Differences in the structure of the lower trophic levels of pelagic ecosystems in the eastern and western subarctic Pacific

To illustrate areal differences in the structure of lower trophic levels of the pelagic ecosystems in the subarctic Pacific, data collected in the quasi-steady state summer/fall conditions were analysed for five areas, i.e. the Bering Basin, Western Subarctic Gyre, the area south of the Aleutians, the Gulf of Alaska, and the Oyashio Region. Average values of stock size of four components of the lower trophic levels showed a clear difference between areas with ranges of 7.5-fold for nitrate, 3.0 for chlorophyll a, 9.9 for microzooplankton, and 2.4 for mesozooplankton. Such differences were more striking when the structure of the lower trophic levels was expressed as a biomass pyramid. In the Gulf of Alaska, Western Subarctic Gyre, and south of the Aleutians, the relative biomass of microzooplankton to phytoplankton is large and large amounts of nitrate remained unused. In addition to possible iron limitation, grazing control by the microzooplankton on small phytoplankton must be substantial in these areas. Conversely, in the Oyashio Region, the nitrate stock is very small indicating higher efficiency of nitrate consumption by phytoplankton. However, since phytoplankton and zooplankton stocks are not particularly large, their products are likely to be transferred, also efficiently, to the higher trophic levels such as planktivorous pelagic fish. The situation in the Bering Basin is intermediate between the Oyashio Region and the other three areas. Inter-annual fluctuations in stock size of the planktivorous fish which migrate into the Oyashio Region in summer/fall were quite large. However, the inter-annual variation of mesozooplankton biomass was small, suggesting the existence of certain mechanisms to stabilize plankton abundance under increasing predation pressure. As a result, the increasing fish stocks likely keep the transfer efficiency from nitrate through to fish higher, at least in the Oyashio Region.     

High feeding rates on large particles by Neocalanus flemingeri and N. plumchrus,and consequences for phytoplankton community structure in the subarctic Pacific Ocean

The open subarctic Pacific Ocean is a high nitrate low chlorophyll (HNLC) system characterized by low concentrations of phytoplankton, a community dominated by small cells, and iron-limited growth of, especially, the larger phytoplankton. In such systems the main energy and material flow is through the microbial web, with large copepods considered primarily to be grazers on the larger microzooplankton occupying the top of this web. Consistent with this is the recognition that much of the nutrition of the dominant copepods in this system, Neocalanus flemingeri, N. plumchrus and N. cristatus, is derived from microzooplankton. Also, these copepods consume only a small fraction of the total phytoplankton production. In this paper, we show that the contribution made by N. flemingeri and N. plumchrus to establishing and maintaining the community structure of this ecosystem should be re-evaluated. Our experiments indicate these grazers have high clearance rates on large particles, including both large phytoplankton and microzooplankton, and this selective removal contributes to establishment and maintenance of the observed foodweb structure in the Gulf of Alaska. These high feeding rates combined with large populations of these two Neocalanus species concentrated in the upper layer of the ocean, result in population-based feeding rates approximately equal to the growth rates of large phytoplankton under iron-limited conditions. We conclude that N. flemingeri and N. plumchrus populations (a) directly prevent the accumulation of large phytoplankton cells by selectively feeding on them at high rates, and (b) indirectly stimulate the accumulation of the smaller phytoplankton by consumption of their major grazers, the microzooplankton.     

Modeling iron limitation of primary production in the coastal Gulf of Alaska

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 phytoplankton-associated 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, model-data 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.     

Modelling the response of the planktonic food web to iron fertilization and warming in the NE subarctic Pacific

