Does biodiversity of estuarine phytoplankton depend on hydrology?

Phytoplankton growth in estuaries is controlled by factors such as flushing, salinity tolerance, light, nutrients and grazing. Here, we show that biodiversity of estuarine phytoplankton is related to flushing, and illustrate this for some European estuaries.The implications for the definition of reference conditions for quality elements in estuaries of different types are examined, leading to the conclusion that constraints on the number of estuarine and coastal types that may be defined for management purposes require that quality classes take into account natural variability within types, in order to be ecologically meaningful. We develop a screening model to predict the growth rate required for a phytoplankton species to be present under different flushing conditions and apply it to estuaries in the EU and US to show how changes in physical forcing may alter biodiversity. Additional results are presented on the consequences for eutrophication, showing that changes in residence time may interact with species-specific nutrient uptake rates to cause shifts in species composition, potentially leading to effects such as harmful algal blooms.We discuss applications for integrated coastal zone management, and propose an approach to normalization of estuarine phytoplankton composition as regards species numbers.     

A hydrodynamic and water quality modeling study of spatial and temporal patterns of phytoplankton growth in a stratified lake with buoyant incoming flow

A three-dimensional hydrodynamic and water quality model was applied to Lake Paldang, a lake in South Korea that is stratified by incoming flows. The spatial and temporal patterns of phytoplankton growth in this lake were determined from the model. The model was calibrated and verified using data measured under different hydrological conditions. The model results were in reasonable agreement with the field measurements, in both the calibration and verification phases. The distributions of water quality and residence time in the lake and phytoplankton response to changes in nutrient loads were examined with the model, and the influence of the hydrodynamics on phytoplankton response was analyzed. The simulation results indicated that Lake Paldang is an essentially phosphorus-limited system, but that phytoplankton growth is limited by low water temperature and short residence time during the winter and the summer monsoon period, respectively. The results of sensitivity analyses also suggested that the hydrodynamics within the lake may have an indirect influence on phytoplankton responses to changes in the limiting nutrient loads, and that reducing phosphorus loading from Kyoungan Stream should be a high priority policy for controlling algal blooms during the pre- and post-monsoon periods. From this study, it was concluded that the three-dimensional water quality model incorporating hydrodynamic processes could successfully simulate phytoplankton response to changes in nutrient loads and that it could become a useful tool for identifying the essential factors determining phytoplankton growth and for developing the best management policy for algal blooms in Lake Paldang.     

A spatial model of phytoplankton patchiness

The one-dimensional theory of critical-length scales of phytoplankton patchiness is developed to include phytoplankton growth and herbivore grazing as functions of time and space. The critical-length scale L _c for the pathch is then determined by the initial spatial distribution and concentration of the limiting nutrient and herbivores in addition to the daily averaged values of the growth and loss processes. The response of an initial phytoplankton patch to the stresses of turbulent diffusion, nutrient depletion, light periodicity, and nocturnal or continuous herbivore grazing is investigated numerically for several oceanic conditions. Nocturnal grazing, while less stressful on primary production than continous grazing, results in lower phytoplankton standing stocks. Increase in biomass of vertically migrating zooplankton results in a net loss of nutrient which might otherwise be egested, recycled, and utilized in the euphotic zone under continuous grazing conditions. The Ivlev constant is shown via sensitivity analysis to be a significant parameter ultimately influencing phytoplankton production. It is demonstrated numerically that diffusion of phytoplankton cells from areas of high concentration to low concentration prevents the local extinction of the standing stock, thereby rendering a positive herbivore grazing-threshold unnecessary for ecosystem stability.     

Spring phenological responses of marine and freshwater plankton to changing temperature and light conditions

Shifts in the timing and magnitude of the spring plankton bloom in response to climate change have been observed across a wide range of aquatic systems. We used meta-analysis to investigate phenological responses of marine and freshwater plankton communities in mesocosms subjected to experimental manipulations of temperature and light intensity. Systems differed with respect to the dominant mesozooplankton (copepods in seawater and daphnids in freshwater). Higher water temperatures advanced the bloom timing of most functional plankton groups in both marine and freshwater systems. In contrast to timing, responses of bloom magnitudes were more variable among taxa and systems and were influenced by light intensity and trophic interactions. Increased light levels increased the magnitude of the spring peaks of most phytoplankton taxa and of total phytoplankton biomass. Intensified size-selective grazing of copepods in warming scenarios affected phytoplankton size structure and lowered intermediate (20–200?μm)-sized phytoplankton in marine systems. In contrast, plankton peak magnitudes in freshwater systems were unaffected by temperature, but decreased at lower light intensities, suggesting that filter feeding daphnids are sensitive to changes in algal carrying capacity as mediated by light supply. Our analysis confirms the general shift toward earlier blooms at increased temperature in both marine and freshwater systems and supports predictions that effects of climate change on plankton production will vary among sites, depending on resource limitation and species composition.     

