The electron transport chain in anaerobically functioning eukaryotes

AbstractMany lower eukaryotes can survive anaerobic conditions via a fermentation pathway that involves the use of the reduction of endogenously produced fumarate as electron sink. This fumarate reduction is linked to electron transport in an especially adapted, anaerobically functioning electron-transport chain.An aerobic energy metabolism with Krebs cycle activity is accompanied by electron transfer from succinate to ubiquinone via complex II of the respiratory chain. On the other hand, in an anaerobic metabolism, where fumarate functions as terminal electron acceptor, electrons are transferred from rhodoquinone to fumarate, which is the reversed direction. Ubiquinone cannot replace rhodoquinone in the process of fumarate reduction in vivo, as ubiquinone can only accept electrons from complex II and cannot donate them to fumarate. Rhodoquinone, with its lower redox potential than ubiquinone, is capable of donating electrons to fumarate. Eukaryotic fumarate reductases were shown to interact with rhodoquinone (a benzoquinone), whereas most prokaryotic fumarate reductases interact with the naphtoquinones menaquinone and demethylmenaquinone.Fumarate reductase, the enzyme essential for the anaerobic functioning of many eukaryotes, is structurally very similar to succinate dehydrogenase, the Krebs cycle enzyme catalysing the reverse reaction. In prokaryotes these enzymes are differentially expressed depending on the external conditions. Evidence is now emerging that also in eukaryotes two different enzymes exist for succinate oxidation and fumarate reduction that are differentially expressed

Multiple origins of hydrogenosomes:functional and phylogenetic evidence from the ADP/ATP carrier of the anaerobic chytrid Neocallimastix sp.

A mitochondrial-type ADP/ATP carrier (AAC) has been identified in the hydrogenosomes of the anaerobic chytridiomycete fungus Neocallimastix sp. L2. Biochemical and immunocytochemical studies revealed that this ADP/ATP carrier is an integral component of hydrogenosomal membranes. Expression of the corresponding cDNA in Escherichia coli confers the ability on the bacterial host to incorporate ADP at significantly higher rates than ATP - similar to isolated mitochondria of yeast and animals. Phylogenetic analysis of this AAC gene (hdgaac ) confirmed with high statistical support that the hydrogenosomal ADP/ATP carrier of Neocallimastix sp. L2 belongs to the family of veritable mitochondrial-type AACs. Hydrogenosome-bearing anaerobic ciliates possess clearly distinct mitochondrial-type AACs, whereas the potential hydrogenosomal carrier Hmp31 of the anaerobic flagellate Trichomonas vaginalis and its homologue from Trichomonas gallinae do not belong to this family of proteins. Also, phylogenetic analysis of genes encoding mitochondrial-type chaperonin 60 proteins (HSP 60) supports the conclusion that the hydrogenosomes of anaerobic chytrids and anaerobic ciliates had independent origins, although both of them arose from mitochondria

Surprising variety in energy metabolism within Trypanosomatidae

The metabolism of Trypanosomatidae differs significantly between distinct species and can even be completely different between various life-cycle stages of the same species. It has been proposed that differences in energy metabolism are related to differences in nutrient supply in the environments of the various trypanosomatids. However, the literature shows that availability of substrates does not dictate the type of energy metabolism of trypanosomatids, as Trypanosoma theileri, Trypanosoma lewisi and African trypanosomes all live in the bloodstream of their mammalian host, but have surprisingly large differences in metabolism. Furthermore, in trypanosomatids no obvious relationship exists between energy metabolism and phylogeny or mode of transmission. We provide an overview of the metabolic capacities in the energy metabolism of distinct trypanosomatids, and suggest that these can be divided into four different metabolic categories of increasing complexity

Phospholipids in parasitic protozoa

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Trypanosomatidae produce acetate via a mitochondrial acetate:succinate CoA transferase

Hydrogenosome-containing anaerobic protists, such as the trichomonads, produce large amounts of acetate by an acetate:succinate CoA transferase (ASCT)/succinyl CoA synthetase cycle. The notion that mitochondria and hydrogenosomes may have originated from the same α-proteobacterial endosymbiont has led us to look for the presence of a similar metabolic pathway in trypanosomatids because these are the earliest-branching mitochondriate eukaryotes and because they also are known to produce acetate. The mechanism of acetate production in these organisms, however, has remained unknown. Four different members of the trypanosomatid family: promastigotes of Leishmania mexicana mexicana, L. infantum and Phytomonas sp., and procyclics of Trypanosoma brucei were analyzed as well as the parasitic helminth Fasciola hepatica. They all use a mitochondrial ASCT for the production of acetate from acetyl CoA. The succinyl CoA that is produced during acetate formation by ASCT is recycled presumably to succinate by a mitochondrial succinyl CoA synthetase, concomitantly producing ATP from ADP. The ASCT of L. mexicana mexicana promastigotes was further characterized after partial purification of the enzyme. It has a high affinity for acetyl CoA (K(m) 0.26 mM) and a low affinity for succinate (K(m) 6.9 mM), which shows that significant acetate production can occur only when high mitochondrial succinate concentrations prevail. This study identifies a metabolic pathway common to mitochondria and hydrogenosomes, which strongly supports a common origin for these two organelles

Schistosome biology and proteomics: Progress and challenges

The recent availability of schistosomal genome-sequence information allows protein identification in schistosome-derived samples by mass spectrometry (proteomics). Over the last few years, several proteome studies have been performed that addressed important questions in schistosome biology. This review summarizes the applied experimental approaches that have been used so far, it provides an overview of the most important conclusions that can be drawn from the performed studies and finally discusses future challenges in this research area

The ability to incorporate functional plastids by the sea slug Elysia viridis is governed by its food source

Functional kleptoplasty in sacoglossan sea slugs is among the most curious photosynthetic associations known. One member of these marine molluscs, Elysia viridis, is known to incorporate plastids from a variety of different algae food sources, but with apparently different outcomes and differences in the time span of the retention of functional kleptoplasts. While it was previously shown that kleptoplasts that stem from Codium tomentosum are kept functional for several weeks (long-term retention, LtR), those that stem from Bryopsis hypnoides or Cladophora rupestris are thought to be of limited use regarding photosynthetic capacity (short-term retention, StR). This is important, because it touches upon the popular yet controversial question of how important photosynthesis is for the thriving of these slugs. The aim of the present study was to determine to what degree the plastid source determines retention time. We, therefore, compared E. viridis feeding on either Cladophora sp. or B. hypnoides. We show that kleptoplasts of B. hypnoides incorporate 14CO2, but with rapidly declining efficiency throughout the first week of starvation, while the plastids of Cladophora sp. are, surprisingly, not incorporated to begin with. The radulae of the different samples showed adjustment to the food source, and when feeding on Cladophora sp., E. viridis survived under laboratory conditions under both starvation and non-starvation conditions. Our results demonstrate that (i) the ability to incorporate plastids by E. viridis differs between the food sources B. hypnoides and Cladophora sp., and (ii) photosynthetic active kleptoplasts are not an inevitable requirement for survival