Phytoplankton Growth Research Articles

Electrochemical ocean iron fertilization and alkalinity enhancement approach toward CO2 sequestration

Achieving net-zero emissions by 2050 requires the development of effective negative emission techniques, including ocean-based approaches for CO2 sequestration. However, the implementation and testing of marine CO2 removal (mCDR) techniques such as ocean iron fertilization (OIF) or ocean alkalinity enhancement (OAE) face significant challenges. Herein, a novel self-operating electrochemical technology is presented that not only combines OIF and OAE, but also recovers hydrogen gas (H2) from seawater, hence offering a promising solution for achieving quantifiable and transparent large-scale mCDR. Experimental results show that the electrochemical OIF (EOIF) can not only increase the concentration of ferrous iron (Fe+2) by 0–0.5 mg/L, but also significantly increases the seawater pH by 8% (i.e., a 25% decrease in the hydrogen ions concentration). The release of iron (Fe+2/Fe+3) can be regulated by adjusting the magnitude of the electric current and its form (e.g., pulsed current and polarity reversal), as well as by optimizing the electrode material and geometry. In certain ocean regions, enhanced iron concentrations stimulate the naturally occurring biological carbon pump (BCP), leading to increased phytoplankton growth, CO2 uptake, and subsequent export of carbon to the deep ocean. Simultaneously, the system increases seawater alkalinity and the buffer capacity, enhancing CO2 solubility and storage in the shallow ocean through the solubility pump. The obtained measurements demonstrate the scalability of EOIF and its ability to operate using solar energy at a lower cost. Overall, the proposed EOIF technology offers a practical, effective, and sustainable solution for addressing climate change on a large scale.

The effects of turbulence on the growth of three different diatom species

The effects of turbulence on phytoplankton growth have received considerable attentions. However, the complexity of turbulence poses a significant challenge to its systematic characterization in the laboratory, resulting in relatively limited data on the effects of turbulence on several algal species. Here, a laboratory turbulence simulation system was set up to systematically investigate the growth of three common diatom species (Thalassiosira pseudonana, Skeletonema costatum, and Phaeodactylum tricornutum) under stationary and turbulent conditions (at 60, 120, 180 rpm), and measurements were taken for the algal biomass, algal photosynthetic activity, and nutrients consumption. By comparing the growth of three algae species under different turbulence exposure intensities, this study found that different algae exhibit varying sensitivities to turbulence, and therefore have different shear thresholds. Meanwhile, cell morphology is the key factor influencing the different shear threshold values observed in the three diatom species. Additionally, turbulence could impact algal aggregation and light availability, and dramatically improve nutrient uptake by phytoplankton. Our study will provides theoretical support for future endeavors in using turbulence to cultivate phytoplankton or combat algal blooms.

Impact of typhoons on the ecological environment of the Pearl River Estuary in the summer of 2021—a study of an algal bloom event

In the summer of 2021, two typhoons, Typhoon Cempaka and Typhoon Lupit, passed through the Pearl River Estuary (PRE), resulting in a noticeable chlorophyll-a (Chla) bloom event on the west side of the PRE. Utilizing a coupled hydrodynamic-ecological model (ROMS-CoSiNE), we investigated the changes in the marine environment during the two-typhoon period and explored the underlying mechanisms involved. Our model effectively reproduces the spatial characteristics of the typhoon-induced algal blooms. The results suggest that the westward invasion of the Pearl River plume has a predominant impact on the alterations in biochemical elements. For further quantitative analyses, we utilized the model to determine the Chla concentration budget in algal bloom region to assess the dominant mechanism, particularly focusing on horizontal advection. Our findings indicated that algal bloom events are mainly attributed to a straightforward physical mechanism, namely, phytoplankton transportation, rather than to nutrient-induced phytoplankton growth. The study also revealed that the different growth processes of the two simulated phytoplankton species did not contribute positively to the increased Chla concentrations. However, further research is needed to investigate the impacts of different phytoplankton functional groups on the marine ecological environment under the influence of typhoons. This study enhances the understanding of the response of ecological environments in coastal areas to typhoons.

Pollution, spatial distribution, and health risks assessment of nutrient concentration in surface water resources of Maroon-Jarahi Basin in southwestern Iran.

