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 in anaerobic conditions. [1]
Hydrogenosomes were discovered in 1973 by D. G. Lindmark and M. Müller. Because hydrogenosomes hold evolutionary lineage significance for organisms living in anaerobic or oxygen-stressed environments, many research institutions have since documented their findings on how the organelle differs in various sources. [2]
Hydrogenosomes were isolated, purified, biochemically characterized and named in the early 1970s by Lindmark and Müller at Rockefeller University. In addition to this seminal study on hydrogenosomes, they also demonstrated for the first time the presence of pyruvate:ferredoxin oxido-reductase and hydrogenase in eukaryotes. [2] Further studies were subsequently conducted on the biochemical cytology and subcellular organization of several anaerobic protozoan parasites (ex: Trichomonas vaginalis, Tritrichomonas foetus, Giardia lamblia, and Entamoeba sp.). [1]
Using information obtained from hydrogenosomal and biochemical cytology studies these researchers determined the mode of action of metronidazole (Flagyl) . Today, metronidazole is recognized as a standard chemotherapeutic agent for the treatment of anaerobic infections. [3] [4]
Since their discovery, hydrogenosomes have been found in a variety of anaerobic unicellular ciliates, flagellates, and fungi. The most notable amongst these is the parasitic Trichomonas vaginalis. [5]
Hydrogenosomes are organelles that are speculated to have evolved from mitochondria to provide a different mechanism for anaerobic ATP synthesis utilizing pyruvate. The reaction results in the production of molecular hydrogen, from which the organelle receives its name. [2]
Hydrogenosomes range from 0.5-2 micrometers and are bound by a double membrane. They are most often dumb-bell-shaped and found in large complexes of stacked hydrogenosomes. These stacks range from 4 or 5 (called juvenile complexes) to 20 or more hydrogenosomes. [1]
In most cases, hydrogenosomes are genomeless, as a majority of the mitochondrial genome was transferred to the nucleus; because of this, all hydrogenosomal proteins are imported to the organelle. [6] [7] However, a hydrogenosomal genome has been detected in the cockroach ciliate Nyctotherus ovalis, and the stramenopile Blastocystis . [8]
Due to the fact that many organisms have evolved to fit their anaerobic environments, a multitude of organisms have independently evolved hydrogenosomes or structures with similar functions. The similarity between Nyctotherus and Blastocystis, which are only distantly related, is believed to be the result of convergent evolution, and calls into question whether there is a clear-cut distinction between mitochondria, hydrogenosomes, and mitosomes (another kind of degenerate mitochondria). [1] [8]
A non-exhaustive list of organisms containing hydrogenosomes includes:
The vast variety of source organisms can be accredited to the theorized convergent evolution of hydrogenosomes from mitochondria to fit an anaerobic environment. [1] [6] [8]
In 2010, scientists have also reported their discovery of the first known anaerobic metazoans with hydrogenosome-like organelles. Three multicellular species of Loricifera — Spinoloricus nov. sp. , Rugiloricus nov. sp. and Pliciloricus nov. sp. — have been found deep in Mediterranean sediments, and use hydrogenosomes in their anaerobic metabolism cycle. [9]
The hydrogenosomes of trichomonads (the most studied of the hydrogenosome-containing microorganisms) produce molecular hydrogen, acetate, carbon dioxide and ATP by the combined actions of pyruvate:ferredoxin oxido-reductase, hydrogenase, acetate:succinate CoA transferase and succinate thiokinase. Superoxide dismutase, malate dehydrogenase (decarboxylating), ferredoxin, adenylate kinase and NADH:ferredoxin oxido-reductase are also localized in the hydrogenosome. [10]
Excavata is a former supergroup of unicellular organisms belonging to the domain Eukaryota. It was first suggested by Simpson and Patterson in 1999 and introduced by Thomas Cavalier-Smith in 2002 as a formal taxon. It contains a variety of free-living and symbiotic forms, 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 classified based on their flagellar structures, and were considered to be the most basal flagellate lineage.
Giardia duodenalis, also known as Giardia intestinalis and Giardia lamblia, is a flagellated parasitic microorganism of the genus Giardia that colonizes the small intestine, causing a diarrheal condition known as giardiasis. The parasite attaches to the epithelium by a ventral adhesive disc or sucker, and reproduces via binary fission. Giardiasis does not spread via the bloodstream, nor does it spread to other parts of the gastrointestinal tract, but remains confined to the lumen of the small intestine. Giardia has an outer membrane that makes it possible to retain life, even when outside of the host body, and which can make it tolerant to chlorine disinfection. Giardia trophozoites absorb their nutrients from the lumen, and are anaerobes. If the organism is split and stained, its characteristic pattern resembles the familiar "smiley face" symbol.
Gardnerella vaginalis is a species of Gram-variable-staining facultative anaerobic bacteria. The organisms are small non-spore-forming, nonmotile coccobacilli.