A one-dimensional ecosystem model with two explicit size classes of phytoplankton was developed for the NE subarctic Pacific to investigate variations in the export of organic particles to the ocean interior due to potential changes in the environment. Specifically, the responses of the planktonic ecosystem to permanent removal of iron limitation and to warming (of 2 and 5 °C) were explored. The ecosystem model consists of five components (small and large phytoplankton, microzooplankton, detritus and nitrogen), and includes grazing by mesozooplankton that varies in time according to long-term observations at Ocean Station Papa (OSP). The model addresses the role of iron limitation on phytoplankton growth and includes temperature dependence of physiological rates. The ecosystem model was forced with annual wind and solar heating from OSP. The model best reproduced the low chlorophyll high nitrate conditions of the NE subarctic Pacific when both small and large phytoplankton were limited by iron such that their maximum specific growth rate was reduced by 10 and 70%, respectively. Sensitivity analysis showed that model results depended on the value of the iron limitation parameter of large phytoplankton (L_Fe-L) and the grazing parameters of micro- and mesozooplankton. To explore the effect of iron limitation, simulations were carried out varying the iron limitation parameters while maintaining the nitrogen flux at the base of the model constant and the grazing pressure by mesozooplankton unchanged. In the warming case, simulations were carried out increasing ocean temperatures by 2° and 5 °C applied only to the ecological components, the flux of nitrate at the base of the model was increased to obtain a steady annual cycle, and grazing by mesozooplankton remained constant. When compared with the standard case, model simulations indicated that both permanent removal of iron limitation and warming cause changes in food web structure and the carbon cycle. The response was more dramatic in the iron-replete case where the phytoplankton community structure in spring changed from one dominated by pico- and nanoplankton to one dominated by large phytoplankton, and primary production increased until it consumed all the external nutrient (N) supply to the upper layer. However, reducing iron deficiency actually led to lower annual primary production due to a decrease in the regeneration of nitrogen in the euphotic zone. These changes in food web structure influenced the magnitude, composition and seasonal cycle of sinking particles.     

Iron regeneration and organic iron(III)-binding ligand production during in situ zooplankton grazing experiment

To elucidate iron regeneration and organic iron(III)-binding ligand formation during microzooplankton and copepod grazing on phytoplankton, incubation experiments were conducted in the western subarctic Pacific. During 8 days of dark incubation of ambient water and that amended with plankton concentrate, dissolved iron and organic iron(III)-binding ligands accumulated, approximately proportionally to the decrease in chlorophyll a. The observed increases in dissolved iron concentration were much greater than those expected from the consumption of phytoplankton biomass and previously reported Fe:C value of cultured algal cells, suggesting resolution from colloidal or particulate iron adsorbed onto the algal cell surface. When copepods were added to the ambient water, organic iron(III)-binding ligands accumulated more rapidly than in the control receiving no copepod addition, although consumed phytoplankton biomass was comparable between the two treatments. Bioassay experiment using filtrates collected from the incubation experiment showed that organic ligands formed during microzooplankton grazing reduced the iron bioavailability to phytoplankton and suppressed their growth. Moreover, picoplankton Synechococcus sp. and Micromonas pusilla were more suppressed by the organic ligands than the diatom Thalassiosira weissflogii. In conclusion, through microzooplankton and copepod grazing on phytoplankton, organic iron(III)-binding ligands as well as regenerated iron are released into the ambient seawater. Because the ligands lower iron bioavailability to phytoplankton through complexation and the degree of availability reduction varies among phytoplankton species, grazing by zooplankton can shift phytoplankton community structure in iron-limited waters.     

Response of microzooplankton (protists and small copepods) to an iron-induced phytoplankton bloom in the Southern Ocean (EisenEx)

The dynamics, composition and grazing impact of microzooplankton were studied during the in situ iron fertilisation experiment EisenEx in the Antarctic Polar Frontal Zone in austral spring (November 2000). During the 21 day experiment, protozooplankton and small metazooplankton were sampled from the mixed layer inside and outside the patch using Niskin bottles. Aplastidic dinoflagellates increased threefold in abundance and biomass in the first 10 days of the experiment, but decreased thereafter to values twofold higher than pre-fertilisation values. The decline after day 10 is attributed to increasing grazing pressure by copepods. They also constrained ciliate abundances and biomass which were higher inside the fertilised patch than outside but highly variable. Copepod nauplii abundance remained stable whereas biomass doubled. Numbers of copepodites and adults of small copepod species (<1.5 mm) increased threefold inside the patch, but doubled in surrounding waters. Grazing rates estimated using the dilution method suggest that microzooplankton grazing constrained pico- and nanoplankton populations, but species capable of feeding on large diatoms (dinoflagellates and small copepods including possibly nauplii) were selectively predated by the metazoan community. Thus, iron fertilisation of a developing spring phytoplankton assemblage resulted in a trophic cascade which favoured dominance of the bloom by large diatoms.     

