Proto-mitochondrion | |
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Scientific classification | |
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Class: | Alphaproteobacteria (or sister to) |
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The proto-mitochondrion is the hypothetical ancestral bacterial endosymbiont from which all mitochondria in eukaryotes are thought to descend, after an episode of symbiogenesis which created the aerobic eukaryotes.
The phylogenetic analyses of the few genes that are still encoded in the genomes of modern mitochondria suggest an alphaproteobacterial origin for this endosymbiont, in an ancient episode of symbiogenesis early in the history of the eukaryotes.
Although the order Rickettsiales has been proposed as the alphaproteobacterial sister-group of mitochondria [1] [2] [3] , no definitive evidence pinpoints the alphaproteobacterial group from which the proto-mitochondrion emerged, and some contradictory evidence, especially in the early, sparse genome samplings. Martijn et al found mitochondria are a possible sister-group to all other alphaproteobacteria. [4] The phylogenetic tree of the Rickettsidae has been inferred by Ferla et al. from the comparison of 16S + 23S ribosomal RNA sequences. [5] Geiger et alii (2023) propose placing the recently-discovered (2016) genus Iodidimonas , found in a sister clade to Rickettsidae, the Caulobacteridae [5] , as the closest free-living relative of mitochondria, as it possesses more metabolic products matching that of mitochondria today, such as cardiolipins and sphingolipids, and important genetic markers such as the COX operon and a counterpart to mitochondrial complex III, the bc1 complex. [6]
Alphaproteobacteria |
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Gabaldón & Huynen (2003) reconstructed the proteome (the entire set of proteins expressed by a genome) and corresponding metabolism of the proto-mitochondrion by comparing extant alpha-proteobacterial and eukaryotic genomes. They concluded that this organism was an aerobic alpha-proteobacterium respiring lipids, glycerol and other compounds provided by the host. At least 630 gene families derived from this organism can still be found in the 9 eukaryotic genomes analyzed in the study. [7]
A mitochondrion (pl. mitochondria) 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. Meaning a thread-like granule, the term mitochondrion was coined by Carl Benda in 1898. The mitochondrion is popularly nicknamed the "powerhouse of the cell", a phrase popularized by Philip Siekevitz in a 1957 Scientific American article of the same name.
Symbiogenesis is the leading evolutionary theory of the origin of eukaryotic cells from prokaryotic organisms. The theory holds that mitochondria, plastids such as chloroplasts, and possibly other organelles of eukaryotic cells are descended from formerly free-living prokaryotes taken one inside the other in endosymbiosis. Mitochondria appear to be phylogenetically related to Rickettsiales bacteria, while chloroplasts are thought to be related to cyanobacteria.
Mitochondrial DNA is the DNA located in the mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the DNA contained in a eukaryotic cell; most of the DNA is in the cell nucleus, and, in plants and algae, the DNA also is found in plastids, such as chloroplasts.
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.
The hydrogen hypothesis is a model proposed by William F. Martin and Miklós Müller in 1998 that describes a possible way in which the mitochondrion arose as an endosymbiont within a prokaryotic host in the archaea, giving rise to a symbiotic association of two cells from which the first eukaryotic cell could have arisen (symbiogenesis).
In biology, Archezoa is a term that has been introduced by several authors to refer to a group of organisms. Authors include Josef Anton Maximilian Perty, Ernst Haeckel and in the 20th century by Thomas Cavalier-Smith in his classification system. Each author used the name to refer to different arrays of organisms. This reuse by later authors of the same taxon name for different groups of organisms is widely criticized in taxonomy because the inclusion of the name in a sentence does not make sense unless the particular usage is specified. Nonetheless, all uses of 'Archezoa' are now obsolete.
The metamonads are a large group of flagellate amitochondriate microscopic eukaryotes. They include the retortamonads, diplomonads, parabasalids, oxymonads, and a range of more poorly studied taxa, most of which are free-living flagellates. All metamonads are anaerobic, and most members of the four groups listed above are symbiotes or parasites of animals, as is the case with Giardia lamblia which causes diarrhea in mammals.
The Rickettsiales, informally called rickettsias, are an order of small Alphaproteobacteria. They are obligate intracellular parasites, and some are notable pathogens, including Rickettsia, which causes a variety of diseases in humans, and Ehrlichia, which causes diseases in livestock. Another genus of well-known Rickettsiales is the Wolbachia, which infect about two-thirds of all arthropods and nearly all filarial nematodes. Genetic studies support the endosymbiotic theory according to which mitochondria and related organelles developed from members of this group.
