right shows three ascomata of Ascobolus : the dark dots are unitunicate-operculate asci (see next section) containing dark spores, arranged in a flat superficial hymenium.or it may permit discharge of only one ascus, or a few asci, at a time,. The left hand scanning electron micrograph shows what many perithecial ascomata look like. . The diagram in the middle is of a vertical section through a perithecial ascoma and its neck. . The right hand photomicrograph is of a translucent ascoma inside which the asci, with their rows of dark ascospores, can be seen. And (3) pseudothecial ascomata below, which contain bitunicate asci (see the next section of the chapter).or they may lack an opening entirely in (4) cleistothecial ascomata (below). The asci in these are often spherical, as in the illustrations below, and no longer shoot their spores: the fungus has evolved a new dispersal strategy. That may have happened because the fungus fruits in a confined space (for example, under bark, or below the surface of the ground) where airborne dispersal cannot operate. We often find that the spores of such fungi are dispersed by animals. . Both diagram and photomicrograph show sections through a cleistothecial ascoma: the globose asci are clearly visible, and it is equally clear that they are not arranged in a layer or hymenium, as they were in the other three kinds of ascoma. Four kinds of Asci. .

It builds up to about 3 atmospheres before the apex blows. Mature asci of the dung-inhabiting Ascobolus (below) project above the hymenium and point toward the light before discharging their spores. In this way they working ensure that the spores will not run into any obstacles on their upward flight (see chapter 8). Four kinds of Ascoma. The multicellular structures (ascomata) that produce the asci, and act as the platforms from which the spores are launched, come in four main designs, sectional views of which are shown in the diagrams below. (1) apothecial ascomata below The construction of the ascoma may allow several or many asci to discharge simultaneously because the entire fertile layer or hymenium is exposed. The diagram on the left shows a vertical section through a very small apothecial ascoma.

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In this way the products of a single encounter are first multiplied, then meiosis generates a lot of genetic diversity. Not table only is the dikaryon itself an unusual phenomenon, but during the dikaryophase an effectively diploid mycelium is growing within, and drawing nourishment from, the haploid ascoma tissue. This phenomenon has interesting parallels in the red algae, though molecular evidence doesn't support a close relationship between the two groups (both lack motile gametes and appear to have simply hit upon the same solution to the problem posed by the rarity of sexual encounters). Ascospores are not motile, in the sense of self-propelling, but most ascomycetes nevertheless send their ascospores on their way with a burst of kinetic essay energy. The ascus is a tiny spore-gun, which works by building up internal pressure, then releasing it through the tip (see animation below). The job of most asci is to get their ascospores into the turbulent airflow above the ascoma. Before the ascus bursts open at the tip and shoots its spores skywards, the pressure inside the ascus is cranked up by osmolytes such as glycerol. .

by comparing the two sets of diagrams above, note how similar the developmental processes are until the final stages. Teleomorph life cycle now let's examine the sexual (teleomorphic) part of the ascomycete life cycle from the beginning (left). When an ascospore germinates, it establishes a haploid mycelium. In heterothallic ascomycetes, this can't undergo sexual reproduction until it meets another compatible haploid mycelium. When this rare event takes place, the fungus cleverly maximizes the ensuing potential for genetic recombination. One would expect a single sexual fusion, resulting in a single zygote. But most ascomycetes interpolate a dikaryophase, during which the number of pairs of compatible nuclei is multiplied, often enormously, as dikaryotic hyphae (often called ascogenous hyphae, as in the diagram above) grow and branch within a mass of monokaryotic (haploid) tissue which is the framework.   eventually, the ultimate branches of the dikaryotic hyphae, of which there may be millions in larger ascomata, reach their ordained positions in the future hymenium and the long-delayed sexual fusions take place. The genome is reshuffled during the ensuing meiosis in each ascus (this genetic recombination is due to crossing-over, which is explained in Chapter 10)Each meiosis will produce a somewhat different arrangement of the genome.

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Some dikaryan fruit bodies are blue microscopic (as in many ascomycetes but often (especially among the basidiomycetes they are large and complex, and most of the common names applied to fungi refer to the visible teleomorphs of basidiomycetes, and in a few cases, ascomycetes. You may already be acquainted with about some of these: I will introduce you to many more in the pages ahead. Phylum ascomycota - characteristics of Teleomorphs Most dikaryan fungi share a number of important features - (1) chitinous cell walls; (2) hyphae with regular cross-walls called septa (centrally perforated to allow movement of cytoplasm, and sometimes nuclei, between compartments (note that many yeasts are unicellular. After sexually compatible nuclei from different mycelia have been brought together by anastomosis, they pair off, but don't fuse immediately to form a diploid zygote. Instead, they go on dividing synchronously to populate what are called dikaryotic hyphae, in which each compartment has two sexually compatible haploid nuclei. Oh yes, they do fuse eventually, but not before some remarkable developments have taken place, and in basidiomycetes, perhaps not for years.