Diet of sardine Sardinops sagax in the southern Benguela upwelling ecosystem

The diet of sardine Sardinops sagax in the southern Benguela was investigated by microscopic examination of stomach contents. The relative dietary importance of prey size and prey type was assessed by calculating the carbon content of prey items. Sardine is an omnivorous clupeoid, ingesting both phytoplankton and zooplankton, with the relative importance of these two food types varying both spatially and temporally. Stomach contents were numerically dominated by small prey items, principally dinoflagellates, followed by crustacean eggs, cyclopoid copepods, calanoid copepods and diatoms. Virtually all prey items ingested by sardine were <1.2 mm maximum dimension, the particle size below which sardine only filter-feed. Despite the numerical dominance by phytoplankton, zooplankton contributed the major portion to sardine dietary carbon, small calanoid and cyclopoid copepods, anchovy eggs and crustacean eggs being the primary prey types. These results indicate that, like anchovy Engraulis capensis, sardine in the southern Benguela are primarily zoophagous, and contrast with earlier dietary studies on sardine in the region. However, the two species appear to partition their prey on the basis of size; sardine consume small zooplankton, whereas anchovy consume large zooplankton. This difference has been observed in other upwelling ecosystems where the two genera co-exist and is likely to contribute to the regime shifts observed between sardine and anchovy.     

Structure, biomass distribution and trophodynamics of the pelagic ecosystem in the Oyashio region, western subarctic Pacific

Biomass distribution and trophodynamics in the oceanic ecosystem in the Oyashio region are presented and analyzed, combining the seasonal data for plankton and micronekton collected at Site H since 1996 with data for nekton and other animals at higher trophic levels from various sources. The total biomass of biological components including bacteria, phytoplankton, microzooplankton, mesozooplankton, micronekton, fishes/squids and marine birds/mammals was 23 g C m⁻², among which the most dominant component was mesozooplankton (34% of the total), followed by phytoplankton (28%), bacteria (15%) and microzooplankton (protozoans) (14%). The remainder (9%) was largely composed of micronekton and fish/squid. Marine mammals/birds are only a small fraction (0.14%) of the total biomass. Large/medium grazing copepods (Neocalaus spp., Eucalanus bungii and Metridia spp.) accounted for 77% of the mesozooplankton biomass. Based on information about diet composition, predators were assigned broadly into mean trophic level 3–4, and carbon flow through the grazing food chain was established based on the estimated annual production/food consumption balance of each trophic level. From the food chain scheme, ecological efficiencies as high as 24% were calculated for the primary/secondary production and 21% for the secondary/tertiary production. Biomass and production of bacteria were estimated as 1/10 of the respective values for phytoplankton at Site H, but the role of the microbial food chain remains unresolved in the present analysis. As keystone species in the oceanic Oyashio region, Neocalanus spp. are suggested as a vital link between primary production and production of pelagic fishes, mammals and birds.     

Eddy-resolving simulation of plankton ecosystem dynamics in the California Current System

We study the dynamics of the planktonic ecosystem in the coastal upwelling zone within the California Current System using a three-dimensional (3-D), eddy-resolving circulation model coupled to an ecosystem/biogeochemistry model. The physical model is based on the Regional Oceanic Modeling System (ROMS), configured at a resolution of 15 km for a domain covering the entire US West Coast, with an embedded child grid covering the central California upwelling region at a resolution of 5 km. The model is forced with monthly mean boundary conditions at the open lateral boundaries as well as at the surface. The ecological/biogeochemical model is nitrogen based, includes single classes for phytoplankton and zooplankton, and considers two detrital pools with different sinking speeds. The model also explicitly simulates a variable chlorophyll-to-carbon ratio. Comparisons of model results with either remote sensing observations (AVHRR, SeaWiFS) or in-situ measurements from the CalCOFI program indicate that our model is capable of replicating many of the large-scale, time-averaged features of the coastal upwelling system. An exception is the underestimation of the chlorophyll levels in the northern part of the domain, perhaps because of the lack of short-term variations in the atmospheric forcing. Another shortcoming is that the modeled thermocline is too diffuse, and that the upward slope of the isolines toward the coast is too small. Detailed time-series comparisons with observations from Monterey Bay reveal similar agreements and discrepancies. We attribute the good agreement between the modeled and observed ecological properties in large part to the accuracy of the physical fields. In turn, many of the discrepancies can be traced back to our use of monthly mean forcing. Analysis of the ecosystem structure and dynamics reveal that the magnitude and pattern of phytoplankton biomass in the nearshore region are determined largely by the balance of growth and zooplankton grazing, while in the offshore region, growth is balanced by mortality. The latter appears to be inconsistent with in situ observations and is a result of our consideration of only one zooplankton size class (mesozooplankton), neglecting the importance of microzooplankton grazing in the offshore region. A comparison of the allocation of nitrogen into the different pools of the ecosystem in the 3-D results with those obtained from a box model configuration of the same ecosystem model reveals that only a few components of the ecosystem reach a local steady-state, i.e. where biological sources and sinks balance each other. The balances for the majority of the components are achieved by local biological source and sink terms balancing the net physical divergence, confirming the importance of the 3-D nature of circulation and mixing in a coastal upwelling system.     

