The role of the "Thiodendron" consortium in postulating the karyomastigont chimaera of the endosymbiosis theory by Lynn Margulis.

The endosymbiosis theory of the origin of eukaryotic cell was first proposed more than a hundred years ago. In the second half of the 20th century, Lynn Margulis suggested a new interpretation of the origin of the nucleus in modern eukaryotes. The background was the study of the consortium "Thiodendron", a symbiotic bacterial community, which includes anaerobic aerotolerant motile spirochaetes and sulfidogenic bacteria (sulfidogens) of vibrioid form with a fermentation type of metabolism. Spirochaetes supply sulfidogens with metabolites (pyruvate and, probably, organic nitrogenous products of cell lysis) and get hydrogen sulfide from sulfidogens that helps to maintain a low redox potential. At low oxygen concentrations, spirochaetes are able to assimilate glucose more efficiently. Margulis hypothesized about the symbiotic origin of the nucleus by adding the bacterium Spirochaeta to the Thermoplasma-like archaea. She considered the "Thiodendron"-like consortium to be an intermediate stage in evolution. According to Margulis, the conversion of carbohydrates and the oxidation of Н2S to S0 by the bacterium provided the archaea with electron acceptors for anaerobic respiration, as shown for modern thermoplasmas and products saturated with carbon. The use of carbon sources increased by attaching the floating bacterium to the archaea. More efficient microaerobic oxidation of glucose pre-adapted the spirochaetes for association with Thermoplasma. However, modern "Thiodendron"-like consortia are not in stable symbiosis and a sulfidogenic component of the consortium is capable for fermentation, rather than anaerobic respiration, which makes the theory by Margulis disputable.

Genomics and Biochemistry of Metabolic Pathways for the C1 Compounds Utilization in Colorless Sulfur Bacterium Beggiatoa leptomitoformis D-402.

The metabolic pathways of one-carbon compounds utilized by colorless sulfur bacterium Beggiatoa leptomitoformis D-402 were revealed based on comprehensive analysis of its genomic organization, together with physiological, biochemical and molecular biological approaches. Strain D-402 was capable of aerobic methylotrophic growth with methanol as a sole source of carbon and energy and was not capable of methanotrophic growth because of the absence of genes of methane monooxygenases. It was established that methanol can be oxidized to CO2 in three consecutive stages. On the first stage methanol was oxidized to formaldehyde by the two PQQ (pyrroloquinolinequinone)-dependent methanol dehydrogenases (MDH): XoxF and Mdh2. Formaldehyde was further oxidized to formate via the tetrahydromethanopterin (H4MPT) pathway. And on the third stage formate was converted to CO2 by NAD+-dependent formate dehydrogenase Fdh2. Finally, it was established that endogenous CO2, formed as a result of methanol oxidation, was subsequently assimilated for anabolism through the Calvin-Benson-Bassham cycle. The similar way of one-carbon compounds utilization also exists in representatives of another freshwater Beggiatoa species-B. alba.

Genomic insights into metabolic versatility of a lithotrophic sulfur-oxidizing diazotrophic Alphaproteobacterium Azospirillum thiophilum.

Diazotrophic Alphaproteobacteria of the genus Azospirillum are usually organotrophs, although some strains of Azospirillum lipoferum are capable of hydrogen-dependent autotrophic growth. Azospirillum thiophilum strain was isolated from a mineral sulfide spring, a biotope highly unusual for azospirilla. Here, the metabolic pathways utilized by A. thiophilum were revealed based on comprehensive analysis of its genomic organization, together with physiological and biochemical approaches. The A. thiophilum genome contained all the genes encoding the enzymes of carbon metabolism via glycolysis, tricarboxylic acid cycle and glyoxylate cycle. Genes for a complete set of enzymes responsible for autotrophic growth, with an active Calvin-Benson-Bassham cycle, were also revealed, and activity of the key enzymes was determined. Microaerobic chemolithoautotrophic growth of A. thiophilum was detected in the presence of thiosulfate and molecular hydrogen, being in line with the discovery of the genes encoding the two enzymes involved in dissimilatory thiosulfate oxidation, the Sox-complex and thiosulfate dehydrogenase and Ni-Fe hydrogenases. Azospirillum thiophilum utilizes methanol and formate, producing CO2 that can further be metabolized via the Calvin cycle. Finally, it is capable of anaerobic respiration, using tetrathionate as a terminal electron acceptor. Such metabolic versatility is of great importance for adaptation of A. thiophilum to constantly changing physicochemical environment.

Expansion of ability of denitrification within the filamentous colorless sulfur bacteria of the genus Thiothrix.

Filamentous sulfur bacteria of the genus Thiothrix are able to respire nitrate (NO3-→NO2-) under anaerobic growth. Here, Thiothrix caldifontis (G1(T), G3), Thiothrix unzii (A1(T), TN) and Thiothrix lacustris AS were shown to be capable of further reduction of nitrite and/or nitrous oxides (denitrification). In particular, in the genomes of these strains, excluding T. unzii TN, the nirS gene encoding periplasmic respiratory nitrite reductase was detected, and for T. lacustris AS the nirS expression was confirmed during anaerobic growth. The nirK gene, coding for an alternative nitrite reductase, and the nrfA gene, encoding nitrite reduction to ammonia, were not found in any investigated strains. All Thiothrix species capable of denitrification possess the cnorB gene encoding cytochrome c-dependent NO reductase but not the qnorB gene coding for quinol-dependent NO reductase. Denitrifying capacity ('full' or 'truncated') can vary between strains belonging to the same species and correlates with physical-chemical parameters of the environment such as nitrate, hydrogen sulfide and oxygen concentrations. Phylogenetic analysis revealed the absence of recent horizontal transfer events for narG and nirS; however, cnorB was subjected to gene transfer before the separation of modern species from a last common ancestor of the Thiothrix species.

Ferrovibrio denitrificans gen. nov., sp. nov., a novel neutrophilic facultative anaerobic Fe(II)-oxidizing bacterium.

A neutrophilic Fe(II)-oxidizing bacterium was isolated from the redox zone of a low-salinity spring in Krasnodar krai (Russia), at the FeS-Fe(OH)(3) interface deposited at the sediment surface. The cells of strain Sp-1 were short, thin motile vibrioids with one polar flagellum dividing by binary fission. The optimal values and ranges for pH and temperature were pH 6.2 (5.5-8) and 35 °C (5-45 °C), respectively. The organism was a facultative anaerobe. Strain Sp-1 was capable of organotrophic, lithoheterotrophic and mixotrophic growth with Fe(II) as an electron donor. The denitrification chain was 'disrupted'. Oxidation of Fe(II) was coupled to reduction of NO3 - to NO2 - or of N(2) O to N(2) , as well as under microaerobic conditions, with O(2) as an electron acceptor. The DNA G+C content was 64.2 mol%. According to the results of phylogenetic analysis, the strain was 10.6-12% remote from the closest relatives, members of the genera Sneathiella, Inquilinus, Oceanibaculum and Phaeospirillum within the Alphaproteobacteria. Based on its morphological, physiological and taxonomic characteristics, together with the results of phylogenetic analysis, strain Sp-1 is described as a member of a new genus Ferrovibrio gen. nov., with the type species Ferrovibrio denitrificans sp. nov. and the type strain Sp-1(T) (= LMG 25817(T)  = VKM B-2673(T) ).