Henneguya zschokkei | |
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Henneguya zschokkei in salmon | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Cnidaria |
Class: | Myxosporea |
Order: | Bivalvulida |
Family: | Myxobolidae |
Genus: | Henneguya |
Species: | H. zschokkei |
Binomial name | |
Henneguya zschokkei (Gurley, 1894) | |
Synonyms [1] | |
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Henneguya zschokkei or Henneguya salminicola is a species of a myxosporean endoparasite. It afflicts several salmon in the genera Oncorhynchus and Salmo. [2] [3] It causes milky flesh or tapioca disease. [1] H. zschokkei is notable for its lack of mitochondria, mitochondrial DNA, aerobic respiration and its reliance on an exclusively anaerobic metabolism. [4]
Henneguya zschokkei is found in fish as an ovoid spore with two anterior polar capsules and two long caudal appendages. [5] Individuals are very small (about 10 micrometers in diameter), [6] but are found aggregated into cysts 3–6 mm in diameter at any place in the animal's musculature. [7]
H. zschokkei is a highly unusual among multicellular animals because it does not rely on the aerobic respiration of oxygen. The creature instead relies on an exclusively anaerobic metabolism. [8] [7] It lacks a mitochondrial genome and therefore mitochondria, making it one of the only known members of the eukaryotic animal kingdom to shun oxygen as the foundation of its metabolism.
H. zschokkei is ultimately a highly derived cnidarian and is distantly related to jellyfish, sea anemone and corals. However, this obligate internal parasite so little resembles other multicellular animals (let alone cnidarians) that it, along with many other species in class Myxosporea, were initially categorized as protozoa. It is nevertheless most closely related to jellyfish. This species, like most myxosporeans, lacks many of the diagnostic criteria that identify cnidarians. Indeed, it is without nervous, epithelial, gut or muscle cells of any kind. [9]
This parasite has not only lost its mitochondria and the mitochondrial DNA residing in them, but also the nuclear genes that code for mitochondrial reproduction. What genetic instructions for these functions that remain lie in useless pseudogenes. [4]
The origin and cause of H. zschokkei's highly reduced genome are not yet known. While eukaryotes are known for aerobic respiration, a few unicellular lineages native to hypoxic environments have also lost this capacity. In the absence of oxygen these single-celled organisms lose the portions of their genome that anticipate and govern aerobic respiration. These unusual eukaryotes have developed mitochondria-related organelles (MROs) that fulfill many of the functions of conventional mitochondria. However there is no evidence of such an adaptation in the multicellular H. zschokkei. [4]
One theory put forward to explain the highly unusual habit of H. zschokkei and its fellow myxosporeans invokes the cancers of cnidarians. On this explanation, animals such as H. zschokkei were originally cancerous growths in free-swimming jellyfish that escaped their parent organism, thereafter becoming a separate species that parasitized other animals. Such an origin is referred to as a SCANDAL, a loose acronym of the phrase speciated by cancer development in animals. [10]
Known hosts of Henneguya zschokkei include: [11]
Cnidaria is a phylum under kingdom Animalia containing over 11,000 species of aquatic animals found both in fresh water and marine environments, including jellyfish, hydroids, sea anemones, corals and some of the smallest marine parasites. Their distinguishing features are a decentralized nervous system distributed throughout a gelatinous body and the presence of cnidocytes or cnidoblasts, specialized cells with ejectable flagella used mainly for envenomation and capturing prey. Their bodies consist of mesoglea, a non-living, jelly-like substance, sandwiched between two layers of epithelium that are mostly one cell thick. Cnidarians are also some of the only animals that can reproduce both sexually and asexually.
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 throughout the cell as a source of chemical energy. They were discovered by Albert von Kölliker in 1857 in the voluntary muscles of insects. The term mitochondrion was coined by Carl Benda in 1898. The mitochondrion is popularly nicknamed the "powerhouse of the cell", a phrase coined by Philip Siekevitz in a 1957 article of the same name.
Salmon is the common name for several commercially important species of euryhaline ray-finned fish from the genera Salmo and Oncorhynchus of the family Salmonidae, native to tributaries of the North Atlantic (Salmo) and North Pacific (Oncorhynchus) basins. Other closely related fish in the same family include trout, char, grayling, whitefish, lenok and taimen, all coldwater fish of the subarctic and cooler temperate regions with some sporadic endorheic populations in Central Asia.
Mitochondrial DNA is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA is only a small portion of the DNA in a eukaryotic cell; most of the DNA can be found in the cell nucleus and, in plants and algae, also in plastids such as chloroplasts.
An anaerobic organism or anaerobe is any organism that does not require molecular oxygen for growth. It may react negatively or even die if free oxygen is present. In contrast, an aerobic organism (aerobe) is an organism that requires an oxygenated environment. Anaerobes may be unicellular or multicellular. Most fungi are obligate aerobes, requiring oxygen to survive. However, some species, such as the Chytridiomycota that reside in the rumen of cattle, are obligate anaerobes; for these species, anaerobic respiration is used because oxygen will disrupt their metabolism or kill them. Deep waters of the ocean are a common anoxic environment.
Cellular respiration is the process by which biological fuels are oxidized in the presence of an inorganic electron acceptor, such as oxygen, to drive the bulk production of adenosine triphosphate (ATP), which contains energy. Cellular respiration may be described as a set of metabolic reactions and processes that take place in the cells of organisms to convert chemical energy from nutrients into ATP, and then release waste products.
