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The fluorophore [2‐(4‐pyridyl)‐5{[4‐dimethylaminoethyl‐aminocarbamoyl‐methoxy]phenyl}oxazole], in short PDMPO, is incorporated in newly polymerized silica in diatom frustules and thereby provides a tool to estimate Si uptake, study diatom cell cycles but also determine mortality‐independent abundance‐based species specific‐growth rates in cultures and natural assemblages. In this study, the theoretical framework and applicability of the PDMPO staining technique to estimate diatom species specific‐growth rates were investigated. Three common polar diatom species, Pseudo‐nitzschia subcurvata, Chaetoceros simplex, and Thalassiosira sp., chosen in order to cover a broad range of species specific frustule and life‐cycle characteristics, were incubated over 24 h in control (no PDMPO) and with 0.125 and 0.6 μM PDMPO addition, respectively. Results indicate that specific‐growth rates of the species tested were not affected in both treatments with PDMPO addition. The specific‐growth rate estimates based on the PDMPO staining patterns (μPDMPO) were comparable and more robust than growth rates estimated from the changes in cell concentrations (μcc). This technique also allowed to investigate and highlight the importance of the illumination cycle (light and dark phases) on cell division in diatoms. [ABSTRACT FROM AUTHOR]

Photosynthesis and respiration cause distinct chemical microenvironments within cyanobacterial aggregates. Here, we used microsensors and a diffusion–reaction model to characterize gradients in carbonate chemistry and investigate how these are affected by ocean acidification in Baltic vs. Pacific aggregates (Nodularia and Dolichospermum vs. Trichodesmium). Microsensor measurements of O2 and pH were performed under in situ and expected future pCO2 levels on Nodularia and Dolichospermum aggregates collected in the Baltic Sea. Under in situ conditions, O2 and pH levels within the aggregates covered ranges of 80–175% air saturation and 7.7–9.4 in dark and light, respectively. Carbon uptake in the light was predicted to reduce HCO3− by 100–150 μmol L−1 and CO2 by 3–6 μmol L−1 in the aggregate center compared to outside, inducing strong CO2 depletion (down to 0.5 μmol L−1 CO2 remaining in the center) even when assuming that HCO3− covered 80–90% of carbon uptake. Under ocean acidification conditions, enhanced CO2 availability allowed for significantly lower activity of carbon concentrating mechanisms, including a reduction of the contribution of HCO3− to carbon uptake by up to a factor of 10. The magnification of proton gradients under elevated pCO2 that was predicted based on a lower buffer capacity was observed in measurements despite a concurrent decrease in photosynthetic activity. In summary, we provide a quantitative image of the inorganic carbon environment in cyanobacterial aggregates under present‐day and expected future conditions, considering both the individual and combined effects of the chemical and biological processes that shape these environments. [ABSTRACT FROM AUTHOR]

Summary: In the marine realm, microorganisms are responsible for the bulk of primary production, thereby sustaining marine life across all trophic levels. Longhurst provinces have distinct microbial fingerprints; however, little is known about how microbial diversity and primary productivity change at finer spatial scales. Here, we sampled the Atlantic Ocean from south to north (~50°S–50°N), every ~0.5° latitude. We conducted measurements of primary productivity, chlorophyll‐a and relative abundance of 16S and 18S rRNA genes, alongside analyses of the physicochemical and hydrographic environment. We analysed the diversity of autotrophs, mixotrophs and heterotrophs, and noted distinct patterns among these guilds across provinces with high and low chlorophyll‐a conditions. Eukaryotic autotrophs and prokaryotic heterotrophs showed a shared inter‐province diversity pattern, distinct from the diversity pattern shared by mixotrophs, cyanobacteria and eukaryotic heterotrophs. Additionally, we calculated samplewise productivity‐specific length scales, the potential horizontal displacement of microbial communities by surface currents to an intrinsic biological rate (here, specific primary productivity). This scale provides key context for our trophically disaggregated diversity analysis that we could relate to underlying oceanographic features. We integrate this element to provide more nuanced insights into the mosaic‐like nature of microbial provincialism, linking diversity patterns to oceanographic transport through primary production. [ABSTRACT FROM AUTHOR]

