Subsurface Chlorophyll Maximum Layer Research Articles

Revealing the seasonal cycles of Arctic phytoplankton: insights from year-round chlorophyll monitoring

Rapid Arctic Ocean warming has caused severe sea ice decline, impacting light distribution, phytoplankton blooms, and primary production. We investigated Arctic phytoplankton bloom timing using continuous chlorophyll-a fluorescence data obtained from three Korea Arctic Mooring Systems (KAMSs) deployed north of the East Siberian Sea (KAMS1), north of the Chukchi Sea (KAMS2), and the middle of the Northwind Ridge (KAMS4). Our findings revealed that the bloom initiation times were June 4 (±28 d) in KAMS1, June 24 in KAMS2, and May 21 (±6 d) in KAMS4, when the sea ice concentration (SIC) was >90% and the ice thickness was 1–2 m, indicating that the under-ice phytoplankton blooms (UIBs) developed 1–2 months before the sea ice retreated (mid-July, when SIC was <80%). Peak bloom and termination times were consistently observed in early August and mid-October, respectively. The average phytoplankton bloom lasted for approximately four months, longer than the open water periods at the mooring sites. However, the timing of the phytoplankton blooms from the biogeochemical model-based reconstructions was, on average, 6–10 weeks later than that deduced from the observed data. Furthermore, the maximum chlorophyll-a concentration observed during the bloom peak was approximately ten-times higher than that indicated by the biogeochemical model-based reconstructions (1.81 vs. 0.17 mg −3). The differences in chlorophyll-a concentrations and bloom timings indicate that biogeochemical models remain insufficient for simulating the phytoplankton dynamics of the Arctic Ocean, such as UIBs and the subsurface chlorophyll maximum layer. Based on the continuously observed chlorophyll-a concentrations, we gained a precise understanding of the seasonal cycles of Arctic phytoplankton, including UIBs. These valuable data will contribute to improving the accuracy of biogeochemical models of the Arctic Ocean.

Environmental characteristics and major factors controlling chlorophyll a in three seamounts in the Subtropical Western Pacific Ocean

Seamounts interacting with oceanic currents created “seamount effect”, which could enhance the primary productivity in the euphotic zone. However, a large number of seamounts have not been investigated. In this study, the sectional distributions of temperature, salinity, dissolved inorganic nutrients, dissolved oxygen (DO), pH and chlorophyll a (Chl a) in three seamounts in the Subtropical Western Pacific Ocean (SWPO), named NA, NLG and MP4, were investigated. Consistent with no uplifts of isotherms and isohalines above three seamounts, the uplifts of the concentrations of dissolved inorganic nutrients and Chl a were unobvious, indicating that there was no “seamount effect” in these seamounts. Comparing with NA and MP4 seamounts, NLG seamount had higher salinity and lower temperature. However, the concentrations of inorganic nutrients and Chl a from surface to the subsurface chlorophyll maximum layer of NA and MP4 seamount were higher than those of NLG seamount. Meanwhile, due to the presence of tropic storm and cold eddy during the investigated period, the hydrological conditions there became more complex. Furthermore, Generalized Additive Models (GAMs) were used to infer the underlying factors controlling bottom-up effect on phytoplankton biomass in three seamounts. This study investigated the mechanism about the bottom-up effect on phytoplankton in seamount environment and provided basic information for future research on seamount ecosystem.