Excavata is an extensive and diverse but paraphyletic group of unicellular Eukaryota. The group was first suggested by Simpson and Patterson in 1999 and the name latinized and assigned a rank by Thomas Cavalier-Smith in 2002. It contains a variety of free-living and symbiotic protists, and includes some important parasites of humans such as Giardia and Trichomonas. Excavates were formerly considered to be included in the now obsolete Protista kingdom. They were distinguished from other lineages based on electron-microscopic information about how the cells are arranged. They are considered to be a basal flagellate lineage.
A multicellular organism is an organism that consists of more than one cell, in contrast to unicellular organism. All species of animals, land plants and most fungi are multicellular, as are many algae, whereas a few organisms are partially uni- and partially multicellular, like slime molds and social amoebae such as the genus Dictyostelium.
Myxozoa is a subphylum of aquatic cnidarian animals – all obligate parasites. 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 and an annelid worm or a bryozoan. The average size of a myxosporean spore usually ranges from 10 μm to 20 μm, whereas that of a malacosporean spore can be up to 2 mm. Myxozoans can live in both freshwater and marine habitats.
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 and an ectothermic vertebrate, usually a fish. Each host releases a different type of spore. The two forms of spore are so different that until relatively recently they were treated as belonging to different classes within the Myxozoa.
A facultative anaerobic organism is an organism that makes ATP by aerobic respiration if oxygen is present, but is capable of switching to fermentation if oxygen is absent.
A mitosome is a mitochondrion-related organelle (MRO) found in a variety of parasitic unicellular eukaryotes, such as members of the supergroup Excavata. The mitosome was first discovered in 1999 in Entamoeba histolytica, an intestinal parasite of humans, and mitosomes have also been identified in several species of Microsporidia and in Giardia intestinalis.
Protozoan infections are parasitic diseases caused by organisms formerly classified in the kingdom Protozoa. These organisms are now classified in the supergroups Excavata, Amoebozoa, Harosa, and Archaeplastida. They are usually contracted by either an insect vector or by contact with an infected substance or surface.
Holozoa is a clade of organisms that includes animals and their closest single-celled relatives, but excludes fungi and all other organisms. Together they amount to more than 1.5 million species of purely heterotrophic organisms, including around 300 unicellular species. It consists of various subgroups, namely Metazoa and the protists Choanoflagellata, Filasterea, Pluriformea and Ichthyosporea. Along with fungi and some other groups, Holozoa is part of the Opisthokonta, a supergroup of eukaryotes. Choanofila was previously used as the name for a group similar in composition to Holozoa, but its usage is discouraged now because it excludes animals and is therefore paraphyletic.
Proteromonas is a genus of single-celled biflagellated microbial eukaryotes belonging to the Superphylum Stramenopiles which are characterized by the presence of tripartite, hair-like structures on the anteriorly-directed larger of the two flagella. Proteromonas on the other hand are notable by having tripartite hairs called somatonemes not on the flagella but on the posterior of the cell. Proteromonas are closely related to Karotomorpha and Blastocystis, which belong to the Opalines group.
Like humans and other animals, fish suffer from diseases and parasites. Fish defences against disease are specific and non-specific. Non-specific defences include skin and scales, as well as the mucus layer secreted by the epidermis that traps microorganisms and inhibits their growth. If pathogens breach these defences, fish can develop inflammatory responses that increase the flow of blood to infected areas and deliver white blood cells that attempt to destroy the pathogens.
Diseases and parasites in salmon, trout and other salmon-like fishes of the family Salmonidae are also found in other fish species. The life cycle of many salmonids is anadromous, so such fish are exposed to parasites in fresh water, brackish water and saline water.
Aerobic fermentation or aerobic glycolysis is a metabolic process by which cells metabolize sugars via fermentation in the presence of oxygen and occurs through the repression of normal respiratory metabolism. Preference of aerobic fermentation over aerobic respiration is referred to as the Crabtree effect in yeast, and is part of the Warburg effect in tumor cells. While aerobic fermentation does not produce adenosine triphosphate (ATP) in high yield, it allows proliferating cells to convert nutrients such as glucose and glutamine more efficiently into biomass by avoiding unnecessary catabolic oxidation of such nutrients into carbon dioxide, preserving carbon-carbon bonds and promoting anabolism.
Monocercomonoides is a genus of flagellate Excavata belonging to the order Oxymonadida. It was established by Bernard V. Travis and was first described as those with "polymastiginid flagellates having three anterior flagella and a trailing one originating at a single basal granule located in front of the anteriorly positioned nucleus, and a more or less well-defined axostyle". It is the first eukaryotic genus to be found to completely lack mitochondria, and all hallmark proteins responsible for mitochondrial function. The genus also lacks any other mitochondria related organelles (MROs) such as hydrogenosomes or mitosomes. Data suggests that the absence of mitochondria is not an ancestral feature, but rather due to secondary loss. Monocercomonoides sp. was found to obtain energy through an enzymatic action of nutrients absorbed from the environment. The genus has replaced the iron-sulfur cluster assembly pathway with a cytosolic sulfur mobilization system, likely acquired by horizontal gene transfer from a eubacterium of a common ancestor of oxymonads. These organisms are significant because they undermine assumptions that eukaryotes must have mitochondria to properly function. The genome of Monocercomonoides exilis has approximately 82 million base pairs, with 18 152 predicted protein-coding genes.
Henneguya is a genus of myxosporean parasites belonging to the family Myxobolidae.
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