Modeling phytoplankton growth rates

Mathematical models of planktonic ecosystems use a variety ofdifferent formulations to relate phytoplankton growth ratesto environmental conditions. Does the formulation influencethe model result? We have modified the model of Fasham, Ducklowarid McKelvie (J. Mar. Res., 34,591–639, 1990) to testhow its results would respond to changes in algal growth rateformulations. The original model uses a Monod relationship betweennutrient concentration and relative growth rate, and a multiplicativerule to combine light and nutrient effects. Use of a Droop formulationfor algal growth rate or a threshold (Blackman's law) mechanismto combine light and nutrient limitation produced significantchanges in simulation results. One important effect was to increasezooplankton population and, as a result, the regenerated production.While there are aesthetic reasons to prefer these alternateformulations, a more accurate formulation will require morelaboratory work on algal physiology. Such laboratory work shouldbe encouraged as an adjunct to modeling work.     

Ocean acidification with (de)eutrophication will alter future phytoplankton growth and succession

Human activity causes ocean acidification (OA) though the dissolution of anthropogenically generated CO₂ into seawater, and eutrophication through the addition of inorganic nutrients. Eutrophication increases the phytoplankton biomass that can be supported during a bloom, and the resultant uptake of dissolved inorganic carbon during photosynthesis increases water-column pH (bloom-induced basification). This increased pH can adversely affect plankton growth. With OA, basification commences at a lower pH. Using experimental analyses of the growth of three contrasting phytoplankton under different pH scenarios, coupled with mathematical models describing growth and death as functions of pH and nutrient status, we show how different conditions of pH modify the scope for competitive interactions between phytoplankton species. We then use the models previously configured against experimental data to explore how the commencement of bloom-induced basification at lower pH with OA, and operating against a background of changing patterns in nutrient loads, may modify phytoplankton growth and competition. We conclude that OA and changed nutrient supply into shelf seas with eutrophication or de-eutrophication (the latter owing to pollution control) has clear scope to alter phytoplankton succession, thus affecting future trophic dynamics and impacting both biogeochemical cycling and fisheries.     

The role of functional traits and trade-offs in structuring phytoplankton communities: scaling from cellular to ecosystem level

Trait-based approaches to community structure are increasingly used in terrestrial ecology. We show that such an approach, augmented by a mechanistic analysis of trade-offs among functional traits, can be successfully used to explain community composition of marine phytoplankton along environmental gradients. Our analysis of literature on major functional traits in phytoplankton, such as parameters of nutrient-dependent growth and uptake, reveals physiological trade-offs in species abilities to acquire and utilize resources. These trade-offs, arising from fundamental relations such as cellular scaling laws and enzyme kinetics, define contrasting ecological strategies of nutrient acquisition. Major groups of marine eukaryotic phytoplankton have adopted distinct strategies with associated traits. These diverse strategies of nutrient utilization can explain the distribution patterns of major functional groups and size classes along nutrient availability gradients.     

Effects of nutrient recycling by zooplankton and fish on phytoplankton communities

In laboratory experiments we tested the hypothesis that nutrients supplied by fish and zooplankton affect the structure and dynamics of phytoplankton communities. As expected from their body size differences, fish released nutrients at lower mass-specific rates than Daphnia. On average, these consumers released nutrients at similar N:P ratios, although the ratios released by Daphnia were more variable than those released by fish. Nutrient supply by both fish and Daphnia reduced species richness and diversity of phytoplankton communities and increased algal biomass and dominance. However, nutrient recycling by fish supported a more diverse phytoplankton community than nutrient recycling by Daphnia. We conclude that nutrient recycling by zooplankton and fish have different effects on phytoplankton community structure due to differences in the quality of nutrients released. Received: 21 December 1998 / Accepted: 31 May 1999     

Phytoplankton growth and stoichiometric responses to warming,nutrient addition and grazing depend on lake productivity and cell size