Assessment of global detection capability of oceanographic lidar

The profiling of marine bio-optical properties by oceanographic lidar is of great significance to the monitoring of the marine environment and the study of the global carbon cycle. In this study, the global detection capability of oceanographic lidar is analyzed with different specifications. To evaluate the detection performance, a simulator on multi-platform is developed. Based on the quasi-single elastic scattering approximation lidar equation and the bio-optical models, the simulator can generate different kinds of outputs, such as lidar signal profiles, signal-to-noise ratio profiles, maximum detectable depth distributions. The optical properties profiles of the global ocean could be derived from the global chlorophyll-a concentration profile data set, which is used as the input to the simulator. The simulation results are compared with the measurements from airborne lidar in the South China Sea, which verifies the reliability of the simulator. The global detection capability of spaceborne lidar with different power aperture products is simulated, and the influence of laser wavelength is analyzed. The results show that, for the simulated spaceborne lidar with typical specifications at 486 nm, there are probabilities ranging from 0.66 to 0.85 that the upper boundary of the thermocline can be detected on a global scale. In the cleanest open ocean, the lidar has a good capability to penetrate the upper boundary of the thermocline, while it cannot penetrate the subsurface chlorophyll maximum layers, in which the global penetration ratio is 0.93 under the simulation specifications at 486 nm. Finally, the influence of repetition frequency is analyzed, and the results show that reducing the pulse repetition frequency at the same average power can effectively reduce the background light of cumulative profiles and improve the signal-to-noise ratio.

Subsurface phytoplankton responses to ocean eddies can run counter to satellite-based inference from surface properties in subtropical gyres

Phytoplankton contribute approximately half of the Earth’s primary production, playing a crucial role in marine ecosystem functioning and global carbon cycling. However, our understanding of how ocean dynamics regulate the distribution of phytoplankton remains limited due to the scarcity of depth-resolved observations. Here, we compared the simultaneous measurements of two biogeochemical-Argo floats that tracked a cyclonic eddy (CE) and an anticyclonic eddy (AE) in the subtropical South Indian Ocean for a period of 3 months. The observations revealed a decoupling of subsurface phytoplankton biomass from surface chlorophyll. The CE exhibited a subsurface phytoplankton bloom initiated by the springtime shoaling of mixed layer to euphotic layer, accompanied by a counter-intuitive decrease in surface chlorophyll due to physiological adaptation. In comparison, the AE exhibited vertically homogeneous phytoplankton biomass, with a weak subsurface chlorophyll maximum layer, as the mixed layer remained at a deep level in the spring. Despite a higher concentration of surface chlorophyll in the AE than in the CE, subsurface and depth-integrated phytoplankton chlorophyll, biomass, and productivity computed with a bio-optical model are greater in the CE than in the AE. Statistics on global biogeochemical-Argo measurements over the past decade confirm the generality of this phenomenon in subtropical gyres. Our findings demonstrate that high concentrations of chlorophyll near the surface in these regions are not necessary indicative of high phytoplankton abundance at the subsurface layer. The results have important implications for inferring subsurface phytoplankton abundance and productivity from satellite measurements.

Distinct vertical behavior of key Arctic copepods following the midnight sun period in the East Siberian continental margin region, Arctic Ocean

Diel vertical migration (DVM) of zooplankton plays a vital role in biological carbon pump and food web interactions. However, there is considerable debate about the DVM of zooplankton in response to environmental changes in the Arctic Ocean. We investigated DVM behavior in the key Arctic copepods Calanus glacialis, Calanus hyperboreus, and Metridia longa following the midnight sun period in the East Siberian continental margin region. The two Calanus species showed non-DVM behaviors, whereas M. longa showed a typical DVM pattern consistent with the solar radiation cycle. Additionally, these species showed different vertical distributions. Calanus glacialis was distributed at depths above 20 m in the warm fresh water, where the highest density gradient was observed. Calanus hyperboreus was distributed at depths between 30 and 55 m in the cold salty water, where a high contribution of micro phytoplankton and the subsurface chlorophyll maximum (SCM) layer were observed. M. longa was found across a broader range of temperature and salinity than both Calanus species, and it was distributed in the upper water column, where the SCM layer was observed at night and at depths between 100 and 135 m in the daytime. These results imply that M. longa can be well adapted to the changing Arctic Ocean environment, where sea ice loss and ocean warming are ongoing, whereas C. hyperboreus can be the most vulnerable to these changes. These findings provide important information for understanding variations in the vertical distributions of key copepod species in the rapidly changing Arctic marine environment.

