Iron fertilization and the structure of planktonic communities in high nutrient regions of the Southern Ocean

In this review article, plankton community structure observations are analyzed both for artificial iron fertilization experiments and also for experiments dedicated to the study of naturally iron-fertilized systems in the Atlantic, Indian and Pacific sectors of the Southern Ocean in the POOZ (Permanently Open Ocean Zone) and the PFZ (Polar Frontal Zone). Observations made in natural systems are combined with those from artificially perturbed systems, in order to evaluate the seasonal evolution of pelagic communities, taking into account controlling factors related to the life cycles and the ecophysiology of dominant organisms. The analysis considers several types of planktonic communities, including both autotrophs and heterotrophs. These communities are spatially segregated owing to different life strategies. A conceptual general scheme is proposed to account for these observations and their variability, regardless of experiment type. Diatoms can be separated into 2 groups: Group 1 has slightly silicified fast growing cells that are homogeneously distributed in the surface mixed layer, and Group 2 has strongly silicified slowly growing cells within discrete layers. During the growth season, Group 1 diatoms show a typical seasonal succession of dominant species, within time windows of development that are conditioned by physical factors (light and temperature) as well as endogenous specific rhythms (internal clock), and biomass accumulation is controlled by the availability of nutrients. Group 1 diatoms are not directly grazed by mesozooplankton which is fed by protozooplankton, linking the microbial food web to higher trophic levels. Instead, successive dominant species of Group 1 are degraded via bacterial activity at the end of their growth season. Organic detritus fragments feed protozooplankton and mesozooplankton. The effective silicon pump leads to the progressive disappearance of silicic acid in surface waters. In contrast, Group 2 is resistant to grazing due to its strong silicification, and its biomass accumulates continuously but relatively slowly throughout the productive period. Group 2 diatoms are concentrated at or near the seasonal pycnocline and thus benefit from upward nutrient fluxes by diapycnal mixing. The decrease in light and the deep convective mixing in the fall produce both light and nutrient limitation leading to a massive carbon export of Group 2 diatoms, a major annual event of the biological pump. This scheme describes the seasonal evolution of plankton communities in surface waters of the Southern Ocean. The scheme could probably be extended to ecosystems that are characterized by a seasonal bloom under influence of iron or other nutrients.     

Nutrient Cycling and a Stoichiometric Model in Epidavros, a Deep Basin in the Aegean Sea

Abstract. The nutrient cycling of Epidavros, a deep basin in the Saronikos Gulf, was studied in relation to various environmental factors during 1973–1976 at a station characterized by stagnant conditions. The regeneration of nutrients was related to the consumption of oxygen, and a seasonal nutrient cycle occurred with low nutrient concentrations in the spring and summer, followed by high nutrient levels in autumn and winter. In addition high values of nitrate and silicate were observed in the deeper waters, which tended to be anoxic, although the water masses were renewed during spring 1974. The distribution pattern of nutrients together with nutrient ratios were compared with previous studies of the same and neighbouring areas as well as of other isolated basins. A stoichiometric model indicates that plankton organisms in the Epidavros basin have approximate atomic ratios for C: N: P of 150: 14:1, while the ratio of change for nitrogen and phosphorus in the water is only 8.8:1 by atoms. This is probably because of the slow rate of regeneration of nitrogenous material and/or assimilation and regeneration in organic forms. The water/plankton relation in the Epidavros basin appears to be very similar to that in the Baltic Sea.     