Phytoplankton growth and microzooplankton grazing in the subarctic Pacific Ocean and the Bering Sea during summer 1999

Phytoplankton growth and microzooplankton grazing rates were measured by the dilution technique in the subarctic North Pacific Ocean along a west–east transect during summer 1999. Average phytoplankton growth rates without added nutrients (μ₀) were 0.33, 0.41, 0.20 and 0.49 d⁻¹ for the four regions sampled: the Western Gyre, the Bering Sea, the Gulf of Alaska gyre and stations along the Aleutian Trench. Average grazing mortality rates (m) were 0.34, 0.27, 0.20 and 0.49 d⁻¹. Limitation of phytoplankton growth by macronutrients, such as NO₃ and SiO₂, was identified only at a few stations, with overall μ₀/μ_n (μ_n is nutrient-enhanced growth rate) averaging 0.9. Phytoplankton growth and microzooplankton grazing were approximately balanced, as indicated by high m/μ₀ ratio, except in the Bering Sea, where the m/μ₀ ratio was 0.65, indicating the relative importance of the diatom-macrozooplankton grazing food chain and possible higher export flux to the deep layer. Flow cytometric analysis revealed that the growth rates of picoplankton (Synechococcus and picoeukaryotes) were usually much lower than the total phytoplankton community growth rates estimated from chlorophyll a, except for stations in the Gulf of Alaska Gyre, where the growth rates for different populations were about the same. Lower than community-average growth rate for picoplankton indicates larger phytoplankters, presumably diatoms, were growing at a much faster rate. Suppressed phytoplankton growth in the Gulf of Alaska was probably a result of iron limitation.     

Mesozooplankton in the eastern and western subarctic Pacific: community structure, seasonal life histories, and interannual variability

The zooplankton community of the subarctic Pacific is relatively simple, and contains a similar set of major species in all deep water areas of the subarctic Pacific. Their role in the food web varies considerably between coastal and offshore locations. In the oceanic gyres, microzooplankton and other mesozooplankton taxa replace phytoplankton as the primary food source for the dominant mesozooplankton species. Micronekton and larger zooplankton probably replace pelagic fish as major direct predators. Productivity and upper ocean biomass concentrations are intensely seasonal, in part because of seasonality of the physical environment and food supply, but also because of life history patterns involving seasonal vertical migrations (400–2000 m range) and winter dormancy. During the spring–summer season of upper ocean growth, small scale horizontal and vertical patchiness is intense. This can create local zones of high prey availability for predators such as planktivorous fish, birds, and marine mammals. On average, the cores of the subarctic gyres have lower biomass and productivity than the margins of the gyres. There is also some evidence that the Western Gyre is more productive than the Alaska Gyre, but more research is needed to confirm whether this east–west gradient is permanent. There is increasing evidence that the pattern of zooplankton productivity is changing over time, probably in response to interdecadal ocean climate variability. These changes include 2–3 fold shifts in total biomass, 30–60 day shifts in seasonal timing, and 10–25% changes in average body length.     

Modeling the long-term variability of phytoplankton functional groups and primary productivity in the South China Sea

Primary productivity (PP) and phytoplankton structure play an important role in regulating oceanic carbon cycle. The unique seasonal circulation and upwelling pattern of the South China Sea (SCS) provide an ideal natural laboratory to study the response of nutrients and phytoplankton dynamics to climate variation. In this study, we used a three-dimensional (3D) physical–biogeochemical coupled model to simulate nutrients, phytoplankton biomass, PP, and functional groups in the SCS from 1958 to 2009. The modeled results showed that the annual mean carbon composition of small phytoplankton, diatoms, and coccolithophores was 33.7, 52.7, and 13.6 %, respectively. Diatoms showed a higher seasonal variability than small phytoplankton and coccolithophores. Diatoms were abundant during winter in most areas of the SCS except for the offshore of southeastern Vietnam, where diatom blooms occurred in both summer and winter. Higher values of small phytoplankton and coccolithophores occurred mostly in summer. Our modeled results indicated that the seasonal variability of PP was driven by the East Asian Monsoon. The northeast winter monsoon results in more nutrients in the offshore area of the northwestern Luzon Island and the Sunda Shelf, while the southwest summer monsoon drives coastal upwelling to bring sufficient nutrients to the offshore area of southeastern Vietnam. The modeled PP was correlated with El Niño/Southern Oscillation (ENSO) at the interannual scale. The positive phase of ENSO (El Niño conditions) corresponded to lower PP and the negative phase of ENSO (La Niña conditions) corresponded to higher PP.     

