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Presentation by F.Pinzari atThe World Congress of Soil Science 2022, which took place in Glasgow from 31st July - 5th August 2022 Abstract: Effects of a fungal invasion on soil bacteria Pinzari F.1,2, Clark M.D.1, Misra R. 3, Chooneea D.3, Xu X.-M.4, Jungblut A.D.1 1Life Sciences Department, Natural History Museum, Cromwell Road, SW7 5BD London, UK 2Institute for Biological Systems, Council of National Research of Italy (CNR), Monterotondo (RM), Italy 3Core Research Laboratories, Molecular Biology, Natural History Museum, London, United Kingdom 4National Institute of Agricultural Botany, East Malling Research Station (EMR), East Malling, UK Fungal bioinoculants have a vast potential in agriculture because they can help increase crop yields and quality and reduce the application of chemicals. Their effectiveness has been widely tested (Malusà et al., 2016). However, little is known about the effect of bioinoculants on microbial assemblages in non-rhizospheric soil. A sudden artificial introduction of a fungal species in soil could theoretically impact the biodiversity of local microbial communities and lead to changes in nutrient availability (van Elsas et al., 2012). We assessed the impact of a competitive fungal inoculum, the globally-used biofertiliser Trichoderma afroharzianum T22, on soil microcosms to understand 1) to what extent the native microbial community richness and relative abundance are influenced by a fungal strain introduced to soil; 2) whether microbial taxa are resilient to the disturbance caused by the fungus; 3) how far the bioinoculant impacts the soil microorganisms functions. We used bacterial 16S rRNA gene amplicon sequencing (Illumina) and a shotgun metagenomic analysis (Oxford Nanopore Sequencing) to analyse the microbial communities in bioreactors after seven weeks of incubation with and without the fungus. The presence of the fungus had a negative impact on the abundance of some groups of bacteria, such as the genus Pseudomonas, and it stimulated the presence of species metabolically linked to the fungus, including chitin degrading Chitinophagaceae. In conclusion, the results suggest that more than an impact on bacteria's overall biodiversity, the fungus has favoured some groups at the expense of others, even creating new food webs and trophic niches. References Malusà E, Pinzari F, Canfora L (2016) Efficacy of Biofertilizers: Challenges to Improve Crop Production. In: D.P. Singh et al. (eds.), Microbial Inoculants in Sustainable Agricultural Productivity: Microbial Inoculants in Sustainable Agricultural Productivity, pp.17-40 Springer India doi.org/10.1007/978-81-322-2644-4_2 van Elsas JD, Chiurazzi M, Mallon CA, Elhottova D, Kristufek V, Salles JF. (2012) Microbial diversity determines the invasion of soil by a bacterial pathogen. Proc Natl Acad Sci U S A. 24;109(4):1159-64. doi: 10.1073/pnas.1109326109. &nbsp

The environmental availability of mineral nutrients plays a role in the adaptation and evolution of soil fungi and fungus-plant symbiotic systems. As nutrient availability is different from soil to soil, it is expected that the most adapted fungi can express multiple genes to promote different mechanisms of mineral attack and of metabolic routes to maximize the use of nutrient resources at the lowest energy cost. Microcosm experiments of Paxillus involutus growing on muscovite, phlogopite and K-exchanged vermiculite as K sources were carried out for 21 days. Gene expression analysis indicated that the level of K-deprivation stress was muscovite > phlogopite >> K-vermiculite. Genes and functions overexpressed in the experiments indicated meaningful metabolic activities including K extraction and transport. SEM-EDS and micro-XRD (∼1.15 mm resolution) analysis of mineral surfaces and chemical analysis of agar and mycelium indicated weathering processes and K uptake in good agreement with gene expression analysis. In many areas, P. involutus caused physical damage to the minerals without any apparent chemical attack, consisting of tracks of different depth, from

Bacterial communities form the basis of biogeochemical processes and determine plant growth and health. Mosses harbour diverse bacterial communities that are involved in nitrogen fixation and carbon cycling. Global climate change is causing changes in aboveground plant biomass and shifting species composition in the Arctic, but little is known about the response of moss microbiomes in these environments. Here, we studied the total and potentially active bacterial communities associated with Racomitrium lanuginosum in response to a 20-yr in situ warming in an Icelandic heathland. We evaluated the effect of warming and warming-induced shrub expansion on the moss bacterial community composition and diversity, and nifH gene abundance. Warming changed both the total and the potentially active bacterial community structure, while litter abundance only affected the total bacterial community structure. The abundance of nifH genes was negatively affected by litter abundance. We also found shifts in the potentially nitrogen-fixing community, with Nostoc decreasing and noncyanobacterial diazotrophs increasing in relative abundance. Our data suggest that the moss microbial community and potentially nitrogen fixing taxa will be sensitive to future warming, partly via changes in litter and shrub abundance.