Nowadays, the introduction of nutrients caused by human activities is considered an environmental issue and a significant problem in river basins and coastal ecosystems. In this study, the concentration of nutrients ( and ) in the surface water sources of the Maroon-Jarahi watershed in the southwest of Iran was determined, and the pollution status and health risk assessment were done. The average concentration of nitrate and phosphate in Ludab, Maroon, Zard, Allah, Jarahi rivers, and Shadegan wetland were obtained at 2.25-0.59, 4.59-1.84, 4.07-2.02, 5.40-2.81, 11.51-4.67, 21.63 and 6.20 (mg/l), respectively. A comparison of the results with the World Health Organization (WHO) limit showed that nitrate was lower than in all stations, but phosphate was higher than the limit in some stations of the Maroon, Allah, Jarahi rivers, and Shadegan wetland. Calculation of linear regression analysis showed significant positive relationships between nitrate and phosphate in all surface water sources (except Ludab) and based on the N/P ratio, nitrogen was estimated as the limiting factor in phytoplankton growth (N/P < 16). The evaluation of the status of the Nutrient pollution index (NPI) was observed as: Shadegan > Jarahi > Allah > Maroon > Zard > Ludab that the Jarahi River and Shadegan wetland were in the medium pollution class (1 < NPI ≤ 3) and other waterbodies were in the non-polluted to low pollution state (NPI < 1). Calculation of the chronic daily intake (CDI) showed that water body nutrients cause more non-carcinogenic health risks through the oral route than dermal exposure, and according to HI, children's health is more at risk than adults. Findings showed that surface water resources especially downstream of the Maroon-Jarahi watershed are at eutrophication risk, and to control the nearby human activities and as a result increase the nutrients in these water resources, measures should be taken.

Predicting river phytoplankton blooms and community succession using ecological niche modeling

AbstractExcessive phytoplankton concentrations in rivers can result in the loss of plant and invertebrate communities, and threaten drinking water supplies. Whilst the physicochemical controls on algal blooms have been identified previously, how these factors combine to control the initiation, size, and cessation of blooms in rivers is not well understood. We applied flow cytometry to quantify diatom, chlorophyte, and cyanobacterial group abundances in the River Thames (UK) at weekly intervals from 2011 to 2022, alongside physicochemical data. A niche modeling approach was used to identify thresholds in water temperature, flow, solar radiation, and soluble reactive phosphorus (SRP) concentrations required to produce periods of phytoplankton growth, with blooms only occurring when all thresholds were met. The thresholds derived from the 2011 to 2018 dataset were applied to a test data set (2019–2022), which predicted the timing and duration of blooms at accuracies of &gt; 80%. Diatoms and nano‐chlorophyte blooms were initiated by flow and water temperature, and usually terminated due to temperature and flow going out of the threshold range, or SRP and Si becoming limiting. Cyanobacterial bloom dynamics were primarily controlled by water temperature and solar radiation. This simple methodology provides a key understanding of phytoplankton community succession and inter‐annual variation and can be applied to any river with similar water quality and phytoplankton data. It provides early warnings of algal and cyanobacterial bloom timings, which support future catchment management decisions to safeguard water resources, and provides a basis for modeling changing phytoplankton bloom risk due to future climate change.

Clustering evaluation of water quality for various classes of in-flow rivers in connected brackish lakes.

Brackish waters and estuaries at the lower reaches of rivers accumulate organic matter and nutrients from various sources in the watershed. Sufficient light and shallow water depth stimulate phytoplankton growth, resulting in a more diversified ecosystem with higher trophic levels. For effective watershed management, it is crucial to characterize the water quality of all rivers, including small and medium-sized ones. Our field survey assessed water quality parameters in 26 inflow rivers surrounding Lakes Shinji and Nakaumi, two consolidated brackish lakes in Japan. The parameters included water temperature, salinity, chlorophyll-a, and nutrients. The study used hierarchical clustering. The Silhouette Index was used to assess clustering outcomes and identify any difficulties in dispersion across clusters. The 26 rivers surrounding Lakes Shinji and Nakaumi were classified into six groups based on their water quality characteristics. This classification distinguishes itself from earlier subjective methods that relied on geographical factors. The new approach identifies a need for improved management of river water quality. The results of the cluster analysis provide valuable insights for future management initiatives. It is important to consider these findings alongside established watershed criteria.

Proton-pumping rhodopsins promote the growth and survival of phytoplankton in a highly variable ocean.

