Neocallimastigomycetes
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Neocallimastigomycota is a phylum containing
anaerobic Anaerobic means "living, active, occurring, or existing in the absence of free oxygen", as opposed to aerobic which means "living, active, or occurring only in the presence of oxygen." Anaerobic may also refer to: * Anaerobic adhesive, a bonding a ...
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 ...
, which are symbionts found in the digestive tracts of larger herbivores. Anaerobic fungi were originally placed within phylum
Chytridiomycota Chytridiomycota are a division of zoosporic organisms in the kingdom Fungi, informally known as chytrids. The name is derived from the Ancient Greek ('), meaning "little pot", describing the structure containing unreleased zoöspores. Chytri ...
, within Order Neocallimastigales but later raised to phylum level, a decision upheld by later phylogenetic reconstructions. It encompasses only one
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
.


Discovery

The fungi in Neocallimastigomycota were first recognised as fungi by Orpin in 1975, based on motile cells present in the
rumen The rumen, also known as a paunch, is the largest stomach compartment in ruminants and the larger part of the reticulorumen, which is the first chamber in the alimentary canal of ruminant animals. The rumen's microbial favoring environment al ...
of
sheep Sheep or domestic sheep (''Ovis aries'') are domesticated, ruminant mammals typically kept as livestock. Although the term ''sheep'' can apply to other species in the genus '' Ovis'', in everyday usage it almost always refers to domesticated ...
. Their zoospores had been observed much earlier but were believed to be flagellate
protist A protist () is any eukaryotic organism (that is, an organism whose cells contain a cell nucleus) that is not an animal, plant, or fungus. While it is likely that protists share a common ancestor (the last eukaryotic common ancestor), the exc ...
s, but Orpin demonstrated that they possessed a chitin cell wall. It has since been shown that they are fungi related to the core chytrids. Prior to this, the microbial population of the rumen was believed to consist only of
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one Cell (biology), biological cell. They constitute a large domain (biology), domain of prokaryotic microorganisms. Typically a few micrometr ...
and protozoa. Since their discovery they have been isolated from the digestive tracts of over 50 herbivores, including
ruminant Ruminants (suborder Ruminantia) are hoofed herbivorous grazing or browsing mammals that are able to acquire nutrients from plant-based food by fermenting it in a specialized stomach prior to digestion, principally through microbial actions. The ...
and non-ruminant (hindgut-fermenting) mammals and herbivorous reptiles. Neocallimastigomycota have also been found in humans.


Circumscription


Reproduction and growth

These fungi reproduce in the rumen of ruminants through the formation of
zoospore A zoospore is a motile asexual spore that uses a flagellum for locomotion. Also called a swarm spore, these spores are created by some protists, bacteria, and fungi to propagate themselves. Diversity Flagella types Zoospores may possess one or ...
s which are released from sporangia. These zoospores bear a kinetosome but lack the nonflagellated centriole known in most chytrids, and have been known to utilize
horizontal gene transfer Horizontal gene transfer (HGT) or lateral gene transfer (LGT) is the movement of genetic material between unicellular and/or multicellular organisms other than by the ("vertical") transmission of DNA from parent to offspring (reproduction). H ...
in their development of
xylanase Endo-1,4-β-xylanase (EC 3.2.1.8, systematic name 4-β-D-xylan xylanohydrolase) is any of a class of enzymes that degrade the linear polysaccharide xylan into xylose, thus breaking down hemicellulose, one of the major components of plant cell ...
(from bacteria) and other
glucanase Glucanases are enzymes that break down large polysaccharides via hydrolysis. The product of the hydrolysis reaction is called a glucan, a linear polysaccharide made of up to 1200 glucose monomers, held together with glycosidic bonds. Glucans are ab ...
s.C.J. Alexopolous, Charles W. Mims, M. Blackwell, ''Introductory Mycology, 4th ed.'' (John Wiley and Sons, Hoboken NJ, 2004) The
nuclear Nuclear may refer to: Physics Relating to the nucleus of the atom: * Nuclear engineering *Nuclear physics *Nuclear power *Nuclear reactor *Nuclear weapon *Nuclear medicine *Radiation therapy *Nuclear warfare Mathematics *Nuclear space *Nuclear ...
envelopes of their cells are notable for remaining intact throughout mitosis. Sexual reproduction has not been observed in anaerobic fungi. However, they are known to be able to survive for many months in aerobic environments, a factor which is important in the colonisation of new hosts. In ''Anaeromyces'', the presence of putative resting spores has been observed but the way in which these are formed and germinate remains unknown.


Metabolism

Neocallimastigomycota lack mitochondria but instead contain
hydrogenosome A hydrogenosome is a membrane-enclosed organelle found in some anaerobic ciliates, flagellates, and fungi. Hydrogenosomes are highly variable organelles that have presumably evolved from protomitochondria to produce molecular hydrogen and ATP i ...
s in which the oxidation of
NADH Nicotinamide adenine dinucleotide (NAD) is a coenzyme central to metabolism. Found in all living cells, NAD is called a dinucleotide because it consists of two nucleotides joined through their phosphate groups. One nucleotide contains an aden ...
to NAD+, leading to formation of H2.


Polysaccharide-degrading activity

Neocallimastigomycota play an essential role in fibre-digestion in their host species. They are present in large numbers in the digestive tracts of animals which are fed on high fibre diets. The polysaccharide degrading enzymes produced by anaerobic fungi can hydrolyse the most recalcitrant plant polymers and can degrade unlignified plant cell walls entirely. Orpinomyces sp. exhibited the capacity of xylanase, CMCase, lichenase, amylase, β-xylosidase, β-glucosidase, α-Larabinofuranosidase and minor amounts of β-cellobiosidase production by utilizing avicel as the sole energy source. The polysaccharide degrading enzymes are organised into a multiprotein complex, similar to the bacterial
cellulosome Cellulosomes are multi-enzyme extracellular complexes. Cellulosomes are associated with the cell surface and mediate cell attachment to insoluble substrates and degrade them to soluble products which are then absorbed. Cellulosome complexes are in ...
.


Spelling of name

The Greek termination, "-mastix", referring to "whips", i.e. the many flagella on these fungi, is changed to "-mastig-" when combined with additional terminations in Latinized names. The family name Neocallimastigaceae was originally incorrectly published as "Neocallimasticaceae" by the publishing authors which led to the coinage of the misspelled, hence incorrect "Neocallimasticales", an easily forgiven error considering that other "-ix" endings such as ''
Salix Willows, also called sallows and osiers, from the genus ''Salix'', comprise around 400 speciesMabberley, D.J. 1997. The Plant Book, Cambridge University Press #2: Cambridge. of typically deciduous trees and shrubs, found primarily on moist s ...
'' goes to
Salicaceae The Salicaceae is the willow family of flowering plants. The traditional family (Salicaceae ''sensu stricto'') included the willows, poplar, aspen, and cottonwoods. Genetic studies summarized by the Angiosperm Phylogeny Group (APG) have greatly ...
. Correction of these names is mandated by the
International Code of Botanical Nomenclature The ''International Code of Nomenclature for algae, fungi, and plants'' (ICN) is the set of rules and recommendations dealing with the formal botanical names that are given to plants, fungi and a few other groups of organisms, all those "trad ...
, Art. 60. The corrected spelling is used by Index Fungorum.Suprafamilial Names
/ref> Both spellings occur in the literature and on the WWW as a result of the spelling in the original publication.


References


External links

*
The Anaerobic Fungi Network
{{Taxonbar, from=Q941929 Fungus phyla Fungi by classification