Following the ubiquitous autotrophic ammonia-oxidizing archaea (AOA), heterotrophic representatives of the marine Nitrososphaerota (HMN) form the second most abundant group within this archaeal phylum. However, their eco-evolutionary strategies remain poorly understood. Previous studies have reported a consistent co-occurrence of HMN with marine AOA (MAOA), prompting a detailed investigation into their potential interaction. Through large-scale (meta)genomic and metatranscriptomic analyses, we reveal that HMN possess ultra-streamlined genomes and globally co-occur with marine AOA. The absence of most B vitamin biosynthesis pathways, incomplete citrate cycle and glycolysis, along with the essential requirement for exogenous amino acids, suggest their potential metabolic dependency on AOA. Meanwhile, catalyzed reporter deposition fluorescence in situ hybridization supports a close physical association between HMN and AOA. The nearly synchronous origins of HMN and AOA after oxygen rise, coupled with HMN's dispersive microhabitats (evidenced by dense, shallow subclades) and extensive horizontal gene transfer between these groups, further support their close relationship-although HMN likely acquired heterotrophic capabilities from bacteria. This study reveals a previously unrecognized association between HMN and AOA, implying a tight coupling between autotrophic and heterotrophic processes in deep-sea habitats.

Anaerobic ammonium oxidation (anammox) plays a critical role in nitrogen loss in estuarine and marine environments. However, the mechanisms underlying the formation and maintenance of the anammox bacterial community remain unclear. This study analyzed the anammox bacterial diversity, community structure, and interspecific relationships in three estuaries along the Chinese coastline —the Changjiang Estuary (CJE), the Oujiang Estuary (OJE), and the Jiulong River Estuary (JLE) — as well as the South China Sea (SCS) to elucidate their community assembly mechanisms. The results indicated that the anammox bacterial community exhibited the highest ammonium concentration as well as the Shannon’s diversity index reflecting both species richness and evenness in the JLE. The lowest Shannon index was observed in the SCS. However, the anammox bacterial species richness was greatest in the CJE. Candidatus Scalindua was the predominant anammox bacteria identified in the coastal sediments, especially in the SCS sediments. Candidatus Brocadia and Candidatus Kuenenia were more abundant in the estuarine sediments, particularly in JLE, than in the SCS. Phylogenetic analysis revealed distinct differentiation among Candidatus Scalindua, Candidatus Brocadia, and Candidatus Kuenenia, with the former exhibiting a greater level of diversity. There was significant spatial heterogeneity in the anammox bacteria across the four regions, characterized by distinct distribution patterns for rare species. Low-abundance (rare) bacteria thrived in their native habitats, whereas abundant taxa displayed greater dispersal capabilities. An analysis of the community assembly mechanism suggested that ecological drift predominantly shaped the overall anammox bacterial community in the coastal sediments. Rare species were more susceptible to dispersal limitations and environmental selection. Co-occurrence network analysis identified Candidatus Scalindua as a keystone genus and highlighted that rare species may play a crucial role in maintaining the ecological stability of the anammox bacterial community in coastal sediments.

Nitrogen fixation is a vital new nitrogen source in the oligotrophic ocean. Although our knowledge of the controlling factors of marine nitrogen fixation have increased rapidly, the physical controls, particularly eddies-induced upwelling and light intensity, remain elusive. In this study, conducted in the Subtropical Northwestern Pacific, we measured nitrogen fixation rates (NFR) in two cyclonic eddies (CEs). Our observations in one CE revealed that depth-integrated NFR (INFR) in core stations were significantly higher than in edge stations, indicating that CEs-induced upwelling might enhance nitrogen fixation. However, more intense upwelling in another CE resulted in lower INFR in core stations compared to edge stations. The INFR distributions in CEs were driven by the upwelling intensity, showing a unimodal response, i.e., the maximum INFR appeared at optimal upwelling intensity. This finding reconciles the debate about whether CEs inhibit nitrogen fixation. Additionally, results from light manipulation incubations proved that light intensity is a key driver for the vertically unimodal pattern of NFR, i.e., peaks at the subsurface layer with an optimum light intensity of 20% to 50% of surface PAR. Furthermore, molecular evidence showed that UCYN-A dominated in the upwelling area, while UCYN-B dominated in the non-upwelling area, indicating that CEs-induced physical perturbation regulates the niches of diazotrophs. Taken together, these results suggest that physical dynamics exert profound controls on the spatial heterogeneity of diazotrophic distribution and activity in the Subtropical Northwestern Pacific, providing new insights into the physical drivers of nitrogen fixation on mesoscale hydrodynamics..

