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Chytridiomycota is a division of organisms in the kingdom , informally known as chytrids. The name is derived from the χυτρίδιον chytridion, meaning "little pot", describing the structure containing unreleased . Chytrids are one of the early diverging fungal lineages, and their membership in kingdom Fungi is demonstrated with chitin cell walls, a posterior whiplash flagellum, absorptive nutrition, use of as an energy storage compound, and synthesis of by the α-amino adipic acid (AAA) pathway.Kendrick, Bryce. 2000. The Fifth Kingdom. 3rd edition Focus Publishing: Newburyport, MA.

Chytrids are , degrading refractory materials such as and , and sometimes act as .Sparrow FK. 1960. Aquatic Phycomyetes. The University of Michigan Press:Ann Arbor. 2nd edition There has been a significant increase in the research of chytrids since the discovery of Batrachochytrium dendrobatidis, the causal agent of .Blackwell M. 2011. The Fungi: 1,2,3 … million species? American Journal of Botany 98:426-438.Longcore JE, AP Pessier & DK Nichols. 1999. Batrachochytirum dendrobatidis gen. et sp. nov., a chytrid pathogenic to amphibians. Mycologia 91(2):219-227.


Classification
Species of Chytridiomycota have traditionally been delineated and classified based on development, morphology, substrate, and method of zoospore discharge. However, single spore isolates (or isogenic lines) display a great amount of variation in many of these features; thus, these features cannot be used to reliably classify or identify a species.Blackwell WH, PM Letcher, & MJ Powell. 2006. Thallus development and the systematics of Chytridiomycota: an additional developmental pattern represented by Podochytrium. Mycotaxon 97: 91-109. Currently, taxonomy in Chytridiomycota is based on molecular data, zoospore and some aspects of thallus morphology and development.

In an older and more restricted sense (not used here), the term "chytrids" referred just to those fungi in the class . Here, the term “chytrid” refers to all members of Chytridiomycota.Alexopoulos CJ, Mims CW, Blackwell M. 1996. Introductory Mycology. 4th edition. John Wiley & Sons, Inc.

The chytrids have also been included among the , but are now regularly classed as fungi.

In older classifications, chytrids, except the recently established order Spizellomycetales, were placed in the class under the subphylum Myxomycophyta of the kingdom Fungi. Previously, they were placed in the as the class Chytridiomycetes.The Fungi: An Advanced Treatise Vol IVB A Taxonomic Review with Keys: Basidiomycetes and Lower Fungi. 1973. Edited by Ainsworth, Sparrow & Sussman. Academic Press: New York. The other classes of the Mastigomycotina, the Hyphochytriomycetes and , were removed from the fungi to be classified as .

The class has over 750 chytrid species distributed among ten orders.

(2018). 9783642553172, Springer. .
Additional classes include the Monoblepharidomycetes, with two orders, and the Hyaloraphidiomycetes with a single order.
(2012). 9781578087235, CRC Press. .

Molecular phylogenetics, and other techniques such as analysis, has greatly increased the understanding of chytrid phylogeny, and led to the formation of several new phyla:

  • The order , originally within the Chytridiomycota, are now classified as a separate phylum, the Blastocladiomycota.
  • The Neocallimastigales, originally an order of anaerobic of the class Chytridiomycetes, found in the digestive tracts of herbivores, was later raised to a separate phylum, the Neocallimastigomycota.
  • The , including the , formerly classified in the order , were raised to a separate phylum, the .


Life cycle and body plan

Chytridiomycota are unusual among the Fungi in that they reproduce with zoospores.Hibbett et al. 2007. A higher-level phylogenetic classification of the Fungi. Mycologia 111(5): 509-547. For most members of Chytridiomycetes, sexual reproduction is not known. Asexual reproduction occurs through the release of zoospores (presumably) derived through .

