Structural basis for the assembly and electron transport mechanisms of the dimeric photosynthetic RC-LH1 supercomplex

The reaction center (RC) and light-harvesting complex 1 (LH1) form a RC-LH1 core supercomplex that is vital for the primary reactions of photosynthesis in purple photosynthetic bacteria. Some species possess the dimeric RC-LH1 complex with an additional polypeptide PufX, representing the largest photosynthetic complex in anoxygenic phototrophs. However, the details of the architecture and assembly mechanism of the RC-LH1 dimer are unclear. Here we report seven cryo-electron microscopy (cryo-EM) structures of RC-LH1 supercomplexes from Rhodobacter sphaeroides. Our structures reveal that two PufX polypeptides are positioned in the center of the S-shaped RC-LH1 dimer, interlocking association between the components and mediating RC-LH1 dimerization. Moreover, we identify a new transmembrane peptide, designated PufY, which is located between the RC and LH1 subunits near the LH1 opening. PufY binds a quinone molecule and prevents LH1 subunits from completely encircling the RC, creating a channel for quinone/quinol exchange. Genetic mutagenesis, cryo-EM structures, and computational simulations enable a mechanistic understanding of the assembly and electron transport pathways of the RC-LH1 dimer and elucidate the roles of individual components in ensuring the structural and functional integrity of the photosynthetic supercomplex.

Taxonomical and functional diversity of microbial communities in two hot springs of the Baikal rift zone

The Baikal Rift Zone hosts many hot springs with a wide range of temperature and physical-chemical conditions, which may harbour different niches for the distribution of microbial communities. We investigated microbial community composition and their functional activity in two alkaline hot springs with a temperature range of 34.4 to 73.6°C. Comparative analysis of the composition of the dominant taxa showed significant differences depending on the collection sites. In the community of high-temperature zones with a water temperature of 55-64°C, a high proportion of thermophilic bacteria Acetothermia (up to 57.9%), Deinococcus-Thermus (up to 50%), and Aquificae (up to 10.8%). Proteobacteria (29-77%) and Firmicutes (15-26%) dominate in the sulphide-free Garga spring (73-75°C). The functional analysis of the microbial community showed that the primary producers are cyanobacteria, anoxygenic phototrophs, and chemolithotrophic bacteria. At the terminal stages of the mineralization of organic matter, sulphate-reducing bacteria are the main destructors in the microbial communities in hot springs. The cyano-bacterial and sulfidogenic microbial communities play an important role in the formation of geochemical barriers and mineral formation.

Redox-Controlled Ammonium Storage and Overturn in Ediacaran Oceans

As a key nutrient, nitrogen can limit primary productivity and carbon cycle dynamics, but also evolutionary progress. Given strong redox-dependency of its molecular speciation, environmental conditions can control nitrogen localization and bioavailability. This particularly applies to periods in Earth history with strong and frequent redox fluctuations, such as the Neoproterozoic. We here report on chlorophyll-derived porphyrins and maleimides in Ediacaran sediments from Oman. Exceptionally light δ15N values (< –10‰) in maleimides derived from anoxygenic phototrophs point towards ammonium assimilation at the chemocline, whereas the isotopic offset between kerogens and chlorophyll-derivatives indicates a variable regime of cyanobacterial and eukaryotic primary production in surface waters. Biomarker and maleimide mass balance considerations imply shallow euxinia during the terminal Ediacaran and a stronger contribution of anoxygenic phototrophs to primary productivity, possibly as a consequence of nutrient ‘lockup’ in a large anoxic ammonium reservoir. Synchronous δ13C and δ15N anomalies at the Ediacaran–Cambrian boundary may reflect one in a series of overturn events, mixing ammonium and isotopically-light DIC into oxic surface waters. By modulating access to nitrogen, environmental redox conditions may have periodically affected Ediacaran primary productivity, carbon cycle perturbations, and possibly played a role in the timing of the metazoan radiation across the terminal Ediacaran and early Cambrian.