Proton-pumping rhodopsins (PPRs) utilize sunlight to produce cellular energy. They are widely distributed in marine phytoplankton and were recently shown to occur in the vacuolar membrane of a marine diatom, making the vacuole a second light transducing organelle. Of course, the first, the chloroplast, is where photosynthesis occurs. However, the two light-driven sources of cellular energy are quite different and, in many ways, complement one another. Photosynthesis works best at low to intermediate light intensities, and is inhibited at high light, while PPR is predicted to work best at high light intensities. And photosynthetic rates decrease with decreasing temperature and are subject to iron limitation, while PPR photochemistry is not directly limited by iron, and is unaffected by temperature. Thus, the two phototrophic systems are favored under different sets of conditions. Placing PPR in the vacuole may benefit this complementary situation where one or the other phototrophic process is favored depending on the environmental conditions. And here, the presence of PPR in the vacuole may be especially beneficial for growth and survival as that organelle often acts as a storage site for cellular energy in the form of the phosphate anhydride bonds of polyphosphates. We hypothesize that this complementary behavior, along with the ability to store excess energy produced by PPR in the vacuole as high energy polyphosphates, represents an important survival strategy in the ocean, where light, iron levels, and temperature vary widely on a variety of spatial and temporal scales.

Contribution of Smoke Aerosols From Wildfires in Indo‐China Peninsula to the Western Pacific Ocean Carbon Sink

AbstractSmoke aerosols from wildfires play a vital role in marine ecosystems and the ocean carbon cycle. This study analyzed a severe wildfires incident occurred in the Indo‐China Peninsula on 26 March 2019, and assessed its impact on ocean carbon sink. The satellite observations showed that the wildfires covered an area of about 56% of the entire peninsula, burning an area of 1.17 × 106 km2. Consequently, the wildfires discharged about 0.43 × 109 kg carbon. Moreover, a substantial amount of smoke released by severe wildfires was carried downstream by the wind and reached the western Pacific Ocean within 2 days. These smoke aerosols contribute to the ocean carbon sink through the mechanisms of the biological pump and their own carbon deposition. Model simulation results showed smoke aerosols contribute 6.44 × 104 kg black carbon to surface ocean by their own carbon deposition. Moreover, during the period of smoke aerosol's deposition, the flourishing growth of phytoplankton resulted in an increased carbon export of 0.25 ± 0.09 × 109 kg carbon (the biological pump mechanism), which represented approximately 57.97% ± 20.30% of the carbon emissions originating from the wildfire‐affected source region. Our research showed a positive impact of large wildfires on the ocean carbon sink, indicating the potential of smoke aerosols to alleviate the climate pressures resulting from carbon dioxide released by wildfires through improving the capacity of ocean carbon sink.

Modelling phytoplankton-virus interactions: phytoplankton blooms and lytic virus transmission

A dynamic reaction–diffusion model of four variables is proposed to describe the spread of lytic viruses among phytoplankton in a poorly mixed aquatic environment. The basic ecological reproductive index for phytoplankton invasion and the basic reproduction number for virus transmission are derived to characterize the phytoplankton growth and virus transmission dynamics. The theoretical and numerical results from the model show that the spread of lytic viruses effectively controls phytoplankton blooms. This validates the observations and experimental results of Emiliana huxleyi-lytic virus interactions. The studies also indicate that the lytic virus transmission cannot occur in a low-light or oligotrophic aquatic environment.

The Identification of Upwelling in the Occurrence of Tropical Cyclone Charlotte in the Waters of Southern Java (Case Study: March 17-28, 2022)

Tropical cyclones are one of the meteorological phenomena that can have negative impacts on humans. These impacts can include heavy rainfall leading to floods and even storm surges. However, tropical cyclones can also have positive effects as they can trigger upwelling. Upwelling affects the fisheries aspect because in areas where upwelling occurs, sea surface temperatures are lower, and nutrients (such as phosphate and nitrate) are lifted from deeper layers of the ocean, which can affect the growth of phytoplankton. The objective of this research is to identify and analyze the upwelling phenomenon in the waters south of Java Island during the occurrence of Tropical Cyclone Charlotte. The data used in this research includes the intensity and movement direction of Tropical Cyclone Charlotte from the zoom.earth website and sea surface temperature, chlorophyll-a, mixed layer depth (MLD), and sea level anomaly data from the E.U. Copernicus Marine Service Information (CMEMS)-Global Monitoring and Forecasting Center during March 17-28, 2022. The results of this research indicate that upwelling was not identified during the period before or after the cyclone, as evidenced by sea surface temperatures between 28.5-29.5°C, chlorophyll-α concentration of &lt;0.15 mg/m3, and a sea level rise of 0.1 m

