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Microzooplankton communities in the coastal Gulf of Alaska (CGOA) were studied during two contrasting years: 2011, with a greatly reduced spring phytoplankton bloom, and 2013, with a robust spring bloom. Other sampling contrasts were season (spring, summer, fall) and region (eastern versus western shelf waters). Ciliates and dinoflagellates comprised nearly all microzooplankton in the ≥ 15 µm size class. Many of the strongest contrasts in the biomass and taxonomic composition of the microzooplankton community were regional. The east had generally lower microzooplankton biomass levels and a greater proportion of ciliates than the west, even in the face of basin-wide seasonal and interannual contrasts. This difference is likely a consequence of the narrower shelf in the east, which leads to a lower productivity environment. Interannual differences in spring bloom intensity were reflected in microzooplankton biomass (higher in spring 2013, especially in the east), while interannual differences in taxonomic composition persisted throughout the year, with a greater representation of ciliates in 2011. Ciliate dominance could reflect adaptations to lower productivity conditions, with many of the largest taxa likely retaining chloroplasts as a strategy (mixotrophy) for survival during times of prey scarcity. Microzooplankton: phytoplankton biomass ratios varied widely over time and space. These ratios indicate that lower productivity regions (east) and seasons (summer) also tend to be locations and times of reduced trophic transfer efficiency from phytoplankton to ciliates and dinoflagellates. Ciliates and dinoflagellates can be preferred prey of mesozooplankton, including dominant CGOA copepod species. Thus multiple mechanisms conspire to reduce the flow of matter and energy to higher trophic levels in low productivity locations and time periods in the CGOA.

The size structure of the phytoplankton community strongly affects energy flow through food webs; hence, monitoring the spatial and temporal dynamics of phytoplankton size structure is crucial to evaluate variability in marine ecosystems. In this study, the spatiotemporal changes in phytoplankton size structure and their impact on benthic macrofaunal distribution were examined in the Pacific Arctic as part of the international Distributed Biological Observatory effort. The exponent of the chlorophyll-a (Chla) size distribution (hereafter, CSD slope), which is an index of synoptic size structure of a phytoplankton community, was estimated based on spectral shape of the phytoplankton absorption coefficient using satellite ocean color data. The CSD slope during the post-bloom period was significantly correlated with sediment Chla concentration, suggesting that the amount of organic carbon input to the seafloor was strongly related to surface phytoplankton size structure after the spring bloom. Average macrofaunal biomass in the Pacific Arctic was persistently high (>20 g C m−2) throughout 2000–2013 in the regional hotspots, and the biomass-based latitudinal centroid of macrofaunal habitat in the region shifted significantly (p

Three cruises were undertaken along a meridional transect in the East/Japan Sea (EJS) in spring (May 2007), summer (July 2009), and fall (October 2012) to determine the geographic variations in phytoplankton biomass and community composition. This study revealed a gradient of surface temperature and a fluctuation of hydrographic conditions along the transect. Although a subpolar front (SPF) formed between the warm- and cold-water masses (37–40°N), no significant differences in phytoplankton biomass and community composition were detected between the southern and northern parts of the EJS. These results disprove our initial hypothesis that different water masses may contain differently structured phytoplankton communities. In the present study, isothermal layers (≤ 12 °C) fluctuated over a depth of 50 m in both warm- and cold-water masses, depending on the SPF. In contrast, the nitracline (i.e. 2.5 μM nitrate isopleth) depth was recorded within a limited range of 20–40 m in spring, 30–50 m in summer, and 40–60 m in fall. The chlorophyll a concentrations at the subsurface chlorophyll maxima (SCM) were significantly higher in spring and summer (356 ± 233 and 270 ± 182 ng L –1 , respectively) than in fall (117 ± 89 ng L –1 ). The relative contributions of individual phytoplankton groups to the depth-integrated chlorophyll a concentration conformed to the composition of the phytoplankton community in the SCM layer, showing a dominance of diatoms (58 ± 19% in spring, 48 ± 11% in summer, and 30 ± 20% in fall). Canonical correspondence analysis revealed that the geographic structures of phytoplankton communities were strongly associated with the vertical structures of water temperature and nutrient concentration in the water column rather than with horizontal gradients of hydrographic conditions. Finally, our findings suggest that water column stability and light–nutrient availability in the euphotic zone play a key role in determining geographical consistency of the biomass and the community structure of phytoplankton in the EJS.

Mesozooplankton (copepods and euphausiids) grazing rates and prey preferences were determined during a series of three research cruises to the eastern Bering Sea in spring 2008, 2009, and 2010. Chlorophyll was dominated by large cells (>5 µm), especially at bloom locations where they usually comprised greater than 90% of the total chlorophyll biomass. The relative importance of microzooplankton to the prey field biomass decreased with increasing chlorophyll concentration, and was less than 10% of the total prey biomass in ice-edge bloom regions. Overall, microzooplankton was the preferred prey of the mesozooplankton, although phytoplankton/ice algae were the dominant component of the diet because of their much greater biomass, especially during blooms. There were differences between mesozooplankton species in their prey preferences: Metridia pacifica, Pseudocalanus spp. and Calanus spp. had the strongest preference for microzooplankton prey, while euphausiids (Thysanoessa spp.) and Neocalanus flemingeri/plumchrus appeared to feed non-selectively on all prey items. Mesozooplankton exhibited a saturating feeding response to chlorophyll concentration (Holling׳s type II) that could be modeled by Michaelis–Menten equations. Taxa-specific maximum ingestion rates generally followed allometric theory, with smaller zooplankton having higher feeding rates than larger zooplankton, and ranged from about 4–30% body carbon day−1. Trophic cascades during grazing experiments could result in a substantial underestimate of chlorophyll ingestion rates, especially for those taxa that had a strong preference for microzooplankton. Grazing impacts by mesozooplankton on the integrated chlorophyll biomass and primary production were 2.7±4.4 and 26±48% day−1, respectively. Impacts increased significantly with increasing mesozooplankton biomass, which increased from early to late spring. However, grazing impacts were extremely low in ice-edge bloom regions. Our findings suggest that even when grazing by microzooplankton is included in our grazing impact estimates, about 50% of the primary production in phytoplankton blooms during spring on the eastern Bering Sea shelf is not grazed and is available for direct export to the benthic community.

Two extended pulsed feeding experiments, following the spring bloom period, investigated lipid retention in the prominent Bering Sea euphausiid (krill) Thysanoessa raschii. These experiments occurred during late spring and early summer of 2010. Concurrent taxonomic analysis of the natural algal community allowed prey type to be linked to lipid composition of the natural communities. In late spring, experimental periods of feeding followed by starvation showed an overall decrease in total lipid for T. raschii. In early summer, no consistent trend was observed for total lipid with the visible presence of storage lipid in some animals. Polar lipids, as phospholipids, were the dominant krill lipid class in both experiments constituting ≥88% of total lipid, and triacylglycerols reached a maximum of 5% of total lipid. The sterols cholesterol and brassicasterol+desmosterol comprised 98–99% of total sterol abundances in T. raschii throughout both experiments, even after feeding periods when alternative sterols (i.e. the algal sterol 24-methylenecholesterol) accounted for up to 39% of sterols in potential food particles. Cholesterol abundance and concentration increased during both incubations, likely due to the metabolism of dietary sterols. Major fatty acids observed in krill included C14:0n, C16:0n, C16:1(n-7), C18:1(n-7), C18:1(n-9), C20:5(n-3), and C22:6(n-3) with the diatom-attributed C16:1(n-7) decreasing in abundance and concentration during starvation. Low concentrations of the dinoflagellate-derived sterol and a novel C28:8 PUFA, typically found in dinoflagellates and prymnesiophytes, indicated predation on protozooplankton in early summer when diatom abundances were low. The stability of lipid distributions over periods of starvation and intermittent feeding suggest that fatty acid and sterol biomarkers present in this polar euphausiid principally reflect long-term diet history rather than short-term feeding episodes.

Observations indicate that spring and fall phytoplankton blooms on the Eastern Bering Sea (EBS) continental shelf tend to co-vary on inter-annual scales – that is, a year with a strong spring bloom also tends to have a strong fall bloom. Similar co-variability of primary production is also seen in the multi-year (1987–2007) integration of a coupled physical–biological model. Moreover, the modeled seasonal amplitudes of 10-meter chlorophyll-a concentrations at the EBS middle shelf mooring locations, computed using the canonical Redfield ratio and a mean carbon-to-chlorophyll-a ratio, are generally consistent with the in situ mooring measurements. The coupled physical–biological model simulation is used to examine the relative contributions of wind mixing, local nutrient recycling/regeneration, horizontal nutrient advection, and water-column stability to this co-variability. There is no significant correlation between the spring and fall surface wind mixing. Although wind mixing is an important mechanism for bringing nutrients in the lower water column to the surface layers, it is not the mechanism tying the two seasons׳ productivity together. Local regeneration/recycling of the nutrients initially fueling spring production is an important mechanism for spring-to-fall nutrient accumulation in the bottom layers at the middle shelf. Horizontal advection does not appear to be the dominant factor for supplying nutrients to the middle shelf during the spring-to-fall period. Fall primary production in the model is strongly influenced by the lower water-column stability/stratification. Taken together, these results highlight the importance of local recycling/regeneration of nutrients assimilated by spring phytoplankton bloom in linking together the spring and fall primary productions on EBS middle shelf.