Climate change effects on phytoplankton depend on cell size and food web structure

We investigated the effects of warming on a natural phytoplankton community from the Baltic Sea, based on six mesocosm experiments conducted 2005–2009. We focused on differences in the dynamics of three phytoplankton size groups which are grazed to a variable extent by different zooplankton groups. While small-sized algae were mostly grazer-controlled, light and nutrient availability largely determined the growth of medium- and large-sized algae. Thus, the latter groups dominated at increased light levels. Warming increased mesozooplankton grazing on medium-sized algae, reducing their biomass. The biomass of small-sized algae was not affected by temperature, probably due to an interplay between indirect effects spreading through the food web. Thus, under the higher temperature and lower light levels anticipated for the next decades in the southern Baltic Sea, a higher share of smaller phytoplankton is expected. We conclude that considering the size structure of the phytoplankton community strongly improves the reliability of projections of climate change effects.     

Phytoplankton carbon dynamics during a winter-spring diatom bloom in an enclosed marine ecosystem: primary production,biomass and loss rates

Phytoplankton production, standing crop, and loss processes (respiration, sedimentation, grazing by zooplankton, and excretion) were measured on a daily basis during the growth, dormancy and decline of a winter-spring diatom bloom in a large-scale (13 m³) marine mesocosm in 1987. Carbonspecific rates of production and biomass change were highly correlated whereas production and loss rates were unrelated over the experimental period when the significant changes in algal biomass characteristic of phytoplankton blooms were occurring. The observed decline in diatom growth rates was caused by nutrient limitation. Daily phytoplankton production rates calculated from the phytoplankton continuity equation were in excellent agreement with rates independently determined using standard ¹⁴C techniques. A carbon budget for the winter bloom indicated that 82.4% of the net daytime primary production was accounted for by measured loss processes, 1.3% was present as standing crop at the end of the experiment, and 16.3% was unexplained. Losses via sedimentation (44.8%) and nighttime phytoplankton respiration (24.1%) predominated, while losses due to zooplankton grazing (10.7%) and nighttime phytoplankton excretion (2.8%) were of lesser importance. A model simulating daily phytoplankton biomass was developed to demonstrate the relative importance of the individual loss processes.     

Phytoplankton blooms are strongly impacted by microzooplankton grazing in coastal North Pacific waters

Phytoplankton growth and microzooplankton grazing were measured in two productive coastal regions of the North Pacific: northern Puget Sound and the coastal Gulf of Alaska. Rates of phytoplankton growth (range: 0.09–2.69 day⁻¹) and microzooplankton grazing (range: 0.00–2.10 day⁻¹) varied seasonally, with lowest values in late fall and winter, and highest values in spring and summer. Chlorophyll concentrations also varied widely (0.19–13.65 μg l⁻¹). Large (>8 μm) phytoplankton cells consistently dominated phytoplankton communities under bloom conditions, contributing on average 65% of total chlorophyll biomass when chlorophyll exceeded 2 μg l⁻¹. Microzooplankton grazing was an important loss process affecting phytoplankton, with grazing rates equivalent to nearly two-thirds (64%) of growth rates on average. Both small and large phytoplankton cells were consumed, with the ratio of grazing to growth (g:μ) for the two size classes averaging 0.80 and 0.42, respectively. Perhaps surprisingly, the coupling between microzooplankton grazing and phytoplankton growth was tighter during phytoplankton blooms than during low biomass periods, with g:μ averaging 0.78 during blooms and 0.49 at other times. This tight coupling may be a result of the high potential growth and ingestion rates of protist grazers, some of which feed on bloom-forming diatoms and other large phytoplankton. Large ciliates and Gyrodinium-like dinoflagellates contributed substantially to microzooplankton biomass at diatom bloom stations in the Gulf of Alaska, and microzooplankton biomass overall was strongly correlated with >8 μm chlorophyll concentrations. Because grazing tended to be proportionally greater when phytoplankton biomass was high, the absolute amount of chlorophyll consumed by microzooplankton was often substantial. In nearly two-thirds of the experiments (14/23), more chlorophyll was ingested by microzooplankton than was available for all other biological and physical loss processes combined. Microzooplankton were important intermediaries in the transfer of primary production to higher trophic levels in these coastal marine food webs. Received: 12 November 1999 / Accepted: 4 October 2000     

