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Lakes in permafrost regions are dynamic landscape components and play an important role for climate change feedbacks. Lake processes such as mineralization and flocculation of dissolved organic carbon (DOC), one of the main carbon fractions in lakes, contribute to the greenhouse effect and are part of the global carbon cycle. These processes are in the focus of climate research, but studies so far are limited to specific study regions. In our synthesis, we analyzed 2167 water samples from 1833 lakes across the Arctic in permafrost regions of Alaska, Canada, Greenland, and Siberia to provide first pan-Arctic insights for linkages between DOC concentrations and the environment. Using published data and unpublished datasets from the author team, we report regional DOC differences linked to latitude, permafrost zones, ecoregions, geology, near-surface soil organic carbon contents, and ground ice classification of each lake region. The lake DOC concentrations in our dataset range from 0 to 1130 mg L−1 (10.8 mg L−1 median DOC concentration). Regarding the permafrost regions of our synthesis, we found median lake DOC concentrations of 12.4 mg L−1 (Siberia), 12.3 mg L−1 (Alaska), 10.3 mg L−1 (Greenland), and 4.5 mg L−1 (Canada). Our synthesis shows a significant relationship between lake DOC concentration and lake ecoregion. We found higher lake DOC concentrations at boreal permafrost sites compared to tundra sites. We found significantly higher DOC concentrations in lakes in regions with ice-rich syngenetic permafrost deposits (yedoma) compared to non-yedoma lakes and a weak but significant relationship between soil organic carbon content and lake DOC concentration as well as between ground ice content and lake DOC. Our pan-Arctic dataset shows that the DOC concentration of a lake depends on its environmental properties, especially on permafrost extent and ecoregion, as well as vegetation, which is the most important driver of lake DOC in this study. This new dataset will be fundamental to quantify a pan-Arctic lake DOC pool for estimations of the impact of lake DOC on the global carbon cycle and climate change.

Coccolithophores are globally important marine calcifying phytoplankton that utilize a haplo-diplontic life cycle. The haplo-diplontic life cycle allows coccolithophores to divide in both life cycle phases and potentially expands coccolithophore niche volume. Research has, however, to date largely overlooked the life cycle of coccolithophores and has instead focused on the diploid life cycle phase of coccolithophores. Through the synthesis and analysis of global scanning electron microscopy (SEM) coccolithophore abundance data (n=2534), we find that calcified haploid coccolithophores generally constitute a minor component of the total coccolithophore abundance (≈ 2 %–15 % depending on season). However, using case studies in the Atlantic Ocean and Mediterranean Sea, we show that, depending on environmental conditions, calcifying haploid coccolithophores can be significant contributors to the coccolithophore standing stock (up to ≈30 %). Furthermore, using hypervolumes to quantify the niche of coccolithophores, we illustrate that the haploid and diploid life cycle phases inhabit contrasting niches and that on average this allows coccolithophores to expand their niche by ≈18.8 %, with a range of 3 %–76 % for individual species. Our results highlight that future coccolithophore research should consider both life cycle stages, as omission of the haploid life cycle phase in current research limits our understanding of coccolithophore ecology. Our results furthermore suggest a different response to nutrient limitation and stratification, which may be of relevance for further climate scenarios. Our compilation highlights the spatial and temporal sparsity of SEM measurements and the need for new molecular techniques to identify uncalcified haploid coccolithophores. Our work also emphasizes the need for further work on the carbonate chemistry niche of the coccolithophore life cycle.

The shallower oxygen-poor water masses of the ocean confine a majority of the microbial communities that can produce up to 90 % of oceanic N2. This effective N2-yielding section encloses a suspended small-particle layer, inferred from particle backscattering (bbp) measurements. It is thus hypothesized that this layer (hereafter, the bbp-layer) is linked to microbial communities involved in N2 yielding such as nitrate-reducing SAR11 as well as sulfur-oxidizing, anammox, and denitrifying bacteria – a hypothesis yet to be evaluated. Here, data collected by three BGC-Argo floats deployed in the Black Sea are used to investigate the origin of this bbp-layer. To this end, we evaluate how the key drivers of N2-yielding bacteria dynamics impact the vertical distribution of bbp and the thickness of the bbp-layer. In conjunction with published data on N2 excess, our results suggest that the bbp-layer is at least partially composed of the bacteria driving N2 yielding for three main reasons: (1) strong correlations are recorded between bbp and nitrate; (2) the top location of the bbp-layer is driven by the ventilation of oxygen-rich subsurface waters, while its thickness is modulated by the amount of nitrate available to produce N2; and (3) the maxima of both bbp and N2 excess coincide at the same isopycnals where bacteria involved in N2 yielding coexist. We thus advance that bbp and O2 can be exploited as a combined proxy to delineate the N2-yielding section of the Black Sea. This proxy can potentially contribute to refining delineation of the effective N2-yielding section of oxygen-deficient zones via data from the growing BGC-Argo float network.

