Interlinking diatom frustule diversity from the abyss of the central Arabian Sea to surface processes: physical forcing and oxygen minimum zone.

This study analyzed the diversity and abundance of diatom frustules including the ancillary parameters using the core top sediments from five locations (21, 19, 15, 13, and 11°N) along the central Arabian Sea (64°E), an area profoundly influenced by atmospheric forcing (monsoons) and oxygen minimum zone (OMZ) with high spatial variability. Significantly higher organic carbon (0.97 ± 0.05%) and diatom frustules (5.92 ± 0.57 × 104 valves g-1) were noticed in the north (21, 19, 15°N) where natural nutrient enrichment via open-ocean upwelling, winter convection, and lateral advection support large diatom-dominated phytoplankton blooms and intense OMZ. Conversely, the south (13, 11°N) depicted significantly lower organic carbon (0.74 ± 0.08%) as well as frustules (4.02 ± 0.87 × 104 valves g-1) as this area mostly remains nutrient-poor dominated by small-medium-sized phytoplankton. The north was dominated by large-sized diatoms like Coscinodiscus that could escape grazing and sink consequently due to higher ballasting. Furthermore, the presence of the intense OMZ in the north might reduce grazing pressure (low zooplankton stock) and mineralization speed facilitating higher phytodetritus transport. Relatively smaller chain-forming centric (Thalassiosira) and pennate diatoms (Pseudo-nitzschia, Fragilaria, Nitzschia, etc.) were found throughout the transect with higher abundance in the south. The euphotic diatom diversity from the existing literature was compared with the frustule diversity from the sediments suggesting not all diatoms make their way to the abyss. Such distinct spatial north-south variability in diatom frustule size as well as abundance could be attributed to cell size, grazing, and water column mineralization rates related to OMZ.

Phytoplankton growth and community shift over a short-term high-CO2 simulation experiment from the southwestern shelf of India, Eastern Arabian Sea (summer monsoon).

The southwestern shelf water of India (eastern Arabian Sea) experiences high seasonality. This area is one of the understudied regions in terms of phytoplankton response to the projected ocean acidification, particularly, during the summer monsoon when phytoplankton abundance is high. Here we present the results of a short-term simulated ocean acidification experiment (ambient CO2 424 µatm; high CO2, 843, 1138 µatm) on the natural phytoplankton assemblages conducted onboard (R. V. Sindhu Sadhana) during the summer monsoon (Aug 2017). Among the dissolved inorganic nutrients, dissolved silicate (DSi) and nitrate + nitrite levels were quite low (< 2 µM). Phytoplankton biomass did not show any net enhancement after the incubation in any treatment. Both marker pigment analysis and microscopy revealed the dominance of diatoms in the phytoplankton community, and a significant restructuring was noticed over the experimental period. Divinyl chlorophylla (DVChla) containing picocyanobacteria and 19'-hexanoyloxyfucoxanthin (19'HF) containing prymnesiophytes did not show any noticeable change in response to CO2 enrichment. A CO2-induced positive growth response was noticed in some diatoms (Guinardia flaccida, Cylindrotheca closterium, and Pseudo-nitzschia sp.) and dinoflagellates (Protoperidinium sp. and Peridinium sp.) indicating their efficiency to quickly acclimatize at elevated CO2 levels. This is important to note that the positive growth response of toxigenic pennate diatoms like Pseudo-nitzschia as well as a few dinoflagellates at elevated CO2 levels can be expected in the future-ocean scenario. The proliferation of such non-palatable phytoplankton may impact grazing, the food chain, and carbon cycling in this region.

Experimental strategies to assess the biological ramifications of multiple drivers of global ocean change-A review.

Marine life is controlled by multiple physical and chemical drivers and by diverse ecological processes. Many of these oceanic properties are being altered by climate change and other anthropogenic pressures. Hence, identifying the influences of multifaceted ocean change, from local to global scales, is a complex task. To guide policy-making and make projections of the future of the marine biosphere, it is essential to understand biological responses at physiological, evolutionary and ecological levels. Here, we contrast and compare different approaches to multiple driver experiments that aim to elucidate biological responses to a complex matrix of ocean global change. We present the benefits and the challenges of each approach with a focus on marine research, and guidelines to navigate through these different categories to help identify strategies that might best address research questions in fundamental physiology, experimental evolutionary biology and community ecology. Our review reveals that the field of multiple driver research is being pulled in complementary directions: the need for reductionist approaches to obtain process-oriented, mechanistic understanding and a requirement to quantify responses to projected future scenarios of ocean change. We conclude the review with recommendations on how best to align different experimental approaches to contribute fundamental information needed for science-based policy formulation.