The cyclonic eddy uplifts nutrient-rich seawater into the euphotic zone, typically directly enhancing phytoplankton abundance and primary production. However, its impact on heterotrophic prokaryotic production (HPP) remains unclear due to the complex interplay of multiple indirect factors governing this process. Here, we conducted a comprehensive investigation of the distribution of picophytoplankton and heterotrophic prokaryotes, prokaryotic community structure, and HPP within a cyclonic eddy in the western North Pacific subtropical gyre. The results indicated that despite the higher abundance of picophytoplankton accompanied by nutrient upwelling at the eddy core compared to the edge, higher levels of HPP were observed at the eddy edge between 100 and 200 m, consistent with the distribution of the low nucleic acid content (LNA) prokaryotes. The significant positive correlation between HPP and the proportion of LNA group in total heterotrophic prokaryotes suggested a primary contribution from the LNA group over the high nucleic acid content (HNA) group. SAR11, a typical member of the LNA group, may primarily contribute to the elevated HPP observed at the eddy edge. The changes in temperature, nutrients, and light intensity induced by the cyclonic eddy may significantly influence the distribution and activity of HNA and LNA groups, potentially exerting a greater impact on HPP compared to phytoplankton-related factors. These findings contribute to understanding the underlying mechanisms of HPP responses to cyclonic eddies in the oligotrophic open ocean.

Small-molecule compounds that elicit mRNA-selective translation repression have attracted interest due to their potential for expansion of druggable space. However, only a limited number of examples have been reported to date. Here, we show that desmethyl desamino pateamine A (DMDA-PatA) represses translation in an mRNA-selective manner by clamping eIF4A, a DEAD-box RNA-binding protein, onto GNG motifs. By systematically comparing multiple eIF4A inhibitors by ribosome profiling, we found that DMDA-PatA has unique mRNA selectivity for translation repression. Unbiased Bind-n-Seq reveals that DMDA-PatA-targeted eIF4A exhibits a preference for GNG motifs in an ATP-independent manner. This unusual RNA binding sterically hinders scanning by 40S ribosomes. A combination of classical molecular dynamics simulations and quantum chemical calculations, and the subsequent development of an inactive DMDA-PatA derivative reveals that the positive charge of the tertiary amine on the trienyl arm induces G selectivity. Moreover, we identified that DDX3, another DEAD-box protein, is an alternative DMDA-PatA target with the same effects on eIF4A. Our results provide an example of the sequence-selective anchoring of RNA-binding proteins and the mRNA-selective inhibition of protein synthesis by small-molecule compounds. Here the authors report that DMDA-PatA, an anti-tumor compound, functions as a mRNA-selective translational inhibitor. This drug clamps the target eIF4A and DDX3 RNA-binding proteins on the GNG RNA motif, providing steric hindrance for scanning ribosomes.