We combined field data and the output from a climate-to-fish coupled biophysical model to calculate weekly climatologies and 1971–2009 time series of physical and biological drivers for 16 distinct regions of the eastern Bering Sea shelf and slope. We focus on spatial trends and physical-biological interactions as a framework to compare model output to localized or season-specific observations. Data on pollock (≥8 cm) diet were used to evaluate energy flows and zooplankton dynamics predicted by the model. Model validation shows good agreement to sea-ice cover albeit with a one month delay in ice retreat. Likewise, the timing of spring phytoplankton blooms in the model were delayed approximately one month in the south and extend further into summer, but the relative timing between the spring and fall bloom peaks was consistent with observations. Ice-related primary producers may shift the timing of the spring bloom maximum biomass earlier in years when sea ice was still present after mid-March in the southern regions. Including the effects of explicit, dynamic fish predation on zooplankton in the model shifts the seasonal spring peak and distribution of zooplankton later in the year relative to simulations with implicit predation dependent only on zooplankton biomass and temperature; the former capturing the dynamic demand on zooplankton prey by fish. Pollock diets based on stomach samples collected in late fall and winter from 1982–2013 show overwintering euphausiids and small pollock as key prey items in the outer and southern Bering Sea shelf; a characteristic not currently present in the model. The model captured two large-scale gradients, supported by field data, characterizing the overall dynamics: 1) inshore to off-shelf physical and biological differences with a gradient in inter-annual variability from higher frequency inshore to lower frequency offshore; and 2) latitudinal gradients in the timing of events. The combined effects of length of day, bathymetry, and tides, which are consistent from year to year, and the two large-scale gradients, characterize the environment on which regional differences were based and restrict their inter-annual and seasonal variability. Thus, the relative timing and sequence of events remained consistent within regions. The combination of model outputs and observational data revealed specific ecosystem processes: (1) The spatial progression in the timing, peaks and sequence of events over the shelf is driven by wind, sea ice, and stratification and creates a seasonal expansion and contraction of the warmer pelagic and bottom habitat suitable to pollock. (2) The seasonal warming of air temperature and the spring-summer expansion of the warm pelagic and bottom habitats influence the ice retreat and the associated ice edge and open water spring blooms, as well as subsequent production/abundance of copepods and euphausiids. (3) These warmer conditions favor pelagic energy flows to pollock (≥10 cm) and allow their distribution to expand shoreward and northward along the shelf break. (4) The fall-winter expansion of the seasonal ice cover drives the contraction of warmer waters towards the outer and southwest shelf and favors benthic energy flows over most of the shelf. There, fall blooms allow for additional lipid storage by large copepods and euphausiids that sink close to the bottom where they either go into diapause or have a restricted diel migration over winter. (5) During these cold months, the preferred pollock habitat shifts and contracts towards the outer and southwest shelf where their increased density and reduced prey availability leads to winter pollock cannibalism and consumption of overwintering euphausiids. Our project highlights the benefits of linking continuous and long-term field work with the development and implementation of highly complex models. In the face of uncertainty, simulations such as these, tightly coupled to field programs, will be instrumental as testbeds for process exploration and management evaluation, increasing their relevance for future fisheries and ecosystem management and strategic planning.

We synthesize recent research on variation in annual production of copepods (Calanus spp.), euphausiids (Thysanoessa spp.), and juvenile walleye pollock (Gadus chalcogrammus) in the southeastern Bering Sea. We reach five conclusions: 1) the timing of the spring bloom is more important than the amount of annual primary production for the transfer of primary to secondary production (i.e., timing matters); 2) summer and fall, not just spring, matter: organisms must maximize energy intake devoted to somatic growth and storage of lipids and minimize energy expenditures during each season; 3) stored lipids are important for the overwinter survival of both zooplankton and age-0 walleye pollock; 4) variation in ice extent and timing of ice retreat affect the spatial distributions of phytoplankton, zooplankton, and age-0 walleye pollock; when these spatial distributions match in late-ice-retreat years, the annual production of copepods, euphausiids, and juvenile walleye pollock often increases (i.e., location matters); 5) if years with late ice retreat, which favor copepod, euphausiid, and juvenile walleye pollock production, occur in succession, top–down control increases. These conclusions help to explain annual variation in production of copepods, euphausiids and juvenile walleye pollock. Copepods and euphausiids often are more abundant in cold years with late ice retreat than in warm years with early ice retreat due to bloom timing and the availability of ice algae during years with late ice retreat. As a consequence, age-0 walleye pollock consume lipid-enriched prey in cold years, better preparing them for their first winter and their overwinter survival is greater. In addition, there is a spatial match of primary production, zooplankton, and age-0 walleye pollock in cold years and a mismatch in warm years.

This paper serves as an introduction to the Deep-Sea Research II special issue on the impacts of a severe lack of sea ice in the northern Bering Sea (NBS) in the winter of 2017-2018. To complement the papers in this volume, we provide a brief synopsis of the physical oceanography of the NBS, including information on the characteristics of the dominant water masses, and the patterns of sea-ice formation and duration in years past. The 17 papers in this volume provide a series of snapshots of the northern Bering Sea in the spring and summer of 2018, which followed an unusual warming in February 2018 and record-breaking low sea-ice concentrations. The lack of ice in spring affected the temperatures and structure of the water column, impacted the timing of the spring bloom, and affected the marine food webs from the amounts and kinds of phytoplankton and zooplankton, to the distribution of fish and the reproduction and survival of marine birds and mammals. As the Arctic and sub-Arctic warm, events like those observed in 2018 may well become the norm. If so, we may anticipate major changes in the NBS marine ecosystem and those dependent upon its resources.

Primary production associated with the spring bloom in high-latitude seas constitutes a substantial fraction of annual total in those environments, and provides a seasonally timed bounty to benthic and pelagic organisms at higher trophic levels. Light is widely believed to regulate the timing and magnitude of spring production in the coastal Gulf of Alaska, although few data are available on the relationship between light and photosynthesis or primary production in that region. In two cruises to the coastal waters of southeast Alaska (May 2011 and April 2013), we observed strongly contrasting early-season phytoplankton communities, although environmental conditions were largely similar. An early, diatom-dominated spring bloom occurred in April 2013, while the anomalously low chlorophyll conditions of spring 2011 were widely observed in May of that year. Integrated chlorophyll a (Chl) was substantially higher in April 2013 than in May 2011 (median 62 versus 29 mg m−2), and April 2013 saw a greater contribution by large cells (>20 µm size fraction) to total Chl. The P M B ) were higher in the high-Chl month of April 2013 than in May 2011, averaging 5.3 versus 3.4 µg C µg Chl−1 h−1 across all depths and both size classes. In May 2011, P M B showed a strong negative correlation with salinity, while several photosynthetic parameters were correlated to light exposure history. In contrast, we found no environmental correlates to photosynthetic parameters in April 2013. Estimated primary production was considerably higher in April 2013 than in May 2011 (medians 2.5 versus 0.9 g C m−2 d−1). Characteristics of the May 2011 phytoplankton community were broadly consistent with a role for iron limitation on this narrow shelf, where cross-shelf exchange processes with nearby iron-limited open ocean waters may vary with larger-scale environmental conditions. Understanding the factors that regulate spring phytoplankton biomass, composition, and photophysiology will be key to understanding the large interannual variations in spring production in the coastal Gulf of Alaska.

The Gulf of Alaska (GoA) shelf is a highly productive regime bordering the nitrate-rich, iron (Fe)-limited waters of the central GoA. The exchange between nitrate-limited, Fe-replete coastal waters and nitrate-rich, Fe-deplete offshore waters, amplified by mesoscale eddies, is key to the productivity of the region. Previous summer field studies have observed the partitioning of Fe in the coastal GoA as being heavily dominated by the particulate phase due to the high suspended particulate loads carried by glacial rivers into these coastal waters. Here we present new physico-chemical iron data and nutrient data from the continental shelf of the GoA during spring and late summer 2011. The late summer data along the Seward Line showed variable surface dissolved iron (DFe) concentrations (0.052 nM offshore to 4.87 nM inshore), within the range of previous observations. Relative to available surface nitrate, DFe was in excess (at Fe:C=50 μmol:mol) inshore, and deficient (at Fe:C=20 μmol:mol) offshore. Summer surface total dissolvable iron (TDFe, acidified unfiltered samples) was dominated by the acid-labile particulate fraction over the shelf (with a median contribution of only 3% by DFe), supporting previously observed Fe partitioning in the GoA. In contrast, our spring data from southeast GoA showed TDFe differently partitioned, with surface DFe (0.28–4.91 nM) accounting on average for a much higher fraction (~25%) of the TDFe pool. Spring surface DFe was insufficient relative to available nitrate over much of the surveyed region (at Fe:C=50 μmol:mol). Organic Fe-binding ligand data reveal excess concentrations of ligands in both spring and summer, indicating incomplete titration by Fe. Excess concentrations of an especially strong-binding ligand class in spring surface waters may reflect in-situ ligand production. Due to anomalous spring conditions in 2011, river flow and phytoplankton biomass during our spring sampling were lower than the expected May average. We argue our samples are likely more representative of early spring pre-bloom conditions, providing an idea of the possible physico-chemical partitioning of Fe in coastal GoA waters relevant to initial spring bloom dynamics.