Impact of Dynamic Phytoplankton Stoichiometry on Global Scale Patterns of Nutrient Limitation, Nitrogen Fixation, and Carbon Export

AbstractPhytoplankton stoichiometry modulates the interaction between carbon, nitrogen and phosphorus cycles. Environmentally driven variations in phytoplankton C:N:P can alter biogeochemical cycling compared to expectations under fixed ratios. In fact, the assumption of fixed C:N:P has been linked to Earth System Model (ESM) biases and potential misrepresentation of responses to future change. Here we integrate key elements of the Adaptive Trait Optimization Model (ATOM) for phytoplankton stoichiometry with the Carbon, Ocean Biogeochemistry and Lower Trophics (COBALT) ocean biogeochemical model. Within a series of global ocean‐ice‐ecosystem retrospective simulations, ATOM‐COBALT reproduced observations of phytoplankton N:P, and compared to static ratios, exhibited reduced phytoplankton P‐limitation, enhanced N‐fixation, and increased low‐latitude export, improving consistency with observations and highlighting the biogeochemical implications of dynamic N:P. We applied ATOM‐COBALT to explore the impacts of different physiological mechanisms hypothesized to underlie N:P variation, finding that two mechanisms together drove the observed patterns: proportionality of P‐rich ribosomes in phytoplankton cells to growth rates and reductions in P‐storage during scarcity. A third mechanism which linked temperature with phytoplankton biomass allocations to non‐ribosomal proteins, led only to relatively modest impacts because this mechanism decreased the temperature dependence of phytoplankton growth rates, compensating for changes in N:P. We find that there are quantitative response differences that associate distinctive biogeochemical footprints with each mechanism, which are most apparent in highly productive low‐latitude regions. These results suggest that variable phytoplankton N:P makes phytoplankton productivity and export resilient to environmental changes, and support further research on the physiological and environmental drivers of phytoplankton stoichiometry and biogeochemical role.

Atlas of phytoplankton phenology indices in selected Eastern Mediterranean marine ecosystems.

Phytoplankton is a fundamental component of marine food webs and play a crucial role in marine ecosystem functioning. The phenology (timing of growth) of these microscopic algae is an important ecological indicator that can be utilized to observe its seasonal dynamics, and assess its response to environmental perturbations. Ocean colour remote sensing is currently the only means of obtaining synoptic estimates of chlorophyll-a (a proxy of phytoplankton biomass) at high temporal and spatial resolution, enabling the calculation of phenology metrics. However, ocean colour observations have acknowledged weaknesses compromising its reliability, while the scarcity of long-term in situ data has impeded the validation of satellite-derived phenology estimates. To address this issue, we compared one of the longest available in situ time series (20 years) of chlorophyll-a concentrations in the Eastern Mediterranean Sea (EMS), along with concurrent remotely-sensed observations. The comparison revealed a marked coherence between the two datasets, indicating the capability of satellite-based measurements in accurately capturing the phytoplankton seasonality and phenology metrics (i.e., timing of initiation, duration, peak and termination) in the studied area. Furthermore, by studying and validating these metrics we constructed a satellite-derived phytoplankton phenology atlas, reporting in detail the seasonal patterns in several sub-regions in coastal and open seas over the EMS. The open waters host higher concentrations from late October to April, with maximum levels recorded during February and lowest during the summer period. The phytoplankton growth over the Northern Aegean Sea appeared to initiate at least a month later than the rest of the EMS (initiating in late November and terminating in late May). The coastal waters and enclosed gulfs (such as Amvrakikos and Maliakos), exhibit a distinct seasonal pattern with consistently higher levels of chlorophyll-a and prolonged growth period compared to the open seas. The proposed phenology atlas represents a useful resource for monitoring phytoplankton growth periods in the EMS, supporting water quality management practices, while enhancing our current comprehension on the relationships between phytoplankton biomass and higher trophic levels (as a food source).

