Time delay as a key factor of model plankton dynamics

Studies of the mechanisms underlying complex dynamics of ecological systems at various spatial and time scales bring increasing awareness that complexity is an intrinsic feature of ecological functioning. This paper is to investigate the role of such an ecologically significant parameter as the time delay due to maturation processes in the complex plankton dynamics. We show that the time lag T1, associated with the zooplankton maturation period can lead to essential changes in the plankton dynamics. Particularly, we show that the coexistence of limit cycle and chaotic attractor we have recently found to be typical of the system at T1 = 0 [A.B. Medvinsky, I.A. Tikhonova, R.R. Aliev, B.-L. Li, Z.-S. Lin, H. Malchow, Patchy environment as a factor of complex plankton dynamics, Phys. Rev. E 64 (2001) 021915] is replaced by pure chaotic plankton dynamics as T1 becomes more than a critical value. The results obtained imply that chaos is a rather common phenomenon in the plankton functioning.     

Seasonal plankton dynamics along a cross-shelf gradient

Much interest has recently been devoted to reconstructing the dynamic structure of ecological systems on the basis of time-series data. Using 10 years of monthly data on phyto- and zooplankton abundance from the Bay of Biscay (coastal to shelf-break sites), we demonstrate that the interaction between these two plankton components is approximately linear, whereas the effects of environmental factors (nutrients, temperature, upwelling and photoperiod) on these two plankton population growth rates are nonlinear. With the inclusion of the environmental factors, the main observed seasonal and inter-annual dynamic patterns within the studied plankton assemblage also indicate the prevalence of bottom-up regulatory control.     

Modelling of autumn plankton bloom dynamics

A simple system of parametrically forced ordinary differentialequations is used to model autumn phytoplankton blooms in temperateoceans by a mechanism involving deepening of the upper mixedlayer. Blooms are triggered provided the increase in nutrientsin the mixed layer is rapid within the first few days of deepeningand provided light-limited phytoplankton growth rate is relativelyhigh. Blooms exist as transient trajectories between quasi-equilibriumstates, rather than as bifurcations of steady states; thereforevery gradual deepening cannot trigger blooms. Very rapid deepeningalso prevents blooms due to the deleterious effect on phytoplanktongrowth rate. The mechanisms identified by this simple modelare vindicated by considering alternative grazing and deepeningregimes and by comparison with a more ecologically complex model(Fasham, 1993, in The Global Carbon Cycle, Springer-Verlag).Modelled estimates of primary productivity from both the simplemodel and the complex model parameterized for Ocean WeatherStation ‘India’ are around 0.5 g C m^–2 day^–1during the autumn bloom, therefore comprising a significantcomponent of annual production in temperate areas.     

The effects of abrupt topography on plankton dynamics

Plankton population dynamics in the upper layer of the ocean depends on upwelling processes that bring nutrients from deeper waters. In turn, these depend on the structure of the vertical velocity field. In coastal areas and in oceanic regions characterized by the presence of strong submarine topographic features, the variable bottom topography induces significant effects on vertical velocities and upwelling/downwelling patterns. As a consequence, large plankton and fish abundances are frequently observed above seamounts, canyons and steep continental shelves. In this work, the dynamics of an NPZ (nutrient-phytoplankton-zooplankton) system is numerically studied by coupling the ecosystem model with a quasi two-dimensional (2D) fluid model with topography. At variance with classical 2D approaches, this formulation allows for an explicit expression of the vertical motions produced when fluid columns are squeezed and stretched as they experience changes of depth. Thus, input or output of nutrients at the surface are associated with fluid motion over the bottom topography. We examine the dynamics of a cyclonic vortex over two basic topographies: a steep escarpment and a submarine mountain. We show that plankton abundance over the escarpment is modulated by the passing of topographic Rossby waves, generated by the vortex-topography interaction. In such configuration, advection effects driven by the flow over the escarpment are of limited relevance for the dynamics of biological fields. By contrast, we find that the flow resulting from the interaction of a vortex with a seamount is sufficiently strong and persistent to allow for a remarkable increase of nutrients, and a corresponding enhancement of phytoplankton and zooplankton concentrations. Over the seamount, advection effects associated with trapped flow perturbations around the summit play an essential role.     

