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Myxozoa (
etymology Etymology ()The New Oxford Dictionary of English (1998) – p. 633 "Etymology /ˌɛtɪˈmɒlədʒi/ the study of the class in words and the way their meanings have changed throughout time". is the study of the history of the Phonological chan ...
:
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
: μύξα ''myxa'' "slime" or "mucus" + thematic vowel o + ζῷον ''zoon'' "animal") is a subphylum of aquatic
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
n
animal Animals are multicellular, eukaryotic organisms in the Kingdom (biology), biological kingdom Animalia. With few exceptions, animals Heterotroph, consume organic material, Cellular respiration#Aerobic respiration, breathe oxygen, are Motilit ...
s – all obligate
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s. It contains the smallest animals ever known to have lived. Over 2,180 species have been described and some estimates have suggested at least 30,000 undiscovered species. Many have a two-host lifecycle, involving a
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
and an
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
worm or a
bryozoa Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a l ...
n. The average size of a
myxosporea Myxosporea is a class of microscopic parasites, belonging to the Myxozoa clade within Cnidaria. They have a complex life cycle which comprises vegetative forms in two hosts, an aquatic invertebrate (generally an annelid but sometimes a bryozoan) ...
n
spore 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, f ...
usually ranges from 10 μm to 20 μm, whereas that of a malacosporean (a subclade of the Myxozoa) spore can be up to 2 mm. Myxozoans can live in both freshwater and marine habitats. Myxozoans are highly derived
cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
ns that have undergone dramatic evolution from a free swimming, self-sufficient jellyfish-like creature into their current form of
obligate parasite An obligate parasite or holoparasite is a parasitic organism that cannot complete its life-cycle without exploiting a suitable host. If an obligate parasite cannot obtain a host it will fail to reproduce. This is opposed to a facultative parasite, ...
s composed of very few cells – sometimes only a single cell. As myxozoans evolved into microscopic parasites, they lost many genes responsible for multicellular development, coordination, cell-cell communication, and even, in some cases, aerobic respiration. The genomes of some myxozoans are now among the smallest genomes of any known animal species.


Life cycle and pathology

Myxozoans are endoparasitic animals exhibiting complex life cycles that, in most of the documented cases, involve an
intermediate host In biology and medicine, a host is a larger organism that harbours a smaller organism; whether a parasitic, a mutualistic, or a commensalist ''guest'' (symbiont). The guest is typically provided with nourishment and shelter. Examples include a ...
, usually a fish, but in rare cases amphibians, reptiles, birds, and mammals; and a definitive host, usually an
annelid The annelids (Annelida , from Latin ', "little ring"), also known as the segmented worms, are a large phylum, with over 22,000 extant species including ragworms, earthworms, and leeches. The species exist in and have adapted to various ecol ...
or an
ectoproct Bryozoa (also known as the Polyzoa, Ectoprocta or commonly as moss animals) are a phylum of simple, aquatic invertebrate animals, nearly all living in sedentary colonies. Typically about long, they have a special feeding structure called a ...
. Only about 100 life cycles have been resolved and it is suspected that there may be some exclusively terrestrial. The mechanism of infection occurs through valve spores that have many forms, but their main morphology is the same: one or two sporoplasts, which are the real infectious agent, surrounded by a layer of attenuated cells called valve cells, which can secrete a layer protective coating and form float appendages. Integrated into the layer of valve cells are two to four specialized capsulogenic cells (in a few cases, one or even 15), each carrying a polar capsule containing coiled polar filaments, an extrudable organelle used for recognition, contact and infiltration. Myxospores are ingested by annelids, in which the polar filaments extrude to anchor the spore to the gut epithelium. Opening of the shell valves allows the sporoplasms to penetrate into the epithelium. Subsequently, the parasite undergoes reproduction and development in the gut tissue, and finally produces usually eight actinosporean spore stages (actinospores) within a pansporocyst. After mature actinospores are released from their hosts they float in the water column. Upon contact with skin or gills of fish, sporoplasms penetrate through the epithelium, followed by development of the myxosporean stage. Myxosporean trophozoites are characterized by cell-in-cell state, where the secondary (daughter) cells develop in the mother (primary) cells. The presporogonic stages multiply, migrate via nervous or circulatory systems, and develop into sporogonic stages. At the final site of infection, they produce mature spores within mono- or di-sporic pseudoplasmodia, or poly-sporic plasmodia. Relationships between myxosporeans and their hosts are often highly evolved and do not usually result in severe diseases of the natural host. Infection in
fish Fish are aquatic, craniate, gill-bearing animals that lack limbs with digits. Included in this definition are the living hagfish, lampreys, and cartilaginous and bony fish as well as various extinct related groups. Approximately 95% of li ...
hosts can be extremely long-lasting, potentially persisting for the lifetime of the host. However, an increasing number of myxosporeans have become commercially important pathogens of fish, largely as a result of
aquaculture Aquaculture (less commonly spelled aquiculture), also known as aquafarming, is the controlled cultivation ("farming") of aquatic organisms such as fish, crustaceans, mollusks, algae and other organisms of value such as aquatic plants (e.g. lot ...
bringing new species into contact with myxosporeans to which they had not been previously exposed, and to which they are highly susceptible. The economic impact of such
parasite Parasitism is a close relationship between species, where one organism, the parasite, lives on or inside another organism, the host, causing it some harm, and is adapted structurally to this way of life. The entomologist E. O. Wilson has ...
s can be severe, especially where
prevalence In epidemiology, prevalence is the proportion of a particular population found to be affected by a medical condition (typically a disease or a risk factor such as smoking or seatbelt use) at a specific time. It is derived by comparing the number o ...
rates are high; they may also have a severe impact on wild fish stocks. The economically most significant diseases worldwide caused by myxosporeas in cultured fishes are
proliferative kidney disease ''Tetracapsuloides bryosalmonae'' is a myxozoan parasite of salmonid fish. It is the only species currently recognized in the monotypic genus ''Tetracapsuloides''. It is the cause of proliferative kidney disease (PKD), one of the most serious par ...
(PKD) caused by the malacosporean ''T. bryosalmonae'', and
whirling disease ''Myxobolus cerebralis'' is a myxosporean parasite of salmonids (salmon and trout species) that causes whirling disease in farmed salmon and trout and also in wild fish populations. It was first described in rainbow trout in Germany in 1893, b ...
, caused by a
myxosporea Myxosporea is a class of microscopic parasites, belonging to the Myxozoa clade within Cnidaria. They have a complex life cycle which comprises vegetative forms in two hosts, an aquatic invertebrate (generally an annelid but sometimes a bryozoan) ...
n ''M. cerebralis''; both diseases affect
salmon Salmon () is the common name for several list of commercially important fish species, commercially important species of euryhaline ray-finned fish from the family (biology), family Salmonidae, which are native to tributary, tributaries of the ...
. Enteromyxosis is caused by ''E. leei'' in cultured marine sparids, while proliferative gill disease (or “hamburger disease”) is caused by ''H. ictaluri'' in
catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...
and ''S. renicola'' infections occur in
common carp The Eurasian carp or European carp (''Cyprinus carpio''), widely known as the common carp, is a widespread freshwater fish of eutrophic waters in lakes and large rivers in Europe and Asia.Fishbase''Cyprinus carpio'' Linnaeus, 1758/ref>Arkive The ...
.