Plants often generate secondary metabolites as defense mechanisms against parasites. Although some fungi may potentially overcome the barrier presented by antimicrobial compounds, only a limited number of examples and molecular mechanisms of resistance have been reported. Here, we found an Aglaia plant-parasitizing fungus that overcomes the toxicity of rocaglates, which are translation inhibitors synthesized by the plant, through an amino acid substitution in a eukaryotic translation initiation factor (eIF). De novo transcriptome assembly revealed that the fungus belongs to the Ophiocordyceps genus and that its eIF4A, a molecular target of rocaglates, harbors an amino acid substitution critical for rocaglate binding. Ribosome profiling harnessing a cucumber-infecting fungus, Colletotrichum orbiculare , demonstrated that the translational inhibitory effects of rocaglates were largely attenuated by the mutation found in the Aglaia parasite. The engineered C. orbiculare showed a survival advantage on cucumber plants with rocaglates. Our study exemplifies a plant–fungus tug-of-war centered on secondary metabolites produced by host plants.

Nitrite, an intermediate product of the oxidation of ammonia to nitrate (nitrification), accumulates in upper oceans, forming the primary nitrite maximum (PNM). Nitrite concentrations in the PNM are relatively low in the western North Pacific subtropical gyre (wNPSG), where eddies are frequent and intense. To explain these low nitrite concentrations, we investigated nitrification in cyclonic eddies in the wNPSG. We detected relatively low half-saturation constants (i.e., high substrate affinities) for ammonia and nitrite oxidation at 150 to 200 meter water depth. Eddy-induced displacement of high-affinity nitrifiers and increased substrate supply enhanced ammonia and nitrite oxidation, depleting ambient substrate concentrations in the euphotic zone. Nitrite oxidation is more strongly enhanced by the cyclonic eddies than ammonia oxidation, reducing concentrations and accelerating the turnover of nitrite in the PNM. These findings demonstrate a spatial decoupling of the two steps of nitrification in response to mesoscale processes and provide insights into physical-ecological controls on the PNM.

Dinitrogen (N2) fixation provides nitrogen sources supporting active chemoautotrophic processes in submarine hydrothermal systems. Here, the diazotrophic phylogenetic diversity and N2 fixation profile were investigated in depth along a sharp biogeochemical gradient from the hydrothermal vents to ambient seawater in a shallow submarine hydrothermal system (SHS) off Kueishantao Islet, Taiwan. Diazotrophic community compositions in the SHS covered all four known nifH clusters (I, II, III, and IV) and contained both characteristics of diazotrophic communities in deep-sea hydrothermal systems and terrestrial geothermal systems. The seawater around the vents was primarily dominated by nifH phylotypes affiliated with chemolithotrophic sulfur-oxidizing and iron-reducing bacteria. In contrast, in the seawater directly above the hydrothermal vents, the nifH community markedly contained diverse heterotrophic bacteria. The analysis, which focused on amino acid composition, protein spatial structure, and the possible co-occurrence of alternative nitrogenases in the presence of molybdenum-nitrogenase, further revealed that diazotrophic nitrogenase found around the hydrothermal vents exhibits potential molecular adaptations to high temperature, low pH, and sulfur-rich conditions. The relatively low N2 fixation rates (NFRs) were observed around the vents, where high concentrations of inorganic nitrogen compounds were present. However, notably, high NFRs were detected in seawater directly above the hydrothermal vents, which were 2–5 times higher than those at the reference site. The abundant iron, along with the depletion of nitrogen nutrients potentially resulting from nitrogen assimilation and loss processes, may have led to a high iron:nitrogen ratio, inducing the observed high NFRs. The co-occurrence of diverse nifH-containing heterotrophic bacteria and high NFRs suggests that these heterotrophs may play a major role in N2 fixation in this environment. This study suggests that the SHSs may be potential hotspots for marine N2 fixation, contributing significantly to the global ocean's N2 fixation budget and warranting further investigation.