Development of subsurface chlorophyll maximum layer and its contribution to the primary productivity of water column in a large subtropical reservoir

The phenomenon of subsurface chlorophyll maximum (SCM) layer emerging at a certain water depth is commonly found in stratified water bodies. Also, it is a crucial contributing region to the primary productivity of the water column. Currently, there is a lack of concern about the occurrence of SCM phenomena in studies targeting inland water bodies such as natural lakes and artificial reservoirs. This led to a significant underestimation of the level of primary productivity in these water bodies and their trophic state. In this study, a subtropical reservoir (the Xinanjiang Reservoir, XAJR) was investigated, to understand the characteristics of SCM layer in deep-large reservoir and its contribution to the primary productivity of the water column. Water sampling were conducted from September 2020 to August 2021, and in September 2022. Buoy station data for this reservoir between 2019 and 2021 were also collected. Based on the detailed observations of the water column profile in riverine area (X1), transitional area (X2), and central area (X3 and X4) of this reservoir, it was found that there was an obvious SCM phenomenon, which was closely related to the characteristics of seasonal thermal stratification. The SCM layer of XAJR appeared at depth around 3–5 m underwater from May to August, and as the thermal stratification strength increased, so did the depth and thickness of the SCM layer. It was estimated that gross primary productivity of euphotic layer of XAJR ranged from 347.9 to 4508.6 mgC·m−2·d−1. The average primary productivity level of the SCM layer reached 1411.7mgC·m−2·d−1, accounting for about 40–90% of the gross primary productivity of euphotic layer. This study contributes to a better understanding of the factors influencing changes in the development of the SCM layer in large reservoirs, as well as its critical role in the inland water carbon cycle.

Trophic Positions of Sympatric Copepods across the Subpolar Front of the East Sea during Spring: A Stable Isotope Approach

We investigated the trophic relationship between particulate organic matter (POM) and sympatric copepods within the epipelagic zone (~200 m depth) in the East Sea during spring based on stable isotope analysis (SIA). The SIA indicated that interspecific differences in the prey size and vertical segregation of feeding migration range among copepods may promote niche partitioning among sympatric copepods in each region of the subpolar front (SPF). Additionally, our results showed remarkable differences in the copepod community structure and resource utilization across the SPF. The south region of the East Sea showed higher species richness of copepods than the north region, while copepods that fed mainly on POM in the surface and subsurface chlorophyll maximum layers showed smaller body sizes in the south region. These results revealed that the food chain between primary producers and higher trophic levels was longer in the south region than in the north region. Additionally, δ13C and δ15N values of copepods increased gradually with the body size increase whereas δ15N values in the north region showed the reverse trend. Latter results could be attributed to the consumption of deep-layer POM in small copepods. Therefore, we suggest that northward shifts in the distribution of copepods under global warming may decrease energy efficiency in the pelagic ecosystem of the East Sea.

Appendicularians and marine snowin situvertical distribution in Argentinean Patagonia

AbstractDetailed in situ vertical and temporal distribution of appendicularians, marine snow, fecal pellets, nano- and microplankton were recorded simultaneously with environmental data in the San Jorge Gulf, Argentinean Patagonia (45°–47°S). Data were taken at a fixed station over 36 h in February 2014 with an autonomous Video Plankton Recorder and a FlowCAM®. The water column was thermally stratified with a pycnocline at ~ 40 m. Appendicularians dominated in the upper 65 m with a condensed pattern above the pycnocline at high chlorophyll a concentrations, matching the subsurface chlorophyll maximum layer at ~ 20 m. Our results suggest the absence of vertical migration of appendicularians. Marine snow, strongly correlated with appendicularians, showed high concentrations above the pycnocline, whereas fecal pellets from krill were distributed throughout the water column. Discarded houses of appendicularians or their mucus fragments were the main components of marine snow aggregates, with phytoplankton, detritus and krill pellets also contributing. Nanoplankton dominated over microplankton, with vertical distribution patterns that might depend on local grazing pressure and advective processes. Our study, the first one in the region using underwater imagery, emphasizes the leading contribution of appendicularians to marine snow aggregates in the San Jorge Gulf and their potential implications in the bentho-pelagic coupling.