Where it has been described, sexual reproduction of Chytridomycetes occurs via a variety of methods. It is generally accepted that the resulting zygote forms a resting spore, which functions as a means of surviving adverse conditions. In some members, sexual reproduction is achieved through the fusion of (gametes of the same size and shape). This group includes the notable plant pathogens . Some algal parasites practice : a motile male gamete attaches itself to a nonmotile structure containing the female gamete. In another group, two produce tubes that fuse and allow the gametes to meet and fuse. In the last group, rhizoids of compatible strains meet and fuse. Both nuclei migrate out of the zoosporangium and into the conjoined rhizoids where they fuse. The resulting zygote germinates into a resting spore.

Sexual reproduction is common and well known among members of the Monblepharidomycetes. Typically, these chytrids practice a version of : the male is motile and the female is stationary. This is the first occurrence of oogamy in kingdom Fungi. Briefly, the monoblephs form oogonia, which give rise to eggs, and antheridia, which give rise to male gametes. Once fertilized, the zygote either becomes an encysted or motile oospore, which ultimately becomes a resting spore that will later germinate and give rise to new zoosporangia.

Upon release from the germinated resting spore, zoospores seek out a suitable substrate for growth using or . Some species encyst and germinate directly upon the substrate; others encyst and germinate a short distance away. Once germinated, enzymes released from the zoospore begin to break down the substrate and utilize it produce a new thallus. Thalli are and usually form no true (having instead).

Chytrids have several different growth patterns. Some are holocarpic, which means they only produce a and . Others are eucarpic, meaning they produce other structures, such as , in addition to the zoosporangium and zoospores. Some chytrids are monocentric, meaning a single zoospore gives rise to a single zoosporangium. Others are polycentric, meaning one zoospore gives rise to many zoosporangium connected by a rhizomycelium. Rhizoids do not have nuclei while a rhizomycelium can.

Growth continues until a new batch of zoospores are ready for release. Chytrids have a diverse set of release mechanisms that can be grouped into the broad categories of operculate or inoperculate. Operculate discharge involves the complete or incomplete detachment of a lid-like structure, called an operculum, allowing the zoospores out of the sporangium. Inoperculate chytrids release their zoospores through pores, slits, or papillae.


Habitats
Chytrids are , though those that thrive in the capillary network around soil particles are typically considered terrestrial.Barr DJS. 1990. Phylum Chytridiomycota. In: Handbook of Protoctista. Eds Margulis, Corliss, Melkonian, & Chapman. Jones & Barlett, Boston. Pgs. 454-466. The zoospore is primarily a means of thoroughly exploring a small volume of water for a suitable substrate rather than a means of long range dispersal.Carlile MJ. 1986. The zoospore and its problems. IN: Ayres, Peter G., and Lynne Boddy, eds. Water, Fungi, and Plants. Vol. 11. Cambridge University Press, 1986.

Chytrids have been isolated from a variety of aquatic habitats, including peats, bogs, rivers, ponds, springs, and ditches, and terrestrial habitats, such as acidic soils, alkaline soils, temperate forest soils, rainforest soils, arctic and Antarctic soils. This has led to the belief that many chytrid species are ubiquitous and cosmopolitan. However, recent taxonomic work has demonstrated that this ubiquitous and cosmopolitan morphospecies hide cryptic diversity at the genetic and ultrastructural levels.Letcher PM et al. 2008. Rhizophlyctidales--a new order in Chytridiomycota. Mycological Research 112: 1031-1048.Simmons DR. 2011. Phylogeny of Powellomycetacea fam. Nov. and description of Geranomyces variabilis gen. et comb. nov. Mycologia 103(6): 1411-1420. It was first thought aquatic chytrids (and other zoosporic fungi) were primarily active in fall, winter, and spring. However, recent molecular inventories of lakes during the summer indicate that chytrids are an active, diverse part of the eukaryotic microbial community.Lefèvre E, PM Letcher & MJ Powell. 2012. Temporal variation of the small eukaryotic community in two freshwater lakes: emphasis on zoosporic fungi. Aquatic Microbial Ecology 67: 91-105.