The effects of abrupt topography on plankton dynamics

Plankton population dynamics in the upper layer of the ocean depends on upwelling processes that bring nutrients from deeper waters. In turn, these depend on the structure of the vertical velocity field. In coastal areas and in oceanic regions characterized by the presence of strong submarine topographic features, the variable bottom topography induces significant effects on vertical velocities and upwelling/downwelling patterns. As a consequence, large plankton and fish abundances are frequently observed above seamounts, canyons and steep continental shelves. In this work, the dynamics of an NPZ (nutrient-phytoplankton-zooplankton) system is numerically studied by coupling the ecosystem model with a quasi two-dimensional (2D) fluid model with topography. At variance with classical 2D approaches, this formulation allows for an explicit expression of the vertical motions produced when fluid columns are squeezed and stretched as they experience changes of depth. Thus, input or output of nutrients at the surface are associated with fluid motion over the bottom topography. We examine the dynamics of a cyclonic vortex over two basic topographies: a steep escarpment and a submarine mountain. We show that plankton abundance over the escarpment is modulated by the passing of topographic Rossby waves, generated by the vortex-topography interaction. In such configuration, advection effects driven by the flow over the escarpment are of limited relevance for the dynamics of biological fields. By contrast, we find that the flow resulting from the interaction of a vortex with a seamount is sufficiently strong and persistent to allow for a remarkable increase of nutrients, and a corresponding enhancement of phytoplankton and zooplankton concentrations. Over the seamount, advection effects associated with trapped flow perturbations around the summit play an essential role.     

春季和夏季巢湖浮游生物群落组成及其动态分析

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Seasonal nutrient and plankton dynamics in a physical-biological model of Crater Lake

A coupled 1D physical-biological model of Crater Lake is presented. The model simulates the seasonal evolution of two functional phytoplankton groups, total chlorophyll, and zooplankton in good quantitative agreement with observations from a 10-year monitoring study. During the stratified period in summer and early fall the model displays a marked vertical structure: the phytoplankton biomass of the functional group 1, which represents diatoms and dinoflagellates, has its highest concentration in the upper 40 m; the phytoplankton biomass of group 2, which represents chlorophyta, chrysophyta, cryptomonads and cyanobacteria, has its highest concentrations between 50 and 80 m, and phytoplankton chlorophyll has its maximum at 120 m depth. A similar vertical structure is a reoccurring feature in the available data. In the model the key process allowing a vertical separation between biomass and chlorophyll is photoacclimation. Vertical light attenuation (i.e., water clarity) and the physiological ability of phytoplankton to increase their cellular chlorophyll-to-biomass ratio are ultimately determining the location of the chlorophyll maximum. The location of the particle maxima on the other hand is determined by the balance between growth and losses and occurs where growth and losses equal. The vertical particle flux simulated by our model agrees well with flux measurements from a sediment trap. This motivated us to revisit a previously published study by Dymond et al. (1996). Dymond et al. used a box model to estimate the vertical particle flux and found a discrepancy by a factor 2.5–10 between their model-derived flux and measured fluxes from a sediment trap. Their box model neglected the exchange flux of dissolved and suspended organic matter, which, as our model and available data suggests is significant for the vertical exchange of nitrogen. Adjustment of Dymond et al.’s assumptions to account for dissolved and suspended nitrogen yields a flux estimate that is consistent with sediment trap measurements and our model.     

Pectin: a critical component in cell-wall-mediated immunity

Contrary to the classical cell-wall model, pectin metabolism may play a crucial role in cell-wall integrity, detection of plant pathogens, and defense response. Here we discuss the evidence and propose a new metabolic and regulatory model linking pectin to cell-wall-mediated immunity, including ripening-associated disease susceptibility in the tomato.     