Lakes in permafrost regions are dynamic landscape components and play an important role for climate change feedbacks. Lake processes such as mineralization and flocculation of dissolved organic carbon (DOC), one of the main carbon fractions in lakes, contribute to the greenhouse effect and are part of the global carbon cycle. These processes are in focus of climate research but studies so far are limited to specific study regions. In our synthesis, we analysed 2,167 water samples from 1,833 lakes across the Arctic in permafrost regions of Alaska, Canada, Greenland, and Siberia to provide first pan-Arctic insights for linkages between DOC concentrations and the environment. Using published data and unpublished datasets from the author team we report regional DOC differences linked to latitude, permafrost zones, ecoregions, geology, near-surface soil organic carbon contents, and ground ice classification of each lake region. The lake DOC concentrations in our dataset range from 0 mg L−1 to 1,130 mg L−1 (10.8 mg L−1 median DOC concentration). Regarding the permafrost regions of our synthesis, we found median lake DOC concentrations of 12.4 mg L−1 (Siberia), 12.3 mg L−1 (Alaska), 10.3 mg L−1 (Greenland), and 4.5 mg L−1 (Canada). Our synthesis shows a significant relationship of lake DOC concentration and ecoregion of the lake. We found higher lake DOC concentrations in boreal permafrost sites compared to tundra sites. About 22 % of the lakes in our extensive dataset are located in regions with ice-rich syngenetic permafrost deposits (yedoma). Yedoma contains large amounts of easily erodible organic carbon and we found significantly higher DOC concentrations in yedoma lakes compared to non-yedoma lakes. Compared to previous studies we found a weak significant relationship of soil organic carbon content and lake DOC concentration as well as between ground-ice content and lake DOC. Our pan-Arctic dataset shows that the DOC concentration of a lake strongly depends on its environmental properties, especially on permafrost extent and ecoregion, as well as vegetation, which is the most important driver of lake DOC in this study. This new dataset will be fundamental to quantify a pan-Arctic lake DOC pool for estimations of the impact of lake DOC on the global carbon cycle and climate change.

Permafrost ground is one of the largest repositories of terrestrial organic carbon and might become or already is a carbon source in response to ongoing global warming. With this study of syngenetically frozen, ice-rich and organic carbon (OC)-bearing Yedoma and associated alas deposits in central Yakutia (Republic of Sakha), we aimed to assess the local sediment deposition regime and its impact on permafrost carbon storage. For this purpose, we investigated the Yukechi alas area (61.76495∘ N, 130.46664∘ E), which is a thermokarst landscape degrading into Yedoma in central Yakutia. We retrieved two sediment cores (Yedoma upland, 22.35 m deep, and alas basin, 19.80 m deep) in 2015 and analyzed the biogeochemistry, sedimentology, radiocarbon dates and stable isotope geochemistry. The laboratory analyses of both cores revealed very low total OC (TOC) contents (<0.1 wt %) for a 12 m section in each core, whereas the remaining sections ranged from 0.1 wt % to 2.4 wt % TOC. The core sections holding very little to no detectable OC consisted of coarser sandy material were estimated to be between 39 000 and 18 000 BP (years before present) in age. For this period, we assume the deposition of organic-poor material. Pore water stable isotope data from the Yedoma core indicated a continuously frozen state except for the surface sample, thereby ruling out Holocene reworking. In consequence, we see evidence that no strong organic matter (OM) decomposition took place in the sediments of the Yedoma core until today. The alas core from an adjacent thermokarst basin was strongly disturbed by lake development and permafrost thaw. Similar to the Yedoma core, some sections of the alas core were also OC poor (<0.1 wt %) in 17 out of 28 samples. The Yedoma deposition was likely influenced by fluvial regimes in nearby streams and the Lena River shifting with climate. With its coarse sediments with low OC content (OC mean of 5.27 kg m−3), the Yedoma deposits in the Yukechi area differ from other Yedoma sites in North Yakutia that were generally characterized by silty sediments with higher OC contents (OC mean of 19 kg m−3 for the non-ice wedge sediment). Therefore, we conclude that sedimentary composition and deposition regimes of Yedoma may differ considerably within the Yedoma domain. The resulting heterogeneity should be taken into account for future upscaling approaches on the Yedoma carbon stock. The alas core, strongly affected by extensive thawing processes during the Holocene, indicates a possible future pathway of ground subsidence and further OC decomposition for thawing central Yakutian Yedoma deposits.

