Abstract
SummaryBlastocystis is a unicellular stramenopile of controversial pathogenicity in humans [1, 2]. Although it is a strict anaerobe, Blastocystis has mitochondrion-like organelles with cristae, a transmembrane potential and DNA [2–4]. An apparent lack of several typical mitochondrial pathways has led some to suggest that these organelles might be hydrogenosomes, anaerobic organelles related to mitochondria [5, 6]. We generated 12,767 expressed sequence tags (ESTs) from Blastocystis and identified 115 clusters that encode putative mitochondrial and hydrogenosomal proteins. Among these is the canonical hydrogenosomal protein iron-only [FeFe] hydrogenase that we show localizes to the organelles. The organelles also have mitochondrial characteristics, including pathways for amino acid metabolism, iron-sulfur cluster biogenesis, and an incomplete tricarboxylic acid cycle as well as a mitochondrial genome. Although complexes I and II of the electron transport chain (ETC) are present, we found no evidence for complexes III and IV or F1Fo ATPases. The Blastocystis organelles have metabolic properties of aerobic and anaerobic mitochondria and of hydrogenosomes [7, 8]. They are convergently similar to organelles recently described in the unrelated ciliate Nyctotherus ovalis[9]. These findings blur the boundaries between mitochondria, hydrogenosomes, and mitosomes, as currently defined, underscoring the disparate selective forces that shape these organelles in eukaryotes.
Highlights
NADH dehydrogenase subunit NDUFS8 NADH dehydrogenase subunit NDUFA2 NADH dehydrogenase subunit NDUFA5 NADH dehydrogenase subunit NDUFS7 NADH dehydrogenase subunit NDUFA9 NADH dehydrogenase subunit NDUFV1
SDH oxidizes succinate into fumarate while passing electrons via FAD to ubiquinone, FRD: reduction of fumarate to succcinate
Description of symbols and abbreviations used: +, Presence of a signal peptide as predicted by two or more of all four analyses; -, Absence of a signal peptide despite full length N-terminus including a Met start codon; ?, Only one out of the four programs predicted an N-terminal signal peptide. ??, No full length Nterminus because no 5’ Met available; orgDNA, organellar DNA; orgDNA?, possibly organellar DNA; SP, Sequence most probably contains a signal for the secretory pathway. *: Target signal determined with TargetP, SignalP, iPSORT, and Mitoprot. †: Genes for which full length sequences were obtained in this study and sequences generated by PCR in this study. ‡: Could be E2 subunit of α-ketoglutarate dehydrogenase complex (succinyl transacetylase). §: PDH E3 – homologous to L-protein of glycine cleavage system; possibly α-ketoglutarate dehydrogenase complex E3 subunit. ¶: Very low Blast score for this protein. ||: Clusters show Blast similarities to Graves disease carrier (solute carrier family 25 member 16) and ADP/ATP translocator (solute carrier family 25 member 43)
Summary
Removal of two electrons from NADH and transfer to ubiquinone coupled with moving 4 protons across the membrane to create a proton gradient. EC644822, EC644015 terminal electron acceptor proton translocator electron acceptor/donor anaerobic decarboxylation of pyruvate to acetyl-CoA
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