Mixotricha Paradoxa
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''Mixotricha paradoxa'' is a species of
protozoan Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
that lives inside the gut of the Australian
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
species '' Mastotermes darwiniensis''. It is composed of five different organisms: three
bacterial Bacteria (; : 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 among the ...
ectosymbionts live on its surface for locomotion and at least one
endosymbiont An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
lives inside to help digest
cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
in wood to produce
acetate An acetate is a salt formed by the combination of acetic acid with a base (e.g. alkaline, earthy, metallic, nonmetallic, or radical base). "Acetate" also describes the conjugate base or ion (specifically, the negatively charged ion called ...
for its host(s). ''Mixotricha'' mitochondria degenerated in hydrogenosomes and mitosomes and lost the ability to produce energy aerobically by
oxidative phosphorylation Oxidative phosphorylation(UK , US : or electron transport-linked phosphorylation or terminal oxidation, is the metabolic pathway in which Cell (biology), cells use enzymes to Redox, oxidize nutrients, thereby releasing chemical energy in order ...
. The mitochondria-derived nuclear genes were however conserved.


Discovery

The name was given by the Australian biologist J.L. Sutherland, who first described ''Mixotricha'' in 1933. The name means "the paradoxical being with mixed-up hairs" because this protist has both
cilia The cilium (: cilia; ; in Medieval Latin and in anatomy, ''cilium'') is a short hair-like membrane protrusion from many types of eukaryotic cell. (Cilia are absent in bacteria and archaea.) The cilium has the shape of a slender threadlike proj ...
and
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
, which was not thought to be possible for
protists A protist ( ) or protoctist is any Eukaryote, eukaryotic organism that is not an animal, Embryophyte, land plant, or fungus. Protists do not form a Clade, natural group, or clade, but are a Paraphyly, paraphyletic grouping of all descendants o ...
.


Behavior

''Mixotricha'' is a species of
protozoan Protozoa (: protozoan or protozoon; alternative plural: protozoans) are a polyphyletic group of single-celled eukaryotes, either free-living or parasitic, that feed on organic matter such as other microorganisms or organic debris. Historically ...
that lives inside the gut of the Australian
termite Termites are a group of detritivore, detritophagous Eusociality, eusocial cockroaches which consume a variety of Detritus, decaying plant material, generally in the form of wood, Plant litter, leaf litter, and Humus, soil humus. They are dist ...
species '' Mastotermes darwiniensis'' and has multiple
bacterial Bacteria (; : 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 among the ...
symbiont Symbiosis (Ancient Greek : living with, companionship < : together; and ''bíōsis'': living) is any type of a close and long-term biological interaction, between two organisms of different species. The two organisms, termed symbionts, can fo ...
s. ''Mixotricha'' is a large protozoan long and contains hundreds of thousands of bacteria. It is an
endosymbiont An endosymbiont or endobiont is an organism that lives within the body or cells of another organism. Typically the two organisms are in a mutualism (biology), mutualistic relationship. Examples are nitrogen-fixing bacteria (called rhizobia), whi ...
and digests
cellulose Cellulose is an organic compound with the chemical formula, formula , a polysaccharide consisting of a linear chain of several hundred to many thousands of glycosidic bond, β(1→4) linked glucose, D-glucose units. Cellulose is an important s ...
for the termite. Trichomonads like ''Mixotricha'' reproduce by a special form of longitudinal fission, leading to large numbers of
trophozoite A trophozoite (G. ''trope'', nourishment + ''zoon'', animal) is the activated, feeding stage in the life cycle of certain protozoa such as malaria-causing ''Plasmodium falciparum'' and those of the ''Giardia'' group. The complementary form of the t ...
s in a relatively short time.
Cysts A cyst is a closed Wikt:sac, sac, having a distinct Cell envelope, envelope and cell division, division compared with the nearby Biological tissue, tissue. Hence, it is a cluster of Cell (biology), cells that have grouped together to form a sac ...
never form, so transmission from one host to another is always based on direct contact between the sites they occupy.