Adaptive model of plankton dynamics for the North Atlantic

Plankton ecosystems in the North Atlantic display strong regional and interannual variability in productivity and trophic structure, which cannot be captured by simple plankton models. Additional compartments subdividing functional groups can increase predictive power, but the high number of parameters tends to compromise portability and robustness of model predictions. An alternative strategy is to use property state variables, such as cell size, normally considered constant parameters in ecosystem models, to define the structure of functional groups in terms of both behaviour and response to physical forcing. This strategy may allow us to simulate realistically regional and temporal differences among plankton communities while keeping model complexity at a minimum.We fit a model of plankton and DOM dynamics globally and individually to observed climatologies at three diverse locations in the North Atlantic. Introducing additional property state variables is shown to improve the model fit both locally and globally, make the model more portable, and help identify model deficiencies. The zooplankton formulation exerts strong control on model performance. Our results suggest that the current paradigm on zooplankton allometric functional relationships might be at odds with observed plankton dynamics. Our parameter estimation resulted in more realistic estimates of parameters important for primary production than previous data assimilation studies.Property state variables generate complex emergent functional relationships, and might be used like tracers to differentiate between locally produced and advected biomass. The model results suggest that the observed temperature dependence of heterotrophic growth efficiency [Rivkin, R.B., Legendre, L., 2001. Biogenic carbon cycling in the upper ocean: effects of microbial respiration. Science 291 (5512) 2398-2400] could be an emergent relation due to intercorrelations among temperature, nutrient concentration and growth efficiency. A major advantage of using property state variables is that no additional parameters are required, such that differences in model performance can be directly related to model structure rather than parameter tuning.     

Effects of daily nitrogen and phosphorus input on planktonic community metabolism in a semi-enclosed bay by mesocosm experiment

Planktonic metabolism plays an important role in affecting the energy transportation and carbon cycle of the marine ecosystem. However, its regulation mechanism remains unclear under the continuously exogenous nutrient inputs in nearshore waters. In this study, a mesocosm experiment was conducted in a semi-enclosed bay, the Daya Bay, to explore the responses of plankton metabolic balance and community structure to a concentration gradient of daily nitrogen and phosphorus inputs. The results showed that nutrient enrichments promoted phytoplankton biomass, total primary production, and community respiration, and the promoting effect enhanced alongwith the increase of nutrient concentration. However, the net community production fluctuated more violently between autotrophic and heterotrophic with the increase of nutrient inputs and tended to be more heterotrophic from the 5th day to the 10th day of the experiment. In addition, the daily flux of nitrogen and phosphorus, 2 μmol/(L·d) and 0.066 μmol/(L·d), respectively, could be regarded as a potential threshold for ecosystem stability and health, since most of the ecological characteristics related to plankton structure and function have undergone significant changes when the nutrient level is higher than that. In the nearshore enclosed or semi-enclosed waters, nutrient from continuous terrigenous input is likely to be concentrated and exceed this level, indicating the ecological risks due to the metabolic unbalance and the deterioration of plankton community structure.     

Simulated biogeochemical responses to iron enrichments in three high nutrient, low chlorophyll (HNLC) regions

To fill temporal gaps in iron-enrichment experimental data and gain further understanding of marine ecosystem responses to iron enrichments, we apply a fifteen-compartment ecosystem model to three iron-enrichment sites, namely SEEDS (the Subarctic Pacific Iron Experiment for Ecosystem Dynamics Study; 48.5°N, 165°E) in the western North Pacific, SOIREE (the Southern Ocean Iron RElease Experiment; 61°S, 140°E) in the Southern Ocean, and IronExII (the second mesoscale iron enrichment experiment; 3.5°S, 104°W) in the Equatorial Pacific. The ecological effects of iron in the model are represented by changing two photosynthetic parameters during the iron-enrichment period. The model results successfully reproduce the observed biogeochemical responses inside and outside the iron patch at each site, such as rapid increases in plankton biomass and biological productivity, and decreases in surface nutrients and pCO₂, inside the patch. However, the modeled timing and magnitude of changes differ among the sites because of differences in both physical environments and plankton species. After the iron enrichment, the diatom productivity is strongly controlled by light at SOIREE and by silicate at IronExII and SEEDS. Light limitation due to self-shading by the phytoplankton is significant during the bloom at all sites. Sensitivity analysis of the model results to duration of the iron enrichment reveals that long-term multiple infusions over more than a week would not be effective at SEEDS because of strong silicate limitation on diatom growth. Sensitivity of the model to water temperature shows that export production is higher at lower temperatures, because of slower recycling of particulate organic carbon. Therefore, the e-ratio (the ratio of export production to primary production) is inversely correlated with temperature, and the relationship can be described with a linear function. Through this study, we conclude that ecosystem modeling is a powerful tool to help design future iron-enrichment experiments and observational plans.     