Cyanobacteria form diverse communities and are important primary producers in Antarctic freshwater environments, but their geographic distribution patterns in Antarctica and globally are still unresolved. There are however few genomes of cultured cyanobacteria from Antarctica available and therefore metagenome-assembled genomes (MAGs) from Antarctic cyanobacteria microbial mats provide an opportunity to explore distribution of uncultured taxa. These MAGs also allow comparison with metagenomes of cyanobacteria enriched communities from a range of habitats, geographic locations, and climates. However, most MAGs do not contain 16S rRNA gene sequences, making a 16S rRNA gene-based biogeography comparison difficult. An alternative technique is to use large-scale k-mer searching to find genomes of interest in public metagenomes.This paper presents the results of k-mer based searches for 5 Antarctic cyanobacteria MAGs from Lakes Fryxell and Lake Vanda, assigned the namesPhormidium pseudopriestleyi, aMicrocoleus, aLeptolyngbya, aPseudanabaena, and aNeosynechococcus(Lumian et al., 2021, Lumian et al., 2022, in prep.) in 498,942 unassembled metagenomes from the National Center for Biotechnology Information (NCBI) Sequence Read Archive (SRA). TheMicrocoleusMAG was found in a wide variety of environments,P. pseudopriestleyiwas found in environments with challenging conditions, theNeosynechococcuswas only found in Antarctica, and theLeptolyngbyaandPseudanabaenaMAGs were found in Antarctic and other cold environments. The findings based on metagenome matches and global comparisons suggest that these Antarctic cyanobacteria have distinct distribution patterns ranging from locally restricted to global distribution across the cold biosphere and other climatic zones.

The undulating ice of the McMurdo Ice Shelf, Southern Victoria Land, supports one of the largest networks of ice-based, multiyear meltwater pond habitats in Antarctica, where microbial mats are abundant and contribute most of the biomass and biodiversity. We used 16S rRNA and 18S rRNA gene high-throughput sequencing to compare variance of the community structure in microbial mats within and between ponds with different salinities and pH. Proteobacteria and Cyanobacteria were the most abundant phyla, and composition at OTU level was highly specific for the meltwater ponds with strong community sorting along the salinity gradient. Our study provides the first detailed evaluation of eukaryote communities for the McMurdo Ice Shelf using the 18S rRNA gene. They were dominated by Ochrophyta, Chlorophyta and Ciliophora, consistent with previous microscopic analyses, but many OTUs belonging to less well-described heterotrophic protists from Antarctic ice shelves were also identified including Amoebozoa, Rhizaria and Labyrinthulea. Comparison of 16S and 18S rRNA gene communities showed that the Eukaryotes had lower richness and greater similarity between ponds in comparison with Bacteria and Archaea communities on the McMurdo Ice shelf. While there was a weak correlation between community dissimilarity and geographic distance, the congruity of microbial assemblages within ponds, especially for Bacteria and Archaea, implies strong habitat filtering in ice shelf meltwater pond ecosystems, especially due to salinity. These findings help to understand processes that are important in sustaining biodiversity and the impact of climate change on ice-based aquatic habitats in Antarctica.

Clues to the evolutionary steps producing innovations in oxygenic photosynthesis may be preserved in the genomes of organisms phylogenetically placed between non-photosynthetic Vampirovibrionia (formerly Melainabacteria) and the thylakoid-containing Cyanobacteria. However, only two species with published genomes are known to occupy this phylogenetic space, both within the genus Gloeobacter. Here, we describe nearly complete, metagenome-assembled genomes (MAGs) of an uncultured organism phylogenetically placed near Gloeobacter, for which we propose the name Candidatus Aurora vandensis {Au’ro.ra. L. fem. n. aurora, the goddess of the dawn in Roman mythology; van.de’nsis. N.L. fem. adj. vandensis of Lake Vanda, Antarctica}. The MAG of A. vandensis contains homologs of most genes necessary for oxygenic photosynthesis including key reaction center proteins. Many accessory subunits associated with the photosystems in other species either are missing from the MAG or are poorly conserved. The MAG also lacks homologs of genes associated with the pigments phycocyanoerethrin, phycoeretherin and several structural parts of the phycobilisome. Additional characterization of this organism is expected to inform models of the evolution of oxygenic photosynthesis.

Photoautotrophic organisms face extreme conditions in the Polar Regions including permanently low temperatures, freezing, salinity and low nutrient. Certain microalgae and cyanobacteria are able to withstand these conditions and adaptation mechanisms associated with photophysiology play an important part in overcoming challenges created by variation in irradiance under low temperatures.

Large skeleton specimens are often featured as iconic open displays in Natural History Museums, for example, the blue whale 'Hope' at the Natural History Museum, London. A study on Hope's bone surface was performed to assess the biodeterioration potential of fungi. Fungi were isolated, and a fungal internal transcribed spacer (ITS) clone library survey was performed on dust and bone material. Mineral particles derived from bone and dust were analysed using energy dispersive X-ray spectroscopy, variable pressure scanning electron microscopy (SEM) and high vacuum SEM. Results showed that bone material, although mainly mineral in nature, and therefore less susceptible than organic materials to biodeterioration phenomena in the indoor environments, offers niches for specialized fungi and is affected by unusual and yet not so well-documented mechanisms of alteration. Areas of bone surface were covered with a dense biofilm mostly composed of fungal hyphae, which produced tunnelling and extensive deposition of calcium and iron-containing secondary minerals. Airborne halophilic and xerophilic fungi including taxa grouping into Ascomycota and Basidiomycota, capable of displacing salts and overcome little water availability, were found to dominate the microbiome of the bone surface.