Quantifying cross-shelf and vertical nutrient flux in the Coastal Gulf of Alaska with a spatially nested,coupled biophysical model

The Coastal Gulf of Alaska (CGOA) is productive, with large populations of fish, seabirds, and marine mammals; yet it is subject to downwelling-favorable coastal winds. Downwelling regions in other parts of the world are typically much less productive than their upwelling counterparts. Alternate sources of nutrients to feed primary production in the topographically complex CGOA are poorly known and difficult to quantify. Here we diagnose the output from a spatially nested, coupled hydrodynamic and lower trophic level model of the CGOA, to quantify both horizontal and vertical nutrient fluxes into the euphotic zone. Our nested model includes both nitrogen and iron limitation of phytoplankton production, and is driven by a fine-scale atmospheric model that resolves the effects of local orography on the coastal winds. Results indicate significant “rivers” of cross-shelf nitrogen flux due to horizontal advection, as well as “fountains” of vertical transport over shallow banks due to tidal mixing. Using these results, we constructed a provisional budget of nutrient transport among subregions of the CGOA. Contrary to expectations, this budget reveals substantial upwelling of nutrients over major portions of the shelf, driven by local wind-stress curl. These effects are large enough to overwhelm the smaller downwelling flux at the coast throughout the growing season. Vertical mixing by winds and tides, and horizontal flux from the deep basin, are other substantial contributors to nutrients above the 15-m horizon. These findings help to explain the productivity of this coastal ecosystem.     

The threshold feeding response of microzooplankton within Pacific high-nitrate low-chlorophyll ecosystem models under steady and variable iron input

The equatorial Pacific is an HNLC (High-Nitrate Low-Chlorophyll) region. Modeling and in-situ process studies have confirmed the importance of microzooplankton grazing in this ecosystem. Unfortunately, both the parameters and functions representing microzooplankton grazing within current ecosystem models are poorly constrained. We used a simple 4-component food web model to test the assumption that a lower grazing threshold, which is common in many models, is necessary to achieve the HNLC condition. Without the grazing threshold, the model did not reproduce the HNLC condition. However, by raising the half-saturation constant within the microzooplankton functional response with no threshold, it was possible to reproduce the critical dynamics of the HNLC condition under both steady and moderate seasonal variability in nutrient input. It was also possible to reproduce the HNLC system using a sigmoidal functional response for the microzooplankton, with results somewhere between the other two forms of the model, although this version had the highest sensitivity to changes in its parameters. The three models predicted similar phytoplankton biomass and primary productivity under steady nutrient input, but diverge in these metrics as the amplitude of nutrient input variability increases. These three functional responses also imply certain important differences in the microzooplankton community. Whereas the threshold model had the least sensitivity to parameter choice, the high half-saturation constant, no-threshold model may actually be a better approximation when modeling a community of grazers. Ecosystem models that predict carbon production and export in HNLC regions can be very sensitive to assumptions concerning microzooplankton grazing; future studies need to concentrate on the functional responses of microzooplankton before these models can be used for predicting fluxes in times or regions where forcing is beyond that used to constrain the original model.     