Industrial-scale mining of deep-sea polymetallic nodules will remove nodules in large areas of the sea floor. The regrowth of the nodules by metal precipitation is estimated to take millions of years. Thus, for future mining impact studies, it is crucial to understand the role of nodules in shaping microbial diversity and function in deep-sea environments. Here we investigated microbial-community composition based on 16S rRNA gene sequences retrieved from sediments and nodules of the Peru Basin (4130–4198 m water depth). The nodule field of the Peru Basin showed a typical deep-sea microbiome, with dominance of the classes Gammaproteobacteria, Alphaproteobacteria, Deltaproteobacteria, and Acidimicrobiia. Nodules and sediments host distinct bacterial and archaeal communities, with nodules showing lower diversity and a higher proportion of sequences related to potential metal-cycling Bacteria (i.e. Magnetospiraceae, Hyphomicrobiaceae), bacterial and archaeal nitrifiers (i.e. AqS1, unclassified Nitrosomonadaceae, Nitrosopumilus, Nitrospina, Nitrospira), and bacterial sequences found in the oceanic crust, nodules, hydrothermal deposits, and sessile fauna. Sediment and nodule communities overall shared a low proportion of operational taxonomic units (OTUs; 21 % for Bacteria and 19 % for Archaea). Our results show that nodules represent a specific ecological niche (i.e. hard substrate, high metal concentrations, and sessile fauna), with a potentially relevant role in organic-carbon degradation. Differences in nodule community composition (e.g. Mn-cycling bacteria, nitrifiers) between the Clarion–Clipperton Fracture Zone (CCZ) and the Peru Basin suggest that changes in environmental setting (e.g. sedimentation rates) also play a significant role in structuring the nodule microbiome.

Upper suboxic water masses confine a majority of the microbial communities that can produce up to 90 % of oceanic N2. This effective N2-yielding section encloses a suspended small-particle layer, inferred from particle backscattering (bbp) measurements. It is thus hypothesized that this layer (hereafter, the bbp-layer) is linked to N2-yielding microbial communities such as anammox and denitrifying bacteria – a hypothesis yet to be evaluated. Here, data collected by three BGC-Argo floats deployed in the Black Sea are used to investigate the origin of this bbp-layer. To this end, we evaluate how key drivers of anammox-denitrifying bacteria dynamics impact on the vertical distribution of bbp and the thickness of the bbp-layer. In conjunction with published data on N2 excess, our results suggest that the bbp-layer is at least partially composed of anammox-denitrifying bacteria for three main reasons: (1) strong correlations are recorded between bbp and nitrate; (2) the top location of the bbp-layer is driven by the ventilation of oxygen-rich subsurface waters, while its thickness is modulated by the amount of nitrate available to produce N2; (3) the maxima of both bbp and N2 excess coincide at the same isopycnals where denitrifying-anammox bacteria coexist. We thus advance that bbp and O2 can be exploited as a combined proxy to delineate the N2-yielding section of the Black Sea. This proxy can potentially contribute to refining delineation of the effective N2-yielding section of oxygen-deficient zones via data from the growing BGC-Argo float network.

Coastal systems can act as filters for anthropogenic nutrient input into marine environments. Here, we assess the processes controlling the removal of phosphorus (P) and nitrogen (N) for four sites in the eutrophic Stockholm archipelago. Bottom water concentrations of oxygen (O2) and P are inversely correlated. This is attributed to the seasonal release of P from iron-oxide-bound (Fe-oxide-bound) P in surface sediments and from degrading organic matter. The abundant presence of sulfide in the pore water and its high upward flux towards the sediment surface (∼4 to 8 mmol m−2 d−1), linked to prior deposition of organic-rich sediments in a low-O2 setting (“legacy of hypoxia”), hinder the formation of a larger Fe-oxide-bound P pool in winter. This is most pronounced at sites where water column mixing is naturally relatively low and where low bottom water O2 concentrations prevail in summer. Burial rates of P are high at all sites (0.03–0.3 mol m−2 yr−1), a combined result of high sedimentation rates (0.5 to 3.5 cm yr−1) and high sedimentary P at depth (∼30 to 50 µmol g−1). Sedimentary P is dominated by Fe-bound P and organic P at the sediment surface and by organic P, authigenic Ca-P and detrital P at depth. Apart from one site in the inner archipelago, where a vivianite-type Fe(II)-P mineral is likely present at depth, there is little evidence for sink switching of organic or Fe-oxide-bound P to authigenic P minerals. Denitrification is the major benthic nitrate-reducing process at all sites (0.09 to 1.7 mmol m−2 d−1) with rates decreasing seaward from the inner to outer archipelago. Our results explain how sediments in this eutrophic coastal system can remove P through burial at a relatively high rate, regardless of whether the bottom waters are oxic or (frequently) hypoxic. Our results suggest that benthic N processes undergo annual cycles of removal and recycling in response to hypoxic conditions. Further nutrient load reductions are expected to contribute to the recovery of the eutrophic Stockholm archipelago from hypoxia. Based on the dominant pathways of P and N removal identified in this study, it is expected that the sediments will continue to remove part of the P and N loads.