Active hydrothermal vents create extreme conditions characterized by high temperatures, low pH levels, and elevated concentrations of heavy metals and other trace elements. These conditions support unique ecosystems where chemolithoautotrophs serve as primary producers. The steep temperature and pH gradients from the vent mouth to its periphery provide a wide range of microhabitats for these specialized microorganisms. However, their metabolic functions, adaptations in response to these gradients, and coping mechanisms under extreme conditions remain areas of limited knowledge. In this study, we conducted temperature gradient incubations of hydrothermal fluids from moderate (pH = 5.6) and extremely (pH = 2.2) acidic vents. Combining the DNA-stable isotope probing technique and subsequent metagenomics, we identified active chemolithoautotrophs under different temperature and pH conditions and analyzed their specific metabolic mechanisms. We found that the carbon fixation activities of Nautiliales in vent fluids were significantly increased from 45 to 65 °C under moderately acidic condition, while their heat tolerance was reduced under extremely acidic conditions. In contrast, Campylobacterales actively fixed carbon under both moderately and extremely acidic conditions under 30 − 45 °C. Compared to Campylobacterales, Nautiliales were found to lack the Sox sulfur oxidation system and instead use NAD(H)-linked glutamate dehydrogenase to boost the reverse tricarboxylic acid (rTCA) cycle. Additionally, they exhibit a high genetic potential for high activity of cytochrome bd ubiquinol oxidase in oxygen respiration and hydrogen oxidation at high temperatures. In terms of high-temperature adaption, the rgy gene plays a critical role in Nautiliales by maintaining DNA stability at high temperature. Genes encoding proteins involved in proton export, including the membrane arm subunits of proton-pumping NADH: ubiquinone oxidoreductase, K+ accumulation, selective transport of charged molecules, permease regulation, and formation of the permeability barrier of bacterial outer membranes, play essential roles in enabling Campylobacterales to adapt to extremely acidic conditions. Our study provides in-depth insights into how high temperature and low pH impact the metabolic processes of energy and main elements in chemolithoautotrophs living in hydrothermal ecosystems, as well as the mechanisms they use to adapt to the extreme hydrothermal conditions.

The heavily human-perturbed coastal oceans are hotspots of nitrous oxide (N 2 O) emission to the atmosphere. The processes underpinning the N 2 O flux, however, remain poorly understood, leading to large uncertainties in assessing global N 2 O budgets. Using a suite of nitrogen isotope labeling experiments, we show that multiple processes contribute to N 2 O production throughout the estuarine-coastal gradient, sustaining intensive N 2 O flux to the atmosphere. Unexpectedly, denitrification, rather than ammonia oxidation as previously assumed, constitutes the major source of N 2 O in well-oxygenated coastal waters. Size-fractionated manipulation experiments with gene analysis further reveal niche partitioning of ammonia oxidizers and denitrifiers across the particle size spectrum; denitrification dominated on large particles and ammonia oxidizers on small particles. Total N 2 O production rate increases with substrate and particle concentrations, suggesting a crucial interplay between nutrients and particles in controlling N 2 O production. The controlling factors identified here may help understand climate feedback mechanisms between human activity and coastal oceans.

A long‐standing enigma in oceanography is why terrestrial organic matter is “missing” in the global ocean, despite the considerable discharge into it every year. Although some explanations, such as mineralogical composition, hydrodynamic processes, and priming effect, have been proposed, we hypothesize that the essential mechanism behind the missing organic matter is microbial processing, for which the underlying coupled geochemical, molecular, and genetic evidence is unknown. An ultra‐large‐volume, long‐term river–seawater stratified simulation system was constructed to unravel the microbially driven fate of terrigenous particulate organic matter (POM) in oceans. Analysis of combining the molecular with POM chemical composition data suggests that Bacteroidetes could act as pioneers in the processing of terrigenous POM in oceans, degrading high‐molecular‐weight, high‐carbon compounds such as polysaccharides. Remaining low‐molecular‐weight nitrogenous organic matter is subsequently degraded by Planctomycetes and Proteobacteria. Isotopic signals show that this preferential degradation causes a distinct “aging” effect of POM, and along with nitrification enhanced by remineralization, causes a decrease in the POM C : N ratio. Degradation of terrigenous POM and bacterial biomass biosynthesis leads to positive deviations in δ 15 N and δ 13 C. Relatively refractory hydrocarbons, aromatic compounds, and phenols are accumulated by microbial processes in this system. This study provides mechanistic insights into the missing chemical and isotopic signals and microbially driven fate of terrigenous POM in the ocean, with important implications for how riverine material input affects marine carbon and nitrogen cycling.