Cryoconite holes, supraglacial depressions containing water and microbe-mineral aggregates, are known to be hotspots of microbial diversity on glacial surfaces. Cryoconite holes form in a variety of locations and conditions, which impacts both their structure and the community that inhabits them. Using high-throughput 16S and 18S rRNA gene sequencing, we have investigated the communities of a wide range of cryoconite holes from 15 locations across the Arctic and Antarctic. Around 24 bacterial and 11 eukaryotic first-rank phyla were observed in total. The various biotic niches (grazer, predator, photoautotroph, and chemotroph), are filled in every location. Significantly, there is a clear divide between the bacterial and microalgal communities of the Arctic and that of the Antarctic. We were able to determine the groups contributing to this difference and the family and genus level. Both polar regions contain a “core group” of bacteria that are present in the majority of cryoconite holes and each contribute >1% of total amplicon sequence variant (ASV) abundance. Whilst both groups contain Microbacteriaceae, the remaining members are specific to the core group of each polar region. Additionally, the microalgal communities of Arctic cryoconite holes are dominated by Chlamydomonas whereas the Antarctic cryoconite holes are dominated by Pleurastrum. Therefore cryoconite holes may be a global feature of glacier landscapes, but they are inhabited by regionally distinct microbial communities. Our results are consistent with the notion that cryoconite microbiomes are adapted to differing conditions within the cryosphere.

Sulfide inhibits oxygenic photosynthesis by blocking electron transfer between H2O and the oxygen-evolving complex in the D1 protein of Photosystem II. The ability of cyanobacteria to counter this effect has implications for understanding the productivity of benthic microbial mats in sulfidic environments throughout Earth history. In Lake Fryxell, Antarctica, the benthic, filamentous cyanobacterium Phormidium pseudopriestleyi creates a 1–2 mm thick layer of 50 µmol L−1 O2 in otherwise sulfidic water, demonstrating that it sustains oxygenic photosynthesis in the presence of sulfide. A metagenome-assembled genome of P. pseudopriestleyi indicates a genetic capacity for oxygenic photosynthesis, including multiple copies of psbA (encoding the D1 protein of Photosystem II), and anoxygenic photosynthesis with a copy of sqr (encoding the sulfide quinone reductase protein that oxidizes sulfide). The genomic content of P. pseudopriestleyi is consistent with sulfide tolerance mechanisms including increasing psbA expression or directly oxidizing sulfide with sulfide quinone reductase. However, the ability of the organism to reduce Photosystem I via sulfide quinone reductase while Photosystem II is sulfide-inhibited, thereby performing anoxygenic photosynthesis in the presence of sulfide, has yet to be demonstrated.

Silicates make up about 90% of the Earth’s crust and constitute the main source of mineral nutrients for microorganisms and plants. Fungi can actively weather silicates to extract nutrients. However, it is unclear whether they are able to obtain the same amounts of nutrients and use the same mechanisms when tapping into different mineral sources. We performed a microcosm experiment using the ectomycorrhizal basidiomycetes Paxillus involutus and the silicates K-vermiculite, muscovite and phlogopite as only potassium sources, as they show a different resistance for the removal of K cations from the mineral structure. A combination of transcriptomic, elemental and SEM analyses showed that different minerals stimulated specific weathering mechanisms and led to a change in fungal genes expression. The differential expression of the fungal genes generated alternative chemical attacks on the minerals, resulting in a tailored dissolution and selective uptake of chemical elements according to the leachability of K from the silicate mineral. The K uptake capacity of the fungus was highest with vermiculite in comparison to growth on phlogopite and muscovite. The findings provide new insights into fungal-mineral interactions that will help to interpret key processes for the homeostasis of soil environments.

Sulfide inhibits oxygenic photosynthesis by blocking electron transfer between H2O and the oxygen-evolving complex in the D1 protein of Photosystem II. The ability of cyanobacteria to counter this effect has implications for understanding the productivity of benthic microbial mats in sulfidic environments throughout Earth history. In Lake Fryxell, Antarctica, the benthic, filamentous cyanobacterium Phormidium pseudopriestleyi creates a 1–2 mm thick layer of 50 µmol L−1 O2 in otherwise sulfidic water, demonstrating that it sustains oxygenic photosynthesis in the presence of sulfide. A metagenome-assembled genome of P. pseudopriestleyi indicates a genetic capacity for oxygenic photosynthesis, including multiple copies of psbA (encoding the D1 protein of Photosystem II), and anoxygenic photosynthesis with a copy of sqr (encoding the sulfide quinone reductase protein that oxidizes sulfide). The genomic content of P. pseudopriestleyi is consistent with sulfide tolerance mechanisms including increasing psbA expression or directly oxidizing sulfide with sulfide quinone reductase. However, the ability of the organism to reduce Photosystem I via sulfide quinone reductase while Photosystem II is sulfide-inhibited, thereby performing anoxygenic photosynthesis in the presence of sulfide, has yet to be demonstrated.