Enhancement/reduction of biological pump depends on ocean circulation in the sea-ice reduction regions of the Arctic Ocean

The biological pump is a central process in the ocean carbon cycle, and is a key factor controlling atmospheric carbon dioxide (CO₂). However, whether the Arctic biological pump is enhanced or reduced by the recent loss of sea ice is still unclear. We examined if the effect was dependent on ocean circulation. Melting of sea ice can both enhance and reduce the biological pump in the Arctic Ocean, depending on ocean circulation. The biological pump is reduced within the Beaufort Gyre in the Canada Basin because freshwater accumulation within the gyre limits nutrient supply from deep layers and shelves hence inhibits the growth of large-bodied phytoplankton. Conversely, the biological pump is enhanced outside the Beaufort Gyre in the western Arctic Ocean because of nutrient supply from shelves and greater light penetration, enhancing photosynthesis, caused by the sea ice loss. The biological pump could also be enhanced by sea ice loss in the Eurasian Basin, where uplifted isohaline surfaces associated with the Transpolar Drift supply nutrients upwards from deep layers. New data on nitrate uptake rates are consistent with the pattern of enhancement and reduction of the Arctic biological pump. Our estimates indicate that the enhanced biological pump can be as large as that in other oceans when the sea ice disappears. Contrary to a recent conclusion based on data from the Canada Basin alone, our study suggests that the biological CO₂ drawdown is important for the Arctic Ocean carbon sink under ice-free conditions.     

Roles of Biogeochemical Processes in the Oceanic Carbon Cycle Described with a Simple Coupled Physical-Biogeochemical Model

To evaluate the contribution of biogeochemical processes to the oceanic carbon cycle and to calculate the ratio of calcium carbonate to organic carbon downward export, we have incorporated biological and alkalinity pumps in the yoked high-latitude exchange/interior diffusion-advection (YOLDA) model. The biogeochemical processes are represented by four parameters. The values of the parameters are tuned so that the model can reproduce the observed phosphate and alkalinity distributions in each oceanic region. The sensitivity of the model to the biogeochemical parameters shows that biological production rates in the euphotic zone and decomposition depths of particulate matters significantly influence horizontal and vertical distributions of biogeochemical substances. The modeled vertical fluxes of particulate organic phosphorus and calcium carbonate are converted to vertical carbon fluxes by the biological pump and the alkalinity pump, respectively. The downward carbon flux from the surface layer to the deep layer in the entire region is estimated to be 3.36 PgC/yr, which consists of 2.93 PgC/yr from the biological pump and 0.43 PgC/yr from the alkalinity pump, which is consistent with previous studies. The modeled rain ratio is higher with depth and higher in the Pacific and Indian Oceans than in the Atlantic Ocean. The global rain ratio at the surface layer is calculated to be 0.14 to 0.15. This value lies between the lower and higher ends of the previous estimates, which range widely from 0.05 to 0.25. This study indicates that the rain ratio is unlikely to be higher than 0.15, at least in the surface waters.     

Linking phytoplankton community size composition with temperature, plankton food web structure and sea-air CO2 flux