The effects of additional food and environmental stochasticity on the asymptotic properties of a nutrient–phytoplankton model

In this paper, we investigate the deterministic and stochastic features of an additional food provided nutrient–phytoplankton model, and discuss the effects of additional food and environmental stochasticity (noise) on its asymptotic properties. A detailed global dynamics analysis of the deterministic model is firstly provided, and then the critical conditions for persistence and extinction of phytoplankton is found. For the stochastic model, we establish some sufficient conditions for the exponential extinction of phytoplankton and the existence of stationary distribution. By solving the Fokker–Planck equation, the explicit expression of local probability density of the stochastic model is further derived. With the help of probability density, we obtain the mean, variance, skewness and kurtosis of the biomass of phytoplankton, and numerically find the additional food and stochasticity have complex effects on these statistical properties. Both theoretical and numerical findings indicate that the additional food source is favorable for the growth of phytoplankton, whereas the stochasticity is adverse to its growth. Moreover, the additional food source and stochasticity play important roles in the formation of phytoplankton bloom.

Bloom dynamics under the effects of periodic driving forces

Phytoplankton bloom received considerable attention for many decades. Different approaches have been used to explain the bloom phenomena. In this paper, we study a Nutrient–Phytoplankton–Zooplankton (NPZ) model consisting of a periodic driving force in the growth rate of phytoplankton due to solar radiation and analyse the dynamics of the corresponding autonomous and non-autonomous systems in different parametric regions. Then we introduce a novel aspect to extend the model by incorporating another periodic driving force into the growth term of the phytoplankton due to sea surface temperature (SST), a key point of innovation. Temperature dependency of the maximum growth rate (μmax) of the phytoplankton is modelled by the well-known Q10 formulation: μmax=μ0∗(Q10)T/10, where μ0 is maximum growth at 0oC. Stability conditions for all three equilibrium points are expressed in terms of the new parameter ρ2, which appears due to the incorporation of periodic driving forces. System dynamics is explored through a detailed bifurcation analysis, both mathematically and numerically, with respect to the light and temperature dependent phytoplankton growth response. Bloom phenomenon is explained by the saddle point bloom mechanism even when the co-existing equilibrium point does not exist for some values of ρ2. Solar radiation and SST are modelled using sinusoidal functions constructed from satellite data. Our results of the proposed model describe the initiation of the phytoplankton bloom better than an existing model for the region 25–35° W, 40–45° N of the North Atlantic Ocean. An improvement of 14 days (approximately) is observed in the bloom initiation time. The rate of change method (ROC) is applied to predict the bloom initiation.

Transcriptome response of diatom Skeletonema marinoi to lower temperature

Temperature is an important environmental factor for phytoplankton. Phytoplankton growth, metabolism, biodiversity, productivity, and distribution are influenced by temperature-driven nutrient stratification and mixing, as well as species’ optimal growth temperatures. There have been a number of studies focused on physiological and biochemical mechanisms of environmental–biological interactions in diatoms, yet the underlying transcriptional regulators remain limited. Here, we performed an RNA-seq-based gene expression analysis to explore the Skeletonema marinoi (isolated from Jiaozhou Bay of Qingdao, 36.13°N, 120.16°E on July 5th, 2013) cellular responses induced by low temperature (12 °C). Digital gene expression profiling of S. marinoi generated 20,319 unigenes, of which 573 differentially expressed genes appeared in the low-temperature treatment group. According to GO and KEGG enrichment analysis, different genes were involved in ten metabolic and biosynthesis pathways: ribosome, lipid, porphyrin, and chlorophyll metabolism showed strong transcriptional cold tolerances. The regulation of genes related to translation processes (e.g., pentatricopeptide repeat), fatty acid metabolism (e.g., acyl-CoA synthetase), and photosynthesis (e.g., porphyrin enzymes) provides new molecular-level insight into cold stress responses in eukaryotic marine phytoplankton. Our study suggests that this Skeletonema species could be a potential candidate for understanding the fate of thermo-sensitive diatom communities and oceanic ecosystems facing climate change.