Significant changes in fisheries resources have occurred in the Gulf of Alaska (GOA) in the mid 1970s, with an increase in groundfish and a decrease in crab and shrimp populations. Increased fishing pressure and such events suggest that the GOA is susceptible to climate variation; however the mechanistic links between ecosystem change and climate remain unclear. At-sea surveys were undertaken during the month of May from 1998 to 2009 to collect data on zooplankton abundance and water mass properties in the northern GOA. Significant changes in temperature, salinity and zooplankton abundance were identified during this period. The euphausiid Thysanoessa inermis and the copepod Calanus marshallae had increased abundances in years when there was a strong phytoplankton spring bloom preceded by anomalously cold winters. The euphausiid Euphausia pacifica and the copepods Pseudocalanus spp., Neocalanus plumchrus/flemingeri, and Oithona spp. were more resilient to relatively high mean water temperatures. High zooplankton abundances in years of substantial cross-shelf mixing suggest that iron and nutrient transport between the shelf and oceanic domains are essential for sustaining high zooplankton populations via phytoplankton blooms. The abundance of zooplankton in the northern GOA is highly influenced by advective processes and changes in temperature. Further understanding of biological and physical mechanisms that control the GOA ecosystem are of major importance to predict the response of zooplankton communities to environmental changes.

Distributions of dissolved iron (dFe) and its reduced form, Fe(II), to a depth of 1000 m were investigated under the seasonal pack ice off East Antarctica during the Sea Ice Physics and Ecosystem experiment (SIPEX-2) sea-ice voyage in September–October 2012. Concentrations of dFe were elevated up to five-fold relative to Southern Ocean background concentrations and were spatially variable. The mean dFe concentration was 0.44±0.4 nM, with a range from 0.09 to 3.05 nM. Profiles of dFe were more variable within and among stations than were macronutrients, suggesting that coupling between these biologically-essential elements was weak at the time of the study. Brine rejection and drainage from sea ice are estimated to be the dominant contributors to elevated dFe concentrations in the mixed layer, but mass budget considerations indicate that estimated dFe fluxes from brine input alone are insufficient to account for all observed dFe. Melting icebergs and shelf sediments are suspected to provide the additional dFe. Fe(II) was mostly below the detection limit but elevated at depth near the continental shelf, implying that benthic processes are a source of reduced Fe in bottom waters. The data indicate that dFe builds up under the seasonal sea-ice cover during winter and that reduction of Fe may be hampered in early spring by several factors such as lack of electron donors, low biological productivity and inadequate light below the sea ice. The accumulated dFe pool in the mixed layer is expected to contribute to the formation of the spring bloom as the ice retreats.

The picoplankton distribution in the East China Sea was investigated during bloom events occurred in spring (June) and summer (August) 2011. In spring, there was no significant difference in picoplankton abundances between areas where bloom conditions were or were not established. In the bloom area, Synechococcus, picoeukaryotes and heterotrophic prokaryotes exhibited at only some stations that abundances were higher than those within the non-bloom area. In summer, the abundances of Synechococcus and picoeukaryotes were significantly higher inside the bloom area than outside. Among the picoplankton components, heterotrophic prokaryotes represented the highest carbon biomass. Factors that most influenced picoplankton distribution under bloom conditions in the East China Sea varied with season. In spring, ciliates and salinity tended to be the main factors, whereas in summer, this role was played by temperature and chlorophyll a concentration. (C) 2015 Elsevier Ltd. All rights reserved.

We investigated the heterotrophic bacterial biomass and production in February 2012, in four habitats (a polynya, sea-ice zone, ice shelf, and the open sea) in the Amundsen Sea to determine the spatial distribution, controlling factors, and ecological role of the bacteria during a late phytoplankton bloom by Phaeocystis antarctica . Bacterial abundance (BA) and production (BP) were highest at the center of the polynya, and both were significantly correlated with phytoplankton biomass. BP accounted for average 17% of the organic carbon produced by phytoplankton primary production (PP), which is higher than the average BP:PP ratio reported in most open ocean. The abundance of heterotrophic nanoflagellates (HNF) was correlated with the BA, and the average bacteria:HNF ratio (260) was lower than the values reported in most marine environments (400–1000), including the Ross Sea Polynya (800). Evidence for a tight coupling of bacteria and phytoplankton activities on the one hand and intense HNF grazing on bacteria on the other could be found in the high BP:PP and low bacteria:HNF ratios, respectively. Interestingly, these data were accompanied by low particulate carbon export fluxes measured during the late Phaeocystis bloom. Together, these results indicated that the microbial loop plays a significant role in the biogeochemical carbon cycle and food web processes in the Amundsen Sea Polynya.

To better understand the impact of climate change on the Southern Ocean (SO) biogeochemistry, we must know the evolution of the seasonal cycle of biological properties. Here we present the first estimate of the seasonal evolution of chlorophyll-a (Chl-a) along oceanic fronts in the Indian sector of SO using Bio-Argo float observations. The present study is based on data collected using three such floats deployed in the frontal zones. The high productivity is usually attributed to the upwelling of nutrient-enriched subsurface water, the supply of dissolved iron to the ocean surface and their horizontal advection. Our analysis shows that the seasonal evolution of Chl-a is similar in all the three frontal regions, namely Sub-Tropical front (STF), Sub-Antarctic Front (SAF) and Polar Front (PF). We also found that shoaling of the Mixed Layer Depth (MLD) above the critical depth, owing to an increase in light exposure during spring, triggers the initiation of the spring bloom.

Based on both physical and biological data collected from cruises in different seasons between 2010 and 2011, we found that the phytoplankton bloom was highly seasonally variable along the longitudinal axis of the Pearl River Estuary (PRE) and was confined to very limited locations because of the variation in the coupled physical–biological forcing of the phytoplankton growth and accumulation. We investigated the relative importance and joint control of the bloom′s variation by freshwater residence time, water column stability, and light limitation that were governed by river discharge, vertical mixing, and turbidity in the PRE, respectively. We found that the bloom occurred in the middle of the well/partially mixed estuary during the dry season when river discharge was relatively small. Although the water nutrient concentration and residence time were favorable for phytoplankton accumulation during this season, the bloom was absent in the upper and lower parts of the estuary likely associated with high turbidity in the upper region and strong vertical mixing in the lower region. In contrast, strong river discharge during the wet season pushed the nutrient-rich river plume farther seaward and formed a salt-wedged estuary. With a stable water column and relatively low turbidity during this season, the bloom could only occur in the lower part of the estuary where the phytoplankton growth rate was greater than the water turnover rate.

The timing and magnitude of phytoplankton blooms in subarctic ecosystems often strongly influence the amount of energy that is transferred through subsequent trophic pathways. In the eastern Bering Sea, spring bloom timing has been linked to ice retreat timing and production of zooplankton and fish. A large part of the eastern Bering Sea shelf (~500 km wide) is ice-covered during winter and spring. Four oceanographic moorings have been deployed along the 70-m depth contour of the eastern Bering Sea shelf with the southern location occupied annually since 1995, the two northern locations since 2004 and the remaining location since 2001. Chlorophyll a fluorescence data from the four moorings provide 37 realizations of a spring bloom and 33 realizations of a fall bloom. We found that in the eastern Bering Sea: if ice was present after mid-March, spring bloom timing was related to ice retreat timing ( p n =11). A fall bloom also commonly occurred, usually in late September (average day 274, SE=4.2, n =33), and its timing was not significantly related to the timing of storms ( p =0.88, df=1, 27) or fall water column overturn ( p =0.49, df=1, 27). The magnitudes of the spring and fall blooms were correlated ( p =0.011, df=28). The interval between the spring and fall blooms varied between four to six months depending on year and location. We present a hypothesis to explain how the large crustacean zooplankton taxa Calanus spp. likely respond to variation in the interval between blooms (spring to fall and fall to spring).

The flow of upper-layer surface water and circulation for the Bay of Biscay, continental slope and in the wider region of the NE Atlantic is presented, as well as the seasonality of flow and internal tides. The marine plankton environments of Biscay Ocean, Biscay Eddies, Biscay Slope and Biscay Shelf are defined. The Shelf region (Armorican and Celtic) is further divided into Stratified Shelf, Frontal and Tidally Mixed. Seasonal distributions of chlorophyll a are given for all environment from in situ measurements and remote sensing data. Mixing and stabilisation of surface water in the euphotic layer for the start of the spring bloom using in situ profiling measurements is examined. Some regional responses for the slope current, dinoflagellate blooms and interannual variations in spring diatom numbers with respect to weather and climate in the Bay of Biscay and around the British Isles are suggested and discussed. An example of the Eastern European Ocean Margin continental slope response to winter weather (sea level atmospheric pressure forcing) resulting in warm winter water in the southern Bay of Biscay (Navidad, with eddy production) and off the Shetland continental slopes (the warm-water supply route to the Arctic) is given from the slope climate observation series.

