Linking population dynamics models with empirically derived models through phytoplankton primary production.
ABSTRACT: There are alternative methods for estimation of phytoplankton primary production (PP) that are fundamentally different in the calculation approach. The process-oriented PP model is a mechanistic, empirically derived method based on the photosynthesis-light relationships. The population dynamics-based PP calculation, which is a synthetic method, provides a production estimate based on population dynamics of phytoplankton. These alternative methods were here compared with regard to production estimates and linked to enhance the performance of the existing models of population dynamics applied to a wide variety of lakes worldwide in terms of morphometry, nutrient status, and light environments. Estimates of PP were shown to be sensitive to changes in phytoplankton sinking and zooplankton grazing rates in both methods. Production estimates in the process-oriented PP model were also sensitive to light-associated parameters such as day length. Although the production estimated from the population dynamics-based PP calculation tended to be lower than that from the process-oriented PP model irrespective of lake morphometry, production estimates calculated from both methods with standard parameterization were comparable when production was estimated on an annual timescale. However, it was also shown that the alternative methods could produce different production estimates when estimated on shorter timescales such as cyanobacterial blooms in summer. Cyanobacteria with low mortality due to grazing and sinking losses have been considered as trophic bottlenecks, but there is increasing evidence that their mortality is, to a considerable extent, due to parasitic pathogens. In the case of cyanobacterial blooms, an addition of parasite-related loss term (19%-33% of standing stock) resulted in a resolution of the difference in production estimates between the methods. These analyses theoretically support the critical role of parasitism and resolve the bottleneck problem in aquatic ecosystem metabolism.
Project description:The impacts of grazing by meso- and microzooplankton on phytoplankton primary production (PP) was investigated in the surface layer of the western North Atlantic during spring. Shipboard experiments were performed on a latitudinal transect at three stations that differed in mixed layer depth, temperature, and mesozooplankton taxonomic composition. The mesozooplankton community was numerically dominated by Calanus finmarchicus at the northern and central station, with Calanus hyperboreus also present at the northern station. The southern station was >10 °C warmer than the other stations and had the most diverse mesozooplankton assemblage, dominated by small copepods including Paracalanus spp. Microzooplankton grazing was detected only at the northern station, where it removed 97% of PP. Estimated clearance rates by C. hyperboreus and C. finmarchicus suggested that at in-situ abundance these mesozooplankton were not likely to have a major impact on phytoplankton abundance, unless locally aggregated. Although mesozooplankton grazing impact on total phytoplankton was minimal, these grazers completely removed the numerically scarce > 10 µm particles, altering the particle-size spectrum. At the southern station, grazing by the whole mesozooplankton assemblage resulted in a removal of 14% of PP, and its effect on net phytoplankton growth rate was similar irrespective of ambient light. In contrast, reduction in light availability had an approximately 3-fold greater impact on net phytoplankton growth rate than mesozooplankton grazing pressure. The low mesozooplankton grazing impact across stations suggests limited mesozooplankton-mediated vertical export of phytoplankton production. The constraints provided here on trophic transfer, as well as quantitative estimates of the relative contribution of light and grazer controls of PP and of grazer-induced shifts in particle size spectra, illuminate food web dynamics and aid in parameterizing modeling-frameworks assessing global elemental fluxes and carbon export.
Project description:In the Southern Ocean, large-scale phytoplankton blooms occur in open water and the sea-ice zone (SIZ). These blooms have a range of fates including physical advection, downward carbon export, or grazing. Here, we determine the magnitude, timing and spatial trends of the biogeochemical (export) and ecological (foodwebs) fates of phytoplankton, based on seven BGC-Argo floats spanning three years across the SIZ. We calculate loss terms using the production of chlorophyll-based on nitrate depletion-compared with measured chlorophyll. Export losses are estimated using conspicuous chlorophyll pulses at depth. By subtracting export losses, we calculate grazing-mediated losses. Herbivory accounts for ~90% of the annually-averaged losses (169?mg?C m<sup>-2</sup> d<sup>-1</sup>), and phytodetritus POC export comprises ~10%. Furthermore, export and grazing losses each exhibit distinctive seasonality captured by all floats spanning 60°S to 69°S. These similar trends reveal widespread patterns in phytoplankton fate throughout the Southern Ocean SIZ.