Data collected at open water stations (depth>400 m) in all major ocean basins in 2006-2008 are used to examine the relationship between the size structure of the phytoplankton community (determined by size fractionated chlorophyll filtration), temperature and inorganic nutrient availability. A significant relationship (p<0.0005) was found between community size structure and temperature, with the importance of large cells in the community decreasing with increase in temperature. Although weaker than the temperature relationship, significant relationships were also noted between community cell size and DIN (nitrate, nitrite and ammonium: p=0.034) and phosphate (p=0.031) concentrations. When the data were divided into two groups of equal size, representing the samples with the highest and lowest DIN/phosphate concentrations, respectively, no difference could be identified between the slopes of the lines representing the relationship between size structure and temperature. There was, however, a difference in the intercepts between the two groups. If the relationship between size and temperature was only a response to nutrient availability, we would expect that the response would be the strongest in the groups with the lowest nutrient concentrations. These results suggest that, in addition to a nutrient effect, temperature may also directly influence the size structure of phytoplankton communities. The size structure of the phytoplankton community in this study correlated to the magnitude of primary production, export production (determined after Laws et al., 2000) and integrated water column chlorophyll. Significant relationships were also found between mesozooplankton production (determined using the proxy of calanoid+cyclopoid nauplii abundance as a percentage of the total number of these copepods) and both temperature and phytoplankton size, with production being the lowest in the warmest waters where phytoplankton were the smallest. In the North Atlantic, export production and community size structure appear to be related to ocean uptake of CO₂ from the atmosphere. The reported results suggest that ocean warming may directly alter plankton community structure. This, in turn, may alter the structure of marine food webs and impact the performance of the open ocean as a natural carbon sink.     

The role of inflow magnitude and frequency on plankton communities from the Guadalupe Estuary,Texas, USA: Findings from microcosm experiments

Changes in magnitude and frequency of inflow results in subsequent alterations in the delivery of nutrients essential for phytoplankton growth and competition producing variations in community composition and nutritional value of phytoplankton. Zooplankton demographics are likely directly influenced by pulsed inflows due to flushing losses, whereas they are also indirectly affected by changes in prey quality. In this study, we report the potential effect of pulsed inflows on the plankton community of the Guadalupe Estuary. Microcosms were used that allowed control of light intensity and photoperiod, turbulence, temperature, nutrient loading, and flushing magnitude and periodicity. Our microcosm experiments were novel as they utilized natural plankton communities, thereby allowing the simultaneous interaction between hydrology, resource availability and grazing. Results show differences in microcosms according to magnitude and frequency of flushing. For example, copepod population density was greatest at the annual mean inflow magnitude. At half the annual mean inflow a decline in prey quality likely resulted in poorer grazer performance, and at double the annual mean inflow magnitude increased flushing losses prevented the incidence of higher copepod densities. Similarly, pulsed inflows resulted in greater copepod population densities, higher overall phytoplankton biomass, and dominance of centric diatoms (known to be faster growing and more edible). While reduced freshwater inflow associated with increasing anthropogenic demands often strains the needs of estuarine systems, the effects of reduced river input may be alleviated with several management options including manipulation of the timing, frequency, and magnitude of freshwater inflows. Before implications for management can be discerned from these findings, however, larger scale experiments are needed focus on the roles of inflow magnitude and frequency.     

黄东海浮游生物群落呼吸率对碳平衡的重要性

黄东海接受长江冲淡水和黑潮带来的大量营养盐和有机物质,其碳循环对陆架海碳源汇格局至关重要.浮游生物群落呼吸是影响碳循环的重要过程.为揭示黄东海浮游生物群落呼吸率(PCR)对碳平衡的贡献,于2011年四季使用黑白瓶培养法测定黄海南部及东海北部浮游群落呼吸率和初级生产力,并同步测定温度、盐度、营养盐、叶绿素和细菌丰度等环境...     