Microbial inoculants can be an efficient tool to manage the soil and plant microbiomes providing direct beneficial effects, and for modulating native soil and plant-associated microbiota. However, the application of soil microbial inoculants as biofertilizers and biopesticides in agriculture is still limited by factors related to their formulation, application method, and the knowledge about the impact and interactions between microbial inoculants and native soil and plant host microbiomes. The review is thus describing and discussing three major aspects related to microbial-based product exploitation, namely: i) the discovery and screening of beneficial microbial strains; ii) the opportunities and challenges associated with strain multifunctional features; iii) the fermentation and formulation strategies also based on the use of wastes as growth substrates and the technical and regulatory challenges faced in their path to field application. All these issues are addressed in activities performed by the EXCALIBUR project (www.excaliburproject.eu), which aims to expand the current concept about microbiomes interactions, acknowledging their interactive network that can impact agricultural practices as well as on all living organisms within an ecosystem.

Freshwater ecosystems are considered hotspots of biodiversity in Antarctic polar deserts. Anticipated warming is expected to change the hydrology of these systems due to increased meltwater and reduction of ice cover, with implications for environmental conditions and physical connectivity between habitats. Using 16S rRNA gene sequencing, we evaluated microbial mat and planktonic communities within a connected freshwater system in the McMurdo Wright Valley, Antarctica, to determine the roles of connectivity and habitat conditions in controlling microbial assemblage composition. We examined communities from glacial Lake Brownworth, the perennially ice-covered Lake Vanda and the Onyx River, which connects the two. In Lake Vanda, we found distinct microbial assemblages occupying sub-habitats at different lake depths, while the communities from Lake Brownworth and Onyx River were structurally similar. Despite the higher physical connectivity and dispersal opportunities between bacterial communities in the shallow parts of the system, environmental abiotic conditions dominated over dispersal in driving community structure. Functional metabolic pathway predictions suggested differences in the functional gene potential between the microbial mat communities located in shallower and deeper water depths. The findings suggest that increasing temperatures and meltwater due to future climate change will affect bacterial diversity and functioning in Antarctic freshwater ecosystems.

Antarctic perennially ice-covered lakes provide a stable low-disturbance environment where complex microbially mediated structures can grow. Lake Untersee, an ultra-oligotrophic lake in East Antarctica, has the lake floor covered in benthic microbial mat communities, where laminated organo-sedimentary structures form with three distinct, sympatric morphologies: small, elongated cuspate pinnacles, large complex cones and flat mats. We examined the diversity of prokaryotes and eukaryotes in pinnacles, cones and flat microbial mats using high-throughput sequencing of 16S and 18S rRNA genes and assessed how microbial composition may underpin the formation of these distinct macroscopic mat morphologies under the same environmental conditions. Our analysis identified distinct clustering of microbial communities according to mat morphology. The prokaryotic communities were dominated by Cyanobacteria, Proteobacteria, Verrucomicrobia, Planctomycetes, and Actinobacteria. While filamentous Tychonema cyanobacteria were common in all mat types, Leptolyngbya showed an increased relative abundance in the pinnacle structures only. Our study provides the first report of the eukaryotic community structure of Lake Untersee benthic mats, which was dominated by Ciliophora, Chlorophyta, Fungi, Cercozoa, and Discicristata. The eukaryote richness was lower than for prokaryote assemblages and no distinct clustering was observed between mat morphologies. These findings suggest that cyanobacterial assemblages and potentially other bacteria and eukaryotes may influence structure morphogenesis, allowing distinct structures to form across a small spatial scale.

The evolution of oxygenic photosynthesis was one of the most transformative evolutionary events in Earth’s history, leading eventually to the oxygenation of Earth’s atmosphere and, consequently, the evolution of aerobic respiration. Previous work has shown that the terminal electron acceptors (complex IV) of aerobic respiration likely evolved after the evolution of oxygenic photosynthesis. However, complex I of the respiratory complex chain can be involved in anaerobic processes and, therefore, may have pre-dated the evolution of oxygenic photosynthesis. If so, aerobic respiration may have built upon respiratory chains that pre-date the rise of oxygen in Earth’s atmosphere. The Melainabacteria provide a unique opportunity to examine this hypothesis because they contain genes for aerobic respiration but likely diverged from the Cyanobacteria before the evolution of oxygenic photosynthesis. Here, we examine the phylogenies of translated complex I sequences from 44 recently published Melainabacteria metagenome assembled genomes and genomes from other Melainabacteria, Cyanobacteria, and other bacterial groups to examine the evolutionary history of complex I. We find that complex I appears to have been present in the common ancestor of Melainabacteria and Cyanobacteria, supporting the idea that aerobic respiration built upon respiratory chains that pre-date the evolution of oxygenic photosynthesis and the rise of oxygen.

