Transitional traits determine the acclimation characteristics of the coccolithophore Chrysotila dentata to ocean warming and acidification.

Ocean warming and acidification interactively affect the coccolithophore physiology and drives major biogeochemical changes. While numerous studies investigated coccolithophore under short-term conditions, knowledge on how different transitional periods over long-exposure could influence the element, macromolecular and metabolic changes for its acclimation are largely unknown. We cultured the coccolithophore Chrysotila dentata, (culture generations of 1st, 10th, and 20th) under present (low-temperature low-carbon-dioxide [LTLC]) and projected (high-temperature high-carbon-dioxide [HTHC]) ocean conditions. We examined elemental and macromolecular component changes and sequenced a transcriptome. We found that with long-exposure, most physiological responses in HTHC cells decreased when compared with those in LTLC, however, HTHC cell physiology showed constant elevation between each generation. Specifically, compared to 1st generation, the 20th generation HTHC cells showed increases in quota carbon (Qc:29%), nitrogen (QN :101%), and subsequent changes in C:N-ratio (68%). We observed higher lipid accumulation than carbohydrates within HTHC cells under long-exposure, suggesting that lipids were used as an alternative energy source for cellular acclimation. Protein biosynthesis pathways increased their efficiency during long-term HTHC condition, indicating that cells produced more proteins than required to initiate acclimation. Our findings suggest that the coccolithophore resilience increased between the 1st-10th generation to initiate the acclimation process under ocean warming and acidifying conditions.

Spatial distribution and origin of organic matters in an Arctic fjord system based on lipid biomarkers (n-alkanes and sterols).

The concentration of n-alkanes (C17-C35) and sterols in marine particulate matter were investigated to trace the origin of organic carbon in Kongsfjorden in early spring (April). The spatial distributions of environmental factors (seawater temperature, salinity, density, turbidity, chlorophyll a (chl. a) and particulate organic carbon (POC) concentrations) and the cell density of phytoplankton differed between the inner and outer fjord regions. In addition, brassicasterol, diatom biomarker, showed a high concentration in the outer fjord and positive correlations with the chl. a and POC concentrations in the water column. In contrast, some sterols originating from terrestrial organic matter (OM), such as stigmasterol and campesterol, showed relatively higher concentrations in the inner fjord than in the outer fjord. Based on the distance-based redundancy analysis (db-RDA) result, the distributions of organic compounds are predominantly controlled by the water density and the POC and chl. a concentrations, and these distributions allowed us to divide the inner and outer fjord regions. However, the hierarchical clustering of principal components (HCPC) results obtained based on principal component analysis (PCA) using lipid biomarkers (C17-C35 alkanes and sterols) and environmental factors indicated that the clusters were distinguished by surface (0 m) and subsurface (>4 m) seawater samples rather than by any regional division. Notably, the concentration of relatively short-chain alkanes (average chain length (ACL): 24.6 ± 3.7) without a carbon preference for odd numbers (carbon preference index (CPI): 0.97 ± 0.11) in the sea surface layer was significantly higher than that of subsurface seawater (ACL: 31.1 ± 0.5 and CPI: 1.06 ± 0.03) in the early spring. This suggests the potential of these compounds as indicators for tidewater glacier-derived OM and freshwater input by snow melt into the fjord system. Hence, these results demonstrate that the distributions of lipid biomarkers in the water column possibly provide important information for a comprehensive understanding of the origin and transport of OM in an Arctic fjord.

Physiological and molecular responses of the Antarctic harpacticoid copepod Tigriopus kingsejongensis to salinity fluctuations - A multigenerational study.

Since Antarctica and the surrounding Southern Ocean are facing global climate change, biota inhabiting those coastal regions is now challenged by environmental fluctuations including coastal freshening. In this study, the effects of salinity range of 0-75 (practical salinity unit, PSU) on the Antarctic harpacticoid copepod Tigriopus kingsejongensis was investigated by measurement of 96 h survival rate, lifespan, and sex ratio with further analysis of multigenerational growth parameters and mRNA expressions under salinity of 15-45. Different stages of the copepods (i.e., nauplius, male, and female) generally showed tolerance to hypo- and hypersalinity, wherein female copepods were more tolerant than males when exposed to salinity fluctuations. Lifespan was significantly shortened by hypo- and hypersalinity compared to control salinity (34), but there was no significant difference in the sex ratio between salinity treatments. Multigenerational experiments across five generations revealed that exposure to salinities of 15 and 45 reduced body length compared to that in control salinity and the first generation of each salinity group. Our results provide evidence regarding T. kingsejongensis on their preferred salinity ranges, physiological limit to salinity fluctuations, and population dynamics in future salinity.

