A potential cyanobacterial ancestor of Viridiplantae chloroplasts

The theory envisaging the origin of plastids from endosymbiotic cyanobacteria is well-established but it is difficult to explain the evolution (spread) of plastids in phylogenetically diverse plant groups. It is widely believed that primordial endosymbiosis occurred in the last common ancestor of all algae^1^, which then diverged into the three primary photosynthetic eukaryotic lineages, viz. the Rhodophyta (red algae), Glaucocystophyta (cyanelle-containing algae) and Viridiplantae (green algae plus all land plants)^2^. Members of these three groups invariably have double membrane-bound plastids^3^, a property that endorses the primary endosymbiotic origin of the organelles. On the other hand, the three or four membrane-bound plastids of the evolutionary complicated Chromalveolates [chromista (cryptophytes, haptophytes, and stramenopiles) and alveolata (dinoflagellates, apicomplexans, and ciliates)] are inexplicable in the light of a single endosymbiosis event, thereby necessitating the postulation of the secondary^4,5^ and tertiary^6^ endosymbiosis theories where a nonphotosynthetic protist supposedly engulfed a red or a green alga^7^ and an alga containing a secondary plastid itself was engulfed^8^ respectively. In the current state of understanding, however, there is no clue about the taxonomic identity of the cyanobacterial ancestor of chloroplasts, even though there is a wide consensus on a single primordial endosymbiosis event. During our metagenomic investigation of a photosynthetic geothermal microbial mat community we discovered a novel order-level lineage of Cyanobacteria that - in 16S rRNA gene sequence-based phylogeny - forms a robust monophyletic clade with chloroplast-derived sequences from diverse divisions of Viridiplantae. This cluster diverged deeply from the other major clade encompassing all hitherto known groups of Cyanobacteria plus the chloroplasts of Rhodophyta, Glaucocystophyceae and Chromalveolates. Since this fundamental dichotomy preceded the origin of all chloroplasts, it appears that two early-diverging cyanobacterial lineages had possibly given rise to two discrete chloroplast descents via two separate engulfment events

Hyperthermophiles: Diversity, Adaptation and Applications

Hyperthermophiles are microorganisms that love to grow optimally in extremely hot environments, with optimum temperatures for growth of 80 °C and above. Most of the hyperthermophiles are represented by archaea; and only a few bacteria, such as Geothermobacterium ferrireducens, and members of the genera Aquifex and Thermotoga have been reported to grow at temperatures closer to 100 °C. Several archaea, on the other hand, such as Methanopyrus kandleri, Geogemma barossii, Pyrolobus fumarii, Pyrococcus kukulkanii, Pyrodictium occultum, etc. isolated from terrestrial hot springs, marine hydrothermal vents, or other hyperthermal environments have been reported to grow optimally even above the boiling point of water. The discovery of this astonishing group of microorganisms has not only provided us with the model systems to study the structural and functional dynamics of the biomolecules, and to understand the molecular mechanisms of their adaptation to such high temperature, not even closer to what can be endured by other life forms, but also have boosted the biotechnological industry to search for new products, particularly enzymes with unique characteristics, from them. This chapter has exhaustively reviewed the different hyperthermal environments on Earth’s surface and the hyperthermophilic microbial diversity in such environments; mechanisms of adaptation of the hyperthermophiles, especially with regard to the adaptations of the membrane structures, maintenance of the structures of the nucleic acids and proteins; and their diverse applications in human welfare.&amp;nbsp;&lt;br&gt;</jats:p