The dynamics of phytoplankton blooms in puget sound a fjord in the Northwestern United States

This paper describes a quantitative investigation of relationships between the growth of phytoplankton, and climatic and hydrodynamci conditions in temperate fjords with marked tides, as exemplified by Puget Sound, Washington (USA). Algal growth in the open waters of the central basin of the Sound is dominated by a number of intense blooms beginning in late April or May and recurring throughout the summer. Rarely, and only briefly, does nitrate become exhausted. The phytoplankton production rate in the central basin of Puget Sound is about 465 g C m^-2 year^-1. During the springs of 1966 and 1967, oceanographic measurements were carried out at a mid-channel station with sufficient frequency to allow investigation of physical and biological processes with time scales of the order of a day. The principal investigative tool is a numerical model in which the hydrodynamical conditions are represented by an approximate analysis of the gravitational convection mode of circulation. Algal concentration is represented as a continous function of space and time in the model which ascribes changes in phytoplankton density to variations in photosynthetic and respiratory activity, algal sinking, grazing by herbivores, and to mixing and advection. Computations adequately reproduce the principal features of phytoplankton concentrations observed during 75 days and 35 days in the springs of 1966 and 1967, respectively. Numerical experiments assess the relative importance of various processes which govern the level of primary production in Puget Sound. It is concluded that phytoplankton growth is limited by a combination of factors, including vertical advection and turbulence, modulation of underwater light intensity by self-shading and inorganic particulates, sinking of algal cells, and occasional rapid horizontal advection of the population from the area by sustained winds. The high primary productivity of the Sound is due to intensive upward transport of nitrate by the estuarine mechanism. These results should be generally applicable to other temperate fjords because of the largely conventional choice of the biological functions.     

Steady states and sensitivities of commonly used pelagic ecosystem model components

Pelagic, coupled ocean circulation-ecosystem models, are widely used in climate research. These tools aim to quantify fluxes of nutrients and carbon in the ocean and are, increasingly, the base of future projections. For this purpose it is crucial to quantify and identify the sources of uncertainties. In contrast to physical models, the underlying equations for ecosystem models are derived from empirical relationships rather than based on first principles. This resulted in the development of a multitude of different ecosystem models - different in respect to both, underlying principles and complexity. Clearly, the question arises, to what extent the sensitivities of these models are comparable.This study focuses on the intrinsic dynamics of some widely used, simple (containing 2-3 prognostic variables) ecosystem models in a 0-D framework (i.e., comprising only the well-mixed oceanic surface layer). A suite of differing model approaches is tuned such that their behavior is similar. The setup resembles the well-mixed oceanic surface layer in the Baltic proper. It is illustrated that strong differences between the model approaches appear due to exemplary, anticipated changes in the external nutrient and light conditions. Herewith, we demonstrate the well-known, but rarely demonstrated fact that, apparent consistency between modeled prognostic variables with today's data bases is not necessarily a good measure of forecast skill. The causes which lead to the different sensitivities are illustrated by considering the steady state solutions. It is pointed out, that apparently small changes in the model formulations can result in very different dynamical behavior and an enormous spread between the model approaches, despite the feasibility to tune a common behavior in a limited range of light and nutrient supply. In our examples, the sensitivity is mainly a function of the formulation of the loss rate of phytoplankton. It is thus, in particular, the formulation of highly unknown heteorotrophic processes that determines the model sensitivity.     