Investigating the Effects of Super Typhoon HAGIBIS in the Northwest Pacific Ocean Using Multiple Observational Data

Various multi-source observational platforms have enabled the exploration of ocean dynamics in the Northwest Pacific Ocean (NPO). This study investigated daily oceanic variables in response to the combined effect of the 2019 super typhoon HAGIBIS and the Kuroshio current meander (KCM), which has caused economic, ecological, and climatic changes in the NPO since August 2017. During HAGIBIS, the six-hourly wind speed data estimated a wind stress power (Pw) which strengthened around the right and left semicircles of the typhoon, and an Ekman pumping velocity (EPV) which intensified at the center of the typhoon track. As a result, firstly, the sea temperature (ST) decreased along a boundary with a high EPV and a strong cyclonic eddy area, and the mixed layer depth (MLD) was shallow. Secondly, a low sea salinity (SS) concentration showed another area where heavy rain fell on the left side of the typhoon track. Phytoplankton bloom (PB) occurred with a large concentration of chlorophyll a (0.641 mg/m3) over a wide extent (56,615 km2; above 0.5 mg/m3) after one day of HAGIBIS. An analysis of a favorable environment of the PB’s growth determined the cause of the PB, and a shift of the subsurface chlorophyll maximum layer (SCML; above 0.7 mg/m3) was estimated by comprehensive impact analysis. This study may contribute to understanding different individually-estimated physical and biological mechanisms and predicting the recurrence of ocean anomalies.

Phytoplankton photophysiology varies depending on nitrogen and light availability at the subsurface chlorophyll maximum in the northern Chukchi Sea

Vertical distributions of phytoplankton in the Arctic Ocean are characterized by a very narrow subsurface chlorophyll maximum (SCM) layer formed every summer after the sea ice retreats. Despite the prevalence of this narrow SCM layer, phytoplankton photosynthetic response to climate change remains to be elucidated. Here, we examined the photophysiological properties of phytoplankton in the SCM layer in the northern Chukchi Sea during the summers of 2015–2018. There was a significant difference in the SCM depth between the northwestern and northeastern Chukchi Sea determined by the distribution of Pacific Summer Water (PSW) around the SCM layer (34 ± 14 m vs. 49 ± 10 m, respectively). The maximum quantum yield of photochemistry in photosystem II (Fv/Fm) in the SCM phytoplankton was high (Fv/Fm ≥ 0.54) and similar in both regions until 2016; however, since then, Fv/Fm in the northeastern Chukchi Sea has decreased by approximately 10%. This decrease was accompanied by a marked decrease in the fraction of microplankton, which are known to be susceptible to nutrient limitation. This result suggests a reduction in nitrogen availability in the SCM layer in the northeastern Chukchi Sea. Meanwhile, the maximum electron transfer rate (ETRmax) did not have a significant relationship with the nitrogen availability and phytoplankton community size structure in the SCM layer; however the improved light conditions (with an approximately two-fold increase in the relative ratio of surface PAR reaching the SCM layer) increased ETRmax by up to 30% in the SCM phytoplankton in the northwestern Chukchi Sea. Therefore, these results provide a better understanding of how changes in nitrogen and light availability could affect phytoplankton photosynthesis and primary production in the Arctic Ocean.