Project description:Top-down grazer control of cyanobacteria is a controversial topic due to conflicting reports of success and failure as well as a bias toward studies in temperate climates with large generalist grazers like Daphnia. In the tropical lowland lakes of Brazil, calanoid copepods of the Notodiaptomus complex dominate zooplankton and co-exist in high abundance with permanent blooms of toxic cyanobacteria, raising questions for grazer effects on bloom dynamics (i.e., top-down control vs. facilitation of cyanobacterial dominance). Accordingly, the effect of copepod grazing on the relative abundance of Microcystis co-cultured with a eukaryotic phytoplankton (Cryptomonas) was evaluated in a series of 6-day laboratory experiments. Grazer effects were tested in incubations where the growth of each phytoplankton in the presence or absence of the copepod Notodiaptomus iheringi was monitored in 1 L co-cultures, starting with a 6-fold initial dominance of Cryptomonas by biomass. Compared to the no grazer controls, N. iheringi reduced the growth of both phytoplankton, but Cryptomonas growth was reduced to negative values while Microcystis growth continued positively despite grazers. Hence, in a matter of 6 days selective grazing by N. iheringi increased the biomass of Microcystis relative to Cryptomonas by an order of magnitude compared to controls, and thus, facilitated the dominance of this cyanobacterium. To account for the potential effect of allelopathy, we performed a secondary experiment comparing the abundance and growth rate of Microcystis and Cryptomonas in single and mixed co-cultures in the absence of grazers. The growth rate of Microcystis was unaffected by the presence or relative abundance of Cryptomonas, and vice versa, indicating no allelopathic effects. Our results suggest that selectively grazing zooplankton may facilitate cyanobacteria blooms by grazing on their eukaryotic phytoplankton competitors in nature. Given that selective grazers predominate zooplankton biomass in warmer waters, grazer facilitation of blooms may be a common but poorly understood regulator of plankton dynamics in a warmer and more eutrophic world.
Project description:Phytoplankton anti-grazer traits control zooplankton grazing and are associated with harmful blooms. Yet, how morphological versus chemical phytoplankton defenses regulate zooplankton grazing is poorly understood. We compared zooplankton grazing and prey selection by contrasting morphological (filament length: short vs. long) and chemical (saxitoxin: STX- vs. STX+) traits of a bloom-forming cyanobacterium (Raphidiopsis) offered at different concentrations in mixed diets with an edible phytoplankton to a copepod grazer. The copepod selectively grazed on the edible prey (avoidance of cyanobacteria) even when the cyanobacterium was dominant. Avoidance of the cyanobacterium was weakest for the "short STX-" filaments and strongest for the other three strains. Hence, filament size had an effect on cyanobacterial avoidance only in the STX- treatments, while toxin production significantly increased cyanobacterial avoidance regardless of filament size. Moreover, cyanobacterial dominance reduced grazing on the edible prey by almost 50%. Results emphasize that the dominance of filamentous cyanobacteria such as Raphidiopsis can interfere with copepod grazing in a trait specific manner. For cyanobacteria, toxin production may be more effective than filament size as an anti-grazer defense against selectively grazing zooplankton such as copepods. Our results highlight how multiple phytoplankton defensive traits interact to regulate the producer-consumer link in plankton ecosystems.
Project description:While considerable effort has been devoted to understanding the factors regulating the development of phytoplankton blooms, the mechanisms leading to bloom decline and termination have received less attention. Grazing and sedimentation have been invoked as the main routes for the loss of phytoplankton biomass, and more recently, viral lysis, parasitism and programmed cell death (PCD) have been recognized as additional removal factors. Despite the importance of bloom declines to phytoplankton dynamics, the incidence and significance of various loss factors in regulating phytoplankton populations have not been widely characterized in natural blooms. To understand mechanisms controlling bloom decline, we studied two independent, inshore blooms of Alexandrium fundyense, paying special attention to cell mortality as a loss pathway. We observed increases in the number of dead cells with PCD features after the peak of both blooms, demonstrating a role for cell mortality in their terminations. In both blooms, sexual cyst formation appears to have been the dominant process leading to bloom termination, as both blooms were dominated by small-sized gamete cells near their peaks. Cell death and parasitism became more significant as sources of cell loss several days after the onset of bloom decline. Our findings show two distinct phases of bloom decline, characterized by sexual fusion as the initial dominant cell removal processes followed by elimination of remaining cells by cell death and parasitism.
Project description:The factors regulating phytoplankton community composition play a crucial role in structuring aquatic food webs. However, consensus is still lacking about the mechanisms underlying the observed biogeographical differences in cell size composition of phytoplankton communities. Here we use a trait-based model to disentangle these mechanisms in two contrasting regions of the Atlantic Ocean. In our model, the phytoplankton community can self-assemble based on a trade-off emerging from relationships between cell size and (1) nutrient uptake, (2) zooplankton grazing, and (3) phytoplankton sinking. Grazing 'pushes' the community towards larger cell sizes, whereas nutrient uptake and sinking 'pull' the community towards smaller cell sizes. We find that the stable environmental conditions of the tropics strongly balance these forces leading to persistently small cell sizes and reduced size diversity. In contrast, the seasonality of the temperate region causes the community to regularly reorganize via shifts in species composition and to exhibit, on average, bigger cell sizes and higher size diversity than in the tropics. Our results raise the importance of environmental variability as a key structuring mechanism of plankton communities in the ocean and call for a reassessment of the current understanding of phytoplankton diversity patterns across latitudinal gradients.