This paper provides an in-depth investigation and analysis of volleyball training patterns using a sensing network composed of flexible strain sensors. This paper models the motion of the hand and leg joints based on an improved human posture estimation technique. The pattern recognition of human motion used the fusion of acceleration sensor and spiral meter data, the output of human motion information from the spiral meter, combined with GA-BP neural network algorithm to repair and fuse the output information, effectively improving the accuracy of posture angle measurement. Based on wireless positioning technology, an improved RFID positioning algorithm combining the LANDMARC algorithm and weighted center-of-mass algorithm in wireless sensor network is proposed, and the positioning accuracy reaches 89.2%; and the error feedback mechanism is introduced to improve the improved algorithm, and the positioning accuracy is improved again by 3.2%. Finally, the two-way dynamic time regularization algorithm is used to align the input action with the standard action, and the action pose evaluation index is compared and analyzed for the action sequence after alignment to obtain the final action quality comparison analysis results. In this paper, the proposed method is applied in practical training, which can effectively locate the stance of athletes and evaluate their movement quality without disturbing them and can provide valuable analysis information to support coaches’ guidance.

Background Geothermal systems have contributed greatly to both our understanding of the functions of extreme life and the evolutionary history of life itself. Shallow-sea hydrothermal systems are ecological intermediates of deep-sea systems and terrestrial springs, harboring unique and complexed ecosystems, which are well-lit and present physicochemical gradients. The microbial communities of deep-sea and terrestrial geothermal systems have been well-studied at the population genome level, yet little is known about the communities inhabiting the shallow-sea hydrothermal systems and how they compare to those inhabiting other geothermal systems. Results Here, we used genome-resolved metagenomic and metaproteomic approaches to probe into the genetic potential and protein expression of microorganisms from the shallow-sea vent fluids off Kueishantao Island. The families Nautiliaceae and Campylobacteraceae within the Epsilonbacteraeota and the Thiomicrospiraceae within the Gammaproteobacteria were prevalent in vent fluids over a 3-year sampling period. We successfully reconstructed the in situ metabolic modules of the predominant populations within the Epsilonbacteraeota and Gammaproteobacteria by mapping the metaproteomic data back to metagenome-assembled genomes. Those active bacteria could use the reductive tricarboxylic acid cycle or Calvin-Benson-Bassham cycle for autotrophic carbon fixation, with the ability to use reduced sulfur species, hydrogen or formate as electron donors, and oxygen as a terminal electron acceptor via cytochrome bd oxidase or cytochrome bb3 oxidase. Comparative metagenomic and genomic analyses revealed dramatic differences between submarine and terrestrial geothermal systems, including microbial functional potentials for carbon fixation and energy conversion. Furthermore, shallow-sea hydrothermal systems shared many of the major microbial genera that were first isolated from deep-sea and terrestrial geothermal systems, while deep-sea and terrestrial geothermal systems shared few genera. Conclusions The metabolic machinery of the active populations within Epsilonbacteraeota and Gammaproteobacteria at shallow-sea vents can mirror those living at deep-sea vents. With respect to specific taxa and metabolic potentials, the microbial realm in the shallow-sea hydrothermal system presented ecological linkage to both deep-sea and terrestrial geothermal systems.