Survey of Bacterial Phylogenetic Diversity During the Glacier Melting Season in an Arctic Fjord.

To understand bacterial biogeography in response to the hydrographic impact of climate change derived from the Arctic glacier melting, we surveyed bacterial diversity and community composition using bacterial 16S rRNA gene metabarcoding in the seawaters of Kongsfjorden, Svalbard, during summer 2016. In the present study, bacterial biogeography in the Kongsfjorden seawaters showed distinct habitat patterns according to water mass classification and habitat transition between Atlantic and fjord surface waters. Moreover, we estimated phylogenetic diversity of bacterial communities using the net relatedness, nearest taxon, and beta nearest taxon indices. We found the influence of freshwater input from glacier melting in shaping bacterial assemblage composition through the stochastic model. We further evaluated bacterial contributions to phytoplankton-derived dimethylsulfoniopropionate (DMSP) using a quantitative PCR (qPCR) measurement with demethylation (dmdA) and cleavage (dddP) genes of two fundamentally different processes. Our qPCR results imply that bacterial DMSP degradation follows the Atlantic inflow during summer in Kongsfjorden. These findings suggest that the Atlantic inflow and glacial melting influence bacterial community composition and assembly processes and thus affect the degradation of phytoplankton-derived organic matter in an Arctic fjord.

Unraveling prokaryotic diversity distribution and functional pattern on nitrogen and methane cycling in the subtropical Western North Pacific Ocean.

Prokaryotes play an important role in marine nitrogen and methane cycles. However, their community changes and metabolic modifications to the concurrent impact of ocean warming (OW), acidification (OA), deoxygenation (OD), and anthropogenic­nitrogen-deposition (AND) from the surface to the deep ocean remains unknown. We examined here the amplicon sequencing approach across the surface (0-200 m; SL), intermediate (200-1000 m; IL), and deep layers (1000-2200 m; DL), and characterized the simultaneous impacts of OW, OA, OD, and AND on the Western North Pacific Ocean prokaryotic changes and their functional pattern in nitrogen and methane cycles. Results showed that SL possesses higher ammonium oxidation community/metabolic composition assumably the reason for excess nitrogen input from AND and modification of their kinetic properties to OW adaptation. Expanding OD at IL showed hypoxic conditions in the oxygen minimum layer, inducing higher microbial respiration that elevates the dimerization of nitrification genes for higher nitrous oxide production. The aerobic methane-oxidation composition was dominant in SL presumably the reason for adjustment in prokaryotic optimal temperature to OW, while anaerobic oxidation composition was dominant at IL due to the evolutionary changes coupling with higher nitrification. Our findings refocus on climate-change impacts on the open ocean ecosystem from the surface to the deep-environment integrating climate-drivers as key factors for higher nitrous-oxide and methane emissions.

Capture of decline in spring phytoplankton biomass derived from COVID-19 lockdown effect in the Yellow Sea offshore waters.

The Yellow Sea, characterized as a high-productivity ecosystem, is considered to be significantly attributable to high nutrient supply via atmospheric deposition. We observed a significant decline in phytoplankton biomass (~30%) over the Yellow Sea during February-May 2020 (period of COVID-19 lockdown effect) compared to the same period in 2015-2019 (period of no effect of COVID-19 lockdown). Several possible factors, such as variations in irradiance, vertical mixing, and river discharges, were not major contributors. Through the analysis of transportation and the constituents of atmospheric pollutants from Northern China (main source regions) to the Yellow Sea, we suggest that the decline in phytoplankton biomass over the Yellow Sea is mainly attributed to decreased atmospheric nutrient deposition due to the COVID-19 lockdown effect, because of decreased anthropogenic emissions in Northern China. Thus, attention should be focused on the Yellow Sea ecosystem response to increasing anthropogenic activities by lifting the COVID-19 lockdown restrictions.

Acclimation traits determine the macromolecular basis of harmful dinoflagellate Alexandrium minutum in response to changing climate conditions.

Ocean warming and acidification are expected to have profound impacts on the marine ecosystem, although the dinoflagellate Alexandrium minutum is reported to be acclimated to such conditions. However, it is unknown on the transition time scale how this species physiologically adjusts their element accumulation and associated resource allocation for this process. We designed a set of experiments to examine how different culture generations (1st, 5th, and 10th) change their cell physiology, cellular quotas and macromolecular cellular contents related to functional processes in A. minutum grown with future (pCO2, 1000 ppm; 25°C) and present (pCO2, 400 ppm; 21°C) ocean conditions. The differing cell sizes and storage capacity at different generations confirmed that compared to ancestors (1st generation), acclimation cells (10th generation) gained increases in quota carbon (QC; 55%; [p < 0.05]) and quota phosphate (QP; 23% [ p < 0.05]). This variation in C:P and N:P influences was transition-specific and largely determined by phosphate-based molecules. It was observed that A. minutum was initially dependent on P molecules, which help cells act as alternative lipids for quick acclimation until N molecules resume carbon-based lipids for their long-term acclimation. Our study demonstrated that rising temperature and pCO2 concentrations in ocean may increase A. minutum based on the comprehensive analysis of different physiological modifications, including its growth, element accumulation, transformation, and functional allocation.