Project description:Wintertime convective mixing plays a pivotal role in the sub-polar North Atlantic spring phytoplankton blooms by favoring phytoplankton survival in the competition between light-dependent production and losses due to grazing and gravitational settling. We use satellite and ocean reanalyses to show that the area-averaged maximum winter mixed layer depth is positively correlated with April chlorophyll concentration in the northern Labrador Sea. A simple theoretical framework is developed to understand the relative roles of winter/spring convection and gravitational sedimentation in spring blooms in this region. Combining climate model simulations that project a weakening of wintertime Labrador Sea convection from Arctic sea ice melt with our framework suggests a potentially significant reduction in the initial fall phytoplankton population that survive the winter to seed the region's spring bloom by the end of the 21st century.
Project description:Turbulence and coherent circulation structures, such as submesoscale and mesoscale eddies, convective plumes and Langmuir cells, play a critical role in shaping phytoplankton spatial distribution and population dynamics. We use a framework of advection-reaction-diffusion equations to investigate the effects of turbulent transport on the phytoplankton population growth and its spatial structure in a vertical two-dimensional vortex flow field. In particular, we focus on how turbulent flow velocities and sinking influence phytoplankton growth and biomass aggregation. Our results indicate that conditions in mixing and growth of phytoplankton can drive different vertical spatial structures in the mixed layer, with the depth of the mixed layer being a critical factor to allow coexistence of populations with different sinking speed. With increasing mixed layer depth, positive growth for sinking phytoplankton can be maintained with increasing turbulent flow velocities, allowing the apparently counter-intuitive persistence of fast sinking phytoplankton populations in highly turbulent and deep mixed layers. These dynamics demonstrate the role of considering advective transport within a turbulent vortex and can help to explain observed phytoplankton biomass during winter in the North Atlantic, where the overturn of deep convection has been suggested to play a critical role in phytoplankton survival.
Project description:Freshwater ecosystems are severely threatened by urban development and agricultural intensification. Increased occurrence of algal blooms is a main issue, and the identification of local dynamics and drivers is hampered by a lack of field data. In this study, data from 13 cities (250 water bodies) were used to examine the capacity of trained community members to assess elevated phytoplankton densities in urban and peri-urban freshwater ecosystems. Coincident nutrient concentrations and land use observations were used to examine possible drivers of algal blooms. Measurements made by participants showed a good relationship to standard laboratory measurements of phytoplankton density, in particular in pond and lake ecosystems. Links between high phytoplankton density and nutrients (mainly phosphate) were observed. Microscale observations of pollution sources and catchment scale estimates of land cover both influenced the occurrence of algal blooms. The acquisition of environmental data by committed and trained community members represents a major opportunity to support agency monitoring programmes and to complement field campaigns in the study of catchment dynamics.
Project description:The phytoplanktonic production and prokaryotic consumption of organic matter significantly contribute to marine carbon cycling. Organic matter released from phytoplankton via three processes (exudation of living cells, cell disruption through grazing, and viral lysis) shows distinct chemical properties. We herein investigated the effects of phytoplanktonic whole-cell fractions (WF) (representing cell disruption by grazing) and extracellular fractions (EF) (representing exudates) prepared from Heterosigma akashiwo, a bloom-forming Raphidophyceae, on prokaryotic communities using culture-based experiments. We analyzed prokaryotic community changes for two weeks. The shift in cell abundance by both treatments showed similar dynamics, reaching the first peak (~4.1×106? ?cells? ?mL-1) on day 3 and second peak (~1.1×106? ?cells? ?mL-1) on day 13. We classified the sequences obtained into operational taxonomic units (OTUs). A Bray-Curtis dissimilarity analysis revealed that the OTU-level community structure changed distinctively with the two treatments. Ten and 13 OTUs were specifically abundant in the WF and EF treatments, respectively. These OTUs were assigned as heterotrophic bacteria mainly belonging to the Alteromonadales (Gammaproteobacteria) and Bacteroidetes clades and showed successive dynamics following the addition of organic matter. We also analyzed the dynamics of these OTUs in the ocean using publicly available metagenomic data from a natural coastal bloom in Monterey Bay, USA. At least two WF treatment OTUs showed co-occurrence with H. akashiwo, indicating that the blooms of H. akashiwo also affect these OTUs in the ocean. The present results strongly suggest that the thriving and dead cells of uninfected phytoplankton differentially influence the marine prokaryotic community.