A key challenge in oceanography is to capture and quantify processes that happen on short time scales, seasonal changes and inter-annual variations. To address this problem the P&O European Ferries Ltd. Ship MV Pride of Bilbao was fitted with a FerryBox from 2002 to 2010 and data returned to NOC in real time providing near continuous measurements between UK (Portsmouth) and Spain (Bilbao) of temperature, salinity, chlorophyll-fluorescence and oxygen. Additional monthly samples were collected on manned crossings. Over 6000 samples were analysed for nitrate (nitrate and nitrite) concentrations. The timing of nitrate concentration increases (with winter mixing) and decreases (with the spring bloom) are different on and off shelf and in autumn nitrate concentrations remain high on the shelf. Off shelf in the Bay of Biscay, the mixed layer depth assessed using Argo floats, was found to vary from 212 m in relatively mild winters (such as 2007/2008) to 476 m in cold winters (2009/2010). Years with deeper mixing were associated with an increase in nitrate concentrations in the surface waters (~3 µmol l-) and the increased vertical nutrient supply resulted in higher productivity the following spring. Bloom progression could be seen through the increase in oxygen anomaly and decrease in nitrate concentrations off shelf prior to changes further north on the shelf and phytoplankton growth was initiated as shoaling begins. The full dataset demonstrates that ships of opportunity, particularly ferries with consistently repeated routes, can deliver high quality in situ measurements over large time and space scales that currently cannot be delivered in any other way.

Some results of the Bay of Biscay regional oceanography presented at ISOBAY are summarized including contributions to physical oceanography, chemical and biological oceanography, marine geology, deep water ecology, marine pollution, fisheries research and cetacean studies. A long-term analysis of the spring bloom of phytoplankton in the area during the last 17 years (1997–2014) is presented as an example of Bay of Biscay climate research. The Spring Bloom presents cycles of 4–6 years reflecting probably the availability of nutrients from the previous winter and has increased in peak intensity during the last decades.

It has been long thought that coccolithophores are a minor component of the phytoplankton assemblage in Arctic waters, with diatoms typically being more dominant. Little is known about how the phytoplankton communities will change, however, as the Arctic warms. We participated in the 2011 Impacts of Climate on EcoSystems and Chemistry of the Arctic Pacific Environment (ICESCAPE) cruise to the western Arctic, performing a combination of discrete measurements (microscopy, calcification, particulate inorganic carbon (PIC), particulate organic carbon (POC), biogenic silica (BSi)) plus continuous surface bio-optical measurements (absorption, scattering, backscattering and acid-labile backscattering; the latter specific for coccolithophores). Here, we report bio-optical and coccolithophore observations from the massive under-ice algal bloom originally described in Arrigo et al. (2012) . The most intense portions of the bloom were centered in cold Winter Water and there was evidence for nitrate drawdown in the top 10–20 m with strong penetration of silicate rich water into the surface waters. Surface chlorophyll a and particulate absorption at 440 nm approached 30 μg L −1 and 1.0 m −1 , respectively. Particulate absorption of detritus ( a p at 412 nm) was highly correlated to a p at 440 nm associated with chlorophyll a and slopes of the absorption spectrum showed that both dissolved and particulate absorption at 412 nm exceeded that at 440 nm, with slopes, S g , of 0.01 nm –1 . Colored dissolved organic matter fluorescence (FDOM) was high in the bloom but the relative fluorescence yields were low, characteristic of phytoplankton-produced FDOM (as opposed to terrestrially-produced FDOM). Coccolithophore backscattering was elevated in the under-ice bloom, but it only accounted for 10% of the total particle backscattering, relatively low compared to typical subpolar waters further to the south. Total particle scattering was significantly elevated in the under-ice bloom (values of almost 2 m −1 ), likely due to the high abundance of large diatoms. Backscattering probabilities in the bloom were ~1%, again characteristic of diatom-dominated populations with few calcifiers. PIC standing stock in the under-ice bloom was low but measurable while biogenic silica molar concentrations were 150 times greater. POC:PON molar ratios were 6–10, characteristic of healthy, rapidly growing phytoplankton, observations further buttressed by carbon:chlorophyll mass ratios of 50–100. Coccolithophore calcification was low but measurable, reaching 1.75 mg C m −3 d −1 in the under-ice bloom, only 0.4% of the photosynthesis. However, the intrinsic carbon-specific growth rate was 0.4 per day for bulk POC and ~1 per day for bulk PIC, close to maximal growth rates expected at these temperatures. SEM and light microscopy results showed mostly diatoms in the bloom. The coccolithophore, Emiliania huxleyi , was observed, providing unequivocal evidence of the presence of coccolithophores in the under-ice algal bloom.

Standard and imaging flow cytometry were used to examine the composition of phytoplankton assemblages in and around a massive under-ice bloom in the Chukchi Sea in 2011. In the core of this bloom, roughly 100 km northwest of Hanna Shoal, diatoms represented roughly 87% of the water column carbon-specific biomass of phytoplankton, while nanophytoplankton contributed ~9%. Picoeukaryotes were also observed in this bloom, as were phycoerythrin-containing cells consistent with Synechococcus spp., but picophytoplankton, dinoflagellates, and prymnesiophytes each represented only ~1% of the bloom׳s phytoplankton biomass. More broadly along this part of the Chukchi shelf, nanophytoplankton typically comprised a larger fraction of phytoplankton biomass in the water column, 22% on average but up to 82% at certain locations. Dinoflagellates and prymnesiophytes contributed at most 2% of water column biomass at any location and were most abundant in the deeper slope stations northeast of Hanna Shoal, east of the bloom. Picophytoplankton were most abundant in these deeper slope stations as well, and also in recently ice-free areas to the south around Hanna Shoal. These cell-derived estimates of phytoplankton carbon biomass, which were computed from imaging and standard cytometric observations of phytoplankton cell sizes and from published carbon:volume relationships, agree well with independent measurements of particulate organic carbon concentration from traditional biochemical assays.

The life cycle of Alexandrium fundyense in the Gulf of Maine includes a dormant cyst stage that spends the winter predominantly in the bottom sediment. Wave-current bottom stress caused by storms and tides induces resuspension of cyst-containing sediment during winter and spring. Resuspended sediment could be transported by water flow to different locations in the Gulf and the redistribution of sediment containing A. fundyense cysts could alter the spatial and temporal manifestation of its spring bloom. The present study evaluates model near-bottom flow during storms, when sediment resuspension and redistribution are most likely to occur, between October and May when A. fundyense cells are predominantly in cyst form. Simulated water column sediment (mud) concentrations from representative locations of the Gulf are used to initialize particle tracking simulations for the period October 2010–May 2011. Particles are tracked in full three-dimensional model solutions including a sinking velocity characteristic of cyst and aggregated mud settling (0.1mms−1). Although most of the material was redeposited near the source areas, small percentages of total resuspended sediment from some locations in the western (~4%) and eastern (2%) Maine shelf and the Bay of Fundy (1%) traveled distances longer than 100km before resettling. The redistribution changed seasonally and was sensitive to the prescribed sinking rate. Estimates of the amount of cysts redistributed with the sediment were small compared to the inventory of cysts in the upper few centimeters of sediment but could potentially have more relevance immediately after deposition.

The development of phytoplankton bloom and its association with physical forcing is examined through an interdisciplinary field-work conducted in the vicinity of the central trough of the southern Yellow Sea during March–April 2009, with the aid of a surface Lagrangian drifter deployed at the bloom site. Bloom patches were detected using an empirical value and two of them were traced by the drifter for a period of several days respectively. Both of them appears as thin-layer subsurface chlorophyll a maximum (SCM) throughout the tracing, although their dominant phytoplankton species are not identical at all. The magnitude as well as the onset of these two blooms is different from each other, but both found to be relevant to local oceanic and meteorological conditions. Both of them demonstrate that the changes in the stability of hydrographical structure, especially at layers around the SCM, take a substantial role in triggering or terminating the blooming processes. Those changes in meteorological conditions, like wind speed and directions, solar radiation, are short and cause daily or synoptic scale variations in phytoplankton concentrations, but the frequency of northerly wind events predating the bloom season has a positive effect on the occurrence of spring blooms. The horizontal advection is another contributing factor indicated by the drifter which accounts for the bloom extinction at the station B20. In addition, due to the weak orbital horizontal movement, the bloom above the central trough persists longer and larger.

Temporal variation of Synechococcus, picoeukaryote and heterotrophic bacteria abundance and depth integrated biomass during three spring blooms in 2007 and 2009 were investigated in the Yellow Sea, China. Synechococcus and picoeukaryote responded differently to different types and course of spring blooms. During the diatom blooms of 2007 and Bloom B20 in 2009, Synechococcus and picoeukaryote abundances decreased sharply during the bloom period. However, during a mixed dinoflagellate and diatom bloom of Bloom B23 in 2009, Synechococcus and picoeukaryote increased in abundance and biomass along the bloom. During all three spring blooms, heterotrophic bacteria biomass had a similar increasing trend. Ciliate and heterotrophic nanoflagellate grazing could be responsible for Synechococcus and picoeukaiyote abundance and biomass decrease during the spring blooms. (C) 2013 Elsevier Ltd. All rights reserved.

