Abstract. There is a growing recognition of the role of particle-attached (PA) and free-living (FL) microorganisms in marine carbon cycle. However, current understanding of PA and FL microbial communities is largely focused on those in the upper photic zone, and relatively fewer studies have focused on microbial communities of the deep ocean. Moreover, archaeal populations receive even less attention. In this study, we determined bacterial and archaeal community structures of both the PA and FL assemblages at different depths, from the surface to the bathypelagic zone along two water column profiles in the South China Sea. Our results suggest that environmental parameters including depth, seawater age, salinity, particulate organic carbon (POC), dissolved organic carbon (DOC), dissolved oxygen (DO) and silicate play a role in structuring these microbial communities. Generally, the PA microbial communities had relatively low abundance and diversity compared with the FL microbial communities at most depths. Further microbial community analysis revealed that PA and FL fractions generally accommodate significantly divergent microbial compositions at each depth. The PA bacterial communities mainly comprise members of Alphaproteobacteria and Gammaproteobacteria, together with some from Planctomycetes and Deltaproteobacteria, while the FL bacterial lineages are also mostly distributed within Alphaproteobacteria and Gammaproteobacteria, along with other abundant members chiefly from Actinobacteria, Cyanobacteria, Bacteroidetes, Marinimicrobia and Deltaproteobacteria. Moreover, there was an obvious shifting in the dominant PA and FL bacterial compositions along the depth profiles from the surface to the bathypelagic deep. By contrast, both PA and FL archaeal communities dominantly consisted of euryarchaeotal Marine Group II (MGII) and thaumarchaeotal Nitrosopumilales, together with variable amounts of Marine Group III (MGIII), Methanosarcinales, Marine Benthic Group A (MBG-A) and Woesearchaeota. However, the pronounced distinction of archaeal community compositions between PA and FL fractions was observed at a finer taxonomic level. A high proportion of overlap of microbial compositions between PA and FL fractions implies that most microorganisms are potentially generalists with PA and FL dual lifestyles for versatile metabolic flexibility. In addition, microbial distribution along the depth profile indicates a potential vertical connectivity between the surface-specific microbial lineages and those in the deep ocean, likely through microbial attachment to sinking particles.

The distribution and succession of microbial communities along the dispersion path of hydrothermal plumes has not been well investigated. In this study, we collected several types of samples from the Longqi hydrothermal field located on the Southwest Indian Ridge, including hydrothermal plumes at different stages of formation, a suite of water column samples across the non-buoyant hydrothermal plumes above this field, and a background seawater column approximately 350 km away from the hydrothermal field. Using CH 4 concentration anomalies, three non-buoyant plume samples between 2,535 and 2,735 m were identified within the water column. Microbial community compositions within these plumes and background seawater samples were examined based on the 16S rRNA genes and revealed significant variations and successions in community composition between different portions of the hydrothermal plumes. Near the vent orifice, representing the initial stage of plume formation, microbial populations were characterized by abundant and diverse putative vent-associated communities including (hyper)thermophiles such as Aquificaceae and Hydrogenothermaceae within the phylum Aquificae, and some epsilonproteobacterial chemolithoautotrophs such as Sulfurovum, Sulfurimonas, and Caminibacter. By contrast, in the rising buoyant plumes and adjacent seawaters, most vent-associated microbial taxa were still present but made only minor contributions to community composition. Some microbial taxa that are common in seawater columns such as alphaproteobacterial Sphingomonadaceae and SAR11 clade, deltaproteobacterial SAR324 clade, and gammaproteobacterial Pseudomonas, together with Sulfurimonas and SUP05 clade, became predominant. Members within the Sulfurimonas and SUP05 clade flourished with considerable abundance in the non-buoyant plumes, although these plumes were mainly composed of alphaproteobacterial Rhodobacteraceae, gammaproteobacterial Alteromonadaceae and Saccharospirillaceae putatively derived from the surrounding ambient seawater. We also analyzed archaeal components in the initial discharge and rising buoyant plume stages, with both primarily consisting of thaumarchaeal Nitrosopumilales

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Microbes in marine sediments play crucial roles in global carbon and nutrient cycling. However, our understanding of microbial diversity and physiology on the ocean floor is limited. Here, we use phylogenomic analyses of thousands of metagenome-assembled genomes (MAGs) from coastal and deep-sea sediments to identify 55 MAGs that are phylogenetically distinct from previously described bacterial phyla. We propose that these MAGs belong to 4 novel bacterial phyla (Blakebacterota, Orphanbacterota, Arandabacterota, and Joyebacterota) and a previously proposed phylum (AABM5-125-24), all of them within the FCB superphylum. Comparison of their rRNA genes with public databases reveals that these phyla are globally distributed in different habitats, including marine, freshwater, and terrestrial environments. Genomic analyses suggest these organisms are capable of mediating key steps in sedimentary biogeochemistry, including anaerobic degradation of polysaccharides and proteins, and respiration of sulfur and nitrogen. Interestingly, these genomes code for an unusually high proportion (~9% on average, up to 20% per genome) of protein families lacking representatives in public databases. Genes encoding hundreds of these protein families colocalize with genes predicted to be involved in sulfur reduction, nitrogen cycling, energy conservation, and degradation of organic compounds. Our findings advance our understanding of bacterial diversity, the ecological roles of these bacteria, and potential links between novel gene families and metabolic processes in the oceans.

Hydrothermal vent sites have been widely studied to investigate novel bacteria and archaea that are adapted to these extreme environments. However, community-level analyses of hydrothermal microbial ecosystems look beyond the presence and activity of particular types of microbes and examine to what extent the entire community of bacteria and archaea is adapted to hydrothermal conditions; these include elevated temperatures, hydrothermally generated carbon sources, and inorganic electron donors and acceptors that are characteristic for hydrothermal environments.