If O2 is available at circumneutral pH, Fe2+ is rapidly oxidized to Fe3+, which precipitates as FeO(OH). Neutrophilic iron oxidizing bacteria have evolved mechanisms to prevent self-encrustation in iron. Hitherto, no mechanism has been proposed for cyanobacteria from Fe2+-rich environments; these produce O2 but are seldom found encrusted in iron. We used two sets of illuminated reactors connected to two groundwater aquifers with different Fe2+ concentrations (0.9 μM vs. 26 μM) in the Aspo Hard Rock Laboratory (HRL), Sweden. Cyanobacterial biofilms developed in all reactors and were phylogenetically different between the reactors. Unexpectedly, cyanobacteria growing in the Fe2+-poor reactors were encrusted in iron, whereas those in the Fe2+-rich reactors were not. In-situ microsensor measurements showed that O2 concentrations and pH near the surface of the cyanobacterial biofilms from the Fe2+-rich reactors were much higher than in the overlying water. This was not the case for the biofilms growing at low Fe2+ concentrations. Measurements with enrichment cultures showed that cyanobacteria from the Fe2+-rich environment increased their photosynthesis with increasing Fe2+ concentrations, whereas those from the low Fe2+ environment were inhibited at Fe2+ > 5 μM. Modeling based on in-situ O2 and pH profiles showed that cyanobacteria from the Fe2+-rich reactor were not exposed to significant Fe2+ concentrations. We propose that, due to limited mass transfer, high photosynthetic activity in Fe2+-rich environments forms a protective zone where Fe2+ precipitates abiotically at a non-lethal distance from the cyanobacteria. This mechanism sheds new light on the possible role of cyanobacteria in precipitation of banded iron formations.

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Oxygenic photosynthesis as a protection mechanism for cyanobacteria against iron-encrustation in environments with high Fe2+ concentrations

Sulfur is required for growth of all organisms and is present in a wide variety of metabolites having distinctive biological functions. Sulfur is cycled in ecosystems in nature where conversion of sulfate to organic sulfur compounds is primarily dependent on sulfate uptake and reduction pathways in photosynthetic organisms and microorganisms. In vascular plant species, transport proteins and enzymes in this pathway are functionally diversified to have distinct biochemical properties in specific cellular and subcellular compartments. Recent findings indicate regulatory processes of sulfate transport and metabolism are tightly connected through several modes of transcriptional and posttranscriptional mechanisms. This review provides up-to-date knowledge in functions and regulations of sulfur assimilation in plants and algae, focusing on sulfate transport systems and metabolic pathways for sulfate reduction and synthesis of downstream metabolites with diverse biological functions.

Sulfur Assimilation in Photosynthetic Organisms: Molecular Functions and Regulations of Transporters and Assimilatory Enzymes

NATURAL oxidation of ferrous to ferric iron by bacteria such as Thiobacillus ferrooxidans or Gallionella ferruginea1, or by chemical oxidation2,3 has previously been thought always to involve molecular oxygen as the electron acceptor. Anoxic photochemical reactions4–6 or a photobiological process involving two photosystems7–9 have also been discussed as mechanisms of ferrous iron oxidation. The knowledge of such processes has implications that bear on our understanding of the origin of Precambrian banded iron formations10–14. The reducing power of ferrous iron increases dramatically at pH values higher than 2–3 owing to the formation of ferric hydroxy and oxyhydroxy compounds1,2,15 (Fig. 1). The standard redox potential of Fe3+/Fe2+ (E0 = +0.77 V) is relevant only under acidic conditions. At pH 7.0, the couples Fe(OH)3/Fe2+ (E′0 = -0.236V) or Fe(OH)3 + HCO−3FeCO3 (E′0 = +0.200 V) prevail, matching redox potentials measured in natural sediments9,16,17. It should thus be possible for Fe(n) around pH 7.0 to function as an electron donor for anoxygenic photosynthesis. The midpoint potential of the reaction centre in purple bacteria is around +0.45 V (ref. 18). Here we describe purple, non-sulphur bacteria that can indeed oxidize colourless Fe(u) to brown Fe(in) and reduce CO2 to cell material, implying that oxygen-independent biological iron oxidation was possible before the evolution of oxygenic photosynthesis.

