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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.

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.

The marine silicon cycle is intrinsically linked with carbon cycling in the oceans via biological production of silica by a wide range of organisms. The stable silicon isotopic composition (denoted by δ30Si) of siliceous microfossils extracted from sediment cores can be used as an archive of past oceanic silicon cycling. However, the silicon isotopic composition of biogenic silica has only been measured in diatoms, sponges and radiolarians, and isotopic fractionation relative to seawater is entirely unknown for many other silicifiers. Furthermore, the biochemical pathways and mechanisms that determine isotopic fractionation during biosilicification remain poorly understood. Here, we present the first measurements of the silicon isotopic fractionation during biosilicification by loricate choanoflagellates, a group of protists closely related to animals. We cultured two species of choanoflagellates, Diaphanoeca grandis and Stephanoeca diplocostata, which showed consistently greater isotopic fractionation (approximately −5 ‰ to −7 ‰) than cultured diatoms (−0.5 ‰ to −2.1 ‰). Instead, choanoflagellate silicon isotopic fractionation appears to be more similar to sponges grown under similar dissolved silica concentrations. Our results highlight that there is a taxonomic component to silicon isotope fractionation during biosilicification, possibly via a shared or related biochemical transport pathway. These findings have implications for the use of biogenic silica δ30Si produced by different silicifiers as proxies for past oceanic change.

Bottom trawling in the deep sea is one of the main drivers of sediment resuspension, eroding the seafloor and altering the content and composition of sedimentary organic matter (OM). The physical and biogeochemical impacts of bottom trawling were studied on the continental slope of the Gulf of Castellammare, Sicily (southwestern Mediterranean), through the analysis of two triplicate sediment cores collected at trawled and untrawled sites (∼550 m water depth) during the summer of 2016. Geochemical and sedimentological parameters (excess 210Pb, excess 234Th, 137Cs, dry bulk density, and grain size), elemental (organic carbon and nitrogen) and biochemical composition of sedimentary OM (proteins, carbohydrates, lipids), as well as its freshness (phytopigments) and degradation rates were determined in both coring locations. The untrawled site had a sedimentation rate of 0.15 cm yr−1 and presented a 6 cm thick surface mixed layer that contained siltier sediment with low excess 210Pb concentrations, possibly resulting from the resuspension, posterior advection, and eventual deposition of coarser and older sediment from adjacent trawling grounds. In contrast, the trawled site was eroded and presented compacted century-old sediment highly depleted in OM components, which were between 20 % and 60 % lower than those in the untrawled site. However, the upper 2 cm of the trawled site consisted of recently accumulated sediments enriched in excess 234Th, excess 210Pb, and phytopigments, while OM contents were similar to those from the untrawled core. This fresh sediment supported protein turnover rates of 0.025 d−1, which doubled those quantified in surface sediments of the untrawled site. The enhancement of remineralization rates in surface sediment of the trawled site was associated with the arrival of fresh particles on a chronically trawled deep-sea region that is generally deprived of OM. We conclude that the detrimental effects of bottom trawling can be temporarily and partially abated by the arrival of fresh and nutritionally rich OM, which stimulate the response of benthic communities. However, these ephemeral deposits are likely to be swiftly eroded due to the high trawling frequency over fishing grounds, highlighting the importance of establishing science-based management strategies to mitigate the impacts of bottom trawling.

Concentrations of dissolved 230Th in the ocean water column increase with depth due to scavenging and downward particle flux. Due to the 230Th scavenging process, any change in the calcium carbonate (CaCO3) fraction of the marine particle flux due to changes in biological CaCO3 hard-shell production as a consequence of progressing ocean acidification would be reflected in the dissolved 230Th activity. Our prognostic simulations with a biogeochemical ocean general circulation model using different scenarios for the reduction of CaCO3 production under ocean acidification and different greenhouse gas emission scenarios – the Representative Concentration Pathways (RCPs) 8.5 to 2.6 – reveal the potential for deep 230Th measurements to detect reduced CaCO3 production at the sea surface. The time of emergence of an acidification-induced signal on dissolved 230Th is of the same order of magnitude as for alkalinity measurements. Interannual and decadal variability in factors other than a reduction in CaCO3 hard-shell production may mask the ocean-acidification-induced signal in dissolved 230Th and make detection of the pure CaCO3-induced signal more difficult so that only really strong changes in marine CaCO3 export would be unambiguously identifiable soon. Nevertheless, the impacts of changes in CaCO3 export production on marine 230Th are stronger than those for changes in POC (particulate organic carbon) or clay fluxes.