Global change involves shifts in multiple environmental factors that act in concert to shape ecological systems in ways that depend on local biotic and abiotic conditions. Little is known about the effects of combined global change stressors on phytoplankton communities, and particularly how these are mediated by distinct community properties such as productivity, grazing pressure and size distribution. Here, we tested for the effects of warming and eutrophication on phytoplankton net growth rate and C:N:P stoichiometry in two phytoplankton cell size fractions (<30 µm and >30 µm) in the presence and absence of grazing in microcosm experiments. Because effects may also depend on lake productivity, we used phytoplankton communities from three Dutch lakes spanning a trophic gradient. We measured the response of each community to multifactorial combinations of temperature, nutrient, and grazing treatments and found that nutrients elevated net growth rates and reduced carbon:nutrient ratios of all three phytoplankton communities. Warming effects on growth and stoichiometry depended on nutrient supply and lake productivity, with enhanced growth in the most productive community dominated by cyanobacteria, and strongest stoichiometric responses in the most oligotrophic community at ambient nutrient levels. Grazing effects were also most evident in the most oligotrophic community, with reduced net growth rates and phytoplankton C:P stoichiometry that suggests consumer‐driven nutrient recycling. Our experiments indicate that stoichiometric responses to warming and interactions with nutrient addition and grazing are not universal but depend on lake productivity and cell size distribution.     

Persistence and coexistence in zooplankton-phytoplankton-nutrient models with instantaneous nutrient recycling

We consider plankton-nutrient interaction models consisting of phytoplankton, herbivorous zooplankton and dissolved limiting nutrient with general nutrient uptake functions and instantaneous nutrient recycling. For the model with constant nutrient input and different constant washout rates, conditions for boundedness of the solutions, existence and stability of non-negative equilibria, as well as persistence are given. We also consider the zooplankton-phytoplankton-nutrient interaction models with a fluctuating nutrient input and with a periodic washout rate, respectively. It is shown that coexistence of the zooplankton and phytoplankton may arise due to positive bifurcating periodic solutions.Research has been supported in part by a University of Alberta Ph.D. Scholarship and is in part based on the author's Ph.D. thesis under the supervision of Professor H. 1. Freedman, to whom the author owes a debt of appreciation and gratitude for his kind advice, helpful comments and continuous encouragement     

Effects of nutrient supply and nutrient ratio on diversity–productivity relationships of phytoplankton in the Cau Hai lagoon,Vietnam

Diversity and productivity of primary producers are known to be influenced simultaneously by resource availability and resource ratio, but the relative importance of these two factors differed among studies and so far only entire phytoplankton communities were investigated which might ignore specific nutrient requirements and stoichiometric plasticity of different functional groups. We measured nutrient availability (DIN, total N [TN], total P [TP]), nutrient imbalance (TN:TP, DIN:TP, N:P_seston), species richness, and abundance of the whole phytoplankton community, as well as those specific for cyanobacteria, diatoms, and dinoflagellates in Cau Hai lagoon in Vietnam. We determined the correlation among these variables, using structural equation modeling. The models applied to the whole phytoplankton community indicated that the nutrient availability (particularly TP and DIN) drove variation in phytoplankton abundance and richness, and that abundance also depended on species richness. The models applied to different functional groups differed considerably from the entire community and among each other, and only a part of the models was significant. The relationship between nutrient availability (mainly TP) and abundance was driven by cyanobacteria, and the relationship between nutrient imbalance (only with N:P_seston) and species richness was driven by diatoms. Remarkably, the positive relationship between species richness and abundance, as consistently observed for the whole phytoplankton community, was only observed for one of the three functional groups (diatoms), indicating that resource complementarity occurs particularly among species of different functional groups. Our results emphasized that nutrient availability (TP and to a lesser extent DIN) as well as nutrient imbalance (albeit only with N:P_seston as proxy) were driving factors for the phytoplankton community in the Cau Hai lagoon and hence alterations in both of these factors leading to a shift in phytoplankton species composition and productivity.     