Anatomy

Species of the order Trichomonadida typically have four to six
flagella A flagellum (; : flagella) (Latin for 'whip' or 'scourge') is a hair-like appendage that protrudes from certain plant and animal sperm cells, from fungal spores ( zoospores), and from a wide range of microorganisms to provide motility. Many pr ...
at the cell's apical pole, one of which is recurrent - that is, it runs along a surface wave, giving the aspect of an undulating membrane. ''Mixotricha paradoxa'' have four weak flagella that serve as rudders. It has four large flagella at the front end, three pointing forwards and one backward. The
basal bodies A basal body (synonymous with basal granule, kinetosome, and in older cytological literature with blepharoplast) is a protein structure found at the base of a eukaryotic undulipodium (cilium or flagellum). The basal body was named by Theodor ...
are also bacteria, not spirochaetes but oval, pill-shaped bacteria. There is a one-to-one relationship between a bracket, a spirochaete, and a basal bacterium. Each bracket has one spirochaete running through it and one pill bacterium at its base as the basal body. It has not been shown definitely, but the basal bodies could also be making cellulases that digest wood.


Endosymbionts for biochemical processes

At least one endosymbiont lives inside the protist to help digest cellulose and
lignin Lignin is a class of complex organic polymers that form key structural materials in the support tissues of most plants. Lignins are particularly important in the formation of cell walls, especially in wood and bark, because they lend rigidit ...
, a major component of the wood the termites eat. The cellulose gets converted to glucose then to acetate, and the lignin is digested directly to acetate. The acetate probably crosses the termite gut membrane to be digested later. ''Mixotricha'' forms a mutualistic relationship with bacteria living inside the termite. There are a total of four species of bacterial symbionts. It has spherical bacteria inside the cell, which function as
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 us ...
, which ''Mixotricha'' lacks. ''Mixotricha'' mitochondria degenerated and lost the ability to produce energy aerobically by
oxidative phosphorylation Oxidative phosphorylation(UK , US : or electron transport-linked phosphorylation or terminal oxidation, is the metabolic pathway in which Cell (biology), cells use enzymes to Redox, oxidize nutrients, thereby releasing chemical energy in order ...
. Mitochondrial relics include
hydrogenosome A hydrogenosome is a membrane-enclosed organelle found in some Anaerobic organism, anaerobic Ciliate, ciliates, Flagellate, flagellates, Fungus, fungi, and three species of Loricifera, loriciferans. Hydrogenosomes are highly variable organelles t ...
s which produce hydrogen and small structures called
mitosome A mitosome (also called a ''crypton'' in early literature) 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 ...
s.


Ectosymbionts for movement

Three surface colonising bacteria are anchored on the surface. The flagella and cilia are actually two different single celled organisms. The ciliate belongs to an archaic group that used to be called archezoa but this term is no longer in fashion. It has four weak flagella, which serve as a rudder. While ''Mixotricha'' has four anterior flagella, it does not use them for locomotion, but more for steering. For locomotion, about 250,000 hairlike '' Treponema''
spirochaete A spirochaete () or spirochete is a member of the phylum Spirochaetota (also called Spirochaetes ), which contains distinctive diderm (double-membrane) Gram-negative bacteria, most of which have long, helically coiled (corkscrew-shaped or ...
s, a species of helical bacteria, are attached to the cell surface and provide the cell with
cilia The cilium (: cilia; ; in Medieval Latin and in anatomy, ''cilium'') is a short hair-like membrane protrusion from many types of eukaryotic cell. (Cilia are absent in bacteria and archaea.) The cilium has the shape of a slender threadlike proj ...
-like movements. The wavelength of the cilia is about and suggests that the spirochaetes are somehow in touch with each other. ''Mixotricha'' also has rod-shaped bacteria arranged in an ordered pattern on the surface of the cell. Each spirochaete has its own little emplacement, called a 'bracket'. Spirochetes move continuously forwards or backwards but when they are attached they move in one direction.
Sperm Sperm (: sperm or sperms) is the male reproductive Cell (biology), cell, or gamete, in anisogamous forms of sexual reproduction (forms in which there is a larger, female reproductive cell and a smaller, male one). Animals produce motile sperm ...
tails might have their origin in spirochaetes. The evidence that cilia (undulipodia) are symbiotic bacteria is found unpersuasive.


Genome

''Mixotricha'' have five
genome A genome is all the genetic information of an organism. It consists of nucleotide sequences of DNA (or RNA in RNA viruses). The nuclear genome includes protein-coding genes and non-coding genes, other functional regions of the genome such as ...
s, as they form very close symbiotic relationships with four types of bacteria. It is a good example organism for
symbiogenesis Symbiogenesis (endosymbiotic theory, or serial endosymbiotic theory) is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibl ...
and nestedness. There are two spirochete and one-rod bacteria on its surface, one endosymbiotic bacteria inside to digest cellulose and the host nucleus.


References

{{Taxonbar, from=Q311708 Metamonads Symbiosis Endosymbiotic events Taxa described in 1933 Metamonad genera