Carotenoid biomarkers in Namibian shelf sediments: Anoxygenic photosynthesis during sulfide eruptions in the Benguela Upwelling System

Aromatic carotenoid-derived hydrocarbon biomarkers are ubiquitous in ancient sediments and oils and are typically attributed to anoxygenic phototrophic green sulfur bacteria (GSB) and purple sulfur bacteria (PSB). These biomarkers serve as proxies for the environmental growth requirements of PSB and GSB, namely euxinic waters extending into the photic zone. Until now, prevailing models for environments supporting anoxygenic phototrophs include microbial mats, restricted basins and fjords with deep chemoclines, and meromictic lakes with shallow chemoclines. However, carotenoids have been reported in ancient open marine settings for which there currently are no known modern analogs that host GSB and PSB. The Benguela Upwelling System offshore Namibia, known for exceptionally high primary productivity, is prone to recurrent toxic gas eruptions whereupon hydrogen sulfide emanates from sediments into the overlying water column. These events, visible in satellite imagery as water masses clouded with elemental sulfur, suggest that the Benguela Upwelling System may be capable of supporting GSB and PSB. Here, we compare distributions of biomarkers in the free and sulfur-bound organic matter of Namibian shelf sediments. Numerous compounds—including acyclic isoprenoids, steranes, triterpanes, and carotenoids—were released from the polar lipid fractions upon Raney nickel desulfurization. The prevalence of isorenieratane and β-isorenieratane in sampling stations along the shelf verified anoxygenic photosynthesis by low-light-adapted, brown-colored GSB in this open marine setting. Renierapurpurane was also present in the sulfur-bound carotenoids and was typically accompanied by lower abundances of renieratane and β-renierapurpurane, thereby identifying cyanobacteria as an additional aromatic carotenoid source.

Draft Genome Sequence of the Aerobic Anoxygenic Phototrophic Bacterium Roseobacter sp. Strain OBYS 0001, Isolated from Coastal Seawater in Otsuchi Bay, Japan

Here, we report the draft genome sequence of the aerobic anoxygenic phototrophic bacterium Roseobacter sp. strain OBYS 0001, isolated from coastal seawater in Ostuchi Bay, Japan. This genome sequence could be useful for our understanding of the variation in photosynthesis-related genes among aerobic anoxygenic phototrophs.

Draft Genome Sequences of Putative Aerobic Anoxygenic Phototrophic Bacterial Strains Jannaschia sp. Strains AI_61 and AI_62, Isolated from Seawater around a Coastal Aquaculture Area

Here, we report the draft genome sequences of putative aerobic anoxygenic phototrophic bacterial strains Jannaschia sp. AI_61 and AI_62, isolated from seawater around a coastal aquaculture in Ainan, Ehime, Japan. These genome sequences could be useful for our understanding of the variation of aerobic anoxygenic phototrophs in the genus.

Syntrophic interspecies electron transfer drives carbon fixation and growth by Rhodopseudomonas palustris under dark, anoxic conditions

In natural anoxic environments, anoxygenic photosynthetic bacteria fix CO2 by photoheterotrophy, photoautotrophy, or syntrophic anaerobic photosynthesis. Here, we describe electroautotrophy, a previously unidentified dark CO2 fixation mode enabled by the electrosyntrophic interaction between Geobacter metallireducens and Rhodopseudomonas palustris. After an electrosyntrophic coculture is formed, electrons are transferred either directly or indirectly (via electron shuttles) from G. metallireducens to R. palustris, thereby providing reducing power and energy for the dark CO2 fixation. Transcriptomic analyses demonstrated the high expression of genes encoding for the extracellular electron transfer pathway in G. metallireducens and the Calvin-Benson-Bassham carbon fixation cycle in R. palustris. Given that sediments constitute one of the most ubiquitous and abundant niches on Earth and that, at depth, most of the sedimentary niche is both anoxic and dark, dark carbon fixation provides a metabolic window for the survival of anoxygenic phototrophs, as well as an as-yet unappreciated contribution to the global carbon cycle.