Role of competition in phytoplankton population for the occurrence and control of plankton bloom in the presence of environmental fluctuations

Termination of harmful algal blooms (HABs) and coexistence of phytoplankton–zooplankton populations are of great importance to human health, ecosystem, environment, tourism and fisheries. In this paper, we propose a three component model consisting of non-toxic phytoplankton (NTP), toxin producing phytoplankton (TPP) and zooplankton (Z). The growth of zooplankton species is assume to reduce due to toxic chemicals released by TPP population. We have extended the model proposed by Chattopadhyay et al. [Chattopadhyay, J., Sarkar, R.R., Pal, S., 2004. Mathematical modelling of harmful algal blooms supported by experimental findings. Ecol. Comp. 1, 225–235] by including competition terms between TPP and NTP. We observe the effect of competition factors both in the presence and absence of the environmental fluctuation. From our field as well as model analysis we observe that competition helps in the coexistence of the species, but if the effect of competition is very high on the TPP population, it results in planktonic bloom. It is shown that the coexistence equilibrium loses its stability when the competition coefficient crosses a critical value and resulting Hopf-bifurcation around the positive equilibrium depicting oscillations phenomena of the populations.     

A simple plankton model for the Oregon upwelling ecosystem: Sensitivity and validation against time-series ocean data

Simple plankton models serve as useful platforms for testing our understanding of the mechanisms underlying ecosystem dynamics. A simple, one-dimensional plankton model was developed to describe the dynamics of nitrate, ammonium, two phytoplankton size-classes, meso-zooplankton, and detritus in the Oregon upwelling ecosystem. Computational simplicity was maintained by linking the biological model to a one-dimensional, cross-shelf physical model driven by the daily coastal upwelling index. The model sacrificed resolution of regional-scale and along-shore (north to south) processes and assumed that seasonal productivity is primarily driven by local cross-shelf Ekman transport of surface waters and upwelling of nutrient-rich water from depth.Our goals were to see how well a simple plankton model could capture the general temporal and spatial dynamics of the system, test system sensitivity to alternate parameter set values, and observe system response to the effective scale of potential retention mechanisms. Model performance across the central Oregon shelf was evaluated against two years (2000-2001) of chlorophyll and copepod time-series observations. While the modeled meso-zooplankton biomass was close in scale to the observed copepod biomass, phytoplankton was overestimated relative to that inferred from the observed surface chlorophyll concentration. Inshore, the system was most sensitive to the nutrient uptake kinetics of diatom-size phytoplankton and to the functional grazing response of meso-zooplankton. Meso-zooplankton was more sensitive to alternate parameter values than was phytoplankton. Reduction of meso-zooplankton cross-shelf advection rates (crudely representing behavioral retention mechanisms) reduced the scale of model error relative to the observed seasonal mean inshore copepod biomass but had little effect of the modeled meso-zooplankton biomass offshore nor upon phytoplankton biomass across the entire shelf.     

Behavior of a simple plankton model with food-level acclimation by herbivores

The acclimation of herbivores to variation in their phytoplankton food source was expressed mathematically and its effect on phytoplankton, herbivore and nutrient cycles explored with a plankton model. The grazing formulation is a modified version of the function experimentally determined by Mayzaud and Poulet. Their function differs from the traditional Ivlev expression for herbivore grazing in that there is no asymptotic limit to the grazing rate. The steady-state solutions of the phytoplankton-herbivorenutrient model were similar with the two grazing formulations, but the time-dependent behaviour of the two models differed markedly. The model with Ivlev grazing showed oscillations when the grazing pressure was high. The model with acclimated herbivore grazing showed only small, highly damped oscillations as it approached steady state. The latter is more similar to the evolution of plankton trophic levels observed in controlled ecosystem experiments.     

The development of a multi-species algal ecodynamic model for urban surface water systems and its application

An ecodynamic model that can simulate four phytoplankton species has been developed to deal with the unique characteristics of urban river systems which has manmade river profile, flow controlled by gates, severe eutrophication status, and fragile aquatic ecosystem. The ecodynamic model was developed referencing two typical models: the water quality simulation model WASP and ecological model CAEDYM. The model can simulate 11 state variables: dissolved oxygen, carbonaceous biochemical oxygen demand, ammonia nitrogen, nitrate nitrogen, organic nitrogen, inorganic phosphorus, organic phosphorus and four phytoplankton species with zooplankton as a boundary condition. The ecodynamic model was applied to Sihai section of the Beijing urban river system, where serious algal blooms broke out in recent years. The dominant phytoplankton species are Cyanophyta, Chlorophyta, Bacillariophyta, and Cryptophyta. Site-specific data on geometry, meteorology, pollution sources, and existing ecosystem parameters were collected and used for model calibration and verification The model results mimic observed trends of water quality and phytoplankton species succession and can be used for forecasting algal blooms as well as assessment of river management measures.     