Influence of nutrient availability on phytoplankton growth and community structure in the Port Adelaide River,Australia: [2pt] bioassay assessment of potential nutrient limitation

We investigated the influence of nutrient availability, specifically nitrogen, phosphorus and silicon on growth and community structure of phytoplankton from the Port Adelaide River estuary, South Australia. Two bioassay experiments were conducted. The first, Nutrich1, involved addition of nutrients in vitro to samples of the natural phytoplankton community from a single location in the upper estuary. The second, Nutrich2, involved nutrient addition and incubation of water from five locations in the estuary following inoculation with a `standardised' phytoplankton assemblage derived from laboratory cultures. In Nutrich1, enrichment with silicon led to greatly enhanced phytoplankton biomass due to increased growth of diatoms. Addition of nitrogen or phosphorus had little effect on phytoplankton growth. In Nutrich2, addition of nitrogen resulted in enhanced growth of phytoplankton in water collected from near the mouth the estuary, but there were no differences in growth among nutrient treatments for the remaining locations. Comparison of phytoplankton growth rate among locations revealed a trend of decreasing growth in moving towards the mouth of the estuary. This trend was unaffected by enrichment with nitrate, phosphate or silicate. We suggest that spatial variation in growth potential within the Port Adelaide River estuary may relate to variation in the concentration of nitrogen as ammonium.     

Relevance of various formulations of phytoplankton chlorophyll a:carbon ratio in a 3D marine ecosystem model

Numerous experimental studies showed that the phytoplankton Chla-to-Carbon ratio (Chla:C) is highly variable, whereas most of the marine ecosystem models use a constant ratio. In this work, we tested three different formulations for computing the modelled Chla in a 3D coupled hydrodynamical-biogeochemical model of the Southwest lagoon of New Caledonia. The first formulation considers a constant Chla:C ratio. In the second one, Chla is a diagnostic variable related to the variable phytoplankton nitrogen-to-carbon ratio. In the last formulation, Chla is a state variable of the model, which is dynamically simulated. Results showed important differences between the formulations, the first leading to overestimate the Chla concentration in low nutrients conditions. Thus, this study strengthens the importance of the Chla modelling in a coupled model in order to better estimate a crucial variable for validation of ecosystem models.     

Phosphorus physiological ecology and molecular mechanisms in marine phytoplankton

Phosphorus (P) is an essential nutrient for marine phytoplankton and indeed all life forms. Current data show that P availability is growth‐limiting in certain marine systems and can impact algal species composition. Available P occurs in marine waters as dissolved inorganic phosphate (primarily orthophosphate [Pi]) or as a myriad of dissolved organic phosphorus (DOP) compounds. Despite numerous studies on P physiology and ecology and increasing research on genomics in marine phytoplankton, there have been few attempts to synthesize information from these different disciplines. This paper is aimed to integrate the physiological and molecular information on the acquisition, utilization, and storage of P in marine phytoplankton and the strategies used by these organisms to acclimate and adapt to variations in P availability. Where applicable, we attempt to identify gaps in our current knowledge that warrant further research and examine possible metabolic pathways that might occur in phytoplankton from well‐studied bacterial models. Physical and chemical limitations governing cellular P uptake are explored along with physiological and molecular mechanisms to adapt and acclimate to temporally and spatially varying P nutrient regimes. Topics covered include cellular Pi uptake and feedback regulation of uptake systems, enzymatic utilization of DOP, P acquisition by phagotrophy, P‐limitation of phytoplankton growth in oceanic and coastal waters, and the role of P‐limitation in regulating cell size and toxin levels in phytoplankton. Finally, we examine the role of P and other nutrients in the transition of phytoplankton communities from early succession species (diatoms) to late succession ones (e.g., dinoflagellates and haptophytes).     

Interactions between temperature and nutrients across levels of ecological organization

Temperature and nutrient availability play key roles in controlling the pathways and rates at which energy and materials move through ecosystems. These factors have also changed dramatically on Earth over the past century as human activities have intensified. Although significant effort has been devoted to understanding the role of temperature and nutrients in isolation, less is known about how these two factors interact to influence ecological processes. Recent advances in ecological stoichiometry and metabolic ecology provide a useful framework for making progress in this area, but conceptual synthesis and review are needed to help catalyze additional research. Here, we examine known and potential interactions between temperature and nutrients from a variety of physiological, community, and ecosystem perspectives. We first review patterns at the level of the individual, focusing on four traits – growth, respiration, body size, and elemental content – that should theoretically govern how temperature and nutrients interact to influence higher levels of biological organization. We next explore the interactive effects of temperature and nutrients on populations, communities, and food webs by synthesizing information related to community size spectra, biomass distributions, and elemental composition. We use metabolic theory to make predictions about how population‐level secondary production should respond to interactions between temperature and resource supply, setting up qualitative predictions about the flows of energy and materials through metazoan food webs. Last, we examine how temperature–nutrient interactions influence processes at the whole‐ecosystem level, focusing on apparent vs. intrinsic activation energies of ecosystem processes, how to represent temperature–nutrient interactions in ecosystem models, and patterns with respect to nutrient uptake and organic matter decomposition. We conclude that a better understanding of interactions between temperature and nutrients will be critical for developing realistic predictions about ecological responses to multiple, simultaneous drivers of global change, including climate warming and elevated nutrient supply.     