The horizontal and vertical transport of nutrient-rich water both from the coast and from the seafloor to the overlying water column should play an important role in supplying nutrients required for the periods of vegetative or reproductive growth of phytoplankton. In the present work, radium isotopes (223Ra, 224Ra and 226Ra) in the southern Yellow Sea were measured after a spring bloom in June 2009. The exponential-like decrease of 223Ra away from the coast to offshore waters yielded horizontal eddy diffusivities (Kh) of (2.93±1.47)×107 cm2 s−1 by neglecting the advection. This estimate was smaller than that with advection indicator by as much as 21% when using an analytic model for 223Ra and 226Ra. The corresponding horizontal nutrient fluxes were 1525 µmol m−2 d−1 (DIN), 15.9 µmol m−2 d−1 (DIP) and 826 µmol m−2 d−1 (DSi), which would supply around 16% of N and 3% of P requirements based on the primary productivity. The decrease of 224Ra and 223Ra activities from sediments to the upper water column suggests the vertical eddy diffusion coefficient (Kz) of 6.23±5.58 cm2 s−1 below the thermocline, which was within the Yellow Sea Cold Water Mass (YSCWM). The calculated vertical fluxes of nutrient were 4945 μmol m−2 d−1 (DIN), 236 μmol m−2 d−1 (DIP) and 5315 μmol m−2 d−1 (DSi), accounting for up to 52% of N and 40% of P requirements for the phytoplankton growth. These results demonstrate the role of YSCWM as a relative nutrient-rich pool for the supply of nutrient to the southern Yellow Sea via the vertical diffusion process relative to the horizontal process during the summer season. Such processes will be strengthened during the weak density stratification in spring when algal blooms occur.

Phytoplankton growth and microzooplankton grazing were studied during the 2007 spring bloom in Central Yellow Sea. The surveyed stations were divided to pre-bloom phase (Chl a concentration less than 2 mu g L-1), and bloom phase (Chl a concentration greater than 2 mu g L-1). Shipboard dilution incubation experiments were carried out at 19 stations to determine the phytoplankton specific growth rates and the specific grazing rates of microzooplankton on phytoplankton. Diatoms dominated in the phytoplankton community in surface waters at most stations. For microzooplankton, Myrionecta rubra and tintinnids were dominant, and heterotrophic dinoflagellate was also important in the community. Phytoplankton-specific growth rates, with an average of 0.60 +/- 0.19 d(-1), were higher at pre-bloom stations (average 0.62 +/- 0.17 d(-1)), and lower at the bloom stations (average 0.59 +/- 0.21 d(-1)), but the difference of growth rates between bloom and pre-bloom stations was not statistically significant (t test, p = 0.77). The phytoplankton mortality rate by microzooplankton grazing averaged 0.41 +/- 0.23 d(-1) at prebloom stations, and 0.58 +/- 031 d(-1) during the blooms. In contrast to the growth rates, the statistic difference of grazing rates between bloom and pre-bloom stations was significant (after removal of outliers, t test, p = 0.04), indicating the importance of the top-down control in the phytoplankton bloom processes. Average potential grazing efficiency on primary productivity was 66% at pre-bloom stations and 98% at bloom stations, respectively. Based on our results, the biomass maximum phase (bloom phase) was not the maximum growth rate phase. Both phytoplankton specific growth rate and net growth rate were higher in the pre-bloom phase than during the bloom phase. Microzooplankton grazing mortality rate was positively correlated with phytoplankton growth rate during both phases, but growth and grazing were highly coupled during the booming phase. There was no correlation between phytoplankton growth rate and cell size during the blooms, but they were positive correlated during the pre-bloom phase. Our results indicate that microzooplankton grazing is an important process controlling the growth of phytoplankton in spring bloom period in the Central Yellow Sea, particularly in the "blooming" phase. (C) 2013 Elsevier Ltd. All rights reserved.

The source and significance of three nutrients - nitrogen, phosphorous and silicon - were investigated by a modified dilution method performed on seawater samples from the Central Yellow Sea (CYS), in spring blooming period of 2007. This modified dilution method accounted for the phytoplankton growth rate, microzooplankton grazing mortality rate, the internal and external nutrient pools, as well as nutrients supplied through remineralization by microzooplankton grazing. The results indicate that phytoplankton growth during the bloom is mostly contributed by internal nutrient pools (K-1 = 0.062-1.730). The external nutrient pools (K-E =

Continental shelves are hotspots for sedimentary denitrification, and the loss of N through denitrification can limit primary production in ecosystems. Spatial and seasonal trends in sedimentary denitrification and benthic nutrient fluxes are poorly characterized in the highly productive Bering Sea shelf ecosystem. Through the Bering Sea Ecosystem Study (BEST) program, we measured benthic fluxes of N2 and dissolved inorganic nitrogen (DIN: NH4 þ þNO2 � þNO3 � ), the extent of coupled sedimentary nitrification/denitrification, and the water column DIN deficit relative to phosphate, as indicated by a modified N* parameter (N**), on the Bering Sea shelf in the spring and summer 2009–2010. We found that sedimentary denitrification is widespread over the shelf, it is fueled mostly through coupled nitrification/denitrification, the net balance of sedimentary DIN flux is near zero over the shelf, and that the water column DIN deficit varies widely according to season and year. In the summer, N** in the surface layer appeared to be strongly affected by non-Redfieldian uptake of inorganic nutrients by phytoplankton in the spring bloom; in the winter, N** was strongly affected by sedimentary denitrification. Our findings indicate that the estimate of total N loss in Bering Sea shelf sediments should be revised upwards by at least 50% to 5.2–6.2 Tg N y �1 . In addition, sediments are not a significant source of

Inter-annual variability in the timing of phytoplankton spring bloom and phytoplankton community structure in the central North Atlantic Ocean was quantified using ocean color data and continuous plankton recorder (CPR) data. This variability was related to the North Atlantic Oscillation using correlation analysis and multivariate auto-regression models. The initiation of the spring bloom derived from CPR phytoplankton color index data is similar to that derived from satellite chlorophyll, and exhibits a nominal correlation with the sea surface temperature (SST) and the North Atlantic Oscillation (NAO). The extrapolated spring bloom timing suggested later initiation of blooms in the mid-1980s and earlier initiation of blooms in the 1990s. The climatological phytoplankton community structure in the central North Atlantic is dominated by diatoms, except for a shift in community composition favoring dinoflagellates in August. The ratio of diatoms to total phytoplankton abundance and the ratio of dinoflagellates to total phytoplankton abundance are both closely correlated with the NAO and SST. The extended time series of phytoplankton community structure between 1985 and 2009, deduced from the time series of SST and NAO over the same interval, showed a decadal shift away from diatoms towards dinoflagellates. The linkages between the NAO, and changes in stratification and phytoplankton processes occur over a larger scale than previously observed.

The spawning success of Antarctic krill (Euphausia superba) is generally assumed to depend on substantial winter sea ice extent, as ice biota can serve as a food source during winter/spring and the seasonal ice melt conditions the upper water column for extensive phytoplankton blooms. However, direct observations during spring are rare. Here we studied krill body condition and maturity stage in relation to feeding (i.e. stomach fullness, diet, absorption of individual fatty acids and defecation rate) across the Scotia Sea in November 2006. The phytoplankton concentrations were low at the marginal ice zone (MIZ) in the southern Scotia Sea (Stn. 1, 2, and 3), high in open waters of the Southern Antarctic Circumpolar Current Front (SACCF) in the central Scotia Sea (Stn. 5), and moderate further north (Stn. 6 and 7). Krill had low lipid reserves (∼6.5% of dry mass, DM), low mass:length ratios (∼1.7 mg DM mm−1), and small digestive glands (∼7% of total DM) near the ice edge. The stomachs contained lithogenic particles, diatom debris, and bacterial fatty acids, but low proportions of diatom-indicating fatty acids, which suggest that these krill were feeding on detritus rather than on fresh ice algae. In the SACCF, krill had higher lipid reserves (∼10% of DM), high mass:length ratios (∼2.2 mg DM mm−1), and large digestive glands (∼16% of total DM). Stomach content and tissue composition indicate feeding on diatoms. In the north, moderate food concentrations co-occurred with low lipid reserves in krill, and moderate mass:length ratios and digestive gland sizes. Only in the phytoplankton bloom in the SACCF had the mating season already started and some females were about to spawn. Based on the way krill processed their food at the different stations, we indicate two mechanisms that can lead to fast regeneration of body reserves and oocyte maturation in E. superba. One is “superfluous” feeding at high food concentrations, which maximises the overall nutrient gain. The other is a preferential absorption of crucial fatty acids: 20:5(n−3) and 22:6(n−3) when regaining body condition after the winter, and 14:0 and 16:1(n−7) during accumulation of oocyte yolk. Multi-year time series of phytoplankton distribution show that an early spring bloom, as found in 2006, is not unusual for the central Scotia Sea and coincides with high krill larval concentrations in this area. In conclusion, the Scotia Sea is a main spawning ground of Antarctic krill and this is linked to phytoplankton blooms in open waters of the SACCF rather than to ice edge blooms.