Investigating the impacts of treated effluent discharge on coastal water health (Visakhapatnam, SW coast of Bay of Bengal, India).

The present study investigated the impacts of treated effluent discharge on physicochemical and biological properties of coastal waters from three pharmaceuticals situated along the coast of Visakhapatnam (SW Bay of Bengal). Seawater samples were collected (during the months of December 2013, March 2014 and April 2014) from different sampling locations (Chippada (CHP), Tikkavanipalem (TKP) and Nakkapalli (NKP)) at 0- and 30-m depths within 2-km radius (0.5 km = inner, 1 km = middle and 2 km = outer sampling circles) from the marine outfall points. Physicochemical and biological parameters, which differed significantly within the stations, were likely to be influenced by strong seasonality rather than local discharge. Dissolved oxygen variability was tightly coupled with both physical and biological processes. Phytoplankton cell density and total chlorophyll (TChla) concentrations were significantly correlated with dissolved inorganic nutrient concentrations. CHP (December) represented a diatom bloom condition where the highest concentrations of diatom cells, total chlorophyll (TChla), dissolved oxygen coupled with lower zooplankton abundance and low nutrient levels were noticed. The centric diatom, Chaetoceros sp. (> 50%) dominated the phytoplankton community. TKP (March) represented a post-diatom bloom phase with the dominance of Pseudo-nitzschia seriata; zooplankton abundance and nutrient concentrations were minimum. Conversely, NKP (April) represented a warm well-stratified heterotrophic period with maximum zooplankton and minimum phytoplankton density. Dinoflagellate abundance increased at this station. Relatively higher water temperature, salinity, inorganic nutrients coupled with very low concentrations of dissolved oxygen, TChla and pH were observed at this station. Copepods dominated the zooplankton communities in all stations and showed their highest abundance in the innermost sampling circles. Treated effluent discharge did not seem to have any significant impact at these discharge points.

A coherent status of summer monsoon phytoplankton communities (2017-2018) along the Western Indian continental shelf: Implications for fisheries.

The western Indian continental shelf is characterized by contrasting biogeochemical features from south to north mostly governed by monsoon wind forcing. Here we present the first detailed study on the phytoplankton community (analyzed by marker pigments and microscopy) during the summer monsoon addressing the interannual variability (Aug 2017 and 2018) along the Western Indian Continental Shelf waters (8° N-21° N, at 200 m isobaths). A distinct interannual variability in monsoon impacted the hydrography and nutrient stoichiometry that was further cascaded to the phytoplankton community structure. The timing of the monsoon onset, wind speeds, and the strength of the alongshore wind component were the major factors that governed the interannual variability in the development and progression of the upwelling as well as phytoplankton bloom. The upwelling-dominated southwestern shelf was characterized by cold, nutrient-rich waters dominated by the marker pigment of diatoms, fucoxanthin, and microscopy also revealed the presence of large, chain-forming diatoms (Chaetoceros sp.; Dactyliosolen fragilissimus). In contrast, low nutrient warm waters in the northern shelf housed nanophytoplankton and picophytoplankton along with small diatoms (Thalassiosira sp.) and dinoflagellates (Gymnodinium sp.). Interestingly, in a few stations in the southern shelf, the surface waters were low saline that was intensified in 2018 compared to 2017 inhibiting upwelled water from reaching the surface. Consequently, due to low nutrient levels caused by reduced upwelling and low salinity, picoplanktonic cyanobacteria dominated replacing the larger diatoms that were found plenty in 2017. Likely, such a niche shift adversely impacted the planktivorous fish catch (Indian Oil Sardine) that was seen to be less in 2018 relative to 2017. Any further changes in monsoon variability and freshwater flow to this region may have direct consequences on the phytoplankton community as well as fisheries.