Freshwater ecosystems are considered hotspots of biodiversity in Antarctic polar deserts. Anticipated warming is expected to change the hydrology of these systems due to increased meltwater and reduction of ice cover, with implications for environmental conditions and physical connectivity between habitats. Using 16S rRNA sequencing, we evaluated the structure of microbial mat and planktonic communities within a connected watershed in the McMurdo Wright Valley, Antarctica to determine the roles of connectivity and habitat conditions in controlling microbial assemblage composition. We examined benthic and planktonic samples from glacial Lake Brownworth, the perennially ice-covered Lake Vanda, and the Onyx River, which connects the two. In Lake Vanda, we found distinct microbial assemblages occupying sub-habitats at different lake depths, while the communities from Lake Brownworth and Onyx River were structurally similar between them. Despite the higher connectivity between bacterial communities in the shallow parts of the system, environmental filtering dominated over dispersal in driving bacterial community structure. Functional metagenomics predictions identified genes related to degradation of halogenated aromatic compounds in surface microbial mats exposed to changes in water regimes, which progressively disappeared with increasing depth. Shifting environmental conditions due to increasing connectivity, rather than dispersal, may become the dominant drivers of bacterial diversity and functioning in Antarctic freshwater ecosystems.

Nostoc (Nostocales, Cyanobacteria) has a global distribution in the Polar Regions. However, the genomic diversity of Nostoc is little known and there are no genomes available for polar Nostoc. Here we carried out the first genomic analysis of the Nostoc commune morphotype with a recent sample from the High Arctic and a herbarium specimen collected during the British Arctic Expedition (1875-76). Comparisons of the polar genomes with 26 present-day non-polar members of the Nostocales family highlighted that there are pronounced genetic variations among Nostoc strains and species. Osmoprotection and other stress genes were found in all Nostoc strains, but the two Arctic strains had markedly higher numbers of biosynthetic gene clusters for uncharacterised non-ribosomal peptide synthetases, suggesting a high diversity of secondary metabolites. Since viral-host interactions contribute to microbial diversity, we analysed the CRISPR-Cas systems in the Arctic and two temperate Nostoc species. There were a large number of unique repeat-spacer arrays in each genome, indicating diverse histories of viral attack. All Nostoc strains had a subtype I-D system, but the polar specimens also showed evidence of a subtype I-B system that has not been previously reported in cyanobacteria, suggesting diverse cyanobacteria-virus interactions in the Arctic.

Microcystins (MCN), β-N-methylamino-L-alanine (BMAA) and anatoxin-a were investigated in Antarctic cyanobacterial mats collected from Ross Island and the McMurdo Ice Shelf, East Antarctica during C...

Calcite-rich columnar stromatolites grew in perennially ice-covered Lake Joyce in the McMurdo Dry Valleys, Antarctica, during a period of environmental change associated with rising lake level. Stromatolite calcite contains carbon and oxygen isotope records of changes to microbial activity in response to variable light environments and water chemistry through time. The stromatolites grew synchronously with correlative calcite zones. The innermost (oldest) calcite zone has a wide range of δ13Ccalcite values consistent with variable photosynthetic effects on local DIC 13C/12C. Subsequent calcite zones preserve a progressive enrichment in δ13Ccalcite values of approximately + 2.6‰ through time, with δ13Ccalcite values becoming less variable. This enrichment likely records the removal of 12C by photosynthesis from the DIC reservoir over decades, with photosynthetic effects decreasing as light levels became lower and more consistent through time. Mean δ18Ocalcite values of the innermost calcified zone were at least 1‰ lower than those of the other calcified zones (t test p-level

Cyanobacteria-dominated microbial mats in Lake Vanda grow with pinnacles and ridges separated by prostrate mat. Rocks protrude over microbial mats on the lake bottom to create localized, dm-scale gradients in sedimentation and irradiance. The effects of sedimentation on pinnacle and ridge growth were isolated from photosynthetic activity by contrasting growth across microenvironmental gradients. Sedimentation rate was measured as the mass of sand and mud sized sediment in mat that accumulated over 11 years, and the incident light was modeled near and under rocks by reconstructing topography using Structure from Motion techniques. Morphologically diverse pinnacles and ridges were documented in both exposed and sheltered mat microenvironments, in addition to growing downward from the underside of overhanging rocks. Mat that grew with > 40% irradiance under overhangs did not have consistent differences in pinnacle density or ridge abundance as a function of sedimentation rates or irradiance when compared to exposed mat. However, their morphology did change significantly with changes in the direction of incident irradiance. Where irradiance was

Bryophytes from submerged habitats are particularly difficult to identify because developmental plasticity obscures their characteristic features. A deep-water moss population of uncertain identity was found isolated at a depth of 31 m within the perennially ice-covered Lake Vanda, Wright Valley, Antarctica. Through phylogenetic analysis of the chloroplast region, ribosomal subunit 4 gene, nuclear ribosomal DNA region, and the internal transcribed spacer region, the Lake Vanda moss was identified as the cosmopolitan species Bryum pseudotriquetrum and resolved to a clade containing exclusively specimens from Antarctica, specifically those from the neighbouring Taylor Valley and Granite Harbour. The close genetic similarity of the Lake Vanda population to other populations of B. pseudotriquetrum in Southern Victoria Land suggests that colonisation was likely to have been from local sources, and colonisation likely occurred at least 80–100 years ago, given the position of mosses in the deeper of two convection cells in the lake. Light and scanning electron microscopy of in vitro cultured specimens revealed adaptations to permanent submersion, including very thin cell walls, which may increase CO2 absorption under water. The production of rhizoidal knots in contaminated, low-nutrient media, but not in axenic cultures, might result from interactions between the moss and organisms in the microbial mat from which it was isolated. The absence of mosses around the lake margin or elsewhere in Wright Valley highlights the importance of freshwater ecosystems as refugia for biodiversity in Antarctica.