Extremophiles: An Overview

Earth contains several environmental extremes which are uninhabitable for most of the living beings. But, astonishingly, in the last few decades, several organisms thriving in such extreme environments have been discovered. “Extremophiles”, meaning “Lovers of Extremities” are the entities that are especially adapted to live in such harsh environmental conditions in which other entities cannot live. The discovery of extremophiles has not only boosted the biotech industry to search for new products from them, but also made researchers to think for the existence of extra-terrestrial life. The most inhospitable environments include physical or chemical extremities, like high or low temperatures, radiation, high pressure, water scarcity, high salinity, pH extremes, and limitation of oxygen. Microorganisms have been found to live in all such environmental conditions, like hyperthermophiles and psychrophiles, acidophiles and alkaliphiles. Bacteria like Deinococcus radiodurans, which is able to withstand extreme gamma radiation, and Moritella sp., able to grow at atmospheric pressure of &amp;gt;1000 atm, have been reported. Environments like the Dead Sea, having saturated NaCl concentrations, hold extreme halophiles like Halobacterium salinarum. Highly acidic environments, like the Rio-Tinto River in Spain or Danakil depression in Ethiopia harbour acidophiles with growth optima of pH zero, or close to it. Bacillus alcalophilus, and Microcystis aeruginosa on the other hand inhabit natural alkaline soda lakes where pH can reach about 12.0. A number of anaerobic prokaryotes can live in complete anoxic environments by using terminal electron acceptors other than oxygen. In this chapter, we shall discuss very briefly the diversity of all extremophiles and their mechanism(s) of adaptation.&lt;br&gt;</jats:p

Subject Index

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Diversity and Mechanisms of Adaptation of Predominant Bacterial Chemolithotrophs in Extreme Habitats

&amp;nbsp;Bacterial chemolithotrophy is one of the most ancient metabolisms and is generally defined as the ability of some microorganisms to utilize a wide range of inorganic substrates as an energy or electron source. While lithotrophy can itself be considered as extremophily, as only some microorganisms (the rock-eaters) have the ability to utilize diverse inorganic chemicals as the sole source of energy, the phylogenetically diverse groups of lithotrophs can thrive in a wide range of extreme habitats. Apart from their excellent eco-physiological adaptability, they also possess versatile enzymatic machinery for maintaining their lithotrophic attributes under such extreme environments. In this chapter, we have highlighted the diversity of iron, hydrogen and sulfur lithotrophic extremophilic bacteria in various extreme habitats, and their role in maintaining the primary productivity, ecosystem stability and mineral cycling / mineralogical transformations. Moreover, genetic determinants and different enzymatic systems which are reported to be involved in such lithotrophic metabolism also have been discussed. We hope this article will shed some new light on the field of extremophile lithotrophy, which will eventually improve our understanding of the extended new boundaries of life.&amp;nbsp;&lt;br&gt;</jats:p

Kinetic Enrichment of ³⁴ S during Proteobacterial Thiosulfate Oxidation and the Conserved Role of SoxB in S-S Bond Breaking

During chemolithoautotrophic thiosulfate oxidation, the phylogenetically diverged proteobacteria Paracoccus pantotrophus , Tetrathiobacter kashmirensis , and Thiomicrospira crunogena rendered steady enrichment of 34 S in the end product sulfate, with overall fractionation ranging between −4.6‰ and +5.8‰. The fractionation kinetics of T. crunogena was essentially similar to that of P. pantotrophus , albeit the former had a slightly higher magnitude and rate of 34 S enrichment. In the case of T. kashmirensis , the only significant departure of its fractionation curve from that of P. pantotrophus was observed during the first 36 h of thiosulfate-dependent growth, in the course of which tetrathionate intermediate formation is completed and sulfate production starts. The almost-identical 34 S enrichment rates observed during the peak sulfate-producing stage of all three processes indicated the potential involvement of identical S-S bond-breaking enzymes. Concurrent proteomic analyses detected the hydrolase SoxB (which is known to cleave terminal sulfone groups from SoxYZ-bound cysteine S -thiosulfonates, as well as cysteine S -sulfonates, in P. pantotrophus ) in the actively sulfate-producing cells of all three species. The inducible expression of soxB during tetrathionate oxidation, as well as the second leg of thiosulfate oxidation, by T. kashmirensis is significant because the current Sox pathway does not accommodate tetrathionate as one of its substrates. Notably, however, no other Sox protein except SoxB could be detected upon matrix-assisted laser desorption ionization mass spectrometry analysis of all such T. kashmirensis proteins as appeared to be thiosulfate inducible in 2-dimensional gel electrophoresis. Instead, several other redox proteins were found to be at least 2-fold overexpressed during thiosulfate- or tetrathionate-dependent growth, thereby indicating that there is more to tetrathionate oxidation than SoxB alone. </jats:p