If ascomycetes and basidiomycetes share all these things, how do they differ? Actually, in many ways, and with experience it's usually easy to tell their sexual fructifications apart with the naked eye. But their microscopic, unicellular meiosporangia are most diagnostic of all (compare the two sets of diagrams below). Upper diagrams - the meiosporangia of ascomycetes are asci (singular, ascus). They are cylindrical or sac-like and at maturity usually contain eight haploid spores (ascospores) which are expelled into the air through the top of the ascus. Lower diagrams - the meiosporangia of basidiomycetes are basidia (singular, basidium they usually have four tiny projections called sterigmata, each bearing a haploid spore (basidiospore) which is shot away individually at maturity. The formation of asci or basidia marks the end of the dikaryophase: the paired nuclei have fused and the resulting zygote has undergone meiosis (and a mitosis in ascomycetes) to produce 8 haploid ascospores or 4 haploid basidiospores. .

  However,. Jim Aist, who has spent many hours watching nuclei in living hyphae, assures me that this does not happen very often. Some dikaryan anamorphs (especially coelomycetes ) grow in dead leaves and stems of desert plants, and other moulds are the most drought tolerant of all organisms, able to grow at water activities below.70 (for example, on jams, salt fish and other substrates of extremely. While many zygomycetes can assimilate only 'accessible' substrates like sugars and starch, ascomycetes can often exploit cellulose, and many basidiomycetes can digest both cellulose and lignin, carbon sources that are available to remarkably few other organisms. Though fungi cannot fix atmospheric nitrogen (this talent seems to be restricted to the bacteria dikaryan fungi can use many different forms of combined nitrogen: some ascomycetes even specialize in metabolizing the protein keratin, which is the main component of hair and skin. In case you were wondering if members of this group constitute a health hazard - they.


Some other orders of ascomycetes are obligate parasites of plants. Remember the 'downy mildews' caused by oomycetes? Well, there are also plant diseases called 'powdery mildews' that are caused by ascomycetes. . The similarity of terminology is unfortunate, but try to remember the difference, because although the groups of fungi involved are both obligately biotrophic, the diseases are different in many important ways, such as host ranges and methods of control. This is just one example of how taxonomy has practical implications (see chapter 12 ). Thousands of basidiomycetes, and a quite a few ascomycetes, establish intimate mutualistic symbioses (mycorrhizas) with the roots of trees, especially conifers (see chapter 17 ). . nearly 18,000 ascomycetes, and a few basidiomycetes, have domesticated algae, thus becoming lichens, which can live in some of the world's harshest climates, and colonize the barest and most inhospitable substrates (see chapter 7 ). Some dikaryan fungi have even re-entered the water and, lacking motile cells, have evolved other mechanisms, such as long appendages, to aid spore dispersal. Dikaryan fungi range from unspecialized, almost omnivorous saprobes, to fungi so specialized and ecologically demanding that they are found only on one particular leg of one species of insect.

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Hyphae average about 5 microns in width, but collectively they are very long (sometimes kilometres per gram of soil). Dikaryan hyphae are also septate - they have cross-walls called septa at regular intervals. These miniature bulkheads give the thesis hyphae some physical rigidity, and limit loss of cytoplasm if the hyphal wall is ruptured. . As a result, we find that dikaryan fungi can grow in a wider range of conditions: they can often spread and fruit in drier situations than zygomycetes could tolerate. This picture shows what you would see if you could crawl inside a hypha and look toward the septum at the end of a compartment. . The septal pore is obvious, and you can also see the fine fibres of which the cross-wall is made. Cytoplasm and nuclei can move through the septal pores of ascomycetes, as you can see in this transmission electron micrograph of a short segment. Neurospora crassa hypha - a nucleus is shown in the act of squeezing through the pore.

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Kingdom, eumycota, phylum 6, ascomycota - the, ascomycetes. Hotlinks to: meiosporangia - life cycle ascomata - asci - anamorphs - coelomycetes - hyphomycetes, introduction - basic features, zygomycetes are terrestrial fungi: there's no doubt about that. But they thrive and sporulate only in damp places where the atmosphere is more or less saturated with moisture. For example, rhizopus stolonifer will colonize the moist interior of a loaf of bread, but won't produce its characteristic sporangiophores burden and mitosporangia on the outside of the bread unless the surrounding atmosphere is humid. If we persuade the fungus to sporulate by keeping the loaf in a damp chamber (a plastic bag containing a few drops of water will do) and then take it out of the bag, the sporangiophores will quickly collapse. Hyphae of most zygomycetes are wide, thin-walled, and coenocytic - continuous tubes with no cross-walls. Hyphae of the, dikarya (Phylum. A scomycota plus Phylum, b asidiomycota ) are narrower - although this picture shows ascomycete hyphae alongside what looks like a pine tree, it is actually a human hair. .

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