Effects of disturbance on phosphorus turnover and plankton dynamics

In coastal lagoons, one common type of anthropogenic disturbanceis opening of the sand barriers that isolate these lagoons fromthe sea. We investigated how a disturbance caused by an artificialopening of the sand barrier affects phytoplankton and bacterioplanktondynamics and alkaline phosphatase activities (APA) in a Braziliancoastal lagoon. We propose an index of potential phosphorusturnover time (PPTT) calculated as the ratio of total phosphorus(TP) and APA in an attempt to quantify the effect of disturbanceson phosphorus recycling within the plankton community. Openingthe barrier caused increased phytoplankton biomass, TP concentrations,APA and PPTT, and decreased bacteria to phytoplankton biomassratio. PPTT is easily calculated and may be used to detect planktonresponses to anthropogenic disturbances.     

Meteorological forcing of plankton dynamics in a large and deep continental European lake

The timing of various plankton successional events in Lake Constance was tightly coupled to a large-scale meteorological phenomenon, the North Atlantic Oscillation (NAO). A causal chain of meteorological, hydrological, and ecological processes connected the NAO as well as winter and early spring meteorological conditions to planktonic events in summer leading to a remarkable memory of climatic effects lasting over almost half a year. The response of Daphnia to meteorological forcing was most probably a direct effect of altered water temperatures on daphnid growth and was not mediated by changes in phytoplankton concentrations. High spring water temperatures during ”high-NAO years” enabled high population growth rates, resulting in a high daphnid biomass as early as May. Hence, a critical Daphnia biomass to suppress phytoplankton was reached earlier in high-NAO years yielding an early and longer-lasting clear-water phase. Finally, an earlier summer decline of Daphnia produced in a negative relationship between Daphnia biomass in July and the NAO. Meteorological forcing of the seasonal plankton dynamics in Lake Constance included simple temporal shifts of processes and successional events, but also complex changes in the relative importance of different mechanisms. Since Daphnia plays an important role in plankton succession, a thorough understanding of the regulation of its population dynamics provides the key for predictions of the response of freshwater planktonic food webs to global climate change. Received: 15 February 1999 / Accepted: 23 August 1999     

Chaos and order in plankton dynamics. Complex behavior of a simple model

The role of the diffusive interaction between fish-populated and fish-free habitats in a patchy environment in plankton pattern formation is studied by means of a minimal reaction-diffusion model of the nutrient-plankton-fish food chain. It is shown that such interaction can give rise to spatio-temporal plankton patterns. The fractal dimension of the patterns is shown dependent on the fish predation rate. The spatially averaged plankton dynamics depending on both fish predation rate and distance between fish-populated habitats can exhibit chaotic and regular behavior. The chaotic plankton dynamics is characteristic of a wide parameter range.     

Physico-chemical limnology and plankton dynamics of Mazvikadei,a tropical reservoir in Zimbabwe

The limnology of Mazvikadei Reservoir, northern Zimbabwe, was investigated in 2015 to determine whether it had changed since filling in 1990. The reservoir is characterised by low algal biomass, low nutrients (i.e. N and P) and high water clarity/transparency. Fifty-four species of phytoplankton were recorded, comprising Bacillariophyta, Chlorophyta, Cyanophyta, Desmids, Dinophyta and Euglenophyta. Chlorophyta numerically dominated in the hot dry season, whereas Bacillariophyta, Desmids, Dinophyta and Euglenophyta dominated in the cool dry season. Species richness was highest at the onset of the cool dry season, in response to high nutrient concentrations. Phytoplankton abundance and composition were significantly correlated with temperature, nitrates and total nitrogen. Nineteen zooplankton species were recorded, including Copepoda, Cladocera and Rotifera. Overall, Cladocera were numerically dominant and became most abundant during the cool dry season. Rotifers and copepods dominated during the hot dry season. The zooplankton abundance was correlated with reactive phosphorus and phytoplankton abundance. The trophic state of Mazvikadei Reservoir seems to have stabilised and to have assumed the physico-chemical characteristics and plankton community typical of an oligotrophic lake.     