Future deep-sea mining for polymetallic nodules in abyssal plains will negatively impact the benthic ecosystem, but it is largely unclear whether this ecosystem will be able to recover from mining disturbance and if so, to what extent and at what timescale. During the “DISturbance and reCOLonization” (DISCOL) experiment, a total of 22 % of the seafloor within a 10.8 km2 circular area of the nodule-rich seafloor in the Peru Basin (SE Pacific) was ploughed in 1989 to bury nodules and mix the surface sediment. This area was revisited 0.1, 0.5, 3, 7, and 26 years after the disturbance to assess macrofauna, invertebrate megafauna and fish density and diversity. We used this unique abyssal faunal time series to develop carbon-based food web models for each point in the time series using the linear inverse modeling approach for sediments subjected to two disturbance levels: (1) outside the plough tracks; not directly disturbed by plough, but probably suffered from additional sedimentation; and (2) inside the plough tracks. Total faunal carbon stock was always higher outside plough tracks compared with inside plough tracks. After 26 years, the carbon stock inside the plough tracks was 54 % of the carbon stock outside plough tracks. Deposit feeders were least affected by the disturbance, with modeled respiration, external predation, and excretion rates being reduced by only 2.6 % inside plough tracks compared with outside plough tracks after 26 years. In contrast, the respiration rate of filter and suspension feeders was 79.5 % lower in the plough tracks after 26 years. The “total system throughput” (T..), i.e., the total sum of modeled carbon flows in the food web, was higher throughout the time series outside plough tracks compared with the corresponding inside plough tracks area and was lowest inside plough tracks directly after the disturbance (8.63 × 10−3 ± 1.58 × 10−5 mmol C m−2 d−1). Even 26 years after the DISCOL disturbance, the discrepancy of T.. between outside and inside plough tracks was still 56 %. Hence, C cycling within the faunal compartments of an abyssal plain ecosystem remains reduced 26 years after physical disturbance, and a longer period is required for the system to recover from such a small-scale sediment disturbance experiment.

The North Atlantic Ocean is a major sink region for atmospheric CO2 and contributes to the storage of anthropogenic carbon (Cant). While there is general agreement that the intensity of the meridional overturning circulation (MOC) modulates uptake, transport and storage of Cant in the North Atlantic Subpolar Ocean, processes controlling their recent variability and evolution over the 21st century remain uncertain. This study investigates the relationship between transport, air–sea flux and storage rate of Cant in the North Atlantic Subpolar Ocean over the past 53 years. Its relies on the combined analysis of a multiannual in situ data set and outputs from a global biogeochemical ocean general circulation model (NEMO–PISCES) at 1∕2∘ spatial resolution forced by an atmospheric reanalysis. Despite an underestimation of Cant transport and an overestimation of anthropogenic air–sea CO2 flux in the model, the interannual variability of the regional Cant storage rate and its driving processes were well simulated by the model. Analysis of the multi-decadal simulation revealed that the MOC intensity variability was the major driver of the Cant transport variability at 25 and 36∘ N, but not at OVIDE. At the subpolar OVIDE section, the interannual variability of Cant transport was controlled by the accumulation of Cant in the MOC upper limb. At multi-decadal timescales, long-term changes in the North Atlantic storage rate of Cant were driven by the increase in air–sea fluxes of anthropogenic CO2. North Atlantic Central Water played a key role for storing Cant in the upper layer of the subtropical region and for supplying Cant to Intermediate Water and North Atlantic Deep Water. The transfer of Cant from surface to deep waters occurred mainly north of the OVIDE section. Most of the Cant transferred to the deep ocean was stored in the subpolar region, while the remainder was exported to the subtropical gyre within the lower MOC.

We analyze the response of different ionospheric equivalent current modes to variations in the interplanetary magnetic field (IMF) components By and Bz. Each mode comprises a fixed spatial pattern whose amplitude varies in time, identified by a month‐by‐month empirical orthogonal function separation of surface measured magnetic field variance. Here we focus on four sets of modes that have been previously identified as DPY, DP2, NBZ, and DP1. We derive the cross‐correlation function of each mode set with either IMF By or Bz for lags ranging from −10 to +600 mins with respect to the IMF state at the bow shock nose. For all four sets of modes, the average correlation can be reproduced by a sum of up to three linear responses to the IMF component, each centered on a different lag. These are interpreted as the statistical ionospheric responses to magnetopause merging (15‐ to 20‐min lag) and magnetotail reconnection (60‐min lag) and to IMF persistence. Of the mode sets, NBZ and DPY are the most predictable from a given IMF component, with DP1 (the substorm component) the least predictable. The proportion of mode variability explained by the IMF increases for the longer lags, thought to indicate conductivity feedbacks from substorms. In summary, we confirm the postulated physical basis of these modes and quantify their multiple reconfiguration timescales.