Phytoplankton community shift in response to experimental Cu addition at the elevated CO2 levels (Arabian Sea, winter monsoon).

Understanding phytoplankton community shifts under multiple stressors is becoming increasingly important. Among other combinations of stressors, the impact of trace metal toxicity on marine phytoplankton under the ocean acidification scenario is an important aspect to address. Such multiple stressor studies are rare from the Arabian Sea, one of the highest productive oceanic provinces within the North Indian Ocean. We studied the interactive impacts of copper (Cu) and CO2 enrichment on two natural phytoplankton communities from the eastern and central Arabian Sea. Low dissolved silicate (DSi < 2 µM) favoured smaller diatoms (e.g. Nitzschia sp.) and non-diatom (Phaeocystis). CO2 enrichment caused both positive (Nitzschia sp. and Phaeocystis sp.) and negative (Cylindrotheca closterium, Navicula sp., Pseudo-nitzschia sp., Alexandrium sp., and Gymnodinium sp.) growth impacts. The addition of Cu under the ambient CO2 level (A-CO2) hindered cell division in most of the species, whereas Chla contents were nearly unaffected. Interestingly, CO2 enrichment seemed to alleviate Cu toxicity in some species (Nitzschia sp., Cylindrotheca closterium, Guinardia flaccida, and Phaeocystis) and increased their growth rates. This could be related to the cellular Cu demand and energy budget at elevated CO2 levels. Dinoflagellates were more sensitive to Cu supply compared to diatoms and prymnesiophytes and could be related to the unavailability of prey. Such community shifts in response to the projected ocean acidification, oligotrophy, and Cu pollution may impact trophic transfer and carbon cycling in this region.

A story of resilience: Arctic diatom Chaetoceros gelidus exhibited high physiological plasticity to changing CO2 and light levels.

Arctic phytoplankton are experiencing multifaceted stresses due to climate warming, ocean acidification, retreating sea ice, and associated changes in light availability, and that may have large ecological consequences. Multiple stressor studies on Arctic phytoplankton, particularly on the bloom-forming species, may help understand their fitness in response to future climate change, however, such studies are scarce. In the present study, a laboratory experiment was conducted on the bloom-forming Arctic diatom Chaetoceros gelidus (earlier C. socialis) under variable CO2 (240 and 900 µatm) and light (50 and 100 µmol photons m-2 s-1) levels. The growth response was documented using the pre-acclimatized culture at 2°C in a closed batch system over 12 days until the dissolved inorganic nitrogen was depleted. Particulate organic carbon and nitrogen (POC and PON), pigments, cell density, and the maximum quantum yield of photosystem II (Fv/Fm) were measured on day 4 (D4), 6 (D6), 10 (D10), and 12 (D12). The overall growth response suggested that C. gelidus maintained a steady-state carboxylation rate with subsequent conversion to macromolecules as reflected in the per-cell POC contents under variable CO2 and light levels. A substantial amount of POC buildup at the low CO2 level (comparable to the high CO2 treatment) indicated the possibility of existing carbon dioxide concentration mechanisms (CCMs) that needs further investigation. Pigment signatures revealed a high level of adaptability to variable irradiance in this species without any major CO2 effect. PON contents per cell increased initially but decreased irrespective of CO2 levels when nitrogen was limited (D6 onward) possibly to recycle intracellular nitrogen resources resulting in enhanced C: N ratios. On D12 the decreased dissolved organic nitrogen levels could be attributed to consumption under nitrogen starvation. Such physiological plasticity could make C. gelidus "ecologically resilient" in the future Arctic.

Simulated ocean acidification altered community composition and growth of a coastal phytoplankton assemblage (South West coast of India, eastern Arabian Sea).