Abstract. Nitrous oxide (N2O) is an important ozone-depleting greenhouse gas produced and consumed by microbially mediated nitrification and denitrification pathways. Estuaries are intensive N2O emission regions in marine ecosystems. However, the potential contributions of nitrifiers and denitrifiers to N2O sources and sinks in China's estuarine and coastal areas are poorly understood. The abundance and transcription of six key microbial functional genes involved in nitrification and denitrification, as well as the clade II-type nosZ gene-bearing community composition of N2O reducers, were investigated in four estuaries spanning the Chinese coastline. The results showed that the ammonia-oxidizing archaeal amoA genes and transcripts were more dominant in the northern Bohai Sea (BS) and Yangtze River estuaries, which had low nitrogen concentrations, while the denitrifier nirS genes and transcripts were more dominant in the southern Jiulong River (JRE) and Pearl River estuaries, which had high levels of terrestrial nitrogen input. Notably, the nosZ clade II gene was more abundant than the clade I-type throughout the estuaries except for in the JRE and a few sites of the BS, while the opposite transcript distribution pattern was observed in these two estuaries. The gene and transcript distributions were significantly constrained by nitrogen and oxygen concentrations as well as by salinity, temperature, and pH. The nosZ clade II gene-bearing community composition along China's coastline had a high level of diversity and was distinctly different from that in the soil and in marine oxygen-minimum-zone waters. By comparing the gene distribution patterns across the estuaries with the distribution patterns of the N2O concentration and flux, we found that denitrification may principally control the N2O emissions pattern.

Marine phytoplankton and heterotrophic bacteria share a very close but usually changeable relationship. However, the ultimate fate of their unstable relationship on a long-term scale is unclear. Here, the relationship between Synechococcus and heterotrophic bacterial communities underwent a dramatic shift from antagonism to commensalism and eventually to mutualism during long-term cocultivation. The relationship change is attributed to the different (even opposite) effects of diverse bacterial members on Synechococcus and the ratio of beneficial to harmful bacteria. Different bacterial members also interact with each other (e.g., quorum-sensing communication, hostility, or mutual promotion) and drive a dynamic succession in the entire community structure that corresponds exactly to the shift in its relationship with Synechococcus. In the final mutualism stage, a self-sufficient nitrogen cycle, including nitrogen fixation, denitrification, and organic nitrogen degradation, contributed to the healthy survival of Synechococcus for 2 years without an exogenous nutrient supply. This natural selective trait of Synechococcus and heterotrophic bacteria toward mutualism under long-term coexistence provides a novel clue for understanding the ubiquity and competitive advantage of Synechococcus in global oceans. IMPORTANCE Phytoplankton and heterotrophic bacteria have a close but usually changeable relationship. Uncovering the dynamic changes and driving factors of their interrelationships is of great significance for an in-depth understanding of the ecological processes and functions of marine microorganisms. Here, we observed that Synechococcus and heterotrophic bacterial communities underwent a dramatic change in their relationship from antagonism to mutualism during a long-term cocultivation process. We revealed that the interactions between different members of the bacterial community and the combined effects of different bacterial individuals on Synechococcus promoted the dynamic changes of the Synechococcus-bacterium relationship. In the end, a self-sufficient nutrient cycle (especially nitrogen) established by Synechococcus and bacterial communities supported their long-term survival without any external nutrition supply. This study provides novel insight into the interaction between Synechococcus and heterotrophic bacteria in the ocean and provides a novel clue for understanding the ubiquity and competitive advantage of Synechococcus in global oceans.

Organic carbon produced by nitrifiers plays an important role in maintaining the microbial metabolism in the aphotic ocean layer with carbon and energy scarcity. However, the contribution of nitrifiers to organic carbon processing remains unclear. To explore how nitrification impacts the material cycle in the starved ecosystem, we set up an ultra-large volume, long-term incubation experiment. Seawater collected from the Halifax coastal ocean was pumped into the Aquatron Tower Tank located at Dalhousie University, Canada, and was incubated under dark conditions for 73 days. The results indicated that the relative abundance of nitrifiers increased and nitrification was strengthened in the dark system where energy and organic carbon were scarce. The importance of nitrogenous compounds in particulate materials increased over the course of the incubation. Correlation analysis showed that the relative abundances of nitrifiers and particulate organic compounds containing nitrogen were significantly and positively correlated. Furthermore, network analysis suggested that metabolic processes related to nitrogenous and aromatic compounds are most important to particle associated bacteria. This study suggests that the nitrifiers could produce a series of organic compounds that result in the alteration of organic matter composition by promoting the degradation of recalcitrant aromatic compounds, which has important implications for organic matter processing in the starved dark ecosystem.