Discovery of Siderophore and Metallophore Production in the Aerobic Anoxygenic Phototrophs

Aerobic anoxygenic phototrophs have been isolated from a rich variety of environments including marine ecosystems, freshwater and meromictic lakes, hypersaline springs, and biological soil crusts, all in the hopes of understanding their ecological niche. Over 100 isolates were chosen for this study, representing 44 species from 27 genera. Interactions with Fe3+ and other metal(loid) cations such as Mg2+, V3+, Mn2+, Co2+, Ni2+, Cu2+, Zn2+, Se4+ and Te2+ were tested using a chromeazurol S assay to detect siderophore or metallophore production, respectively. Representatives from 20 species in 14 genera of α-Proteobacteria, or 30% of strains, produced highly diffusible siderophores that could bind one or more metal(loid)s, with activity strength as follows: Fe > Zn > V > Te > Cu > Mn > Mg > Se > Ni > Co. In addition, γ-proteobacterial Chromocurvus halotolerans, strain EG19 excreted a brown compound into growth medium, which was purified and confirmed to act as a siderophore. It had an approximate size of ~341 Da and drew similarities to the siderophore rhodotorulic acid, a member of the hydroxamate group, previously found only among yeasts. This study is the first to discover siderophore production to be widespread among the aerobic anoxygenic phototrophs, which may be another key method of metal(loid) chelation and potential detoxification within their environments.

Diversity of Aerobic Anoxygenic Phototrophs and Rhodopsin-Containing Bacteria in the Surface Microlayer, Water Column and Epilithic Biofilms of Lake Baikal

The diversity of aerobic anoxygenic phototrophs (AAPs) and rhodopsin-containing bacteria in the surface microlayer, water column, and epilithic biofilms of Lake Baikal was studied for the first time, employing pufM and rhodopsin genes, and compared to 16S rRNA diversity. We detected pufM-containing Alphaproteobacteria (orders Rhodobacterales, Rhizobiales, Rhodospirillales, and Sphingomonadales), Betaproteobacteria (order Burkholderiales), Gemmatimonadetes, and Planctomycetes. Rhodobacterales dominated all the studied biotopes. The diversity of rhodopsin-containing bacteria in neuston and plankton of Lake Baikal was comparable to other studied water bodies. Bacteroidetes along with Proteobacteria were the prevailing phyla, and Verrucomicrobia and Planctomycetes were also detected. The number of rhodopsin sequences unclassified to the phylum level was rather high: 29% in the water microbiomes and 22% in the epilithon. Diversity of rhodopsin-containing bacteria in epilithic biofilms was comparable with that in neuston and plankton at the phyla level. Unweighted pair group method with arithmetic mean (UPGMA) and non-metric multidimensional scaling (NMDS) analysis indicated a distinct discrepancy between epilithon and microbial communities of water (including neuston and plankton) in the 16S rRNA, pufM and rhodopsin genes.

The role of anoxygenic phototrophs in the Baltic Sea

<p>Both oxygenic and anoxygenic phototrophic bacteria (OPB and APB, respectively) are widely distributed in the ocean and play significant roles in carbon cycle and marine productivity. These organisms capture light as energy source via chlorophyll or bacteriochlorophylls-based photosystems. While OPB are relatively well studied, information on APB is rather scarce although they have been shown abundant in some ocean ecosystems and may play an important role in oxygen depleted environments. Here, we investigate the spatial profile of OPB and APB, gene abundance and expression of the key functional marker gene <em>pufM </em>(APB specific photosynthetic reaction center subunit M), in one fjord and three basins of the Baltic Sea using 16S rRNA amplicon sequencing and qPCR. Among the microbial community, abundances of OPB and APB were found to be similar thus emphasizing a potential importance of APB, with APB representing 1.6-17.5% and OPB representing 0.5-20%. Among APB, we identified eleven different orders, with <em>Rhodobacterales</em> being quantitatively dominant. The identified seven orders of OPB were dominated by <em>Synechococcales</em>. OPB were more abundant than APB in surface waters (<8m), while APB were comparably more abundant in deeper waters. Besides a depth-dependent distribution, we observed an impact of salinity on the distribution of APB and OPB, both of which being suggestive of distinct niches for those primary producer clades. <em>pufM</em> gene abundance ranged from 10<sup>4</sup> to 10<sup>9</sup> copies/L, with highest counts detectable in the mixed layer (<40m), however, even in deeper waters where gene abundances decreased APB <em>pufM</em> gene expression was high with up to 10<sup>4</sup> copies/L. These results indicate APB may play a more important role in marine primary productivity which has been underestimated before. </p>