Microbial mat growth can produce diagnostic delicate structures in low-energy environments, but mat morphology, and thus preservation of delicate structures as morphological biosignatures, is affected by mud accumulation. In Lake Joyce, Antarctica, varying rates of siliciclastic mud deposition affected the morphology of partially calcified microbial mats. These mats were characterized along a shore-parallel transect from higher- to lower-sedimentation regions of the lake. Mats with high mud accumulation rates were predominantly flat with sparse pinnacles. At lower sedimentation rates, mats developed more pinnacles and more delicate features, including vertical webs of biofilm suspended between pinnacles. Morphological transitions suggest that mud deposition disrupts growth of delicate microbial structures like biofilm webs and inhibits pinnacle growth at higher accumulation rates. The simplification of mat morphology with increasing mud accumulation has important implications for the distribution of stromatolite types in the rock record. Stromatolites with intricate biofilm structures are rare, possibly due to the paucity of appropriate environments rather than the intrinsic properties of microbial communities. Communities capable of forming delicate microbialites likely produced morphologically simpler pinnacles and flat mat in environments with significant siliciclastic mud or micrite influx. Thus, delicate mats have narrow environmental ranges for growth, and temporal or spatial variations in siliciclastic mud or micrite sedimentation rates bias the record of microbial activity preserved in stromatolites.

Ecological communities are regulated by the flow of energy through environments. Energy flow is typically limited by access to photosynthetically active radiation (PAR) and oxygen concentration (O2). The microbial mats growing on the bottom of Lake Fryxell, Antarctica, have well-defined environmental gradients in PAR and (O2). We analyzed the metagenomes of layers from these microbial mats to test the extent to which access to oxygen and light controls community structure. We found variation in the diversity and relative abundances of Archaea, Bacteria and Eukaryotes across three (O2) and PAR conditions: high (O2) and maximum PAR, variable (O2) with lower maximum PAR, and low (O2) and maximum PAR. We found distinct communities structured by the optimization of energy use on a millimeter-scale across these conditions. In mat layers where (O2) was saturated, PAR structured the community. In contrast, (O2) positively correlated with diversity and affected the distribution of dominant populations across the three habitats, suggesting that meter-scale diversity is structured by energy availability. Microbial communities changed across covarying gradients of PAR and (O2). The comprehensive metagenomic analysis suggests that the benthic microbial communities in Lake Fryxell are structured by energy flow across both meter- and millimeter-scales., Energy input affects the composition and diversity of microbial mats growing at the bottom of an Antarctic lake.

Ecological theories posit that heterogeneity in environmental conditions greatly affects community structure and function. However, the degree to which ecological theory developed using plant- and animal-dominated systems applies to microbiomes is unclear. Investigating the metabolic strategies found in microbiomes are particularly informative for testing the universality of ecological theories because microorganisms have far wider metabolic capacity than plants and animals. We used metagenomic analyses to explore the relationships between the energy and physicochemical gradients in Lake Fryxell and the metabolic capacity of its benthic microbiome. Statistical analysis of the relative abundance of metabolic marker genes and gene family diversity shows that oxygenic photosynthesis, carbon fixation, and flavin-based electron bifurcation differentiate mats growing in different environmental conditions. The pattern of gene family diversity points to the likely importance of temporal environmental heterogeneity in addition to resource gradients. Overall, we found that the environmental heterogeneity of photosynthetically active radiation (PAR) and oxygen concentration ([O2]) in Lake Fryxell provide the framework by which metabolic diversity and composition of the community is structured, in accordance with its phylogenetic structure. The organization of the resulting microbial ecosystems are consistent with the maximum power principle and the species sorting model.

Bacterial communities form the basis of biogeochemical processes and determine plant growth and health. Mosses, an abundant plant group in Arctic ecosystems, harbour diverse bacterial communities that are involved in nitrogen fixation and carbon cycling. Global climate change is causing changes in aboveground plant biomass and shifting species composition in the Arctic, but little is known about the response of moss microbiomes.Here, we studied the total and potentially active bacterial community associated with Racomitrium lanuginosum, in response to 20-year in situ warming in an Icelandic heathland. We evaluated the effect of warming and warming-induced shrub expansion on the moss bacterial community composition and diversity, nifH gene abundance and nitrogen-fixation rates.Warming changed both the total and the potentially active bacterial community structure, while litter abundance only affected the total bacterial community structure. The relative abundance of Proteobacteria increased, while the relative abundance of Cyanobacteria and Acidobacteria decreased. NifH gene abundance and nitrogen-fixation rates were negatively affected by litter and Betula nana abundance, respectively. We also found shifts in the potentially nitrogen-fixing community, with Nostoc decreasing and non-cyanobacterial diazotrophs increasing in relative abundance. Our data suggests that the moss microbial community including the potentially nitrogen-fixing taxa is sensitive to future warming.Synthesis. Long-term warming led to a shift in moss-associated bacterial community composition, while the abundance of nitrogen-fixing bacteria and nitrogen-fixation rates were negatively affected by increased litter and Betula nana abundance respectively. Warming and increased shrub abundance as a result of warming can affect moss-associated bacterial communities and nitrogen fixation rates in tundra ecosystems.