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Ferrous iron oxidation by anoxygenic phototrophic bacteria

Cyanobacteria are photoautotrophic prokaryotes which include a large variety of species of widespread occurrence and with diverse morphological, physiological and biochemical properties. Many cyanobacteria are known to be able to synthesise outermost slimy investments and to release polysaccharidic material into the culture medium during cell growth. These released polysaccharides (RPSs), being easily recoverable from the culture medium, are attracting much interest in view of their possible uses in several industrial applications. In this paper, an overview of the current knowledge on both RPS-producing cyanobacterial strains (including the possible roles of the exopolysaccharides) and chemical characteristics of the cyanobacterial RPSs is given, with particular emphasis on RPS properties and possible industrial applications. On the whole, cyanobacterial RPSs are characterised by a great variety in both number (from two to 10) and type of constitutive monosaccharides (various arrangements of acidic and neutral sugars). Most polymers show an anionic nature due to the presence of uronic acids and/or other charged groups such as pyruvyl or sulfate. Polypeptide moieties as well as acetyl substituents have also sometimes been found, causing additional structural complexity. All the cyanobacterial RPSs so far tested showed a pseudoplastic behaviour, but with marked differences in both viscosity values and shear thinning. In terms of RPS production, the responses of cyanobacteria to changes of culture conditions appear strain-dependent. RPS productivities shown by some cyanobacteria are well comparable with those reported for other photosynthetic microorganisms proposed for polysaccharide production, but very low in comparison with those of heterotrophic microorganisms. Nevertheless, cyanobacteria may be regarded as a very abundant source of structurally diverse polysaccharides, some of which may possess unique properties for special applications, not fulfilled by the polymers currently available. However, much work has still to be done to bridge the wide gap existing between data on the biology of the RPS-producer strains and information concerning technological and other useful properties of the cyanobacterial RPS.

Exocellular polysaccharides from cyanobacteria and their possible applications

We imaged reflectance and variable fluorescence in 25 cyanobacterial mats from four distant sites around the globe to assess, at different scales of resolution, spatial variabilities in the physiological parameters characterizing their photosynthetic capacity, including the absorptivity by chlorophyll a (Achl), maximum quantum yield of photosynthesis (Ymax), and light acclimation irradiance (Ik). Generally, these parameters significantly varied within individual mats on a sub-millimeter scale, with about 2-fold higher variability in the vertical than in the horizontal direction. The average vertical profiles of Ymax and Ik decreased with depth in the mat, while Achl exhibited a sub-surface maximum. The within-mat variability was comparable to, but often larger than, the between-sites variability, whereas the within-site variabilities (i.e., between samples from the same site) were generally lowest. When compared based on averaged values of their photosynthetic parameters, mats clustered according to their site of origin. Similar clustering was found when the community composition of the mats' cyanobacterial layers were compared by automated ribosomal intergenic spacer analysis (ARISA), indicating a significant link between the microbial community composition and function. Although this link is likely the result of community adaptation to the prevailing site-specific environmental conditions, our present data is insufficient to identify the main factors determining these patterns. Nevertheless, this study demonstrates that the spatial variability in the photosynthetic capacity and light acclimation of benthic phototrophic microbial communities is at least as large on a sub-millimeter scale as it is on a global scale, and suggests that this pattern of variability scaling is similar for the microbial community composition.

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Spatial patterns and links between microbial community composition and function in cyanobacterial mats

Four different types of adaptation to sulfide among cyanobacteria are described based on the differential toxicity to sulfide of photosystems I and II and the capacity for the induction of anoxygenic photosynthesis. Most cyanobacteria are highly sensitive to sulfide toxicity, and brief exposures to low concentrations cause complete and irreversible cessation of CO2 photoassimilation. Resistance of photosystem II to sulfide toxicity, allowing for oxygenic photosynthesis under sulfide, is found in cyanobacteria exposed to low H2S concentrations in various hot springs. When H2S levels exceed 200 μM another type of adaptation involving partial induction of anoxygenic photosynthesis, operating in concert with partially inhibited oxygenic photosynthesis, is found in cyanobacterial strains isolated from both hot springs and hypersaline cyanobacterial mats. The fourth type of adaptation to sulfide is found at H2S concentrations higher than 1 mM and involves a complete replacement of oxygenic photosynthesis by an effective sulfide-dependent, photosystem II-independent anoxygenic photosynthesis. The ecophysiology of the various sulfide-adapted cyanobacteria may point to their uniqueness within the division of cyanobacteria.

Ricardo Poplawski

Papers

Adaptation to Hydrogen Sulfide of Oxygenic and Anoxygenic Photosynthesis among Cyanobacteria.