Insignificant effects of eddies and typhoons on the biogeochemistry of the tropical northwest Pacific Ocean

We investigated the effects of eddies and typhoons on the biogeochemistry of the tropical northwest Pacific by examining the distribution of nutrients, dissolved oxygen (DO), chlorophyll-a (Chl-a), gross primary production (GPP), dissolved organic carbon (DOC), and fluorescent dissolved organic matter (FDOM). Water samples were collected from anticyclonic and cyclonic eddies in September 2019 and 2020, and before and after the passage of a Category 2 typhoon in 2019. The study region was characterized by a deep nitracline (~150 m), which was deeper than both the pycnocline (~50 m) and the FDOM-depleted layer (~75 m). A subsurface chlorophyll maximum layer was observed at 100–150 m depth. No clear differences in Chl-a, DO, GPP, DOC, and FDOM were observed for the anticyclonic and cyclonic eddies, indicating that the eddies did not have a significant influence on biological production. Similarly, there were no discernable changes in Chl-a concentrations or other biogeochemical parameters after the passage of the typhoon, which induced water mixing to a depth of ~60 m. We conjecture that the nutrient-depleted layer was too deep for any eddy- or typhoon-induced vertical mixing to cause upwelling of nutrients to the euphotic zone. Our results imply that the disturbances caused by mesoscale processes in the upper layer of the highly oligotrophic northwest Pacific may have a smaller effect than in oceans in other parts of the world.

Seasonal Variability of Phytoplankton Biomass Revealed by Satellite and BGC‐Argo Data in the Central Tropical Indian Ocean

AbstractThe variability of phytoplankton biomass related to ocean's biological carbon pump in the central tropical Indian Ocean is not fully understood. Using satellite and biogeochemical Argo data, we found that phytoplankton biomass exhibits significantly different and even opposite changes at the near‐surface (0–10 m) and subsurface layer (50–100 m). Results showed that the shoaling/deepening of the mixed layer and thermocline dominated the phytoplankton biomass change in different layers. In February–May, the shallow mixed‐layer depth (MLD) exacerbated the oligotrophic environment in the mixed layer, thus resulting in low near‐surface phytoplankton biomass (Chl a &lt; 0.1 mg m−3). However, strengthened wind stress and surface cooling induced deep vertical mixing (MLD &gt; 50 m), hence increasing the supply of both nutrients and phytoplankton from the subsurface to maintain the near‐surface phytoplankton blooms (Chl a &gt; 0.2 mg m−3) in June–October. Below the mixed layer, wind‐induced upwelling/downwelling modulated the thermocline, which significantly affects the supply of nutrients for phytoplankton growth in the subsurface layer. The shallower thermocline during February and March provided favorable conditions for the subsurface blooms (Chl a &gt; 0.5 mg m−3) with a prominent subsurface chlorophyll maximum layer (SCML). Conversely, the deeper thermocline in June and July was related to the lowest phytoplankton biomass with the disappearance of the SCML. These results may help explain the response of vertical phytoplankton to changes driven by multiple atmospheric and physical forcing factors in future climate‐change scenarios.

Ecological Responses of Core Phytoplankton by Latitudinal Differences in the Arctic Ocean in Late Summer Revealed by 18S rDNA Metabarcoding

Limited information is available regarding the phytoplankton communities in the Arctic Ocean, especially in the Chukchi Sea. We conducted research cruises in the Arctic Ocean in the late summers of 2015, 2016, 2018, and 2019, and used 18S ribosomal DNA metabarcoding to examine the dynamic changes in core phytoplankton. Environmental factors were divided into three Groups: “surface layers in low latitude”, “subsurface chlorophyll maximum layers in low latitude”, and “high latitude.” The phytoplankton community was divided into two Groups. One Group, “the phytoplankton community in lower latitudes affected by the Pacific Influx “, comprised Dinophyta (56.33%), Bacillariophyta (22.28%), and Chlorophyta (20.77%), while the other Group, “the phytoplankton community in higher latitudes affected by the Arctic Sea “, comprised Dinophyta (57.51%), Bacillariophyta (10.71%), and Chlorophyta (27.57%). The common phytoplankton taxa in Group of lower latitudes included 33 operational taxonomic units (OTUs) (99.04%), while the other Group included nine OTUs (98.80%). In these groups, OTUs #005 (Heterocapsa rotundata), #001 (Micromonas pusilla), and #003 (Chaetoceros gelidus) were core OTUs (&amp;gt;5%). These core OTUs were significantly different between the two Groups; OTU #001 and #003 were distributed at high relative abundance and rapidly increased at higher latitudes. OTU #005 showed a relatively high abundance at lower latitudes. Analysis of the indicator species revealed that 34 OTUs in lower latitudes and only one OTU (#003) in higher latitudes were observed to have a strong value of &amp;gt;0.8. Our results indicate that core phytoplankton taxa and community structures by latitudinal differences are highly dependent on different water masses and show their dynamic ecological responses to extreme environmental conditions.