A multi-disciplinary investigation was undertaken to determine the cause of corrosion of iron nails from a skeletal structure in a museum setting. Corroded iron nails were removed from a large whale skeleton, that had been assembled in the 1930s and exhibited for over 80 years at the Natural History Museum in London, United Kingdom. Oxford Nanopore Technologies sequencing, SEM-EDS imaging and chemical mapping were used to document the fungal and bacterial communities associated with the corrosion of the iron nails. A mechanism for indoor fungal-mediated corrosion (biodeterioration) and biomineralisation of iron is proposed. The mechanism includes the establishment of a biofilm dominated by fungi, a corrosive electrolyte produced by fungal activities, transport of ions and biomineralisation of iron-rich minerals.

Cyanobacteria-based microbial mats are common in Antarctic terrestrial freshwater ecosystems such as the extensive wetland seepages that cover Byers Peninsula on Livingston Island (South Shetland Islands), maritime Antarctica, where they play an important role in biomass generation and productivity. Although cyanobacteria, microfauna and fungal communities have been described for such microbial mats, to date, little is known about trophic interactions within the mats, which are likely important to overcome nutrient constraints in oligotrophic polar freshwater ecosystems. We therefore carried out a biomass assessment of the different taxonomic components and their trophic interactions using DNA analysis and stable isotope analysis, as well as physiological activities such as primary and secondary production and nitrogen uptake within the mat food web throughout an austral spring and summer season. Our results suggested, based on a Bayesian mixing model, that carbon flow from cyanobacteria to upper trophic levels was limited to tardigrades and rotifers, whereas fungal and bacterial activity were likely the main connectors between consumers and producers via a heterotrophic loop. This suggests that homeostatic state displayed in freshwater microbial mats from maritime Antarctica provides stability to the microbial mats under the fluctuating environmental conditions commonly found in permanently cold shallow terrestrial aquatic ecosystems in Antarctica.

Microbial pinnacles in ice-covered Lake Vanda, McMurdo Dry Valleys, Antarctica, extend from the base of the ice to more than 50 m water depth. The distribution of microbial communities, their photosynthetic potential, and pinnacle morphology affects the local accumulation of biomass, which in turn shapes pinnacle morphology. This feedback, plus environmental stability, promotes the growth of elaborate microbial structures. In Lake Vanda, all mats sampled from greater than 10 m water depth contained pinnacles with a gradation in size from

Microbial mats growing under the permanent ice cover of Antarctic lakes occupy an exceptionally low-disturbance regime. Constant temperature, the absence of bioturbation or physical disturbance from wind action or ice formation allow mats to accumulate, as annual growth layers, over many decades or even centuries. In so doing they often assume decimetre scale, three-dimensional morphologies such as elaborate pinnacle structures and conical mounds. Here we combine existing and new information to describe microbial structures in three Antarctic lakes—simple prostrate mats in Lake Hoare, emergent cones in Lake Untersee and elaborate pinnacles in Lake Vanda. We attempt to determine whether structures emerge simply from uncoordinated organism-environment interactions or whether they represent an example of “emergent complexity”, within which some degree of self-organisation occurs to confer a holistic functional advantage to component organisms. While some holistic advantages were evident from the structures—the increase in surface area allows greater biomass and overall productivity and nutrient exchange with overlying water—the structures could also be understood in terms of potential interactions between individuals, their orientation and their environment. The data lack strong evidence of coordinated behaviour directed towards holistic advantages to the structure, though hints of coordinated behaviour are present as non-random distributions of structural elements. The great size of microbial structures in Antarctic lakes, and their relatively simple community composition, makes them excellent models for more focused research on microbial cooperation.

Atmospheric oxygen level rose dramatically around 2.4 billion years ago due to oxygenic photosynthesis by the Cyanobacteria. The oxidation of surface environments permanently changed the future of life on Earth, yet the evolutionary processes leading to oxygen production are poorly constrained. Partial records of these evolutionary steps are preserved in the genomes of organisms phylogenetically placed between non-photosynthetic Melainabacteria, crown-group Cyanobacteria, andGloeobacter, representing the earliest-branching Cyanobacteria capable of oxygenic photosynthesis. Here, we describe nearly complete, metagenome assembled genomes of an uncultured organism phylogenetically placed between the Melainabacteria and crown-group Cyanobacteria, for which we propose the name CandidatusAurora vandensis {au.roraLatin noundawnandvand.ensis, originating from Vanda}.The metagenome assembled genome ofA. vandensiscontains homologs of most genes necessary for oxygenic photosynthesis including key reaction center proteins. Many extrinsic proteins associated with the photosystems in other species are, however, missing or poorly conserved. The assembled genome also lacks homologs of genes associated with the pigments phycocyanoerethrin, phycoeretherin and several structural parts of the phycobilisome. Based on the content of the genome, we propose an evolutionary model for increasing efficiency of oxygenic photosynthesis through the evolution of extrinsic proteins to stabilize photosystem II and I reaction centers and improve photon capture. This model suggests that the evolution of oxygenic photosynthesis may have significantly preceded oxidation of Earth’s atmosphere due to low net oxygen production by early Cyanobacteria.