A pelagic ecosystem model calibrated with BATS and OWSI data

Ocean ecosystems play an important role in the global carbon cycle. In this paper, we develop and calibrate a pelagic ocean ecosystem model by simultaneously fitting data from two sites in the North Atlantic: the site of the Bermuda Atlantic Time-series Study (BATS, 32°N, 64°W) and Ocean Weather Ship Station “I” (OWSI, 59°N, 20°W). These sites differ dramatically in the magnitude and timing of primary production, chlorophyll concentrations, and light and nutrient availabilities. We find that a relatively simple model can be simultaneously fit to ecosystem data at both sites. As in the model of Hurtt and Armstrong (1996, Deep-Sea Research II 43(2–3): 653–683), the fit largely depends on the inclusion of multiple size classes of phytoplankton and detritus and a variable chlorophyll-to-biomass (nitrogen) ratio for phyto-plankton. The inclusion of these features in the model enables it to adjust to predict important biological differences temporally within sites and spatially between sites. In addition, the model also includes a phytoplankton self-shading parameter to capture differences in water clarity between sites, and the parameterization of iron limitation at OWSI as a hypothesis for the moderately HNLC condition there. Finally, we suggest that the model performs reasonably well at the Hawaii Ocean Time-series (HOT, 23°N, 158°W), a site where no model calibration was done.     

Estuarine classification and response to nitrogen loading: Insights from simple ecological models

Estuaries exhibit a large range in their responses to nitrogen loadings determined in part by characteristics of the driver, such as magnitude and frequency, but also by such intrinsic characteristics as physical/chemical factors (e.g., depth, volume, hypsometry, salinity, turbidity) and biological factors (e.g., nature of ecological communities, trophic interactions). To address the richness of estuarine response to driver variables, the aim ultimately is to establish a simple estuarine classification scheme, beginning with a river-dominated subset of estuarine systems and focusing on the role of water residence time in the estuary. Residence time (or flushing time) is related to other drivers (streamflow, nutrient, and sediment loads) and drives much of the biological response of estuaries because of flushing effects on plankton, temperature, nutrients, and light. Toward this goal, nutrient–phytoplankton–zooplankton (NPZ) models have been used to examine a range of subjects including effects of nutrient limitation and zooplankton predation on phytoplankton dynamics and fish predation. This class of model can admit a wide range of behavior, including multiple steady-states and oscillatory behavior. The NPZ equations include terms for nutrient recycling, phytoplankton settling, benthic regeneration, and zooplankton mortality. Analysis of the equations suggests that both the nature of nitrogen loading (i.e., whether it is correlated with discharge or independent of it) and residence time are critical in determining the steady-state response of the system.     

Riverine influence determines nearshore heterogeneity of nutrient (C,N, P) content and stoichiometry in the KwaZulu-Natal Bight,South Africa

Riverine influences on nearshore oceanic habitats often have detrimental consequences leading to algal blooms and hypoxia. In oligo- to mesotrophic systems, however, nutrient delivery via rivers may stimulate production and even be a vital source of nutrients, as may nutrient supplements from upwelling. We investigated the nutrient content (C, N, P) and stoichiometry of sediment, and several pelagic, benthopelagic and benthic species in the KwaZulu-Natal (KZN) Bight, a narrow shelf area on the south-east coast of South Africa, bordering the Agulhas Current. Three suggested nutrient sources to the bight are the Thukela River in the central region of the bight, upwelling in the northern part and a semi-permanent eddy (Durban Eddy) in the southern part. Elemental content of the various groups studied showed significantly higher values for most groups at the site near the Thukela River. C:P and N:P were highest in the southern part of the bight, and lowest near the Thukela Mouth or at Richards Bay in the north, indicating the latter were the P-richer sites. Sediment organic matter showed lowest elemental content, as expected, and zooplankton stoichiometry was highest compared to all other biotic groups. Environmental heterogeneity played a greater role in organismal C, N and P content and stoichiometry compared to phylogeny, with the exception of the differences in C:P and N:P of zooplankton. From this bight-wide study, the higher elemental content and lower ratios at the Thukela Mouth site supported previous findings of the importance of coastal nutrient sources to the bight ecosystem. Reductions in river flow for water use in the catchment areas may therefore have negative consequences for the productivity of the entire ecosystem.     