Anatomy

Myxozoans are very small animals, typically 10–300 μm in length. Like other cnidarians they possess cnidocysts, which were referred to as "polar capsules" before the discovery that myxozoans are cnidarians. These cnidocysts fire tubules as in other cnidarians; some inject substances into the host. However, the tubules lack hooks or barbs, and in some species are more elastic than in other cnidarians. Myxozoans have secondarily lost
epithelial Epithelium or epithelial tissue is one of the four basic types of animal tissue, along with connective tissue, muscle tissue and nervous tissue. It is a thin, continuous, protective layer of compactly packed cells with a little intercellula ...
structures, a
nervous system In biology, the nervous system is the highly complex part of an animal that coordinates its actions and sensory information by transmitting signals to and from different parts of its body. The nervous system detects environmental changes th ...
, gut, and
cilia The cilium, plural cilia (), is a membrane-bound organelle found on most types of eukaryotic cell, and certain microorganisms known as ciliates. Cilia are absent in bacteria and archaea. The cilium has the shape of a slender threadlike projecti ...
. Most lack
muscle Skeletal muscles (commonly referred to as muscles) are organs of the vertebrate muscular system and typically are attached by tendons to bones of a skeleton. The muscle cells of skeletal muscles are much longer than in the other types of muscl ...
s, though these are retained in some members of malacosporea. Those who have lost their muscles move around inside the host using other forms of locomotion, such as the use of
filopodia Filopodia (singular filopodium) are slender cytoplasmic projections that extend beyond the leading edge of lamellipodia in migrating cells. Within the lamellipodium, actin ribs are known as ''microspikes'', and when they extend beyond the lame ...
, spore valve contractions, amoeboid movements, and rapidly creating and reabsorbing folds on the cell membrane. Myxozoans do not undergo embryogenesis during development and have lost true
gamete A gamete (; , ultimately ) is a haploid cell that fuses with another haploid cell during fertilization in organisms that reproduce sexually. Gametes are an organism's reproductive cells, also referred to as sex cells. In species that produce t ...
s. Instead, they reproduce via multicellular spores. These spores contain the polar capsules, which are not typically present in somatic cells.
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 a ...
s do not participate in the nuclear division of myxozoans. Cell division by
binary fission Binary may refer to: Science and technology Mathematics * Binary number, a representation of numbers using only two digits (0 and 1) * Binary function, a function that takes two arguments * Binary operation, a mathematical operation that t ...
is rare, and cells divide instead via
endogeny Endogenous substances and processes are those that originate from within a living system such as an organism, tissue, or cell. In contrast, exogenous substances and processes are those that originate from outside of an organism. For example, es ...
. In 2020, the myxozoan '' Henneguya salminicola'' was found to lack a mitochondrial genome, and thus be incapable of aerobic respiration; it was the first animal to be positively identified as such. Its actual metabolism is currently unknown.