Macrozooplankton and its grazing pressure shape ecosystem structures and carbon pathways in the Southern Ocean. Here, we present the implementation of "polar macrozooplankton" as a plankton functional type and a related fast‐sinking detritus class (fecal pellets) into the biogeochemical model REcoM‐2. We use the model to assess major carbon pathways and ecosystem structure in the Southern Ocean south of 50°S. The model represents zooplankton biomass and its spatial distribution in the Southern Ocean reasonably well in comparison to available biomass data. A distinct difference of our model from previous versions is the seasonal pattern of particle formation processes and ecosystem structures in the Southern Ocean. REcoM‐2 now captures high zooplankton biomass and a typical shift from a dominance of phytodetrital aggregates in spring to zooplankton fecal pellets later in the year. At sites with high biomass of macrozooplankton, the transfer efficiency of particulate organic carbon can be as high as 50%, and the carbon content of the exported material increases. In our simulations, macrozooplankton is an important component of the Southern Ocean plankton community, contributing up to 0.12 Pg C per year (14%) to total modeled carbon export across 100 m depth. Macrozooplankton changes the phytoplankton composition and supports the recycling of macronutrients. These results support the important role of macrozooplankton such as krill in the Southern Ocean and have implications for the representation of Southern Ocean biogeochemical cycles in models. Plain Language Summary: Large zooplankton such as krill is an important component of marine ecosystems, but it is rarely represented in ocean ecosystem models. Large zooplankton grazes on phytoplankton, smaller zooplankton, and sinking particles in the ocean. To understand the role of large zooplankton for carbon cycling in the Southern Ocean, we have implemented this group into our ecosystem model and describe macrozooplankton feeding and particle formation based on observations. We find that large zooplankton supports a stable or even increased growth of phytoplankton, in spite of its grazing. This is because it also returns nutrients to phytoplankton via excretion. Besides, large zooplankton increases the transfer of carbon to the deep ocean. Key Points: A new macrozoplankton parametrization is implemented into the biogeochemical model REcoM‐2Transfer efficiency reaches up to 50% due to the high carbon content and fast sinking of macrozooplankton fecal pelletsMacrozooplankton contributes up to 14% (0.12 Pg C yr−1) to modeled carbon export at 100 m depth in the Southern Ocean south of 50°S [ABSTRACT FROM AUTHOR]

Responses of marine primary production to a changing climate are determined by a concert of multiple environmental changes, for example in temperature, light, pCO2, nutrients, and grazing. To make robust projections of future global marine primary production, it is crucial to understand multiple driver effects on phytoplankton. This meta‐analysis quantifies individual and interactive effects of dual driver combinations on marine phytoplankton growth rates. Almost 50% of the single‐species laboratory studies were excluded because central data and metadata (growth rates, carbonate system, experimental treatments) were insufficiently reported. The remaining data (42 studies) allowed for the analysis of interactions of pCO2 with temperature, light, and nutrients, respectively. Growth rates mostly respond non‐additively, whereby the interaction with increased pCO2 profusely dampens growth‐enhancing effects of high temperature and high light. Multiple and single driver effects on coccolithophores differ from other phytoplankton groups, especially in their high sensitivity to increasing pCO2. Polar species decrease their growth rate in response to high pCO2, while temperate and tropical species benefit under these conditions. Based on the observed interactions and projected changes, we anticipate primary productivity to: (a) first increase but eventually decrease in the Arctic Ocean once nutrient limitation outweighs the benefits of higher light availability; (b) decrease in the tropics and mid‐latitudes due to intensifying nutrient limitation, possibly amplified by elevated pCO2; and (c) increase in the Southern Ocean in view of higher nutrient availability and synergistic interaction with increasing pCO2. Growth‐enhancing effect of high light and warming to coccolithophores, mainly Emiliania huxleyi, might increase their relative abundance as long as not offset by acidification. Dinoflagellates are expected to increase their relative abundance due to their positive growth response to increasing pCO2 and light levels. Our analysis reveals gaps in the knowledge on multiple driver responses and provides recommendations for future work on phytoplankton. [ABSTRACT FROM AUTHOR]