During the RV Polarstern ANT XXIV-2 cruise to the Southern Ocean and the Weddell Sea in 2007/2008, sediment samples were taken during and after a phytoplankton bloom at 52°S 0°E. The station, located at 2960 m water depth, was sampled for the first time at the beginning of December 2007 and revisited at the end of January 2008. Fresh phytodetritus originating from the phytoplankton bloom first observed in the water column had reached the sea floor by the time of the second visit. Absolute abundances of bacteria and most major meiofauna taxa did not change between the two sampling dates. In the copepods, the second most abundant meiofauna taxon after the nematodes, the enhanced input of organic material did not lead to an observable increase of reproductive effort. However, significantly higher relative abundances of meiofauna could be observed at the sediment surface after the remains of the phytoplankton bloom reached the sea floor. Vertical shifts in meiofauna distribution between December and January may be related to changing pore-water oxygen concentration, total sediment fatty acid content, and pigment profiles measured during our study. Higher oxygen consumption after the phytoplankton bloom may have resulted from an enhanced respiratory activity of the living benthic component, as neither meiofauna nor bacteria reacted with an increase in individual numbers to the food input from the water column. Based on our results, we infer that low temperatures and ecological strategies are the underlying factors for the delayed response of benthic deep-sea copepods, in terms of egg and larval production, to the modified environmental situation.

This study focuses on the comparison of oceanic and coastal cold-core eddies with inner-shelf and East Australian Current (EAC) waters at the time of the spring bloom (October 2008). The surface water was biologically characterised by the phytoplankton biomass, composition, photo-physiology, carbon fixation and by nutrient-enrichment experiments. Marked differences in phytoplankton biomass and composition were observed. Contrasted biomarker composition suggests that biomarkers could be used to track water masses in this area. Divinyl chlorophyll a , a biomarker for tropical Prochlorophytes, was found only in the EAC. Zeaxanthin a biomarker for Cyanophytes, was found only within the oceanic eddy and in the EAC, whereas chlorophyll b (Chlorophytes) was only present in the coastal eddy and at the front between the inner-shelf and EAC waters. This study showed that cold-core eddies can affect phytoplankton, biomass, biodiversity and productivity. Inside the oceanic eddy, greater phytoplankton biomass and a more complex phytoplankton community were observed relative to adjacent water masses (including the EAC). In fact, phytoplankton communities inside the oceanic eddy more closely resembled the community observed in the inner-shelf waters. At a light level close to half-saturation, phytoplankton carbon fixation (gC d −1 ) in the oceanic eddy was 13-times greater than at the frontal zone between the eddy and the EAC and 3-times greater than in the inner-shelf water. Nutrient-enrichment experiments demonstrated that nitrogen was the major macronutrient limiting phytoplankton growth in water masses associated with the oceanic eddy. Although the effective quantum yield values demonstrate healthy phytoplankton communities, the phytoplankton community bloomed and shifted in response to nitrogen enrichments inside the oceanic eddy and in the frontal zone between this eddy and the EAC. An effect of Si enrichment was only observed at the frontal zone between the eddy and the EAC. No response to nutrient enrichment was observed in the inner-shelf water where ambient NO x , Si and PO 4 concentrations were up to 14, 4 and 3-times greater than in the EAC and oceanic eddy. Although results from the nutrient-enrichment experiments suggest that nutrients can affect biomass and the composition of the phytoplankton community, the comparison of all sites sampled showed no direct relationship between phytoplankton biomass, nutrients and the depth of the mixed layer. This is probably due to the different timeframe between the rapidly changing physical and chemical oceanography in the separation zone of the EAC.

The composition and dynamics of the phytoplankton communities and hydrographic factors that control them are described for eastern and western Australia with a focus on the Eastern Australian Current (EAC) and Leeuwin Current (LC) between 27.5° and 34.5°S latitude. A total of 1685 samples collected from 1996 to 2010 and analysed for pigments by high performance liquid chromatography (HPLC) showed the average TChla (monovinyl+divinyl chlorophyll a) concentration on the west coast to be 0.28±0.16γgL-1 while it was 0.58±1.4γgL-1 on the east coast. Both coasts showed significant decreases in the proportions of picoplankton and relatively more nanoplankton and microplankton with increasing latitude. On both coasts the phytoplankton biomass (by SeaWiFS) increased with the onset of winter. At higher latitudes (>27.5°S) the southeast coast developed a spring bloom (September) when the mean monthly, surface chlorophyll a (chla) concentration (by SeaWiFS) was 48% greater than on the south west coast. In this southern region (27.5-34.5°S) Synechococcus was the dominant taxon with 60% of the total biomass in the southeast (SE) and 43% in the southwest (SW). Both the SE and SW regions had similar proportions of haptophytes; ~14% of the phytoplankton community. The SW coast had relatively more pelagophytes, prasinophytes, cryptophytes, chlorophytes and less bacillariophytes and dinophytes. These differences in phytoplankton biomass and community composition reflect the differences in seasonality of the 2 major boundary currents, the influence this has on the vertical stability of the water column and the average availability of nutrients in the euphotic zone. Seasonal variation in mixed layer depth and upwelling on the west coast appears to be suppressed by the Leeuwin Current. The long-term depth averaged (0-100m) nitrate concentration on the west coast was only 14% of the average concentration on the east coast. Redfield ratios for NO3:SiO2:PO4 were 6.5:11.9:1 on the east coast and 2.2:16.2:1 on the west coast. Thus new production (nitrate based) on the west coast was likely to be substantially more limited than on the eastcoast. Short term (hourly) rates of vertical mixing were greater on the east coast. The more stable water column on the west coast produced deeper subsurface chlorophyll a maxima with a 25% greater proportion of picoeukaryotes. © 2010 Elsevier Ltd.

Ten years (1998–2007) of sea-level anomalies (SLA) from multiple satellite altimeters and chlorophyll -a concentration from the Sea-viewing Wide Field-of-view Sensor (SeaWiFS) were used to detect eddy pumping/advection of nutrients by the action of propagating Rossby waves in the Kuroshio Extension (KE) region near 35 o N after the seasonal SeaWiFS chlorophyll -a (Chl -a ) concentrations cycles and annual changes of altimeter SLA were eliminated. Spatial structure of Chl -a anomalies to the seasonal cycle is examined in relation to altimeter eddy structure. Eddy propagation speeds and scales by the Rossby waves are also identified. Chl -a structure is evident during the spring bloom period with a scale around 460 km. Cold-core (cyclonic) rings correspond to areas of high Chl -a concentrations. Warm-core (anticyclonic) rings relate to areas of low Chl -a concentration. Chl -a anomalies and SLA have an overall modest negative correlation coefficient of r= −0.45, which may imply the co-existence of both eddy pumping and eddy advection mechanisms in the KE region. Swirl currents between eddies redistribute surface chlorophyll concentrations and can spatially bias maximum and minimum concentration levels off eddy center. The correlation coefficient has seasonal fluctuations. In the western KE region, it varies from high negative correlation ( r= −0.70) in April and September (eddy pumping dominant) to low negative correlation ( r =−0.33) in February and March (eddy advection dominant). In the eastern KE region, it varies from high negative correlation ( r =−0.67) in February to low negative correlation ( r =−0.42) in December. It is noted that the characteristic wavelengths of the SLA and Chl -a features, and the seasonality of the correlation between these two variables have not been previously documented.

We analyzed the near-daily hydrographic data collected from ships at a station southeast of Hokkaido Japan during April 2007. We calculated mixing ratios of Coastal Oyashio Water (COW), Oyashio Water (OYW), and modified Kuroshio Water (MKW). The COW mixing ratio tended to be high in depths shallower than 30 m during April, while the OYW ratio was high at the deeper depths. Diapycnal mixing due to the strong wind was suggested between these layers. It is suggested that stratification of low-salinity COW initiates the spring bloom in early-April, but that it is not important for temporal changes of chlorophyll at each depth during and after the bloom. Despite the intense water exchange at the station, the COW ratio correlated significantly with chlorophyll for the overall data set during April and for the data in 2–4-day blocks. Silicic acid, nitrate, and phosphate were also significantly correlated with the COW ratio for each 2–4-day block. This implies that each of these variables changed simultaneously at a constant COW ratio within the temporal and spatial scales of the phytoplankton bloom, were it not for the occurrence of vigorous vertical mixing and large-scale advective events. Using the linear regression equations for each 2–4-day block, we calculated time-series of chlorophyll and nutrients at the mean COW ratio (46%). Their rates of change in these series reveal the spring bloom dynamics in 2007 as follows: The spring bloom ended due to silicate depletion and grazing by large zooplankton in mid-April. In late-April, the nutrients increased, but chlorophyll decreased, indicating little nutrient utilization.