Effects of Mixing and Stratification on the Vertical Distribution and Size Spectrum of Zooplankton on the Shelf and Slope of the Northern South China Sea

The northern shelf and off-shelf regions of the South China Sea (SCS) present a stark contrast between being eutrophic from terrestrial runoffs of nutrients and biota and being oligotrophic with multiple nutrient limitations due to the distance to land sources and stable permanent stratification. The abundance, size, and trophic structures of plankton in the shelf and off-shelf regions were studied in a joint Sino-German cruise conducted between September 1 and 24, 2018. A laser optical plankton counter was mounted on a water sampler-CTD (conductivity–temperature–depth) system for studying the horizontal and vertical distributions of plankton structures in a size range between 0.1 and 35 mm and their relationships with hydrographic and biological features in the northern SCS and its shelf region. Results revealed the subsurface chlorophyll maximum layer (SCM) below the pycnocline and plankton aggregation near the SCM about a depth of 50–60 m. The distributions of small plankton between 0.1 and 0.5 mm were strongly correlated with stratification and SCM compared to those of large plankton. Analyzing the intercept and slope of a normalized biovolume spectrum (NBVS) as an indicator for abundance and size structure of a plankton community, results revealed that in the shelf region, the slopes exhibited no significant vertical variations in the water column regardless of stratification and SCM. In contrast, in the off-shelf stratified water column, the intercepts and slopes were lower and flatter in the surface layer, higher and steeper in the SCM layer, and the lowest and flattest in the deep layer. Stirring by the typhoon also altered both the abundances and size structures of the plankton communities, with significant regional differences. This study elucidates the variances of plankton abundances, distributions, NBVS slopes, and intercepts among different water column structures in both shelf and off-shelf regions of the northern SCS.

Biological production in two contrasted regions of the Mediterranean Sea during the oligotrophic period: an estimate based on the diel cycle of optical properties measured by BioGeoChemical-Argo profiling floats

Abstract. This study assesses marine community production based on the diel variability of bio-optical properties monitored by two BioGeoChemical-Argo (BGC-Argo) floats. Experiments were conducted in two distinct Mediterranean systems, the northwestern Ligurian Sea and the central Ionian Sea, during summer months. We derived particulate organic carbon (POC) stock and gross community production integrated within the surface, euphotic and subsurface chlorophyll maximum (SCM) layers, using an existing approach applied to diel cycle measurements of the particulate beam attenuation (cp) and backscattering (bbp) coefficients. The diel cycle of cp provided a robust proxy for quantifying biological production in both systems; that of bbp was comparatively less robust. Derived primary production estimates vary by a factor of 2 depending upon the choice of the bio-optical relationship that converts the measured optical coefficient to POC, which is thus a critical step to constrain. Our results indicate a substantial contribution to the water column production of the SCM layer (16 %–42 %), which varies largely with the considered system. In the Ligurian Sea, the SCM is a seasonal feature that behaves as a subsurface biomass maximum (SBM) with the ability to respond to episodic abiotic forcing by increasing production. In contrast, in the Ionian Sea, the SCM is permanent, primarily induced by phytoplankton photoacclimation, and contributes moderately to water column production. These results clearly demonstrate the strong potential for transmissometers deployed on BGC-Argo profiling floats to quantify non-intrusively in situ biological production of organic carbon in the water column of stratified oligotrophic systems with recurring or permanent SCMs, which are widespread features in the global ocean.