Benthic assemblages in relation to planktonic assemblages in a eutrophic, thermally stratified reservoir

Few studies have explored the relationship between benthic and planktonic assemblages in reservoirs, despite their role in food chains and maintaining ecological functions. Macroinvertebrates play a crucial role in food webs and contribute about 42% of whole-lake secondary productivity. Therefore, their status is vital for maintaining good ecological functions. In this study, we selected Nanwan Reservoir, a eutrophic thermally stratified reservoir in China, to evaluate the community of macroinvertebrates in different seasons and explore the relationships between macroinvertebrate assemblages and different planktonic groups, including phytoplankton, protozoans, rotifers, and planktonic crustaceans. Cyanophyta and Bacillariophyta dominated the phytoplankton assemblages, and their proportions varied with seasons. The bad community of macroinvertebrates in summer, autumn, and winter could be attributed to the excessive growth of Cyanophyta or other phytoplankton. Based on the results of partial least squares regression, an algorithm used for prediction, we determined that the excessive growth of algae, crustaceans, protozoans, and Cyanophyta was detrimental to the development of macroinvertebrates, and was indicative of high nutrient loads in the reservoir. However, the growth of Bacillariophyta, Chrysophyta, and Euglenophyta was advantageous to macroinvertebrate assemblages and indicated a better ecological condition of the reservoir. The results of partial least squares structural equation modeling demonstrated close associations between phytoplankton and both zooplankton and macroinvertebrates, indicating their interdependence in this reservoir system. Our study results imply that the status of macroinvertebrates can be predicted by the abundance of some planktonic assemblages, and cost savings from selecting one of the planktonic groups to monitor ecological conditions could be accomplished in future studies.

Distribution of polysaccharidic and proteinaceous gel−like particles in three cyclonic eddies in the Eastern Tropical North Atlantic

Transparent exopolymer particles (TEP) and Coomassie stainable particles (CSP), two prominent classes of gel−like particles in the ocean primarily produced by phytoplankton, play crucial roles in ecological and biogeochemical processes, influencing microbial nutrition, growth, and particle aggregation. The distribution of these particles is intricately linked to the spatiotemporal dynamics of phytoplankton. Mesoscale cyclonic eddies (CEs) are known to stimulate phytoplankton growth and influence particle transport, but their effects on TEP and CSP remain to be determined. In the Eastern Tropical North Atlantic (ETNA), we examined three CEs: one off the Mauritanian coast during summer (Mau), one offshore during winter (Sal), and another near Brava island during winter. Mau and Brava CEs were in their intensification/maturity phase, while the Sal CE was in its decay phase. Both TEP and CSP concentrations correlated with primary productivity, but TEP increased with chlorophyll−a concentration, whereas elevated CSP coincided also with the highest abundance of pico−nanophytoplankton (&amp;lt;20 µm), mainly Synechococcus. Both gels exhibited a positive correlation with bacterial biomass production, indicating their consumption by heterotrophic bacteria. TEP total area in the epipelagic waters of all CEs (Mau, Brava, and Sal) was elevated compared to surrounding waters, with on average 4, 2.5, and 1.6−fold higher values, respectively. However, no significant difference in TEP size distribution was observed within any CEs and their surroundings. Similarly, CSP total area increased in the epipelagic waters of Mau and Brava CEs, with on average 5 and 2.4−fold higher values, respectively, compared to surrounding waters. CSP particles were notably larger in these two eddies, while the Sal CE showed no significant difference from surrounding waters in CSP abundance and size. Overall, TEP and CSP exhibited distinct responses to CEs, with increased concentrations during their intensification/maturation stage and remineralization dominating during their decaying stage.

Exploring the relationship between sea ice and phytoplankton growth in the Weddell Gyre using satellite and Argo float data