Evolutionary dynamics as a component of stage-structured matrix models: an example using Trillium grandiflorum

Evolution by natural selection improves fitness and may therefore influence population trajectories. Demographic matrix models are often employed in conservation studies to project population dynamics, but such analyses have not incorporated evolutionary dynamics. We project evolutionarily informed population trajectories for a population of the perennial plant Trillium grandiflorum, which is declining due to high levels of herbivory by white-tailed deer. Individuals with later flowering times are less often consumed, so there is selection on this trait. We first incorporated selection analyses into a deterministic matrix model in three ways (corresponding to different methods that have been used for analyzing evolution in structured populations). Because it is not clear which of these methods works best for stage-structured models, we compared each with a more realistic, individual-based model. Deterministic models using fitness averaged over the phenotypic distribution gave trajectories that were similar to those of the individual-based model, whereas the deterministic model using fitness at the mean phenotype gave a much faster rate of evolution than that which was observed. This illustrates that subtle differences in the way in which one splices evolution into demographic models can have a large effect on expected outcomes. This study demonstrates that, by combining demographic and selection analyses, one can gauge the potential relevance of evolution to population dynamics and persistence.     

Data transformation as a scaling operation in ordination of plankton

Data transformation is seen here as an aspect of scaling such that we are less interested in the quirks and properties of each transformation but are more concerned with the general scaling properties and trends of suites of transformations.Over two years of daily phytoplankton abundance data for 30 species from a temperate lake (Llyn Maelog, North Wales) were subjected to a series of scale-ordered transformations. Two major classes of transformation were systematically varied: binary and smoothing. Binary transformation scaled the cutoff threshold between presence and absence of a species to various levels of abundance.With successively smaller universes and smoothing windows and successive species exclusion, ordinations of sample dates revealed smaller scaled structures in the order: annual cycles of species turnover, seasonal areas of attraction and uniqueness of individual sample dates. Gradual increases in the length of the smoothing window resulted in gradual shifts in the positions of points in sample data ordination, but not necessarily in the species ordinations. Thus sample data structures are more stable with change in scale than are species data structures. These differences in stability are discussed in the context of filtering characteristics of data collection and data analysis. Transformations producing similar species statistics (means, variances and skews) did not generally give similar ordination results, while some transformations giving similar ordinations differed in species statistical parameters.Acknowledgement: The present research was supported by a grant from the National Science foundation (NSF DEB 78-07546) to the first author.     

Temporal plankton dynamics in an oligotrophic maritime Antarctic lake

• 1 The population density, diversity and productivity of the microbial plankton in an oligotrophic maritime Antarctic lake were studied for a 15‐month period between December 1994 and February 1996.
• 2 In the lake, concentrations of nutrients and dissolved organic carbon were uniformly low, temperature varied over a small annual range of 0.1–3 °C, and the surface was ice‐covered except during a period of approximately 6 weeks in summer.
• 3 The total of 57 morphotypes of protozoa observed during the study is a higher taxonomic diversity than previously reported from continental Antarctic lakes, but lower than that found in more eutrophic maritime Antarctic lakes. Likewise, planktonic abundance and productivity were lower than has been reported in other lakes on Signy Island, but generally higher than those of lakes on the Antarctic continent.
• 4 There were marked seasonal and interannual variations in planktonic population density.
• 5 Chlorophyll a concentrations ranged from undetectable to 4.2 µg L^‐1 and the greatest rate of primary productivity measured was 4.5 mg C m^‐3 h^‐1. The phytoplankton was dominated by small chlorophytes and chrysophytes, with phototrophic nanoflagellate abundance ranging from 1.1 × 10³ to 1.2 × 10⁷ L^‐1.
• 6 Bacterial densities of 3.6 × 10⁸ to 1.9 × 10¹⁰ L^‐1 were recorded and bacterial productivity reached a peak of 0.36 µg C L^‐1 h^‐1. Numbers of heterotrophic nanoflagellates between 5.0 × 10⁴ and 1.8 × 10⁷ L^‐1, and of ciliates from undetectable to 1.1 × 10⁴ L^‐1 were observed. Naked amoebae were usually rare, but occasionally reached peaks of up to 1.5 × 10³ L^‐1.