The extracellular concentration of H2O2 in surface aquatic environments is controlled by a balance between photochemical production and the microbial synthesis of catalase and peroxidase enzymes to remove H2O2 from solution. In any kind of incubation experiment, the formation rates and equilibrium concentrations of reactive oxygen species (ROSs) such as H2O2 may be sensitive to both the experiment design, particularly to the regulation of incident light, and the abundance of different microbial groups, as both cellular H2O2 production and catalase–peroxidase enzyme production rates differ between species. Whilst there are extensive measurements of photochemical H2O2 formation rates and the distribution of H2O2 in the marine environment, it is poorly constrained how different microbial groups affect extracellular H2O2 concentrations, how comparable extracellular H2O2 concentrations within large-scale incubation experiments are to those observed in the surface-mixed layer, and to what extent a mismatch with environmentally relevant concentrations of ROS in incubations could influence biological processes differently to what would be observed in nature. Here we show that both experiment design and bacterial abundance consistently exert control on extracellular H2O2 concentrations across a range of incubation experiments in diverse marine environments. During four large-scale (>1000 L) mesocosm experiments (in Gran Canaria, the Mediterranean, Patagonia and Svalbard) most experimental factors appeared to exert only minor, or no, direct effect on H2O2 concentrations. For example, in three of four experiments where pH was manipulated to 0.4–0.5 below ambient pH, no significant change was evident in extracellular H2O2 concentrations relative to controls. An influence was sometimes inferred from zooplankton density, but not consistently between different incubation experiments, and no change in H2O2 was evident in controlled experiments using different densities of the copepod Calanus finmarchicus grazing on the diatom Skeletonema costatum (<1 % change in [H2O2] comparing copepod densities from 1 to 10 L−1). Instead, the changes in H2O2 concentration contrasting high- and low-zooplankton incubations appeared to arise from the resulting changes in bacterial activity. The correlation between bacterial abundance and extracellular H2O2 was stronger in some incubations than others (R2 range 0.09 to 0.55), yet high bacterial densities were consistently associated with low H2O2. Nonetheless, the main control on H2O2 concentrations during incubation experiments relative to those in ambient, unenclosed waters was the regulation of incident light. In an open (lidless) mesocosm experiment in Gran Canaria, H2O2 was persistently elevated (2–6-fold) above ambient concentrations; whereas using closed high-density polyethylene mesocosms in Crete, Svalbard and Patagonia H2O2 within incubations was always reduced (median 10 %–90 %) relative to ambient waters.

The speciation of dissolved iron (DFe) in the ocean is widely assumed to consist almost exclusively of Fe(III)-ligand complexes. Yet in most aqueous environments a poorly defined fraction of DFe also exists as Fe(II), the speciation of which is uncertain. Here we deploy flow injection analysis to measure in situ Fe(II) concentrations during a series of mesocosm/microcosm/multistressor experiments in coastal environments in addition to the decay rate of this Fe(II) when moved into the dark. During five mesocosm/microcosm/multistressor experiments in Svalbard and Patagonia, where dissolved (0.2 µm) Fe and Fe(II) were quantified simultaneously, Fe(II) constituted 24 %–65 % of DFe, suggesting that Fe(II) was a large fraction of the DFe pool. When this Fe(II) was allowed to decay in the dark, the vast majority of measured oxidation rate constants were less than calculated constants derived from ambient temperature, salinity, pH, and dissolved O2. The oxidation rates of Fe(II) spikes added to Atlantic seawater more closely matched calculated rate constants. The difference between observed and theoretical decay rates in Svalbard and Patagonia was most pronounced at Fe(II) concentrations <2 nM, suggesting that the effect may have arisen from organic Fe(II) ligands. This apparent enhancement of Fe(II) stability under post-bloom conditions and the existence of such a high fraction of DFe as Fe(II) challenge the assumption that DFe speciation in coastal seawater is dominated by ligand bound-Fe(III) species.