Abstract. Some of the highest rates of primary production across the Southern Ocean occur in the seasonal ice zone (SIZ), making this a prominent area of importance for both local ecosystems and the global carbon cycle. There, the annual advance and retreat of ice impacts light and nutrient availability, as well as the circulation and stratification, thereby imposing a dominant control on phytoplankton growth. In this study, the drivers of variability in phytoplankton growth between 2002–2020 in the Weddell Gyre SIZ were assessed using satellite net primary production (NPP) products alongside chlorophyll-a and particulate organic carbon (POC) data from autonomous biogeochemical floats. Although the highest daily rates of NPP are consistently observed in the continental shelf region (water depths shallower than 2000 m), the open-ocean region's larger size and longer ice-free season mean that it dominates biological carbon uptake within the Weddell Gyre, accounting for 93 %–96 % of the basin's total annual NPP. Variability in the summer maximum ice-free area is the strongest predictor of inter-annual variability in total NPP across the Weddell Gyre, with greater ice-free area resulting in greater annual NPP, explaining nearly half of the variance (R2=42 %). In the shelf region, the return of sea ice cover controls the end of the productive season. In the open ocean, however, both satellite NPP and float data show that a decline in NPP occurs before the end of the ice-free season (∼ 80 to 130 d after sea ice retreat). Evidence of concurrent increases in float-observed chlorophyll-a and POC suggest that later in the summer season additional factors such as micro-nutrient availability or top-down controls (e.g. grazing) could be limiting NPP. These results indicate that in a warmer and more ice-free Weddell Gyre, notwithstanding compensating changes in nutrient supply, NPP is likely to be enhanced only up to a certain limit of ice-free days.

Importance of microzooplankton for sustaining high mesozooplankton biomass during post-bloom period in the Oyashio region of the western subarctic Pacific

We investigated the plankton community structure and biomass during the post-bloom season in the Oyashio region of the western subarctic Pacific, including pico-, nano-, microplankton and mesozooplankton. We found that the nitrate, phosphate and silicic acid concentrations remained high at >4.2 μM, >0.77 μM and >7.1 μM, respectively, in the euphotic layer at almost all sampling stations, but that the chlorophyll a concentrations were low (<3 µg Chl. a l−1). These findings indicate high nutrient and low chlorophyll (HNLC)-like conditions. In the phytoplankton community, pennate diatoms, the larger subpopulation of pico-sized eukaryotic phytoplankton, and nano-flagellates substantially contributed to the low biomass of the chain-forming centric diatoms that mainly comprised the spring phytoplankton bloom. The microzooplankton biomass was 2.7–4.4 fold greater than the phytoplankton biomass in the surface layer. Naked ciliates substantially contributed to the microzooplankton community (40 %–87 %). The naked ciliate growth rates during our in situ bottle incubation experiments were significantly greater than the maximum growth rates as calculated from cell volume and water temperature. The mesozooplankton biomass was mainly composed of krill and copepods and was 5.9–9.3 fold higher than the microzooplankton biomass. This inverted biomass pyramid with relatively low microzooplankton and high mesozooplankton biomass may be explained by the high production and growth rates of the microzooplankton. The ratio of phytoplankton growth (µ, d−1) to grazing mortality (m, d−1) by microzooplankton were relatively low at 0.26–0.44 m/µ in our dilution experiments. These low values indicate that microzooplankton grazing does not regulate phytoplankton growth and suggests that microzooplankton feed on an alternative nutritional source, such as heterotrophic prey items, or mixotrophy to fulfill their growth needs. Additional research is needed during the post-bloom period to further evaluate the mechanisms that sustain microzooplankton dominance and production in the Oyashio region under the HNLC-like conditions, especially for naked ciliates.

Oceanographic factors determining the distribution of nutrients and primary production in the subpolar Southern Ocean

To investigate the spatial distributions and determinants of nutrient concentrations, we measured NO3−+NO2−, PO43−, and Si(OH)4 concentrations in the eastern Indian Ocean sector of the Antarctic Ocean (80 − 150°E, south of 60°S) between December 2018 and February 2019. In the region influenced by the Antarctic Circumpolar Current, nutrient concentrations were increased by nutrients supplied from the deep layer and by organic matter decomposition and remineralization within the seasonal pycnocline after the development of strong stratification. Strong stratification also enhanced phytoplankton growth and nutrient consumption by photosynthesis. In contrast, in the subpolar region, nutrient concentrations were increased by nutrients supplied by brine discharged during sea ice formation and decreased by dilution with sea ice meltwater. Although high salinity in the surface and subsurface layers corresponded well to upwelling areas around subpolar subgyres, high salinity was not necessarily correlated with nutrient concentrations. We estimated primary production both from in situ nutrient data and from satellite-acquired chlorophyll-a data. According to both estimation methods, primary production was high in the subpolar region, especially around 120 − 130°E. However, nutrient-based estimation also showed high production in coastal areas where, because of sea ice and cloud cover, estimation based on satellite data was not possible. To understand primary production in seasonal ice areas, the best estimation method should be selected for the research goals or multiple methods should be used in combination.