     

Seasonal plankton dynamics in a Mediterranean hypersaline coastal lagoon: the Mar Menor

The seasonal distribution of plankton in a Mediterranean hypersalinecoastal lagoon has been studied through a dataset, comprisingthe taxonomic composition and the size–abundance distributionof both phyto- and zooplankton, measured by image analysis techniquesduring a one-year time series of weekly samplings. The studiedorganisms ranged from small nanoplanktonic heterotrophic flagellates(2 µm diameter) to fish larvae (>2 µm). The phytoplanktonannual succession was characterized by a winter period dominatedby Rhodomonas spp. and Cryptomonas spp. with Cyclotella spp.as the main diatom represented, a spring phase where diatoms(mainly Cyclotella) were the dominant group with some monospecificblooms of other diatoms (mainly of Chaetoceros sp.), a summerphase characterized by diatoms with blooms of Niztschia closterium,and a post-summer phase where dinoflagellates increased withpeaks of Ceratium furca. High densities of the microbial foodweb elements, flagellates and ciliates, indicate the importanceof the microbial loop in the ecosystem. Meroplankton contributedwidely to the seasonal character of the zooplankton distribution.Copepods, represented by Oithona nana, Centropages ponticusand Acartia spp. (mainly latisetosa), remained relatively constantthroughout the year, exhibiting a lower density in the warmerwater period (July–September). At the end of the samplingperiod, a massive proliferation of copepods (>1000 ind l^–1), mainly due to O. nana, took place. The autotrophsto heterotrophs biovolume ratio (A:H) remained lower than 1throughout the year except when, occasionally, large phytoplanktoncells bloomed. Persistent very low values of A:H suggest thatadditional sources of energy, such as the microbial loop ordetrital pathways, would be needed to sustain the high heterotrophicbiovolume found in the lagoon.     

Short-term dynamics of a Melosira population in the plankton of an impoundment in central Chile

Relationships among the chlorophyll-specific photosyntheticbehaviour of a Melosira-dominated phytoplankton, net variationsin the quantity of Melosira in suspension, the rates of sinkingloss of Melosira cells and did fluctuations in water-columnstability were investigated in Embalse Rapel. central Chile.The short study was carried out during the autumnal equinoxialperiod (March. 1984) of maximal Melosira abundance to determinethe extent to which these various components might contributeto the distinctly bimodal seasonal distribution of phytoplanktonabundance in this reservoir. Diurnal photosynthetic productionwithin the illuminated layer and supposed column respirationwere found to be of a similar order, suggesting that net increasethrough cellular growth was already substantially limited byself-shading. Thus observed net changes in the standing populationmay have depended more on the relative magnitude of sinkingloss rates and recruitment through resuspension, which processesare profoundly influenced by alternations in wind-induced mixingand intense, near-surface, thermal stratification. As the relativefrequency and duration of these episodes vary during the year,so does the capacity of the Melosira to maintain or increaseits standing biomass. High cellular photosynthetic efficiencycoupled with rapid sinking rate appear to be of selective advantagewhen the cycle of alternation is approximately diel. The applicationof these findings to the seasonal distribution of M. granulatain the Rapel reservoir and to the ecology of planktonic Melosirapopulations elsewhere is briefly discussed.