Coral reefs are threatened by global and local stressors. Yet, reefs appear to respond differently to different environmental stressors. Using a global dataset of coral reef occurrence as a proxy for the long‐term adaptation of corals to environmental conditions in combination with global environmental data, we show here how global (warming: sea surface temperature; acidification: aragonite saturation state, Ωarag) and local (eutrophication: nitrate concentration, and phosphate concentration) stressors influence coral reef habitat suitability. We analyse the relative distance of coral communities to their regional environmental optima. In addition, we calculate the expected change of coral reef habitat suitability across the tropics in relation to an increase of 0.1°C in temperature, an increase of 0.02 μmol/L in nitrate, an increase of 0.01 μmol/L in phosphate and a decrease of 0.04 in Ωarag. Our findings reveal that only 6% of the reefs worldwide will be unaffected by local and global stressors and can thus act as temporary refugia. Local stressors, driven by nutrient increase, will affect 22% of the reefs worldwide, whereas global stressors will affect 11% of these reefs. The remaining 61% of the reefs will be simultaneously affected by local and global stressors. Appropriate wastewater treatments can mitigate local eutrophication and could increase areas of temporary refugia to 28%, allowing us to 'buy time', while international agreements are found to abate global stressors. [ABSTRACT FROM AUTHOR]

The haptophyte Phaeocystis antarctica is endemic to the Southern Ocean, where iron supply is sporadic and its availability limits primary production. In iron fertilization experiments, P. antarctica showed a prompt and steady increase in cell abundance compared to heavily silicified diatoms along with enhanced colony formation. Here we utilized a transcriptomic approach to investigate molecular responses to alleviation of iron limitation in P. antarctica. We analyzed the transcriptomic response before and after (14 h, 24 h and 72 h) iron addition to a low‐iron acclimated culture. After iron addition, we observed indicators of a quick reorganization of cellular energetics, from carbohydrate catabolism and mitochondrial energy production to anabolism. In addition to typical substitution responses from an iron‐economic toward an iron‐sufficient state for flavodoxin (ferredoxin) and plastocyanin (cytochrome c6), we found other genes utilizing the same strategy involved in nitrogen assimilation and fatty acid desaturation. Our results shed light on a number of adaptive mechanisms that P. antarctica uses under low iron, including the utilization of a Cu‐dependent ferric reductase system and indication of mixotrophic growth. The gene expression patterns underpin P. antarctica as a quick responder to iron addition. [ABSTRACT FROM AUTHOR]

The Western Antarctic Peninsula (WAP), one of the most productive regions of the Southern Ocean, is currently undergoing rapid environmental changes such as ocean acidification (OA) and increased daily irradiances from enhanced surface‐water stratification. To assess the potential for future biological CO2 sequestration of this region, we incubated a natural phytoplankton assemblage from Ryder Bay, WAP, under a range of pCO2 levels (180 μatm, 450 μatm, and 1000 μatm) combined with either moderate or high natural solar radiation (MSR: 124 μmol photons m−2 s−1 and HSR: 435 μmol photons m−2 s−1, respectively). The initial and final phytoplankton communities were numerically dominated by the prymnesiophyte Phaeocystis antarctica, with the single cells initially being predominant and solitary and colonial cells reaching similar high abundances by the end. Only when communities were grown under ambient pCO2 in conjunction with HSR did the small diatom Fragilariopsis pseudonana outcompete P. antarctica at the end of the experiment. Such positive light‐dependent growth response of the diatom was, however, dampened by OA. These changes in community composition were caused by an enhanced photosensitivity of diatoms, especially F. pseudonana, under OA and HSR, reducing thereby their competitiveness toward P. antarctica. Moreover, community primary production (PP) of all treatments yielded similar high rates at the start and the end of the experiment, but with the main contributors shifting from initially large to small cells toward the end. Even though community PP of Ryder Bay phytoplankton was insensitive to the changes in light and CO2 availability, the observed size‐dependent shift in productivity could, however, weaken the biological CO2 sequestration potential of this region in the future. [ABSTRACT FROM AUTHOR]

Phytoplankton photosynthesis strongly relies on the operation of carbon‐concentrating mechanisms (CCMs) to accumulate CO2 around their carboxylating enzyme ribulose‐1,5‐bisphosphate carboxylase/oxygenase (RuBisCO). Earlier evolved phytoplankton groups were shown to exhibit higher CCM activities to compensate for their RuBisCO with low CO2 specificities. Here, we tested whether earlier evolved phytoplankton groups also exhibit a higher CCM plasticity. To this end, we collected data from literature and applied a Bayesian linear meta‐analytic model. Our results show that with elevated pCO2, photosynthetic CO2 affinities decreased strongest and most consistent for the earlier evolved groups, i.e., cyanobacteria and dinoflagellates, while CO2‐dependent changes in affinities for haptophytes and diatoms were smaller and less consistent. In addition, responses of maximum photosynthetic rates toward elevated pCO2 were generally small and inconsistent across species. Our results demonstrate that phytoplankton groups with an earlier origin possess a high CCM plasticity, whereas more recently evolved groups do not, which likely results from evolved differences in the CO2 specificity of RuBisCO. [ABSTRACT FROM AUTHOR]

