INVITED REVIEW The Hydrogenosome
Abstract
The hydrogenosome is an organelle described in trichomonads, parasitic protists where most studies were carried out and also in organisms phylogenetically distant such as rumen ciliates, some free-living ciliates and chytrid fungí (Chytridiomycota). The organisms presenting hydrogenosomes live in anaerobic habitats and they all lack mitochondria. The hydrogenosome contains enzymes that particípate in the metabolism of pyruvate and is the site of formation of ATP and molecular hydrogen. The origin of the hydrogenosome brought a great debate, since, like mitochondria, it has a double membrane envelope, divides autonomously by fission, imports proteins post-translationally, and produces ATP. How- ever, it differs from mitochondria in that it seems to lack a genome, a respiratory chain, cytochromes, the F F system, the tricarboxylic acid cycle, oxidative phosphorylation, and cardiolipin. The hydrogenosome uses pyruvate: ferredoxin oxidoreductase, a counterpart to the mitochondrial pyruvate dehydrogenase complex and presents hydrogenase, an enzyme typically restricted to anaerobes. Hypotheses have been proposed for the origin of the trichomonad hydrogenosome: an independent endosymbiosis of an anaerobic eubacterium with a eukaryotic host or the conversión of an established mi- mitochondrion adapted to an anaerobic lifestyle. The common origin hypothesis for mitochondria and hydrogenosome was refined to state that both organelles evolved from a common progenitor structure present in eukaryotes before the advent of true mitochondria or hydrogenosome.
Hydrogenosome is a spherical or slightly elongated structure (when in process of división), 0.5-1.0 pm diameter, usually associated with cytoskeletal structures such as the axostyle and costa in trichomonads. The matrix of the hydrogenosome is homogeneously granular. Crista-like invaginations of inner-membrane were described in the hydrogenosome of some rumen ciliates. The matrix also prevents calcium deposits. The hydrogenosome of trichomonads contains a flattened, membrane-bounded vesicle completely dissimilar from the hydrogenosomes matrix. This vesicle contains high levéis of Mg++, Ca++ and P and possibly functions in intracellular calcium regulation. Its membrane presents N-acetyl-glucosamine-containing glycoconjugates. Close proximity and even continuity, between the endoplasmic reticulum and the hydrogenosome were observed. Hydrogenosomal proteins are synthesized on free polyribosomes, protein import occurs post-translationally, is dependent on the presence of a cleavable N-terminal presequence, cytosolic protein (s) and energy in the form of temperature, ATP and a transmembrane electrochemical potential, similar to that of mitochondria and chloroplasts.