NEW ARCHAEAL LINEAGES AND ORIGIN OF MITOCHONDRIA

Nucleus arose in the wake of mitochondria, probably the endomembrane system. The ribosomal proteins
link the archaeal ribosomes found in the cytosol of the hosts to specific lineages in the marine sediment.
The draft genome assembly of the Lokiarchaeum lineage was pieced together from marine segments and it
caused quite a stir since it was reported a complex cell which could provide the missing link between
prokaryotes and eukaryotes, a probable candidate for LECA. Archezoa were early branching eukaryotes
which weren’t amitochondriate as previously hypothesized, but they had reduced forms of mitochondria
called hydrogenosomes and mitosomes. Hydrogenosomes generated hydrogen as the end product of
fermentative ATP synthesis via SLP whereas mitosomes had no role in ATP synthesis but was involved in
sulfur metabolism. The hydrogen hypothesis suggests that the endosymbiotic origins of mitochondria is
due to anaerobic syntrophy rather than phagocytosis. The shorthand for this would be conversion of a
free-living symbiont to endosymbiont, which in turn becomes the ATP producing organelle, thereby the
union of prokaryotes forms a eukaryotic cell. This idea has parallels in the real world where fungi have
bacterial endosymbionts but no fungi are phagotrophic. By the hydrogen hypothesis for phagocytosis to
happen, the need of mitochondria can’t be neglected since it’s an energy intensive process. This forms our
basis for the hypothesis that LECA had mitochondria. This anaerobic syntrophy makes certain predictions
like the host should be hydrogen dependent and it should be an archaeon.

There have been two basic hypotheses regarding the formation of the eukaryotic endomembrane system
like the invagination of plasma membrane or the hypothetical cellular fusion processes involving
symbiosis which doesn’t involve origin of mitochondrion. The recent hypothesis that tries to explain it is
simpler, it states the outer membrane vesicles of the bacterial ancestor of mitochondrion accumulating in
the cytosol of archaeal host, this slowly results in the formation of a primitive ER by fusion of the
vesicles, selection acts on it and slowly nuclear membrane is derived from it. The outward vesicle flux
accounts for the transformation from archaeal lipids to bacterial lipids and the transformation of
chemiosmotic energy conservation to mitochondrial ATP synthesis. A few genes that perhaps could
explain the energy and carbon metabolism are missing in the metagenome, but it’s not clear whether they
are missing in the genome or only in the metagenome assembly. If a new gene crops up, it becomes
difficult to explain whether they stem from the same genome corresponding to rRNA or protein or is just
common in the environment of the surviving lineage. Lokiarchaeum genome has autotrophic carbon
dioxide fixation, pathways that are used for methanogenesis, generating ion gradient for ATP synthesis.
Archaeal fermenters use protons as terminal electron acceptors via a membrane bound hydrogenase
complex.

The proposal also includes a part about Bathyarchaeota, which can use the hydrogen and carbon dioxide
from fermentation for acetogenesis which could be beneficial in wild conditions. If this is happening
where the cell uses hydrogen and carbon dioxide produced by itself, then it’s a case of intracellular
syntrophy where a cell lives off on its own waste products of metabolism. One question raised is about
whether Lokiarchaeum has both fermentative and acetogenic metabolisms. But due to the fact that many
crucial genes like rotor stator ATPase and soluble hydrogenase in many lineages it’s difficult to say
whether they are actually missing or just the metagenome is incomplete. In anaerobic sediments,
Hydrogen partial pressure plays a key role in whether the organism gains energy by carbon dioxide
reducing or carbon dioxide generating reaction. Under anaerobic conditions syntrophic interactions are
pretty common and thermodynamics plays a defining role especially for acetogenesis. The genes only
predict an archaeum as a host and an alpha-proteobacterium as the symbiont at the root of origin of the
mitochondrion. Another question raised is whether sulfur reducers are at the origin of eukaryotes, where it
 NEW ARCHAEAL LINEAGES AND ORIGIN OF MITOCHONDRIA

plays the endosymbiotic origin of nucleus which is pretty much unanswered.