Evaluating bloom potential of the green-tide forming alga Ulva ohnoi under ocean acidification and warming.

The occurrence of green-tides, whose bloom potential may be increased by various human activities and biogeochemical process, results in enormous economic losses and ecosystem collapse. In this study, we investigated the ecophysiology of the subtropical green-tide forming alga, Ulva ohnoi complex (hereafter: U. ohnoi), under simulated future ocean conditions in order to predict its bloom potential using photosynthesis and growth measurements, and stable isotope analyses. Our mesocosm system included four experimental conditions that simulated the individual and combined effects of elevated CO2 and temperature, namely control (450 µatm CO2 & 20 °C), acidification (900 µatm CO2 & 20 °C), warming (450 µatm CO2 & 25 °C), and greenhouse (900 µatm CO2 & 25 °C). Photosynthetic electron transport rates (rETR) increased significantly under acidification conditions, but net photosynthesis and growth were not affected. In contrast, rETR, net photosynthesis, and growth all decreased significantly under elevated temperature conditions (i.e. both warming and greenhouse). These results represent the imbalance of energy metabolism between electron transport and O2 production that may be expected under ocean acidification conditions. This imbalance appears to be related to carbon and nitrogen assimilation by U. ohnoi. In particular, 13C and 15N discrimination data suggest U. ohnoi prefers CO2 and NH4+ over HCO3- and NO3- as sources of carbon and nitrogen, respectively, and this results in increased N content in the thallus under ocean acidification conditions. Together, our results suggest a trade-off in which the bloom potential of U. ohnoi could increase under ocean acidification due to greater N accumulation and through the saving of energy during carbon and nitrogen metabolism, but that elevated temperatures could decrease U. ohnoi's bloom potential through a decrease in photosynthesis and growth.

N2O dynamics in the western Arctic Ocean during the summer of 2017.

The western Arctic Ocean (WAO) has experienced increased heat transport into the region, sea-ice reduction, and changes to the WAO nitrous oxide (N2O) cycles from greenhouse gases. We investigated WAO N2O dynamics through an intensive and precise N2O survey during the open-water season of summer 2017. The effects of physical processes (i.e., solubility and advection) were dominant in both the surface (0-50 m) and deep layers (200-2200 m) of the northern Chukchi Sea with an under-saturation of N2O. By contrast, both the surface layer (0-50 m) of the southern Chukchi Sea and the intermediate (50-200 m) layer of the northern Chukchi Sea were significantly influenced by biogeochemically derived N2O production (i.e., through nitrification), with N2O over-saturation. During summer 2017, the southern region acted as a source of atmospheric N2O (mean: + 2.3 ± 2.7 µmol N2O m-2 day-1), whereas the northern region acted as a sink (mean - 1.3 ± 1.5 µmol N2O m-2 day-1). If Arctic environmental changes continue to accelerate and consequently drive the productivity of the Arctic Ocean, the WAO may become a N2O "hot spot", and therefore, a key region requiring continued observations to both understand N2O dynamics and possibly predict their future changes.

Distribution and Control of Bacterial Community Composition in Marian Cove Surface Waters, King George Island, Antarctica during the Summer of 2018.

Marian Cove is experiencing some of the most rapid environmental changes in the Antarctic region; however, little is known about the response of bacterial communities to these changes. The main purpose of this study was to investigate the spatial variation of physical‒biogeochemical‒bacterial community features in the Marian Cove surface waters and the environmental parameters governing the spatial variation in the bacterial community composition during the summer of 2018. The Marian Cove surface waters are largely composed of two different characteristics of water masses: relatively low-temperature, -salinity, and -nutrient surface glacier water (named SGW) and relatively high-temperature, -salinity, and -nutrient surface Maxwell Bay water (named SMBW). The SGW bacterial communities were dominated by unclassified Cryomorphaceae, Sedimenticola, and Salibacter genera, while the SMBW bacterial communities were dominated by Sulfitobacter, Arcobacter, and Odoribacter genera. Spatial variations in bacterial community composition were mainly attributed to physical and biogeochemical characteristics, suggesting that the bacterial community composition of the Marian Cove surface waters is mainly determined by environmental characteristics. These findings provide a foundation to improve the understanding of bacterial community variations in response to a rapidly changing Marian Cove in the Antarctic.