A combined increase in seawater [CO2] and [H+] was recently shown to induce a shift from photosynthetic HCO3− to CO2 uptake in Emiliania huxleyi. This shift occurred within minutes, whereas acclimation to ocean acidification (OA) did not affect the carbon source., To identify the driver of this shift, we exposed low- and high-light acclimated E. huxleyi to a matrix of two levels of dissolved inorganic carbon (1400, 2800 μmol kg−1) and pH (8.15, 7.85) and directly measured cellular O2, CO2 and HCO3− fluxes under these conditions., Exposure to increased [CO2] had little effect on the photosynthetic fluxes, whereas increased [H+] led to a significant decline in HCO3− uptake. Low-light acclimated cells overcompensated for the inhibition of HCO3− uptake by increasing CO2 uptake. High-light acclimated cells, relying on higher proportions of HCO3− uptake, could not increase CO2 uptake and photosynthetic O2 evolution consequently became carbon-limited., These regulations indicate that OA responses in photosynthesis are caused by [H+] rather than by [CO2]. The impaired HCO3− uptake also provides a mechanistic explanation for lowered calcification under OA. Moreover, it explains the OA-dependent decrease in photosynthesis observed in high-light grown phytoplankton. [ABSTRACT FROM AUTHOR]

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed. [ABSTRACT FROM AUTHOR]

The variability of total alkalinity (TA) and its relationship with salinity in the tropical and subtropical surface ocean were examined using data collected in various marine environments from a ship of opportunity. In the open ocean regions of the Atlantic, Pacific, and Indian Oceans, sea surface TA variability was observed to be mainly controlled by the simple dilution or concentration (SDC) effect of precipitation and evaporation, and the measured concentrations of TA agreed well with those predicted from salinity and temperature. Non-SDC changes in alkalinity in ocean margins and inland seas were examined by comparing the salinity-normalized alkalinity with that of the open ocean end-member. Non-SDC alkalinity additions to the western North Atlantic margin, eastern North Pacific margin, and Mediterranean Sea were identified, which mainly resulted from river inputs and shelf currents. In contrast, removal of TA through formation and sedimentation of calcium carbonate was observed to be an important control in the Red Sea. The concentration of the river end-member can only be reliably derived from the y intercept of TA-S regression (TAS0) in river-dominated systems such as estuaries and river plumes. In coastal regions where other processes (evaporation, shelf currents, upwelling, calcification, etc.) are more influential, TAS0 can significantly deviate from the river water concentration and hence be an unreliable indicator of it. Negative values of TAS0 can result from non-SDC TA removal at the low salinity end (relative to the salinity of the oceanic end-member) and/or non-SDC TA addition at high salinities (as occurs in the Mediterranean Sea). [ABSTRACT FROM AUTHOR]

An Earth System model of intermediate complexity is used to quantify the effects of potential reorganizations of the Atlantic meridional overturning circulation, ranging from a weakening to a complete shutdown, on future uptake of excess CO[sub2] by the ocean and the terrestrial biosphere. Circulation changes significantly reduce oceanic CO[sub2] uptake, with the largest effects occurring under a complete shutdown. The reduction in uptake is caused to a large extent by a decline in marine export of organic particulate matter. On land, soil carbon increases because of reduced soil respiration in the mid-latitudes of the Northern Hemisphere. This gain is partially compensated by loss of vegetation carbon associated with reduced primary production in the northern high latitudes due to a cooler and dryer climate. Altogether, these processes act to modestly enhance CO[sub2] concentration levels in the atmosphere, thereby leading to a small positive feedback. The rise in atmospheric CO[sub2] ranges from 13 ppm to 34 ppm in 2500, depending on the amount of AMOC weakening, the magnitude of the anthropogenic CO[sub2] perturbation, and the future CO[sub2] fertilization effect. [ABSTRACT FROM AUTHOR]