The impact of detritivorous fishes on a mangrove estuarine system

Sundarban mangrove ecosystem in India is one of the largest detritus based ecosystem of the world and it supplies the detritus and nutrients to the adjacent Hooghly-Matla estuarine complex. In this estuary a group of fish completely detritivorous in nature, belonging to the genus Mugil spp. is present. This group of fish is expected to have important effects on the trophic dynamics of ecosystems, but exact nature of these effects is not known. In order to study the impact that detritivory by fish may have on the estuarine food chain, we developed mathematical formulations. We run two models, one with phytoplankton, zooplankton, carnivorous fish, detritus and nutrient and without this group of fish and a second one after including this fish in the system. In our model this group of fish has no major impact on primary productions of the estuarine system but has extensive role in total fish production. Coexistence of detritivorous fish and carnivorous fish occurs within reasonable parameter range. We have tested different growth rates of phytoplankton, grazing rates and predation rates of zooplankton, detritivorous fish and carnivorous fish for total system equilibrium. Carnivorous fish predation rate on detritivorous fish and detritivorous fish grazing rate on detritus are very important. Different foraging ratios are also tested in this study. Foraging preference of carnivorous fish on detritivorous fish appears significant for the system equilibrium.     

The phenology of phytoplankton blooms: Ecosystem indicators from remote sensing

Remote sensing offers many advantages in the development of ecosystem indicators for the pelagic zone of the ocean. Particularly suitable in this context are the indicators arising from time series that can be constructed from remotely sensed data. For example, using ocean-colour radiometry, the phenology of phytoplankton blooms can be assessed. Metrics defined in this way show promise as informative indicators for the entire pelagic ecosystem. A simple phytoplankton–substrate model, with forcing dependent on latitude and day number is used to explore the qualitative features of bloom phenology for comparison with the results observed in a suite of 10-year time series of chlorophyll concentration, as assessed by remote sensing, from the Northwest Atlantic Ocean. The model reveals features of the dynamics that might otherwise have been overlooked in evaluation of the observational data.     

The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

Water management strategies against toxic Microcystis blooms in the Dutch delta.

To prevent flooding of the Dutch delta, former estuaries have been impounded by the building of dams and sluices. Some of these water bodies, however, experience major ecological problems. One of the problem areas is the former Volkerak estuary that was turned into a freshwater lake in 1987. From the early 1990s onward, toxic Microcystis blooms dominate the phytoplankton of the lake every summer. Two management strategies have been suggested to suppress these harmful algal blooms: flushing the lake with fresh water or reintroducing saline water into the lake. This study aims at an advance assessment of these strategies through the development of a mechanistic model of the population dynamics of Microcystis. To calibrate the model, we monitored the benthic and pelagic Microcystis populations in the lake during two years. Field samples of Microcystis were incubated in the laboratory to estimate growth and mortality rates as functions of light, temperature, and salinity. Recruitment and sedimentation rates were measured in the lake, using traps, to quantify benthic-pelagic coupling of the Microcystis populations. The model predicts that flushing with fresh water will suppress Microcystis blooms when the current flushing rate is sufficiently increased. Furthermore, the inlet of saline water will suppress Microcystis blooms for salinities exceeding 14 g/L. Both management options are technically feasible. Our study illustrates that quantitative ecological knowledge can be a helpful tool guiding large-scale water management.     

Development of Lake Ladoga ecosystem models: modeling of the phytoplankton succession in the eutrophication process. I

New models of Lake Ladoga ecosystem and the results of modeling are presented. In the first part the model of phytoplankton succession in the process of anthropogenic eutrophication of the lake is considered under the evolution of the phosphorus loading. The still continued anthropogenic eutrophication of the lake started in 1962 when the phosphorus load began to increase. Since 1962 during the evolution of the lake’s state from oligotrophic to developed mezotrophic one, the structure of phytoplankton community dominating species was significantly changed as well as its total productivity. The system state in the model is described by 14 parameters: nine phytoplankton complexes, zooplankton, dissolved organic matter, detritus, dissolved mineral phosphorus and dissolved oxygen. The number of parameters of this model is noticeably larger than that of previous models created by the authors. The relative dynamics of phytoplankton complexes in the lake’s ecosystem evolution was simulated by the new model. It is shown that the modeling results are adequately corresponding to the observation data. The results of phytoplankton structure modeling allow to estimate the impact of phytoplankton on the water quality as well as give the prediction of the lake’s ecosystem evolution with the changes of the phosphorus loading.