Temporal changes in abundance, growth and mortality of heterotrophic bacteria were investigated from frequent dilution experiments in the Oyashio region to evaluate their response to the spring phytoplankton bloom. Bacterial abundance fluctuated from 1.0 to 2.6×10 5 cells mL −1 . Cell density was not different between the pre-bloom and bloom periods, and showed no clear temporal pattern. Potential growth ( g ′) and mortality ( m ) varied from 1.11 to 2.39 day −1 and from 1.21 to 2.16 day −1 , respectively. Neither g ′ nor m was correlated with water temperature or chlorophyll a . Apparent growth rate in unenriched 100% seawater incubations ( g o ) was not limited by nutrients throughout the study period and was comparable to net growth rate ( g ′ −m ) estimates from dilution experiments. These results suggest that bacterial abundance is nearly steady-state in the pre-bloom and bloom period, because bacterial growth is nearly nutritionally saturated and balanced with high mortality.

Bottle incubations were conducted to investigate silica deposition and growth characteristics of siliceous phytoplankton in the Oyashio region using PDMPO, 2-(4-pyridyl)-5-((4-(2-dimethylaminoethyl-aminocarbamoyl)-methoxy)phenyl)oxazole, a fluorescence probe. This stain shows clear fluorescence in the position of newly produced silica cell walls during their vegetative cell growth. Parmales and diatoms dominated siliceous phytoplankton during the pre-bloom and bloom periods, and were successfully stained with PDMPO, indicating active silica deposition. Low PDMPO labeling of cells of the dominant diatoms relative to their initial abundance was often observed in the later half of the bloom period compared with high PDMPO labeling early in the bloom. That suggests that diatom silica deposition had slowed. Silica deposition in other siliceous plankton, i.e. silicoflagellates and choanoflagellates, and diatom resting spores was also detected. Our results show that the PDMPO technique is appropriate to demonstrate important biological processes in the silicon and carbon biogeochemical and cycles in the Oyashio region.

Despite large diatom blooms occurring in the Oyashio region of the NW Pacific every spring, our knowledge of factors controlling primary productivity in the region during that season remains incomplete. Therefore, we investigated phytoplankton abundance, size structure, and primary productivity from April to June 2007. Significant changes were observed. Chlorophyll a (Chl a) concentrations in surface waters fluctuated between 0.37 and 17 mg m−3. Micro-sized (>10 μm) phytoplankton dominated the phytoplankton community when Chl a was >1 mg m−3. Depth-integrated daily primary production within the euphotic layer ranged between 328 and 3231 mg C m−2 d−1. Higher values of the water-column light utilization index (Ψ) for phytoplankton photosynthesis were observed in May and June. Although no significant relationships were found between surface primary productivity and macronutrient concentrations or photosynthetically available radiation (PAR), surface primary productivity correlated significantly with Chl a concentration during April, indicating that algal productivity depended on phytoplankton biomass. Furthermore, significant linear relationships were found throughout the observations of phytoplankton absorption coefficient to surface primary productivity and of that coefficient to the optimum algal photosynthetic rate normalized by Chl a level (PBopt) in the water column. Modeling PBopt with the empirical equations of Behrenfeld and Falkowski (1997) or Kameda and Ishizaka (2005) did not accurately reproduce in situ PBopt. These results suggest that the phytoplankton absorption properties could become useful indicators for estimating primary productivity in the Oyashio region during spring.

We investigated the nutrient and diatom dynamics during late winter and spring (9-March to 1-May 2007) in the Oyashio region as part of the OECOS-WEST research cruises. Macronutrients, iron, chlorophyll a (Chl- a ) and biogenic silica (BSi) concentrations in the upper mixed layer varied remarkably ranges were 1.88–18.8 μmol L −1 for NO 3 +NO 2 , 0.64–1.85 μmol L −1 for PO 4 , 3.14–35.7 μmol L −1 for Si(OH) 4 , 0.14–0.54 nmol L −1 for D-Fe, 0.64–24.6 nmol L −1 for T-Fe, 0.30–17.4 μg L −1 for Chl- a , and 0.34–14.1 μmol L −1 for BSi. Mixed-layer depth (MLD) varied from 8–190 m during the cruises. The growth rate of in situ phytoplankton communities, dominated by centric diatoms, varied in shipboard culture experiments from 0.55 d −1 for iron-replete to 0.14 d −1 for iron-limited conditions. A relationship between BSi and Chl- a concentrations indicates that the in situ diatom community in the warmer water system (>4 °C) was heavily silicified, probably due to iron-limitation. The in situ macronutrient and dissolved iron concentrations below the MLD and estimated macronutrient concentrations during winter were negatively correlated to temperature (1–6 °C), that is to the relative proportion of warm modified Kuroshio Water mixed into the colder Oyashio water system. The rate of decrease in Si(OH) 4 per °C increase was greater than the rates for NO 3 +NO 2 and PO 4 for both in situ and estimated winter values. These results suggest that the spring bloom in the cold-water system with high macronutrients and iron concentrations would progress rapidly and intensely, and then be terminated by nitrogenous nutrient depletion. However, the diatom bloom in warmer waters with lower macronutrients and iron concentrations would be terminated by Si- and/or iron-limitation of heavily-silicified diatoms. In the OECOS study, variation of macronutrients and iron due to the surface intrusions of several water masses and modification from different chemical conditions during winter were the most important factors regulating the progression, magnitude and probably fate of the spring phytoplankton bloom in the Oyashio region.

We investigated the feeding habits and grazing rates of the ontogenetically migrating copepods in the Oyashio region to evaluate their grazing impacts on the food web during the spring phytoplankton bloom. The bloom was in progress from early to late April, although chlorophyll a concentrations fluctuated considerably with the frequent exchange of different water masses. Biomass of the copepod community reached a maximum in mid-April when late copepodites of Neocalanus cristatus , Neocalanus flemingeri and Eucalanus bungii contributed to the biomass increase. Gut pigment contents of the predominant copepods were much higher during the bloom compared with the levels in March (pre-bloom). The temporal fluctuations were not correlated with those of mean chlorophyll a concentrations in the 0–50 m layer. Feeding experiments indicated that major food items for the copepods were centric diatoms and flagellates. During the period of lower ambient chlorophyll, the copepods changed their heterotrophic prey from naked ciliates to tintinnids. Apparent clearance rates were positive for naked ciliates and negative for heterotrophic nanoplankton, Cryptophyceae and bacteria when chlorophyll was high, suggesting trophic cascade effects from copepod feeding even during the phytoplankton bloom. The carbon demands of the copepod community were estimated to be 156 mgC m −2 day −1 in early March to 797 mgC m −2 day −1 in mid-April. The grazing rates on phytoplankton reached 480 mgC m −2 day −1 , equivalent to as much as 28% of primary production. Non-phytoplankton prey supported 40 to 71% of the copepod carbon requirement. These results suggest that the copepod community does not graze the phytoplankton bloom down, but it does have significant impacts on microbial food webs.

Inter-annual variability in the magnitude and timing of the spring bloom was investigated for the Oyashio region (40 °-48 °N, 143 °E-152 °E) using 10 years (from 1998 to 2007) of satellite ocean-color data. Geostrophic currents were examined using satellite altimeter data. Early spring blooms (>1.5 mg m−3) occurred in early April 2001 and late March 2002. The 2001 bloom continued for one month. Late blooms occurred from mid-May 1999, early June 2004 and late April 2006, continuing for about 1 month, 8 days and 16 days, respectively. A strong bloom (4.7 mg m−3) also occurred in mid-April 1998; however, it terminated in early May. We classified the Oyashio region based on the pattern of temporal variation of Chl-a concentration from March to June. The spatio-temporal variability in Chl-a concentration during spring was different among years. The area where Chl-a concentration was highest in April was more extensive in 2001, 2002 and 2006 than usual. In 1999, the area where Chl-a concentration was highest in May was the widest among the 10 years. Mesoscale eddies and currents with high velocity were frequently observed in the area of high Chl-a concentration east of Hokkaido, propagating Coastal Oyashio Water of low salinity and low density into the oceanic region. That strengthened stratification in the surface layer. We suggest that this seaward transfer of coastal water could be one of the important factors for phytoplankton distribution in two ways: (1) horizontal advection of water with high Chl-a concentration and (2) enhancement of stratification in the oceanic region.

We present a 3-year multidisciplinary biogeochemical data set taken in situ at the Porcupine Abyssal Plain (PAP) time-series observatory in the Northeast Atlantic (49°N, 16.5°W; water depth not, vert, similar4850 m) for the period 2003 to 2005. The high-resolution year-round autonomous measurements include temperature, salinity, chlorophyll-a (derived from in situ chlorophyll-fluorescence) and inorganic nitrate, all at a nominal depth of 30 m on an Eulerian observatory mooring. This study compares these in situ time-series data with satellite chlorophyll-a data, regional data from a ship of opportunity, mixed-layer depth measurements from profiling Argo floats and lateral advection estimates from altimetry. This combined and substantial data set is used to analyse seasonal and inter-annual variability in hydrography and nitrate concentrations in relation to convective mixing and lateral advection. The PAP observatory site is in the inter-gyre region of the North Atlantic where convective mixing ranges from 25 m in the summer to over 400 m in winter when nutrients are supplied to the surface. Small inter-annual changes in the winter mixed layer can result in large changes in nitrate supply and productivity. However the decrease in maximum winter nitrate over the three-year period, from 10 to 7 mmol m?3, cannot be fully explained by convective mixing. Trajectories leading to the PAP site, computed from altimetry-derived geostrophic velocities, confirm that lateral advection cannot be ignored at this site and may be an important process along with convective mixing. Over the three years, there is an associated decrease in new production calculated from nitrate assimilation from 85.4 to 40.3±4.3 gCm?2 a?1. This confirms year-to-year variability in primary production seen in model estimates for the region. The continuous in situ dataset also shows inter-annual variation in the timing of the spring bloom due to variations in heat flux; the 2005 bloom occurred earlier than in 2004.