Marine phytoplankton can be highly sensitive to ocean acidification; however, their responses are diverse and therefore, phytoplankton response study on the regional scale is of high research priority. The present study documented the community shift and growth responses of a natural phytoplankton assemblage from the South West coastal water of India (South Eastern Arabian Sea) under ambient CO2 (A-CO2 ≈ 400 µatm) and high CO2 (H-CO2 ≈ 830 µatm) levels in microcosms during the winter monsoon. A doubling of pCO2 resulted in increased cell density, particulate organic carbon and nitrogen (POC, PON) contents, and C:N ratios. The depleted values of δ13CPOC in the H-CO2-incubated cells indicated a higher diffusive CO2 influx. HPLC marker pigment analysis revealed that the community was microphytoplankton dominated (mostly diatoms); nanoplanktonic prymnesiophytic algae and picoplanktonic cyanobacteria showed insignificant response to the simulated ocean acidification. A high CO2-induced increased growth rate was noticed in 6 diatoms (Leptocylindrus danicus; Rhizosolenia setigera; Navicula sp., Asterionella glacialis, Dactyliosolen fragilissimus, and Thalassiosira sp.). The cell volumes of Thalassionema frauenfeldii, Asterionella glacialis, and Cylindrotheca closterium increased significantly, whereas Rhizosolenia setigera and Thalassiosira sp. showed decreased cell volume at the elevated CO2 levels. These changes in growth rate, cell volume, and elemental stoichiometry could be related to CO2 acquisition and the nutritional status of the cells. Some phytoplankton genera from this region are probably acclimatized to pCO2 fluctuations and are likely to benefit from the future increase in CO2 levels. Higher POC production and increased C:N ratio along with variable cell volume may impact the trophic transfer and cycling of organic carbon in this coastal water. However, a multi-stressor approach in a longer experimental exposure should be considered in future research.

Interannual variability in particulate organic matter distribution and its carbon stable isotope signatures from the western Indian shelf waters.

Particulate organic carbon and nitrogen (POC, PN, collectively particulate organic matter, POM) and the stable isotopic signature of POC (δ13CPOC) are important to delineate its sources and recycling in shelf water. The present study provides insights into the factors responsible for spatial and interannual variability in POM and δ13CPOC values along the western Indian shelf waters (8° N -21° N) during the southwest (SW) monsoon (August) 2017 and 2018. The dominance of phytoplankton-derived POM with a negligible terrestrial influence was evident from the positive correlation between POC and TChla contents, ratios of C: N, and δ13CPOC signatures. Prominent upwelling signatures [cold nutrient-rich water, higher POM, total Chlorophylla (TChla), and δ13CPOC values] were noted in the south (8-12° N), whereas low nutrient warm waters (lower values of POM, TChla, and δ13CPOC) were prevalent in the north (13-21° N). Phytoplankton biomass was significantly higher and matured in 2017 due to an early and stronger upwelling in the south. In 2018, delayed and weak upwelling (evident from Ekman offshore transport and pumping velocity) resulted in the late development of phytoplankton bloom and lower POM. Furthermore, considerably lower nutrient supply within the mixed layers in 2018 compared to 2017 was partially attributed to the enhanced spatial expansion of low salinity waters closer to the surface. In the north, in 2018, higher wind speeds enhanced vertical mixing resulting in increased nutrient supply and TChla compared to 2017. We conclude that monsoon wind speed in the northern shelf and strength as well as the timing of the upwelling, including freshwater flux in the south, can be the key factors in modulating the interannual variability in POM distribution and δ13CPOC signature in the western Indian Shelf waters.

Carbon Dioxide Concentration Mechanisms in Natural Populations of Marine Diatoms: Insights From Tara Oceans.