C : N ratios in the mixed layer during the productive season in the northeast Atlantic Ocean

Redfield stoichiometry has proved a robust paradigm for the understanding of biological production and export in the ocean on a long-term and a large-scale basis. However, deviations of carbon and nitrogen uptake ratios from the Redfield ratio have been reported. A comprehensive data set including all carbon and nitrogen pools relevant to biological production in the surface ocean (DIC, DIN, DOC, DON, POC, PON) was used to calculate seasonal new production based on carbon and nitrogen uptake in summer along 20°W in the northeast Atlantic Ocean. The 20°W transect between 30 and 60°N covers different trophic states and seasonal stages of the productive surface layer, including early bloom, bloom, post-bloom and non-bloom situations. The spatial pattern has elements of a seasonal progression. We also calculated exported production, i.e., that part of seasonal new production not accumulated in particulate and dissolved pools, again separately for carbon and nitrogen. The pairs of estimates of `seasonal new production’ and `exported production’ allowed us to calculate the C : N ratios of these quantities. While suspended particulate matter in the mixed layer largely conforms to Redfield stoichiometry, marked deviations were observed in carbon and nitrogen uptake and export with progressing season or nutrient depletion. The spring system was characterized by nitrogen overconsumption and the oligotrophic summer system by a marked carbon overconsumption. The C : N ratios of seasonal new as well as exported production increase from early bloom values of 5–6 to values of 10–16 in the post-bloom/oligotrophic system. The summertime accumulation of nitrogen-poor dissolved organic matter can explain only part of this shift.     

Impact of the river nutrient load variability on the North Aegean ecosystem functioning over the last decades

The impact of river load variability on the North Aegean ecosystem functioning over the last decades (1980–2000) was investigated by means of a coupled hydrodynamic/biogeochemical model simulation. Model results were validated against available SeaWiFS Chl-a and in situ data. The simulated food web was found dominated by small cells, in agreement with observations, with most of the carbon channelled through the microbial loop. Diatoms and dinoflagellates presented a higher relative abundance in the more productive coastal areas. The increased phosphate river loads in the early 80s resulted in nitrogen and silicate deficiency in coastal, river-influenced regions. Primary production presented a decreasing trend for most areas. During periods of increased phosphate/nitrate inputs, silicate deficiency resulted in a relative decrease of diatoms, triggering an increase of dinoflagellates. Such an increase was simulated in the late 90s in the Thermaikos Gulf, in agreement with the observed increased occurrence of Harmful Algal Blooms. Microzooplankton was found to closely follow the relative increase of dinoflagellates under higher nutrient availability, showing a faster response than mesozooplankton. Sensitivity simulations with varying nutrient river inputs revealed a linear response of net primary production and plankton biomass. A stronger effect of river inputs was simulated in the enclosed Thermaikos Gulf, in terms of productivity and plankton composition, showing a significant increase of dinoflagellates relative abundance under increased nutrient loads.     

The vertical distribution and diurnal migration of copepods at different stations in the Irish Sea

The vertical distribution of plankton and hydrographic factors was studied at five stations within the Irish Sea, differing in depth, strength of tidal currents and location with respect to non-tidal currents. Neither the plankton nor the hydrographic factors showed vertical homogeneity at any station, even when there were strong tidal currents. The day depth of bothPseudocalanus elongatus andMicrocalanus pusillus varied considerably at different stations and at different seasons at the same station.P. elongatus showed diurnal migrations on some but not all occasions. No clear diurnal migration was shown byM. pusillus. Some of the vertical distributions of both species, both by day and over 24 h, could be correlated with physical or chemical factors. These included stratification involving differences in temperature or salinity, but in some cases only differences in nutrient concentration were detected. Although layers of seawater of similar density are unlikely to maintain their identity for very long periods they are probably more common than is generally realized and sufficiently persistent to have important effects on the distribution of plankton. The determination of nutrient concentrations provides a convenient and effective means of detecting such layers.     