The Physical-Biogeochemical Responses to a Subsurface Anticyclonic Eddy in the Northwest Pacific

Due to the unique physical processes of mesoscale eddies, the physical and biogeochemical properties within the subsurface anticyclonic eddy (SSAE) and in the surrounding water are distinct. Analyses using satellite and model data have revealed distinct seasonal variations in the central potential density structure of a long-standing SSAE south of Japan; this SSAE exhibits a normal concave isopycnals structure from January to April and a convex lens isopycnals structure from May to December, and these variations may be related to the subduction of low-potential vorticity (PV) mode water. In contrast to the idea of the self-sustained oscillation mechanism, the strength of the SSAE was enhanced due to the eddy kinetic energy provided by dramatic increasing of the positive baroclinic conversion rate during the Kuroshio path transition period from the non-large meander (NLM) path to the large meander (LM) path. Twofold to threefold enhancement of chlorophyll (CHL) was detected in the subsurface CHL maximum layers at the core of the SSAE, and this enhancement was related to the injection of nutrients into the euphotic layer due to winter mixing and the convex of isopycnals. During the period from May to December, elevated CHL and dissolved oxygen (DO) levels and reduced nitrate levels were observed along the periphery of the eddy below the maximum subsurface CHL anomaly depth. The combined result of these two processes: (1) the central downward displaced isopycnals caused by intensified SSAE, and (2) winter mixing deepened to the nutricline due to the thickened mixed layer depth (MLD) and weakened stratification in winter 2017 (during the NLM period) may have led to numerous nutrients and CHL enrichments throughout the mixed layer, thus generating a CHL bloom in the following April. The SSAE intensified in winter 2018 (during the LM period), whereas the shallower MLD and stronger stratification limited the depth of CHL downward displacement.

Contrasting Community Composition and Co-Occurrence Relationships of the Active Pico-Sized Haptophytes in the Surface and Subsurface Chlorophyll Maximum Layers of the Arctic Ocean in Summer.

Haptophytes (Hacrobia: Haptophyta), which can perform phototrophic, phagotrophic, or mixotrophic nutritional modes, are critical for element cycling in a variety of aquatic ecosystems. However, their diversity, particularly in the changing Arctic Ocean (AO), remains largely unknown. In the present study, the biodiversity, community composition, and co-occurrence networks of pico-sized haptophytes in the surface water and subsurface chlorophyll maximum (SCM) layer of the AO were explored. Our results found higher alpha diversity estimates in the surface water compared with in the SCM based on high-throughput sequencing of haptophyte specific 18S rRNA. The community composition of the surface water was significantly different from that of the SCM, and water temperature was identified as the primary factor shaping the community compositions. Prymnesiales (mostly Chrysochromulina), uncultured Prymnesiophyceae, and Phaeocystis dominated the surface water communities, whereas Phaeocystis dominated the SCM communities, followed by Chrysochromulina, uncultured Prymnesiophyceae, and the remaining taxa. The communities of the surface water and SCM layer developed relatively independent modules in the metacommunity network. Nodes in the surface water were more closely connected to one another than those in the SCM. Network stability analysis revealed that surface water networks were more stable than SCM networks. These findings suggest that SCM communities are more susceptible to environmental fluctuations than those in surface water and that future global changes (e.g., global warming) may profoundly influence the development, persistence, and service of SCM in the AO.

Characteristics of subsurface chlorophyll maxima during the boreal summer in the South China Sea with respect to environmental properties