Marine diatoms, the most successful photoautotrophs in the ocean, efficiently sequester a significant part of atmospheric CO2 to the ocean interior through their participation in the biological carbon pump. However, it is poorly understood how marine diatoms fix such a considerable amount of CO2, which is vital information toward modeling their response to future CO2 levels. The Tara Oceans expeditions generated molecular data coupled with in situ biogeochemical measurements across the main ocean regions, and thus provides a framework to compare diatom genetic and transcriptional flexibility under natural CO2 variability. The current study investigates the interlink between the environmental variability of CO2 and other physicochemical parameters with the gene and transcript copy numbers of five key enzymes of diatom CO2 concentration mechanisms (CCMs): Rubisco activase and carbonic anhydrase (CA) as part of the physical pathway, together with phosphoenolpyruvate carboxylase, phosphoenolpyruvate carboxykinase, and malic enzyme as part of the potential C4 biochemical pathway. Toward this aim, we mined >200 metagenomes and >220 metatranscriptomes generated from samples of the surface layer of 66 globally distributed sampling sites and corresponding to the four main size fractions in which diatoms can be found: 0.8-5 µm, 5-20 µm, 20-180 µm, and 180-2,000 µm. Our analyses revealed that the transcripts for the enzymes of the putative C4 biochemical CCM did not in general display co-occurring profiles. The transcripts for CAs were the most abundant, with an order of magnitude higher values than the other enzymes, thus implying the importance of physical CCMs in diatom natural communities. Among the different classes of this enzyme, the most prevalent was the recently characterized iota class. Consequently, very little information is available from natural diatom assemblages about the distribution of this class. Biogeographic distributions for all the enzymes show different abundance hotspots according to the size fraction, pointing to the influence of cell size and aggregation in CCMs. Environmental correlations showed a complex pattern of responses to CO2 levels, total phytoplankton biomass, temperature, and nutrient concentrations. In conclusion, we propose that biophysical CCMs are prevalent in natural diatom communities.

Impacts of Zn and Cu enrichment under ocean acidification scenario on a phytoplankton community from tropical upwelling system.

Increasing dissolution of CO2 in the surface ocean is rapidly decreasing its pH and changing carbon chemistry which is further affecting marine biota in several ways. Phytoplankton response studies under the combination of elevated CO2 and trace metals are rare. We have conducted two consecutive onboard incubation experiments (R. V. Sindhu Sadhana; August 2017) in the eastern Arabian Sea (SW coast of India) during an upwelling event. A nutrient enriched diatom bloom was initiated onboard and grown under ambient (≈400 µatm, A-CO2) and high CO2 levels (≈1000 µatm; H-CO2) with different zinc (Zn; 1 nM) and copper (Cu) concentrations (1 nM, 2 nM and 8 nM). Phytoplankton community composition and the dominant genera were different during these two experiments. CO2 enrichment alone did not show any significant growth stimulating impact on the experimental community except enhanced cell density in the first experiment. Addition of Zn at A-CO2 level revealed no noticeable responses; whereas, the same treatment under H-CO2 level significantly reduced cell number. Considerably high protein content under H-CO2+Zn treatment was possibly counteracting Zn toxicity which also caused slower growth rate. Cu addition did not show any noticeable impact on growth and biomass production except increased protein content as well as decreased carbohydrate: protein ratio. This can be attributed to relatively higher protein synthesis than carbohydrate to alleviate oxidative stress generated by Cu. The centric diatom Chaetoceros and toxin producing pennate diatom Pseudo-nitzschia showed no significant response to either CO2 or Zn enrichment. Large centric diatom Leptocylindrus and Skeletonema responded positively to Zn addition in both CO2 levels. The former species showed the most sensitive response at the highest Cu and H-CO2 treatment; whereas, the pennate diatoms Nitzschia and Pseudo-nitzschia (toxigenic diatom) showed higher resilience under elevated CO2 and Cu levels. This observation indicated that in future ocean, increasing CO2 concentrations and trace metal pollution may potentially alter phytoplankton community structure and may facilitate toxigenic diatom bloom in the coastal waters.

Physiological responses of coastal phytoplankton (Visakhapatnam, SW Bay of Bengal, India) to experimental copper addition.

Trace amount of copper (Cu) is essential for many physiological processes; however, it can be potentially toxic at elevated levels. The impact of variable Cu concentrations on a coastal phytoplankton community was investigated along a coastal transect in SW Bay of Bengal. A small increase in Cu supply enhanced the concentrations of particulate organic carbon, particulate organic nitrogen, biogenic silica, total pigment, phytoplankton cell and total bacterial count. At elevated Cu levels all these parameters were adversely affected. δ13CPOM and δ15NPOC reflected a visible signature of both beneficial and toxic impacts of Cu supply. Skeletonema costatum, the dominant diatom species, showed higher tolerance to increasing Cu levels relative to Chaetoceros sp. Cyanobacteria showed greater sensitivity to copper than diatoms. The magnitude of Cu toxicity on the phytoplankton communities was inversely related to the distance from the coast. Co-enrichment of iron alleviated Cu toxicity to phytoplankton.