Global warming offsets the ecophysiological stress of ocean acidification on temperate crustose coralline algae.

Dramatic increases in the release of anthropogenic CO2 and global temperatures have resulted in alterations to seawater carbonate chemistry and metabolisms of marine organisms. There has been recent interest in the effects of these stressors on crustose coralline algae (CCA) because photosynthesis and calcification are influenced by all components of carbonate chemistry. To examine this, a mesocosm experiment was conducted to evaluate photosynthesis, calcification and growth in the temperate CCA Chamberlainium sp. under acidification (doubled CO2), warming (+5 °C), and greenhouse (doubled CO2 and +5 °C) conditions compared to present-day conditions. After 47 days of acclimation to these conditions, productivity was lowest under acidification, although photochemical properties were improved, while respiration was highest under warming. Likewise, growth was lowest under acidification, but this negative response was offset by elevated temperature under greenhouse. Together, these results suggest that warming offsets the negative effects of acidification by creating more suitable conditions for photosynthesis and growth.

Latitudinal Distributions and Controls of Bacterial Community Composition during the Summer of 2017 in Western Arctic Surface Waters (from the Bering Strait to the Chukchi Borderland).

The western Arctic Ocean is experiencing some of the most rapid environmental changes in the Arctic. However, little is known about the microbial community response to these changes. Employing observations from the summer of 2017, this study investigated latitudinal variations in bacterial community composition in surface waters between the Bering Strait and Chukchi Borderland and the factors driving the changes. Results indicate three distinctive communities. Southern Chukchi bacterial communities are associated with nutrient rich conditions, including genera such as Sulfitobacter, whereas the northern Chukchi bacterial community is dominated by SAR clades, Flavobacterium, Paraglaciecola, and Polaribacter genera associated with low nutrients and sea ice conditions. The frontal region, located on the boundary between the southern and northern Chukchi, is a transition zone with intermediate physical and biogeochemical properties; however, bacterial communities differed markedly from those found to the north and south. In the transition zone, Sphingomonas, with as yet undetermined ecological characteristics, are relatively abundant. Latitudinal distributions in bacterial community composition are mainly attributed to physical and biogeochemical characteristics, suggesting that these communities are susceptible to Arctic environmental changes. These findings provide a foundation to improve understanding of bacterial community variations in response to a rapidly changing Arctic Ocean.

Temporal and Vertical Variation in Microbial Community Composition in Response to Physicochemical Characteristics in a Water Column of Highly Eutrophied Jinhae Bay, South Korea.

Microbial communities play an essential role in marine biogeochemical cycles. Physical and biogeochemical changes in Jinhae Bay, the most anthropogenically eutrophied bay on the coasts of South Korea, are well described, but less is known about the associated changes in microbial communities. Temporal and vertical variation in microbial communities at three depths (surface, middle, and bottom) at seven time points (June to December) at the J1 sampling site were investigated on the MiSeq platform based on the 16S rRNA gene. Overall, the microbial community was dominated by Proteobacteria, Cyanobacteria, and Bacteroidetes from June to November, whereas Firmicutes were dominant in December, especially in the middle and bottom layers. The results indicate that the microbial community composition strongly varied with temporal changes in the physicochemical water properties. Moreover, the community composition differed markedly between the surface and middle layers and the bottom layer in the summer, when the water column was strongly stratified and bottom water hypoxia developed. A redundancy analysis suggested a significant correlation between physicochemical variables (i.e., temperature, salinity, and oxygen concentration) and microbial community composition. This study indicates that temporal changes in water conditions and eutrophication-induced hypoxia effectively shape the structure of the microbial community.

Chemical oceanography. Increasing anthropogenic nitrogen in the North Pacific Ocean.

The recent increase in anthropogenic emissions of reactive nitrogen from northeastern Asia and the subsequent enhanced deposition over the extensive regions of the North Pacific Ocean (NPO) have led to a detectable increase in the nitrate (N) concentration of the upper ocean. The rate of increase of excess N relative to phosphate (P) was found to be highest (~0.24 micromoles per kilogram per year) in the vicinity of the Asian source continent, with rates decreasing eastward across the NPO, consistent with the magnitude and distribution of atmospheric nitrogen deposition. This anthropogenically driven increase in the N content of the upper NPO may enhance primary production in this N-limited region, potentially leading to a long-term change of the NPO from being N-limited to P-limited.