Seasonal and spatial distributions of phytoplankton biomass associated with monsoon and oceanic environments in the South China Sea

Seasonal variations of phytoplankton/chlorophyll-a （Chl-a） distribution, sea surface wind, sea height anomaly, sea surface temperature and other oceanic environments for long periods are analyzed in the South China Sea （SCS）, especially in the two typical regions off the east coast of Vietnam and off the northwest coast of Luzon, using remote sensing data and other oceanographic data. The results show that seasonal and spatial distributions of phytoplankton biomass in the SCS are primarily influenced by the monsoon winds and oceanic environments. Off the east coast of Vietnam, Chl-a concentration is a peak in August, a jet shape extending into the interior SCS, which is associated with strong southwesterly monsoon winds, the coastal upwetling induced by offshore Ekman transport and the strong offshore current in the western SCS. In December, high Chl-a concentration appears in the upwelling region off the northwest coast of Luzon and spreads southwestward. Strong mixing by the strong northeasterly monsoon winds, the cyclonic circulation, southwestward coastal currents and river discharge have impacts on distribution of phytoplankton, so that the high phytoplankton biomass extends from the coastal areas over the northern SCS to the entire SCS in winter. These research activities could be important for revealing spatial and temporal patterns of phytoplankton and their interactions with physical environments in the SCS.     

A comparison of biological trends from four marine ecosystems: Synchronies, differences, and commonalities

Major features of four marine ecosystems were analyzed based on a broad range of fisheries-associated datasets and a suite of oceanographic surveys. The ecosystems analyzed included the Gulf of Maine/Georges Bank in the Northwest Atlantic Ocean, the Norwegian/Barents Seas in the Northeast Atlantic Ocean, and the eastern Bering Sea and the Gulf of Alaska in the Northeast Pacific Ocean. We examined survey trends in major fish abundances, total system fish biomass, and zooplankton biomasses. We standardized each time series and examined trends and anomalies over time, using both time series and cross-correlational statistical methods. We compared dynamics of functionally analogous species from each of these four ecosystems. Major commonalities among ecosystems included a relatively stable amount of total fish biomass and the importance of large calanoid copepods, small pelagic fishes and gadids. Some of the changes in these components were synchronous across ecosystems. Major differences between ecosystems included gradients in the magnitude of total fish biomass, commercial fish biomass, and the timing of major detected events. This work demonstrates the value of comparative analysis across a wide range of marine ecosystems, suggestive of very few but none-the-less detectable common features across all northern hemisphere ocean systems.     

冬季南海北部海域微型浮游动物及其对浮游植物摄食压力研究

2009年1月在南海北部海域的5个站位,采用稀释法和显微分析技术研究了浮游植物生长率及微型浮游动物对浮游植物的摄食压力,同时测定了微型浮游动物的丰度及类群组成.结果表明:南海北部微型浮游动物类群主要以无壳纤毛虫为主,南海北部微型浮游动物类群细胞丰度为33～529个/dm3.南海北部浮游植物生长率为0.45～1.83 d-1,微型浮游动物摄食率为0.44～1.76 d-1,摄食压力占浮游植物现存量的42.6%～82.8%,占初级生产力的97.3%～225.1%.近岸区摄食压力比陆架区高,表明冬季南海近岸区微型浮游动物摄食能够有效的控制浮游植物的生长,而陆架区浮游植物生长率大于摄食率,浮游植物存在着现存量的积累,微型浮游动物并不能完全控制浮游植物的生长.     

A numerical analysis of carbon dynamics of the Southern Ocean phytoplankton community: the roles of light and grazing in effecting both sequestration of atmospheric CO2 and food availability to larval krill

Reduced ice extent within coastal regions of the Southern Ocean may lead to deeper surface mixed layers (SML), as prevail in offshore areas. A future decline of ice melt-induced stability of the water column may be associated with a shift in dominant food webs, from larger, sun-adapted diatoms grazed by euphausiids to smaller, shade-adapted flagellates consumed by salps. A basically one-dimensional numerical model of three dominant groups of the Antarctic phytoplankton community (diatoms, cryptophytes, and colonial prymnesiophytes) and four types of herbivore (protozoans, salps, copepods, and euphausiids) is used to explore the seasonal importance of both light limitation and grazing pressure on the amount of annual carbon sequestration and larval krill survival within contrasting oceanic and neritic waters, where respective validation data have been gathered during austral spring by the European JGOFS and RACER programs. With imposition of moderate and large grazing stresses, thought to be typical of offshore waters, we were able to replicate the European JGOFS 1992 observations of light penetration, phytoplankton biomass, primary production, pCO₂, bacterial biomass, labile DOC, ammonium, and total particle effluxes at 100 m within the deep SML of our model. The fidelity of such a large set of simulated state variables suggests that multiple limiting factors are indeed operating on different components of the oceanic phytoplankton community — selective grazing losses on the flagellates, but light limitation of diatoms. Release of protozoan grazing pressure in our model instead leads to unobserved spring blooms of cryptophytes, found only in laboratory enclosures. On an annual basis, weak sequestration of atmospheric CO₂ is simulated in a habitat typical of the Polar Front, while evasion of carbon dioxide occurs under biophysical conditions of the Antarctic Circumpolar Current. Stratification in shallow SML and the same absolute grazing demands by krill and copepods allows sun-adapted diatoms of our model to bloom at the expense of shade-adapted cryptophytes and prymnesiophytes, eaten by salps and protozoans. We were also able to replicate RACER-I observations of the same suite of variables in 1986–1987, as well as the observed 10-fold range of detrital fluxes caught by other sediment trap deployments during 1980 and 1983 along the Antarctic Peninsula. In western Bransfield Strait, coastal waters are a strong sink for atmospheric CO₂ within parcels of Bellingshausen Sea origin, but not perhaps in those from the Weddell Sea, which resemble the oceanic regime of deep SML. We conclude that even in shallow neritic SML, some protozoan rivals of larval krill must still crop flagellates to ensure sufficient abundance of diatom food for both euphausiid survival and possible clogging of the mucous nets of other salp rivals.     