The translation inhibitor rocaglamide A (RocA) has shown promising antitumor activity because it uniquely clamps eukaryotic initiation factor (eIF) 4A onto polypurine RNA for selective translational repression. As eIF4A has been speculated to be a unique target of RocA, alternative targets have not been investigated. Here, we reveal that DDX3 is another molecular target of RocA. Proximity-specific fluorescence labeling of an O-nitrobenzoxadiazole-conjugated derivative revealed that RocA binds to DDX3. RocA clamps the DDX3 protein onto polypurine RNA in an ATP-independent manner. Analysis of a de novo-assembled transcriptome from the plant Aglaia, a natural source of RocA, uncovered the amino acid critical for RocA binding. Moreover, ribosome profiling showed that because of the dominant-negative effect of RocA, high expression of eIF4A and DDX3 strengthens translational repression in cancer cells. This study indicates that sequence-selective clamping of DDX3 and eIF4A, and subsequent dominant-negative translational repression by RocA determine its tumor toxicity.

Ammonia oxidation to nitrite and its subsequent oxidation to nitrate provides energy to the two populations of nitrifying chemoautotrophs in the energy-starved dark ocean, driving a coupling between reduced inorganic nitrogen (N) pools and production of new organic carbon (C) in the dark ocean. However, the relationship between the flux of new C production and the fluxes of N of the two steps of oxidation remains unclear. Here, we show that, despite orders-of-magnitude difference in cell abundances between ammonia oxidizers and nitrite oxidizers, the two populations sustain similar bulk N-oxidation rates throughout the deep waters with similarly high affinities for ammonia and nitrite under increasing substrate limitation, thus maintaining overall homeostasis in the oceanic nitrification pathway. Our observations confirm the theoretical predictions of a redox-informed ecosystem model. Using balances from this model, we suggest that consistently low ammonia and nitrite concentrations are maintained when the two populations have similarly high substrate affinities and their loss rates are proportional to their maximum growth rates. The stoichiometric relations between the fluxes of C and N indicate a threefold to fourfold higher C-fixation efficiency per mole of N oxidized by ammonia oxidizers compared to nitrite oxidizers due to nearly identical apparent energetic requirements for C fixation of the two populations. We estimate that the rate of chemoautotrophic C fixation amounts to ∼1 × 10 13 to ∼2 × 10 13 mol of C per year globally through the flux of ∼1 × 10 14 to ∼2 × 10 14 mol of N per year of the two steps of oxidation throughout the dark ocean.

Heterotrophic bacteria such as those from theRoseobactergroup and genusAlteromonasdominate the hadal zones of oceans; however, we know little about the genomic characteristics and potential metabolic adaptations of hadal trench-dwelling bacteria. Here, we report multiple single amplified genomes (SAGs) belonging toRoseobacterandAlteromonas, recovered from the hadal zone of the Mariana Trench. While phylogenetic analyses show that these hadal SAGs cluster with their surface relatives, an analysis of genomic recruitment indicates that they have higher relative abundances in the hadal zone of the Mariana Trench. Comparative genomic analyses between the hadal SAGs and reference genomes of closely related shallow-water relatives indicate that genes involved in the mobilome (prophages and transposons) are overrepresented among the unique genes of the hadalRoseobacterandAlteromonasSAGs; the functional proteins encoded by this category of genes also shows higher amino acid sequence variation than those encoded by other gene sets within theRoseobacterSAGs. We also found that genes involved in cell wall/membrane/envelope biogenesis, transcriptional regulation, and metal transport may be important for the adaptation of hadalRoseobacterandAlteromonaslineages. These results imply that the modification of cell surface-related proteins and transporters is the major direction of genomic evolution inRoseobacterandAlteromonasbacteria adapting to the hadal environment, and that prophages and transposons may be the key factors driving this process.