Characterizing the vertical distribution of chlorophyll a in the water column in each oceanic region is crucial for accurate assessment of depth-integrated phytoplankton biomass. In this study, the characteristics of the subsurface chlorophyll maximum (SCM) in the South China Sea (SCS) during the boreal summer were investigated by using bio-optical and hydrological data collected during four cruises from 2008 to 2015. During the boreal summer, a well-developed SCM layer was found to be a prominent feature in the SCS, with the thickness, depth, and magnitude of the SCM exhibiting large spatial variability. The vertical position of the SCM varied between 11 and 99 m, with an average value of 53 m. Light attenuation played a fundamental role in determining the depth (ZSCM) and magnitude of the SCM (Chlmax), as reflected by their relationships with the euphotic zone depth (Zeu). However, because significant positive correlations were found between ZSCM and the depth of potential density at 23 kg m−3 (Zσ=23), physical processes were inferred to be more important in modulating the fluctuation of ZSCM, especially in open-ocean areas. Anticyclonic eddies, which act to deepen the nutricline by means of isopyncnal displacement, may play a role in aggravating nutrient limitation in the SCM layer, which leads to deepening of ZSCM, weakening of Chlmax, and a fall in the total integrated chlorophyll a within the euphotic layer (ChlintZeu). Knowledge of ocean physical conditions and the variability of ZSCM should be taken into consideration to improve the accuracy of ChlintZeu estimates based on the surface chlorophyll a concentration (Chlsurf), especially when Chlsurf is very low. Our assessment of SCM parameters provides the basis for a better understanding and quantification of their role in primary production estimation within the SCS.

Impact of Shifting Subpolar Front on Phytoplankton Dynamics in the Western Margin of East/Japan Sea

A subpolar front (SPF) generated between the East Korea Warm Current (EKWC) and the North Korea Cold Current (NKCC) in the western margin of the East/Japan Sea has shifted northward in recent decades. This study investigated the biomass and composition of the phytoplankton assemblage in relation to hydrological and biogeochemical features in the shallow shelf and slope off the Korean coast from January to June in 2016 and 2017, to determine the mechanistic effects of SPF on spring–summer phytoplankton bloom dynamics. Monthly average depth-integrated chlorophylla(Chla) levels and the contribution of phytoplankton classes revealed bimodal diatom blooms in early spring and summer in the frontal zone. Canonical correspondence analysis showed that the distribution of high Chlawas associated with cold, low-salinity NKCC water in March 2016. No Chlapeak was observed in March 2017 when the warm saline EKWC water mass invaded. These results suggest that the NKCC intrusion acts as a forcing mechanism leading to enhanced phytoplankton biomass in the frontal zone. In contrast, positive correlations of Chlaconcentration with water density and nutrient concentrations suggest that summer blooms were fed by the subsurface chlorophyll maximum (SCM) driven by shoaling of the pycnocline and nitracline. Varying water-column stratification determined the thickness of the SCM layer, driving year-to-year variability in the magnitude of diatom blooms. These findings further suggest that seasonal/interannual variability in the timing of algal blooms affects regional trophodynamics and hence could be an important factor in explaining ecosystem changes in this region.

Effect of El Niño‐Related Warming on Phytoplankton's Vertical Distribution in the Arabian Sea

AbstractIn this study, we analyzed the effect of warming in ecological responses due to seasonal cycle and 2015/2016 El Niño event on the vertical distribution of phytoplankton within the entire euphotic zone (0–100 m) in the eastern Arabian Sea using BGC‐Argo data during 2015–2019. We found that phytoplankton in different layers had different responses to warming. Phytoplankton biomass (defined as the integrated chlorophyll a concentration in this paper) decreased in 0–50 m layer while it increased in 50–100 m layer during spring (March‐June) and 2015/2016 El Niño event. At the same time, the vertical expansion of warming (temperature &gt;28 °C), together with nitrate and silicate co‐limitation, triggered the formation of subsurface chlorophyll maximum layer, which became thinner and deeper during these warming periods. Results from the generalized additive models showed that the monthly 28 °C isotherm can be used as a first approximation to separate vertical water column where warming had opposite signed effects. For a 0–50 m layer of poor nutrients due to the mixed layer shoaling and reduced atmospheric deposition, a negative effect of warming on phytoplankton was seen if the temperature was higher than 28 °C. But, for a 50–100 m layer of rich nutrients, a positive effect of warming on phytoplankton was seen if the temperature was within the range of 25.5–28.0 °C and the light was enhanced. These results shed light on the effect of global warming on subsurface phytoplankton and may help research of the same topic regarding future warming.