Trichomonas vaginalis is an anaerobic, flagellated protozoan parasite and the causative agent of a sexually transmitted disease called trichomoniasis. It is the most common pathogenic protozoan that infects humans in industrialized countries. Infection rates in men and women are similar but women are usually symptomatic, while infections in men are usually asymptomatic. Transmission usually occurs via direct, skin-to-skin contact with an infected individual, most often through vaginal intercourse. The WHO has estimated that 160 million cases of infection are acquired annually worldwide. The estimates for North America alone are between 5 and 8 million new infections each year, with an estimated rate of asymptomatic cases as high as 50%. Usually treatment consists of metronidazole and tinidazole.
The parabasalids are a group of flagellated protists within the supergroup Excavata. Most of these eukaryotic organisms form a symbiotic relationship in animals. These include a variety of forms found in the intestines of termites and cockroaches, many of which have symbiotic bacteria that help them digest cellulose in woody plants. Other species within this supergroup are known parasites, and include human pathogens.
Trichomonas is a genus of anaerobic excavate parasites of vertebrates. It was first discovered by Alfred François Donné in 1836 when he found these parasites in the pus of a patient suffering from vaginitis, an inflammation of the vagina. Donné named the genus from its morphological characteristics. The prefix tricho- originates from the Ancient Greek word θρίξ (thrix) meaning hair, describing Trichomonas’s flagella. The suffix -monas, describes its similarity to unicellular organisms from the genus Monas.
A mitosome is an organelle found in some unicellular eukaryotic organisms, like in members of the supergroup Excavata. The mitosome was found and named in 1999, and its function has not yet been well characterized. It was termed a crypton by one group, but that name is no longer in use.
In enzymology, a pyruvate synthase is an enzyme that catalyzes the interconversion of pyruvate and acetyl-CoA. It is also called pyruvate:ferredoxin oxidoreductase (PFOR).
Protozoan infections are parasitic diseases caused by organisms formerly classified in the kingdom Protozoa. They are usually contracted by either an insect vector or by contact with an infected substance or surface and include organisms that are now classified in the supergroups Excavata, Amoebozoa, SAR, and Archaeplastida.
Breviata anathema is a single-celled flagellate amoeboid eukaryote, previously studied under the name Mastigamoeba invertens. The cell lacks mitochondria, but has remnant mitochondrial genes, and possesses an organelle believed to be a modified anaerobic mitochondrion, similar to the mitosomes and hydrogenosomes found in other eukaryotes that live in low-oxygen environments.
Trimastix is a genus of excavates, the sole occupant of the order Trimastigida. Trimastix are bacterivorous, free living and anaerobic. When first observed in 1881 by William Kent, the morphology of Trimastix was not well described but over time the oral structure and flagellar organization have become clearer. There are few known species, and the genus's role in the ecosystem is largely unknown. However, it is known that they generally live in marine environments within the tissues of decaying organisms to maintain an anoxic environment. Much interest in this group is related to its close association with other members of Anaeromonadea. These organisms do not have classical mitochondria, and as such, much of the research involving these microbes is aimed at investigating the evolution of mitochondria.
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.
Mastigamoeba is a genus of pelobionts, and treated by some as members of the Archamoebae group of protists. Mastigamoeba are characterized as anaerobic, amitochondriate organisms that are polymorphic. Their dominant life cycle stage is as an amoeboid flagellate. Species are typically free living, though endobiotic species have been described.
Eukaryota, whose members are known as eukaryotes, is a diverse domain of organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms are eukaryotes. They constitute a major group of living things, along with the two groups of prokaryotes, the Bacteria and the Archaea.
Psalteriomonas lanterna is a species of amoebae in the group of Heterolobosea. The cells of the flagellate stage show four nuclei, four ventral grooves and four mastigont systems, each with four flagella. It lacks a Golgi apparatus and reproduction occurs in both stages of its life cycle.
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 overrule the notion 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.
Armophorea is a class of ciliates in the subphylum Intramacronucleata. . It was first resolved in 2004 and comprises three orders: Metopida, Clevelandellida, and Armophorida. Previously members of this class were thought to be heterotrichs because of similarities in morphology, most notably a characteristic dense arrangement of cilia surrounding their oral structures. However, the development of genetic tools and subsequent incorporation of DNA sequence information has led to major revisions in the evolutionary relationships of many protists, including ciliates. Metopids, clevelandellids, and armophorids were grouped into this class based on similarities in their small subunit rRNA sequences, making them one of two so-called "riboclasses" of ciliates, however, recent analyses suggest that Armophorida may not be related to the other two orders.
Metopus is a genus of anaerobic organisms from the family of Metopidae.
Stygiella /ˌstɪ.d͡ʒiˈɛ.lə/ is a genus of free-living marine flagellates belonging to the family Stygiellidae in the jakobids (excavata).
Stygiellidae is a family of free-living marine flagellates belonging to the order Jakobida, a deep-branching lineage within the eukaryotic supergroup Discoba. They are unicellular organisms that commonly inhabit anoxic, sulfide-rich and ammonium-rich marine habitats worldwide.