Nutrient interactions between phytoplankton and bacterioplankton under different carbon dioxide regimes

Light, nutrients, temperature, pH, and salinity are important factors in controlling the growth of phytoplankton and bacterioplankton. Supply of key nutrients to these communities can result in mutualistic or competitive relationships between bacterioplankton and phytoplankton. In this study, we investigated growth and uptake of nutrients by the marine prasinophyte flagellate Tetraselmis chui (strain PLY429) in the presence and absence of a community of bacterioplankton at two pH levels. Growth of PLY429 and total nutrient uptake were calculated for each treatment. The addition of bacterioplankton resulted in lower growth rates of PLY429, but the removal of ammonium was greater in those cultures with bacterioplankton present. The division rate of PLY429 was affected by pH; however, pH changes did not result in different uptake rates of nitrate, ammonium, or phosphate by the mixed algal and bacterial assemblage. These findings suggest that bacterioplankton and phytoplankton were competing for ammonium and that a lower pH resulted in more rapid algal growth. Mention of a trade name does not imply endorsement by the National Marine Fisheries Service.     

A study of phosphate limitation in Lake Maarsseveen: phosphate uptake kinetics versus bioassays

In order to assess possible phosphate limitation for the phytoplankton community of Lake Maarsseveen, two techniques (phosphate uptake experiments and bioassays) were employed simultaneously in February–March 1982. In that period the ambient phosphate concentration of the lake water was less than 0.03 µM P and the diatom Asterionella formosa constituted more than 90% of the phytoplankton population. The phosphate uptake experiments showed relatively high uptake capacities and low cell phosphorus contents for the natural phytoplankton community. This suggested phosphate limitation throughout the test period. The growth stimulation of the phytoplankton after enrichment with phosphate, however, only revealed phosphate limitation from the beginning of March and bioassays may therefore be regarded as a less sensitive method.     

Effects of sea surface warming on marine plankton

Ocean warming has been implicated in the observed decline of oceanic phytoplankton biomass. Some studies suggest a physical pathway of warming via stratification and nutrient flux, and others a biological effect on plankton metabolic rates; yet the relative strength and possible interaction of these mechanisms remains unknown. Here, we implement projections from a global circulation model in a mesocosm experiment to examine both mechanisms in a multi‐trophic plankton community. Warming treatments had positive direct effects on phytoplankton biomass, but these were overcompensated by the negative effects of decreased nutrient flux. Zooplankton switched from phytoplankton to grazing on ciliates. These results contrast with previous experiments under nutrient‐replete conditions, where warming indirectly reduced phytoplankton biomass via increased zooplankton grazing. We conclude that the effect of ocean warming on marine plankton depends on the nutrient regime, and provide a mechanistic basis for understanding global change in marine ecosystems.     

Dynamics of nutrient-phytoplankton interaction in the presence of viral infection

The present paper deals with the problem of a nutrient-phytoplankton (N-P) populations where phytoplankton population is divided into two groups, namely susceptible phytoplankton and infected phytoplankton. Conditions for coexistence or extinction of populations are derived taking into account general nutrient uptake functions and Holling type-II functional response as an example. It is observed that the three component systems persist when the infected phytoplankton population is not able to consume nutrient.     