Phylogenetics

Myxozoans were originally considered to be
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 ...
ns, and were included among other non-motile forms in the group
Sporozoa The Apicomplexa (also called Apicomplexia) are a large phylum of parasitic alveolates. Most of them possess a unique form of organelle that comprises a type of non-photosynthetic plastid called an apicoplast, and an apical complex structure. The ...
. As their distinct nature became clear through 18S
ribosomal Ribosomes ( ) are macromolecular machines, found within all cells, that perform biological protein synthesis (mRNA translation). Ribosomes link amino acids together in the order specified by the codons of messenger RNA (mRNA) molecules to fo ...
DNA (rDNA) sequencing, they were relocated in the
metazoa Animals are multicellular, eukaryotic organisms in the biological kingdom Animalia. With few exceptions, animals consume organic material, breathe oxygen, are able to move, can reproduce sexually, and go through an ontogenetic stage in ...
. Detailed classification within the metazoa was however long hindered by conflicting rDNA evidence: although 18S rDNA suggested an affinity with
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
, other rDNA sampled, and the HOX genes of two species, were more similar to those of the
Bilateria The Bilateria or bilaterians are animals with bilateral symmetry as an embryo, i.e. having a left and a right side that are mirror images of each other. This also means they have a head and a tail (anterior-posterior axis) as well as a belly and ...
. The discovery that ''
Buddenbrockia plumatellae ''Buddenbrockia plumatellae'' is a worm-like parasite of bryozoans whose taxonomic placement long puzzled biologists. It is now classified as one of only three myxozoans in the Malacosporea subclass and its only family, Saccosporidae, on the b ...
'', a worm-like parasite of bryozoans up to 2 mm in length, is a myxozoan initially appeared to strengthen the case for a bilaterian origin, as the body plan is superficially similar. Nevertheless, closer examination reveals that ''Buddenbrockia'' longitudinal symmetry is not twofold, but fourfold, casting doubt on this
hypothesis A hypothesis (plural hypotheses) is a proposed explanation for a phenomenon. For a hypothesis to be a scientific hypothesis, the scientific method requires that one can test it. Scientists generally base scientific hypotheses on previous obse ...
. Further testing resolved the genetic conundrum by sourcing the first three previously identified discrepant HOX genes (''Myx1-3'') to the bryozoan '' Cristatella mucedo'' and the fourth (''Myx4'') to
northern pike The northern pike (''Esox lucius'') is a species of carnivorous fish of the genus '' Esox'' (the pikes). They are typical of brackish and fresh waters of the Northern Hemisphere (''i.e.'' holarctic in distribution). They are known simply as a ...
, the respective hosts of the two corresponding Myxozoa samples. This explained the confusion: the original experiments had used samples contaminated by tissue from host organisms, leading to
false positive A false positive is an error in binary classification in which a test result incorrectly indicates the presence of a condition (such as a disease when the disease is not present), while a false negative is the opposite error, where the test result ...
s for a position among the Bilateria. More careful cloning of 50 coding genes from ''Buddenbrockia'' firmly established the clade as severely modified members of the phylum
Cnidaria Cnidaria () is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in freshwater and marine environments, predominantly the latter. Their distinguishing feature is cnidocytes, specialized cells that th ...
, with
medusozoa Medusozoa is a clade in the phylum Cnidaria, and is often considered a subphylum. It includes the classes Hydrozoa, Scyphozoa, Staurozoa and Cubozoa, and possibly the parasitic Polypodiozoa. Medusozoans are distinguished by having a medusa stage ...
ns as their closest relatives. Similarities between myxozoan polar capsules and cnidarian
nematocyst A cnidocyte (also known as a cnidoblast or nematocyte) is an explosive cell containing one large secretory organelle called a cnidocyst (also known as a cnida () or nematocyst) that can deliver a sting to other organisms. The presence of this ce ...
s had been drawn for a long time, but were generally assumed to be the result of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Taxonomists now recognize the outdated subgroup Actinosporea as a life-cycle phase of
Myxosporea Myxosporea is a class of microscopic parasites, belonging to the Myxozoa clade within Cnidaria. They have a complex life cycle which comprises vegetative forms in two hosts, an aquatic invertebrate (generally an annelid but sometimes a bryozoan) ...
. Molecular clocks suggest that myxozoans and their closest relatives, the polypodiozoa, shared their last common ancestor with medusazoans about 600 million years ago, during the
Ediacaran The Ediacaran Period ( ) is a geological period that spans 96 million years from the end of the Cryogenian Period 635 million years ago (Mya), to the beginning of the Cambrian Period 538.8 Mya. It marks the end of the Proterozoic Eon, and th ...
period.


Taxonomy

Myxozoan taxonomy has undergone great and important changes in its levels of generic, family and suborder classification. Fiala et al. (2015) proposed a new classification based on spores.


References


External links

* * {{Authority control Animal classes