A potential cyanobacterial ancestor of Viridiplantae chloroplasts

AbstractThe theory envisaging the origin of plastids from endosymbiotic cyanobacteria is well-established but it is difficult to explain the evolution (spread) of plastids in phylogenetically diverse plant groups. It is widely believed that primordial endosymbiosis occurred in the last common ancestor of all algae^1^, which then diverged into the three primary photosynthetic eukaryotic lineages, viz. the Rhodophyta (red algae), Glaucocystophyta (cyanelle-containing algae) and Viridiplantae (green algae plus all land plants)^2^. Members of these three groups invariably have double membrane-bound plastids^3^, a property that endorses the primary endosymbiotic origin of the organelles. On the other hand, the three or four membrane-bound plastids of the evolutionary complicated Chromalveolates [chromista (cryptophytes, haptophytes, and stramenopiles) and alveolata (dinoflagellates, apicomplexans, and ciliates)] are inexplicable in the light of a single endosymbiosis event, thereby necessitating the postulation of the secondary^4,5^ and tertiary^6^ endosymbiosis theories where a nonphotosynthetic protist supposedly engulfed a red or a green alga^7^ and an alga containing a secondary plastid itself was engulfed^8^ respectively. In the current state of understanding, however, there is no clue about the taxonomic identity of the cyanobacterial ancestor of chloroplasts, even though there is a wide consensus on a single primordial endosymbiosis event. During our metagenomic investigation of a photosynthetic geothermal microbial mat community we discovered a novel order-level lineage of Cyanobacteria that - in 16S rRNA gene sequence-based phylogeny - forms a robust monophyletic clade with chloroplast-derived sequences from diverse divisions of Viridiplantae. This cluster diverged deeply from the other major clade encompassing all hitherto known groups of Cyanobacteria plus the chloroplasts of Rhodophyta, Glaucocystophyceae and Chromalveolates. Since this fundamental dichotomy preceded the origin of all chloroplasts, it appears that two early-diverging cyanobacterial lineages had possibly given rise to two discrete chloroplast descents via two separate engulfment events.</jats:p

34S enrichment as a signature of thiosulfate oxidation in the “<i>Proteobacteria</i>”

ABSTRACT Kinetics of thiosulfate oxidation, product and intermediate formation, and 34S fractionation, were studied for the members of Alphaproteobacteria Paracoccus sp. SMMA5 and Mesorhizobium thiogangeticum SJTT, the Betaproteobacteria member Pusillimonas ginsengisoli SBO3, and the Acidithiobacillia member Thermithiobacillus sp. SMMA2, during chemolithoautotrophic growth in minimal salts media supplemented with 20 mM thiosulfate. The two Alphaproteobacteria oxidized thiosulfate directly to sulfate, progressively enriching the end-product with 34S; Δ34Sthiosulfate-sulfate values recorded at the end of the two processes (when no thiosulfate was oxidized any further) were −2.9‰ and −3.5‰, respectively. Pusillimonas ginsengisoli SBO3 and Thermithiobacillus sp. SMMA2, on the other hand, oxidized thiosulfate to sulfate via tetrathionate intermediate formation, with progressive 34S enrichment in the end-product sulfate throughout the incubation period; Δ34Sthiosulfate-sulfate, at the end of the two processes (when no further oxidation took place), reached −3.5‰ and −3.8‰, respectively. Based on similar 34S fractionation patterns recorded previously during thiosulfate oxidation by strains of Paracoccus pantotrophus, Advenella kashmirensis and Hydrogenovibrio crunogenus, it was concluded that progressive reverse fractionation, enriching the end-product sulfate with 34S, could be a characteristic signature of bacterial thiosulfate oxidation.</jats:p