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 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 use ...

arose as an

endosymbiont An ''endosymbiont'' or ''endobiont'' is any organism that lives within the body or cells of another organism most often, though not always, in a mutualistic relationship. (The term endosymbiosis is from the Greek: ἔνδον ''endon'' "within ...

within a prokaryotic host in the

archaea Archaea ( ; singular archaeon ) is a domain of single-celled organisms. These microorganisms lack cell nuclei and are therefore prokaryotes. Archaea were initially classified as bacteria, receiving the name archaebacteria (in the Archaeba ...

, giving rise to a

symbiotic Symbiosis (from Greek , , "living together", from , , "together", and , bíōsis, "living") is any type of a close and long-term biological interaction between two different biological organisms, be it mutualistic, commensalistic, or para ...

association of two cells from which the first

eukaryotic cell Eukaryotes () are organisms whose cells have a nucleus. All animals, plants, fungi, and many unicellular organisms, are Eukaryotes. They belong to the group of organisms Eukaryota or Eukarya, which is one of the three domains of life. Bacter ...

could have arisen (

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 pos ...

). According to the hydrogen hypothesis: * The hosts that acquired the mitochondria were

hydrogen Hydrogen is the chemical element with the symbol H and atomic number 1. Hydrogen is the lightest element. At standard conditions hydrogen is a gas of diatomic molecules having the formula . It is colorless, odorless, tasteless, non-to ...

-dependent

, possibly similar in physiology to modern methanogenic archaea, which use hydrogen and

carbon dioxide Carbon dioxide ( chemical formula ) is a chemical compound made up of molecules that each have one carbon atom covalently double bonded to two oxygen atoms. It is found in the gas state at room temperature. In the air, carbon dioxide is t ...

to produce

methane Methane ( , ) is a chemical compound with the chemical formula (one carbon atom bonded to four hydrogen atoms). It is a group-14 hydride, the simplest alkane, and the main constituent of natural gas. The relative abundance of methane ...

; * The future mitochondrion was a

facultatively anaerobic 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. Some examples of facultatively anaerobic bacteria are ''Staphylococcus' ...

eubacterium ''Eubacterium'' is a genus of Gram-positive bacteria in the family Eubacteriaceae. These bacteria are characterised by a rigid cell wall A cell wall is a structural layer surrounding some types of cells, just outside the cell membrane. It ...

which produced hydrogen and carbon dioxide as byproducts of

anaerobic respiration Anaerobic respiration is respiration using electron acceptors other than molecular oxygen (O2). Although oxygen is not the final electron acceptor, the process still uses a respiratory electron transport chain. In aerobic organisms undergoing r ...

; * A symbiotic relationship between the two started, based on the host's hydrogen dependence (anaerobic

syntrophy In biology, syntrophy, synthrophy, or cross-feeding (from Greek ''syn'' meaning together, ''trophe'' meaning nourishment) is the phenomenon of one species feeding on the metabolic products of another species to cope up with the energy limitations b ...

Mechanism

The hypothesis differs from many alternative views within the

endosymbiotic theory 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 possib ...

framework, which suggest that the first eukaryotic

cells Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery w ...

evolved a nucleus but lacked mitochondria, the latter arising as a eukaryote engulfed a primitive

bacterium Bacteria (; singular: 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 am ...

that eventually became the mitochondrion. The hypothesis attaches

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

ary significance to

hydrogenosome 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 i ...

s and provides a rationale for their common ancestry with mitochondria. Hydrogenosomes are anaerobic mitochondria that produce ATP by, as a rule, converting

pyruvate Pyruvic acid (CH3COCOOH) is the simplest of the alpha-keto acids, with a carboxylic acid and a ketone functional group. Pyruvate, the conjugate base, CH3COCOO−, is an intermediate in several metabolic pathways throughout the cell. Pyruvic a ...

into hydrogen, carbon dioxide and

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 ...

. Examples from modern biology are known where methanogens cluster around hydrogenosomes within eukaryotic cells. Most theories within the

framework do not address the common ancestry of mitochondria and hydrogenosomes. The hypothesis provides a straightforward explanation for the observation that eukaryotes are genetic chimeras with genes of archaeal and eubacterial ancestry. Furthermore, it would imply that archaea and eukarya split after the modern groups of archaea appeared. Most theories within the

framework predict that some eukaryotes never possessed mitochondria. The hydrogen hypothesis predicts that no primitively mitochondrion-lacking eukaryotes ever existed. In the 15 years following the publication of the hydrogen hypothesis, this specific prediction has been tested many times and found to be in agreement with observation. In 2015, the discovery and placement of the Lokiarchaeota (an archaeal lineage possessing an expanded genetic repertoire including genes involved in membrane remodeling and

actin Actin is a family of globular multi-functional proteins that form microfilaments in the cytoskeleton, and the thin filaments in muscle fibrils. It is found in essentially all eukaryotic cells, where it may be present at a concentration of ov ...

cytoskeletal structure) as the sister group to eukaryotes called into question particular tenets of the hydrogen hypothesis, as Lokiarchaeota appear to lack methanogenesis.

References

{{reflist Hydrogen biology Biological hypotheses

Mechanism

See also

References