Neocallimastigales
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Neocallimastigomycota is a
phylum In biology, a phylum (; plural: phyla) is a level of classification or taxonomic rank below kingdom and above class. Traditionally, in botany the term division has been used instead of phylum, although the International Code of Nomenclature ...
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 fr ...
, which are symbionts found in the digestive tracts of larger
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
s. 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 Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of ...
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 domesticate ...
. Their zoospores had been observed much earlier but were believed to be
flagellate A flagellate is a cell or organism with one or more whip-like appendages called flagella. The word ''flagellate'' also describes a particular construction (or level of organization) characteristic of many prokaryotes and eukaryotes and thei ...
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 e ...
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 A microorganism, or microbe,, ''mikros'', "small") and ''organism'' from the el, ὀργανισμός, ''organismós'', "organism"). It is usually written as a single word but is sometimes hyphenated (''micro-organism''), especially in olde ...
population of the rumen was believed to consist only of
bacteria Bacteria (; singular: bacterium) are ubiquitous, mostly free-living organisms often consisting of one biological cell. They constitute a large domain of prokaryotic microorganisms. Typically a few micrometres in length, bacteria were am ...
and
protozoa Protozoa (singular: protozoan or protozoon; alternative plural: protozoans) are a group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic tissues and debris. Histo ...
. Since their discovery they have been isolated from the digestive tracts of over 50
herbivore A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthpar ...
s, 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
reptile Reptiles, as most commonly defined are the animals in the class Reptilia ( ), a paraphyletic grouping comprising all sauropsids except birds. Living reptiles comprise turtles, crocodilians, squamates ( lizards and snakes) and rhynchocephalia ...
s. Neocallimastigomycota have also been found in humans.


Circumscription


Reproduction and growth

These fungi reproduce in the rumen of ruminants through the formation of zoospores which are released from sporangia. These zoospores bear a kinetosome but lack the nonflagellated
centriole In cell biology a centriole is a cylindrical organelle composed mainly of a protein called tubulin. Centrioles are found in most eukaryotic cells, but are not present in conifers (Pinophyta), flowering plants (angiosperms) and most fungi, and are ...
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 cel ...
(from bacteria) and other glucanases.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 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 w ...
are notable for remaining intact throughout
mitosis 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 ...
. 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 A mitochondrion (; ) is an organelle found in the cells of most Eukaryotes, such as animals, plants and fungi. Mitochondria have a double membrane structure and use aerobic respiration to generate adenosine triphosphate (ATP), which is used ...
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+ 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 ade ...
, 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.


Spelling of name

The Greek termination, "-mastix", referring to "whips", i.e. the many
flagella A flagellum (; ) is a hairlike appendage that protrudes from certain plant and animal sperm cells, and from a wide range of microorganisms to provide motility. Many protists with flagella are termed as flagellates. A microorganism may have fro ...
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