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PHYSIOLOGY,PHYLOGENY,EARLY EVOLUTION AND GADPH

Biologists have recognized the fact that complexity in eukaryotic cells stems from endosymbiosis,
primarily the acquisition of mitochondria. Bacterial organisms may develop an absolute symbiosis with a
higher organism and can impress a new character in the factors of heredity. One of the major ideas
involved in the endosymbiotic theory is the transfer of genes from organelle to the host chromosome.
Chloroplast cytosol isoenzymes called GADPH was an excellent system to test gene transfer in
endosymbiosis. It is reported that GADPH activity in plants is similar to that of yeast and animal tissues.
GADPH from E.coli were more similar to the ones in eukaryotes than the ones in thermophilic bacteria,
the reasoning was that it was a case of LGT from eukaryotes to prokaryotes which is usually not the case.
The A and B subunits of GADPH were results of nuclear gene duplication early in evolution of green
plant lineage. On closer look at GADPH genes, evidence was found for identical intron positions in
anciently diverged genes. The cyanobacterium Anabena had a Calvin cycle homologue of GADPH that
had branched specifically with the nuclear encoded chloroplast GADPH enzymes of higher plants.
Anabena also had two other GADPH genes one which was a copy of the E.coli version erroneously
thought to be a case of eukaryote-to-prokaryote LGT. The studies had shown eukaryotic GADPH should
derive from archaeal GADPH but the archaeal GADPH for all practical purposes is unrelated to both
bacterial and eukaryotic GADPH. This was interesting because it led to the following conclusion even
though glycolytic GADPH is an acquisition of endosymbiosis and stems from mitochondrion, it showed
the evolution of the host that acquired the plastid but not that of the host that acquired the mitochondrion.
Eukaryotes have a bacterial glycolytic pathway; protein import machinery is also a eukaryotic invention
albeit having prokaryotic components, thus when this machinery wasn’t there all gene products of
endosymbionts were targeted at the cytosol, hence the predominance of bacterial genes in eukaryotes. For
the transformation from autotrophs to heterotrophs would require a new glycolytic pathway to feed the
mitochondria. Many endosymbiotic theories exist to explain this, a controversial and interesting idea
being the bacterial glycolytic pathway moving to the host cytosol to feed the mitochondrion via gene
transfer. But we should never forget that these retargeting of enzymes and pathways is governed by
variation and natural selection, and a single enzyme retargeting without the whole pathway is more of a
burden than help to the cell. Analyses of GADPH helped in explaining the origin of complex algal plastids
and the evolutionary origins of land plants, also providing insights into the early diversification of LUCA
genes.

Life is just a chemical reaction and we are just its by-products whose meaning is to keep the reaction
going.

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