A daily study of the diatom spring bloom at Roscoff (France) in 1985. I. The spring bloom within the annual cycle

The repeated occurrence of a monospecific bloom of the plankton diatom Rhizosolenia delicatula at Roscoff (western English Channel) was made the subject of an interdisciplinary research programme. Samples were taken at daily intervals from April to July and at longer intervals during the remaining part of the year 1985. Routine physical parameters, light transmission, nutrients, dissolved oxygen, particle load, particulate N and P, chlorophyll content, phytoplankton counting, and zooplankton biomass were measured as a basis for more specific studies (to follow as subsequent papers in this series).The area is characterized by: high tidal range, permanent mixing throughout the year, low attenuation coefficients, moderate nutrient supply, and the dominance of benthic algae over phytoplankton. The spring bloom is significantly delayed with respect to the usual model for the temperate seas. Tidal cycles are expected to exert the main influence on bloom dynamics at the time scale of phytoplankton growth.     

Nutrient, sulfur and carbon dynamics in a hypersaline lagoon

We measured benthic and water column fluxes in a hypersaline coastal system (Baffin Bay, Texas) in 1996–1997, a period of decreasing salinity (increased freshwater input) and turbidity. Salinity decreased from a mean of 60 to 32 practical salinity units (psu) and turbidity decreased from a mean of 78 to 25 NTU over the study period. Associated with hydrological changes, there were important changes in nutrient fluxes and metabolism. There was a shift of total respiration from the water column to the sediments and an increased amount of the benthic metabolism (2–67%) was attributed to sulfate reduction in this system when salinity was lowest, perhaps a consequence of increased benthic light levels and photosynthetic production of labile carbon in the sediments. The sediments were a large sink for both N and P. Sediment particulate C:N (9.8) and C:P (119) ratios were lower than those in the water column. However, ammonium:phosphate fluxes increased coincident with increased sulfate reduction rates and porewater sulfide concentrations. Efficient N-retention mediated through dissimilative nitrate reduction to ammonium, and high rates of N-fixation in shallow, hypersaline systems may facilitate transitions from N-limitation to P-limitation. During the most hypersaline period, seston exhibited some of the most extreme nutrient ratios ever reported for a marine ecosystem (C:N 10–37 and C:P 200–1200) and suggest that plankton are likely to be P-limited or are very well adapted to low P availability. When salinity and N:P and C:P ratios were highest, the plankton was dominated by a brown tide alga (Aureoumbra lagunensis), supporting evidence that this organism is adapted to low P, long residence time systems.     

Environmental control of short-term variation in the plankton community of inner Tokyo Bay, Japan

Water temperature, salinity, nutrient concentrations and the composition of the plankton community were recorded at three stations in inner Tokyo Bay over a period of 328 days (from June 8, 1995 to April 30, 1996) with a nominal sampling frequency of once per day. Inspection of the results revealed that the data could be divided into two blocs as an aid to analysis: the period from June to October was characterized by the development of stratification of temperature and salinity (stratification period), and November to March was characterized by uniform temperature and salinity in the water column due to vertical mixing (mixing period). Oxygen-depleted water forms in the bottom layer during the stratification period, but vertical mixing of the water column, due to changing wind and rainfall conditions caused by passing weather fronts, results in the breakdown of the oxygen-depleted water mass. Nutrient loads are high in the surface water due to the freshwater supply, but occasional pulses of primary production cause a depletion of phosphate in the surface water, suggesting that the phosphorus becomes a limiting nutrient for phytoplankton growth in this period. Several short-term peaks of plankton abundance (blooms) occurred as responses to temporal changes in water quality from June to November, with consequent species succession. Significant fluctuations in the densities of the diatom Skeletonema costatum and several species of ciliates corresponded to the daily changes in the physical and chemical characteristics of the coastal environment. During the mixing period, when water temperature and solar radiation decreased, there were no short-term variations in water quality and although nutrient concentrations gradually increased from November to February, primary production remained low. This study shows that the short-term dynamics of the phytoplankton community are closely coupled to fluctuations in environmental forcing, and that the degree of coupling is stronger during periods when solar radiation is greater. The results provide a novel typological understanding of seasonal plankton dynamics in a shallow, eutrophicated marine embayment, and suggest how such systems may be treated in simulation modeling.