Decadal-Scale Climate and Ecosystem Interactions in the North Pacific Ocean

Decadal-scale climate variations in the Pacific Ocean wield a strong influence on the oceanic ecosystem. Two dominant patterns of large-scale SST variability and one dominant pattern of large-scale thermocline variability can be explained as a forced oceanic response to large-scale changes in the Aleutian Low. The physical mechanisms that generate this decadal variability are still unclear, but stochastic atmospheric forcing of the ocean combined with atmospheric teleconnections from the tropics to the midlatitudes and some weak ocean-atmosphere feedbacks processes are the most plausible explanation. These observed physical variations organize the oceanic ecosystem response through large-scale basin-wide forcings that exert distinct local influences through many different processes. The regional ecosystem impacts of these local processes are discussed for the Tropical Pacific, the Central North Pacific, the Kuroshio-Oyashio Extension, the Bering Sea, the Gulf of Alaska, and the California Current System regions in the context of the observed decadal climate variability. The physical ocean-atmosphere system and the oceanic ecosystem interact through many different processes. These include physical forcing of the ecosystem by changes in solar fluxes, ocean temperature, horizontal current advection, vertical mixing and upwelling, freshwater fluxes, and sea ice. These also include oceanic ecosystem forcing of the climate by attenuation of solar energy by phytoplankton absorption and atmospheric aerosol production by phytoplankton DMS fluxes. A more complete understanding of the complicated feedback processes controlling decadal variability, ocean ecosystems, and biogeochemical cycling requires a concerted and organized long-term observational and modeling effort. This revised version was published online in July 2006 with corrections to the Cover Date.     

Copepod grazing during spring blooms: Can Pseudocalanus newmani induce trophic cascades?

During late winter and spring of 2002 and 2003, 24 two- to three-day cruises were conducted to Dabob Bay, Washington State, USA, to examine the grazing, egg production, and hatching success rates of adult female Calanus pacificus and Pseudocalanus newmani. Here, we discuss the results of our grazing experiments for P. newmani. Each week, we conducted traditional microzooplankton dilution experiments and “copepod dilution” experiments, each from two different layers. Grazing was measured by changes in chlorophyll concentration and direct cell counts. Clearance rates on individual prey species, as calculated by cell counts, showed that Pseudocalanus are highly selective in their feeding, and may have much higher grazing rates on individual taxa than calculated from bulk chlorophyll disappearance. The grazing rates of the copepods, however, are typically an order of magnitude lower than the grazing rates of the microzooplankton community, or the growth rates of the phytoplankton. P. newmani ingested diatoms, but, at certain times fed preferentially on microzooplankton, such as ciliates, tintinnids, and larger dinoflagellates. Removal of the microzooplankton may have released the other phytoplankton species from grazing pressure, allowing those species’ abundance to increase, which was measured as an apparent “negative” grazing on those phytoplankton species. The net result of grazing on some phytoplankton species, while simultaneously releasing others from grazing pressure resulted in bulk chlorophyll-derived estimates of grazing which were essentially zero or slightly negative; thus bulk chlorophyll disappearance is a poor indicator of copepod grazing. Whether copepods can significantly release phytoplankton from the grazing pressure by microzooplankton in situ, thus causing a trophic cascade, remains to be verified, but is suggested by our study.