In the northern Baltic Sea, brackish water dinoflagellates Peridiniella catenata, Woloszynskia spp. and Scrippsiella hangoei form a major part of the spring bloom biomass, comparable to, or even exceeding the biomass of diatoms. The life cycle of these dinoflagellates involves a relatively short period of vegetative growth in early spring (2–3 months), followed by encystment during the first part of May, and a resting period in the form of benthic cysts. The bloom intensity of the Woloszynskia/Scrippsiella complex in the Baltic Sea is spatially variable, with peak abundances in the central and eastern Gulf of Finland, while the species are rare in the Gulf of Riga. During a field survey in late May 2004, we investigated the abundance and distribution of benthic cysts of P. catenata and Woloszynskia spp. in surface (5 cm) sediments around the Estonian coast. The broad distribution patterns of benthic cysts reflected the overall knowledge of basin-wide planktonic phase distribution of the species. On a finer scale, sediment properties (percent of clay, organic matter content) demarked the accumulation regions of the cysts. Woloszynskia cyst abundances in the surface 5 cm sediment were up to 3.2×106 cysts cm−2 and Peridiniella cyst abundances were approximately an order of magnitude less, up to 0.3×106 cysts cm−2.

A lower trophic level NPZD ecosystem model with explicit iron limitation on nutrient uptake is coupled to a three-dimensional coastal ocean circulation model to investigate the regional ecosystem dynamics of the northwestern coastal Gulf of Alaska (CGOA). Iron limitation is included in the NPZD model by adding governing equations for two micro-nutrient compartments: dissolved iron and phytoplanktonassociated iron. The model has separate budgets for nitrate (the limiting macro-nutrient in the standard NPZD model) and for iron, with iron limitation on nitrate uptake being imposed as a function of the local phytoplankton realized Fe:C ratio. While the ecosystem model represents a simple approximation of the complex lower trophic level ecosystem of the northwestern CGOA, simulated chlorophyll concentrations reproduce the main characteristics of the spring bloom, high shelf primary production, and ‘‘high-nutrient, low-chlorophyll’’ (HNLC) environment offshore. Over the 1998–2004 period, modeldata correlations based on spatially averaged, monthly mean chlorophyll concentrations are on average 0.7, with values as high as 0.9 and as low as 0.5 for individual years. The model also provides insight on the importance of micro- and macro-nutrient limitation on the shelf and offshore, with the shelfbreak region acting as a transition zone where both nitrate and iron availability significantly impact phytoplankton growth. Overall, the relative simplicity of the ecosystem model provides a useful platform to perform long-term simulations to investigate the seasonal and interannual CGOA ecosystem variability, as well as to conduct sensitivity studies to evaluate the robustness of simulated fields to ecosystem model parameterization and forcing. The ability of the model to differentiate between nitrate-limited, and iron-limited growth conditions, and to identify their spatial and temporal occurrences, is also a first step towards understanding the role of environmental gradients in shaping the complex CGOA phytoplankton community structure.

Modeling the coastal Gulf of Alaska (CGOA) is complicated by the highly diverse physical and biological features influencing productivity and energy flow through the region. The GOA consists of the offshore oceanic environment, characterized by iron limitation, high-nutrients and low-chlorophyll. The coastal environment is consistently downwelling, with high iron levels from glacial melt water and runoff, but lower concentrations of macronutrients, and with a spring bloom, nutrient depletion cycle (low-nutrient, high-chlorophyll). Cross-shelf movement of water masses mixes coastal and oceanic ecosystem elements. Simulations and field data indicate that the minimum model complexity necessary to characterize lower trophic-level production and biomass in the offshore and coastal regions includes 10 boxes: iron, nitrate, ammonium, small phytoplankton, large phytoplankton, small microzooplankton, large microzooplankton, small copepods, large oceanic copepods and detritus, with copepod mortality as a model closure term. We present the model structure, equations required (and initial parameters used) to simulate onshore and offshore lower trophic-level production in the Gulf of Alaska, along with the information from field data and simulations used to construct the model. We show the results of simulations with and without iron, and with and without two size classes of phytoplankton. These simulations indicate that our method of inclusion of iron works well to distinguish the coastal and the oceanic ecosystems, and that the inclusion of two size categories of phytoplankton is also necessary to generate the differences between these two ecosystems.

During a circumnavigation of the Svalbard archipelago in May 2006, simultaneous marine environmental (meteorology, heat flux, ocean turbulence, irradiance) and biological (phytoplankton and zooplankton biomass/species) data were sampled at selected stations. The zooplankton data were supplemented by high-resolution, high-speed VPR sampling down to 100 m depth at most stations. We were able to sample different phases of the phytoplankton spring bloom in Arctic as well as in Atlantic waters, and the stations represented different situations with respect to irradiance, turbulence and water-column stability. Phytoplankton growth and depth distribution were physically controlled, while zooplankton distributions were affected by biological parameters and turbulence. Development of the zooplankton followed the phytoplankton bloom phase, which was progressing in a direction from west to east in the waters north of Svalbard, and southwards in the Barents Sea. Our results also showed that the zooplankton did not avoid Phaeocystis pouchetii colonies, which have earlier been described as toxic. Despite an early retreat of the ice this year there was no apparent mismatch between the phytoplankton bloom and the dominant mesozooplankton, Calanus spp.

The transfer of material through the twilight zone of the ocean is controlled by sinking particles that contain organic matter (OM) and mineral ballast. During the MedFlux field program in the northwestern Mediterranean Sea in 2003, sinking particulate matter was collected in time series (TS) and settling velocity (SV) traps and analyzed for amino acids, lipids, and pigments (along with ballast minerals) [Lee, C., Armstrong, R.A., Wakeham, S.G., Peterson, M.L., Miquel, J.C., Cochran, J.K., Fowler, S.W., Hirschberg, D., Beck, A. Xue, J., 2009b. Particulate matter fluxes in time series and settling velocity sediment traps in the northwestern Mediterranean Sea. Deep-Sea Research II, this volume [ doi:10.1016/j.dsr2.2008.12.003 ]]. The goal was to identify how organic chemical compositions of sinking particles varied as a function of their in-situ settling velocity. The TS record was used to define the biogeochemical character and temporal pattern in flux during the period of SV trap deployment. Temporal variations in organic and mineral compositions are consistent with particle biogeochemistry being driven by the seasonal succession of phytoplankton. Spring diatom bloom conditions led to a high flux of rapidly sinking aggregates and zooplankton fecal matter; summer oligotrophy followed and was characterized by a higher proportion of slowly sinking phytoplankton cells. Bacterial degradation is particularly important during the low-flux summer period. Settling velocity traps show that a large proportion of particulate organic matter sinks at 200–500 m d−1. Organic compositions of this fast-sinking material mirrors that of fecal pellets and aggregated material that sinks as the spring bloom terminates. More-slowly sinking OM bears a stronger signature of bacterial degradation than do the faster-sinking particles. The observation that compositions of SV-sorted fractions are different implies that the particle field is compositionally heterogeneous over a range of settling velocities. Thus physical and biological exchange between fast-sinking and slow-sinking particles as they pass down the water column must be incomplete.

Sediment cores were collected from the shelf, slope, and basin of the Bering, Chukchi, and Beaufort Seas during May–June (under ice cover) and July–August (largely ice-free) 2004. Measurements of chlorophyll a (chl a ), total organic carbon (TOC), and C/N ratios were made in surface and some subsurface core increments. Surface sediment chl a decreased with increasing water depth. Significant positive correlations were found between chl a and TOC and chl a and C/N ratios in the basin (>2000 m), but there were significant negative correlations between chl a and C/N ratios on the shelf (⩽200 m). Chl a values generally declined in down-core profiles, but in some deeper slope and basin cores, measurable inventories of subsurface chl a were present at depth. In some cases, these subsurface chlorophyll inventories coincident with peak activities of the anthropogenic radionuclide 137 Cs were detected, which had maximal deposition following the atmospheric nuclear weapons testing era in the 1960s. A sedimentation rate independently determined for one of these cores using 210 Pb was consistent with the depths of subsurface 137 Cs peaks in slope sediments reflecting steady, relatively undisturbed deposition over a several-decade period. The depth of penetration of 137 Cs in some continental slope sediments, together with detectable chl a, suggests that chl a can be buried in some of these deeper-water sediments under cold conditions for decadal periods in the absence of deposit feeders. Because organic deposition from the water column is episodic at high latitudes and concentrated following the spring bloom, these buried sources of organic materials, whether on the shelf or in deeper basin sediments, may ultimately be important for benthic invertebrates that could utilize this food source during times of the year when primary production flux from the overlying water column is reduced.