Increasing importance of small phytoplankton in a warmer ocean

The macroecological relationships among marine phytoplankton total cell density, community size structure and temperature have lacked a theoretical explanation. The tiniest members of this planktonic group comprise cyanobacteria and eukaryotic algae smaller than 2 μm in diameter, collectively known as picophytoplankton. We combine here two ecological rules, the temperature–size relationship with the allometric size‐scaling of population abundance to explain a remarkably consistent pattern of increasing picophytoplankton biomass with temperature over the ?0.6 to 22 °C range in a merged dataset obtained in the eastern and western temperate North Atlantic Ocean across a diverse range of environmental conditions. Our results show that temperature alone was able to explain 73% of the variance in the relative contribution of small cells to total phytoplankton biomass regardless of differences in trophic status or inorganic nutrient loading. Our analysis predicts a gradual shift toward smaller primary producers in a warmer ocean. Because the fate of photosynthesized organic carbon largely depends on phytoplankton size, we anticipate future alterations in the functioning of oceanic ecosystems.     

Microscale patchiness leads to large and important intraspecific internal nutrient heterogeneity in phytoplankton

Phytoplankton stoichiometry or nutrient content has been shown to vary in a number of dimensions (species, condition, time, space), but the heterogeneity within a species at a given time and location, and the underlying mechanisms and importance have not been explored. There are a number of mechanisms that can create intraspecific heterogeneity, and theory suggests it can affect the population growth rate. We studied heterogeneity in P content of the freshwater diatom Cyclotella meneghiniana in the Charles River in Boston. Single-cell observations using synchrotron-based X-ray fluorescence show that the nutrient status varies from P-starved to P-replete. We simulate individual cells using an agent-based model that accounts for a number of mechanisms that can create heterogeneity, including surface area–based uptake, mortality differentiation, stochastic biological variability in states and behavior, macroscale mixing, and microscale nutrient patch encounter. By performing a number of simulations with various mechanisms turned on/off and comparing to data, we conclude that the heterogeneity is mostly due to microscale patchiness (85%). We explore the importance of accounting for heterogeneity in models by performing a simulation with the growth rate based on the population-average internal nutrient, as is done in conventional population-level models. This shows that ignoring heterogeneity increases the population growth rate by a factor of 1.47. To account for different heterogeneity in the laboratory and field, population-level ecosystem models should reduce maximum growth rates. The magnitude of this correction depends on local conditions, and in our case, it is a factor of 0.72.     

Long Term Effect of Cyprinid Fishes on Phytoplankton and Zooplankton Communities in a Shallow Water Protection Reservoir

The effects of fish kill and different fish stocks on the phytoplankton and zooplankton dynamics were studied in a shallow hypertrophic reservoir system. When fish stock was below 100 kg ha⁻¹, nutrient availability was not the main limiting factor for growth of phytoplankton. Consequently top‐down forces controlled phytoplankton. In the years with high fish stock (>100 kg ha⁻¹) the bottom‐up forces dominated as nutrient availability was the main limiting factor for growth of phytoplankton. We can conclude that significant water quality improvement can be achieved in the reservoir system by decreasing fish stock below 100 kg ha⁻¹. Although clear‐water phase could be stabilised temporary by macrophytes, stabilisation of good water quality requires continuous regulation of fish community. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

Soluble nitrogen and phosphorus excretion of exotic freshwater mussels (Dreissena spp.): potential impacts for nutrient remineralisation in western Lake Erie

1. Recent increases in phytoplankton biomass and the recurrence of cyanobacterial blooms in western Lake Erie, concomitant with a shift from a community dominated by zebra mussels (Dreissena polymorpha) to one dominated by quagga mussels (D. bugensis), led us to test for differences in ammonia‐nitrogen and phosphate‐phosphorus excretion rates of these two species of invasive molluscs. 2. We found significant differences in excretion rate both between size classes within a taxon and between taxa, with zebra mussels generally having greater nutrient excretion rates than quagga mussels. Combining measured excretion rates with measurements of mussel soft‐tissue dry weight and shell length, we developed nutrient excretion equations allowing estimation of nutrient excretion by dreissenids. 3. Comparing dreissenid ammonia and phosphate excretion with that of the crustacean zooplankton, we demonstrated that the mussels add to nitrogen and phosphorus remineralisation, shortening nitrogen and phosphorus turnover times, and, importantly, modify the nitrogen and phosphorus cycles in Lake Erie. The increased nutrient flux from dreissenids may facilitate phytoplankton growth and cyanobacterial blooms in well‐mixed and/or shallow areas of western Lake Erie.