The Rickettsiaceae are a family of bacteria. The genus Rickettsia is the most prominent genus within the family. The bacteria that eventually formed the mitochondrion is believed to have originated from this family. Most human pathogens in this family are in genus Rickettsia. They spend part of their lifecycle in the bodies of arthropods such as ticks or lice, and are then transmitted to humans or other mammals by the bite of the arthropod. It contains Gram-negative bacteria, very sensitive to environmental exposure, thus is adapted to obligate intracellular infection. Rickettsia rickettsii is considered the prototypical infectious organism in the group.
Genome size is the total amount of DNA contained within one copy of a single complete genome. It is typically measured in terms of mass in picograms or less frequently in daltons, or as the total number of nucleotide base pairs, usually in megabases. One picogram is equal to 978 megabases. In diploid organisms, genome size is often used interchangeably with the term C-value.
A nuclear gene is a gene that has its DNA nucleotide sequence physically situated within the cell nucleus of a eukaryotic organism. This term is employed to differentiate nuclear genes, which are located in the cell nucleus, from genes that are found in mitochondria or chloroplasts. The vast majority of genes in eukaryotes are nuclear.
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.
Alphaproteobacteria or α-proteobacteria, also called α-Purple bacteria in earlier literature, is a class of bacteria in the phylum Pseudomonadota. The Magnetococcales and Mariprofundales are considered basal or sister to the Alphaproteobacteria. The Alphaproteobacteria are highly diverse and possess few commonalities, but nevertheless share a common ancestor. Like all Proteobacteria, its members are gram-negative, although some of its intracellular parasitic members lack peptidoglycan and are consequently gram variable.
Paracoccus denitrificans, is a coccoid bacterium known for its nitrate reducing properties, its ability to replicate under conditions of hypergravity and for being a relative of the eukaryotic mitochondrion.
"Candidatus Midichloria" is a candidatus genus of Gram-negative, non-endospore-forming bacteria, with a bacillus shape around 0.45 μm in diameter and 1.2 μm in length. First described in 2004 with the temporary name IricES1, "Candidatus Midichloria" species are symbionts of several species of hard ticks. They live in the cells of the ovary of the females of this tick species. These bacteria have been observed in the mitochondria of the host cells, a trait that has never been described in any other symbiont of animals.
Jakobids are an order of free-living, heterotrophic, flagellar eukaryotes in the supergroup Excavata. They are small, and can be found in aerobic and anaerobic environments. The order Jakobida, believed to be monophyletic, consists of only twenty species at present, and was classified as a group in 1993. There is ongoing research into the mitochondrial genomes of jakobids, which are unusually large and bacteria-like, evidence that jakobids may be important to the evolutionary history of eukaryotes.
Jakoba is a genus in the taxon Excavata, and currently has a single described species, Jakoba libera described by Patterson in 1990, and named in honour of Dutch botanist Jakoba Ruinen.
The eukaryotes constitute the domain of Eukaryota or Eukarya, organisms whose cells have a membrane-bound nucleus. All animals, plants, fungi, and many unicellular organisms are eukaryotes. They constitute a major group of life forms alongside the two groups of prokaryotes: the Bacteria and the Archaea. Eukaryotes represent a small minority of the number of organisms, but given their generally much larger size, their collective global biomass is much larger than that of prokaryotes.
Eukaryogenesis, the process which created the eukaryotic cell and lineage, is a milestone in the evolution of life, since eukaryotes include all complex cells and almost all multicellular organisms. The process is widely agreed to have involved symbiogenesis, in which an archeon and a bacterium came together to create the first eukaryotic common ancestor (FECA). This cell had a new level of complexity and capability, with a nucleus, at least one centriole and cilium, facultatively aerobic mitochondria, sex, a dormant cyst with a cell wall of chitin and/or cellulose and peroxisomes. It evolved into a population of single-celled organisms that included the last eukaryotic common ancestor (LECA), gaining capabilities along the way, though the sequence of the steps involved has been disputed, and may not have started with symbiogenesis. In turn, the LECA gave rise to the eukaryotes' crown group, containing the ancestors of animals, fungi, plants, and a diverse range of single-celled organisms.
Andrew J. Roger is a Canadian-Australian molecular biologist and evolutionary bioinformatician. He is currently a professor in the Department of Biochemistry and Molecular Biology at Dalhousie University and was the founding director of the inter-departmental Centre for Comparative Genomics and Evolutionary Bioinformatics (CGEB).