One of the least expected terrestrial environments the chytrid thrive in are periglacial soils. The population of the Chytridiomycota species are able to be supported even though there is a lack of plant life in these frozen regions due to the large amounts of water in periglacial soil and pollen blowing up from below the timberline.


Ecological functions

Batrachochytrium dendrobatidis

The chytrid Batrachochytrium dendrobatidis is responsible for , a disease of amphibians. Discovered in 1998 in and this disease is known to kill in large numbers, and has been suggested as a principal cause for the worldwide amphibian decline. Outbreaks of the fungus were found responsible for killing much of the Kihansi Spray Toad population in its native habitat of , Saving Tiny Toads Without a Home, by Cornelia Dean. The New York Times. February 1, 2010. as well as the extinction of the in 1989. Chytridiomycosis has also been implicated in the presumed extinction of the Southern Gastric Brooding Frog [http://www.environment.gov.au/cgi-bin/sprat/public/publicspecies.pl?taxon_id=1909 Rheobatrachus silus species profile and threats database, last seen in the wild in 1981, and the Northern Gastric Brooding Frog, last recorded in the wild in March 1985 (http://www.environment.gov.au/cgi-bin/sprat/public/publicspecies.pl?taxon_id=1910 Rheobatrachus eungellensis species profile and threats database). The process leading to frog mortality is thought to be the loss of essential ions through pores made in the epidermal cells by the chytrid during its replication.(Voyles, J., L. Berger, S. Young, et al. 2007. Electrolyte depletion and osmotic imbalance in amphibians with chytridiomycosis. Dis Aquat Organ. 77: 113-118.)

Recent research has revealed that elevating salt levels slightly may be able to cure chytridiomycosis in some Australian frog species (http://www.abc.net.au/news/2018-02-05/salt-curing-frogs-worldwide-from-chytrid-fungus/9390882 A salty cure for a deadly frog disease), although further experimentation is needed.


Other parasites
Chytrids mainly infect algae and other and microbes. The infection can be so severe as to control primary production within the lake.Ibelings BW, de Bruin A, Kagami M, Rijkeboer M, van Donk E. 2004. Host parasite interactions between freshwater phytolankton and chytrid fungi (chytridiomycota). J Phycol 40:457-455. It has been suggested that parasitic chytrids have a large effect on lake and pond food webs.Gleason, Frank H., et al. "The ecology of chytrids in aquatic ecosystems: roles in food web dynamics." Fungal Biology Reviews 22.1 (2008): 17-25. Chytrids may also infect plant species; in particular, Synchytrium endobioticum is an important pathogen.


Saprobes
Arguably, the most important ecological function chytrids perform is decomposition. These ubiquitous and cosmopolitan organisms are responsible for decomposition of refractory materials, such as , , , and . There are also chytrids that live and grow on pollen by attaching threadlike structures, called rhizoids, onto the pollen grains. This mostly occurs during asexual reproduction because the zoospores that become attached to the pollen continuously reproduce and form new chytrids that will attach to other pollen grains for nutrients. This colonization of pollen happens during the spring time when bodies of water accumulate pollen falling from trees and plants.


Fossil record
The earliest of chytrids are from the , a -age lagerstätte with anatomical preservation of and fungi. Among the microfossils are chytrids preserved as on . These fossils closely resemble the modern genus Allomyces. Holocarpic chytrid remains were found in cherts from Combres in central France that date back to the late . These remains were found along with eucarpic remains and are ambiguous in nature although they are thought to be of chytrids. Other chytrid-like fossils were found in cherts from the upper Pennsylvanian in the Basin in , dating between 300 and 350ma.


In fictional media
The novel (2007) features a species of chytrid that feeds on petroleum and oil-based products. In the story the species is modified using nuclear radiation, to increase the rate at which it feeds on oil. It is then used by Islamic extremists in an attempt to destroy the world's oil supplies, thereby taking away the technological advantage of the .
(2018). 9780425218242, Penguin Group.

The by Ethan Kocak features an arc with an evil, anthropomorphic chytrid developed as a Nazi experiment to infect Xolotl, a giant, talking black .


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