Summary Laminated microbial mats are important ecosystem components of perennially ice-covered Antarctic dry valley lakes. In order to understand better their response to changing environment, we made observations and carried out a manipulation experiment to determine their response to variations in irradiance in Lake Hoare (77°38′ S, 162°53′ E). Ice transparency was the most variable parameter that affected benthic light dose, both spatially and between years. Patterns of lamina accrual corresponded to irradiance history, with laminae that were initiated in high transmission years thicker than those from low transmission years. A shading experiment confirmed that accrual of lamina thickness, calcite precipitation and ash-free dry mass were determined by irradiance, but photosynthetic biomass and phototrophic species composition were less affected. Buried laminae decomposed only slowly over time, with potentially viable phototrophs many laminae down into the microbial mat. Decay rate increased only slightly with shading. We conclude that the microbial mats in Lake Hoare are characterised by remarkable stability, with slow accumulation rates and turnover of biomass over time. Photosynthetic biomass and species composition appeared to be stable across long time periods, with interannual variation in lamination pattern due to differential accumulation of extracellular polysaccharide and representing the visible expression of annual growth conditions.

Perennially ice-covered, meromictic lakes in the McMurdo Dry Valleys, Antarctica, are useful models to study the relationship between cyanobacterial and environmental variables. They have rich benthic cyanobacterial mat accumulations and stable stratification of physical and chemical conditions. Here, we evaluated cyanobacteria from benthic mats from multiple depths in three geographically separated ice-covered lakes, Lakes Vanda, Hoare and Joyce, using 16S rRNA gene clone libraries. We identified 19 ribotypes, mostly Oscillatoriales and several Chroococcales, as well as potentially novel cyanobacterial ribotypes. The majority of ribotype diversity was shared between lakes, and only a weak relationship between ribotype community structure and environmental variables was evident. Multivariate analysis of all lake–depth combinations implied that photosynthetically active radiation, dissolved reactive phosphorus and conductivity were potentially important for shaping benthic communities in McMurdo Dry Valley lakes. Cyanobacterial-specific pigment signature analysis by high-performance liquid chromatography showed that the cyanobacterial communities responded to light conditions similarly, irrespective of community composition. The results imply a capability within a suite of cyanobacteria to colonise, adapt and grow across broad environmental ranges and geographic space, and such adaptability may provide a high degree of community resistance and resilience to future climate-driven environmental change in Antarctic terrestrial aquatic ecosystems.

Mineral weathering processes in soils are important controls on soil characteristics and on bio-and geochemical cycling. Elucidation of these processes and their mechanisms is crucial for understanding soil environments and their influence globally. An Umbric Podzol from the Falkland Islands was studied while investigating possible ways to counteract soil degradation and loss. The soil had lost the O, E and Bs horizons through erosion, thus revealing the transitional B/C horizon, which grades into the underlying parent material. Samples were taken from the B/C surface and 5 cm below the surface, then analyzed with X-ray diffraction, scanning electron microscopy with energy-dispersive X-ray spectroscopy, organic C and N analysis, analysis of extractable Fe and Al with the dithionite-citrate-bicarbonate and ammonium oxalate methods, and Fourier-transform infrared analysis. The soil fabric and mineralogy were compatible with derivation from sandstone rock. Clasts of heterogeneous mineral composition as well as loose material from disaggregated clasts were present. The soil had large proportions of quartz and albite, and minor amounts of muscovite, chlorite, plagioclase, feldspar, kaolinite, and non-diffracting Fe oxide (goethite and/or ferrihydrite). The most peculiar characteristic was a large component (~7 wt% of the bulk soil) of an amorphous (non-X-ray diffracting) silicate phase of small particle size (< 1 m), non extractable, with heterogeneous composition. The average composition of this phase is similar to that of the bulk soil and approaches that of Al-Fe-rich smectite. The amorphous phase is not allophane or imogolite by any of the analyses carried out. The amorphous silicate phase is formed partly by the translocation of metals from O, E and Bs horizons and partly by dissolution of the primary minerals of the B/C horizon, both of which precipitated in combination with low water mobility causing rapid saturation of the interstitial water. There are no reports of amorphous silicate phases with these characteristics or abundances from soils or other weathering environments. Thus, our observations indicate the existence of complex, successive weathering steps not yet identified that could be investigated in materials subjected to slow weathering such as the soil described here. Slow weathering of a sandstone-derived Podzol (Falkland Islands) resulting in high content of a non-crystalline silicate