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The idea that 'everything is everywhere, but the environment selects' has been seminal in microbial biogeography, and marine phytoplankton is one of the prototypical groups used to illustrate this. The typical argument has been that phytoplankton is ubiquitous, but that distinct assemblages form under environmental selection. It is well established that phytoplankton assemblages vary considerably between coastal ecosystems. However, the relative roles of compartmentalization of regional seas and site-specific environmental conditions in shaping assemblage structures have not been specifically examined. We collected data from coastal embayments that fall within two different water compartments within the same regional sea and are characterized by highly localized environmental pressures. We used principal coordinates of neighbour matrices (PCNM) and asymmetric eigenvector maps (AEM) models to partition the effects that spatial structures, environmental conditions and their overlap had on the variation in assemblage composition. Our models explained a high percentage of variation in assemblage composition (59-65%) and showed that spatial structure consistent with marine compartmentalization played a more important role than local environmental conditions. At least during the study period, surface currents connecting sites within the two compartments failed to generate sufficient dispersal to offset the impact of differences due to compartmentalization. In other words, our findings suggest that, even for a prototypical cosmopolitan group, everything is not everywhere.

Phytoplankton face environmental nutrient variations that occur in the dynamic upper layers of the ocean. Phytoplankton cells are able to rapidly acclimate to nutrient fluctuations by adjusting their nutrient-uptake system and metabolism. Disentangling these acclimation responses is a critical step in bridging the gap between phytoplankton cellular physiology and community ecology. Here, we analyzed the dynamics of phosphate (P) uptake acclimation responses along different P temporal gradients by using batch cultures of the diatom Phaeodactylum tricornutum. We employed a multidisciplinary approach that combined nutrient-uptake bioassays, transcriptomic analysis, and mathematical models. Our results indicated that cells increase their maximum nutrient-uptake rate (V max ) both in response to P pulses and strong phosphorus limitation. The upregulation of three genes coding for different P transporters in cells experiencing low intracellular phosphorus levels supported some of the observed V max variations. In addition, our mathematical model reproduced the empirical V max patterns by including two types of P transporters upregulated at medium-high environmental and low intracellular phosphorus levels, respectively. Our results highlight the existence of a sequence of acclimation stages along the phosphate continuum that can be understood as a succession of acclimation responses. We provide a novel conceptual framework that can contribute to integrating and understanding the dynamics and wide diversity of acclimation responses developed by phytoplankton.

Fluctuations in nutrient ratios over seasonal scales in aquatic ecosystems can result in overyielding, a condition arising when complementary life-history traits of coexisting phytoplankton species enables more complete use of resources. However, when nutrient concentrations fluctuate under short-period pulsed resource supply, the role of complementarity is less understood. We explore this using the framework of Resource Saturation Limitation Theory (r-strategists vs. K-strategists) to interpret findings from laboratory experiments. For these experiments, we isolated dominant species from a natural assemblage, stabilized to a state of coexistence in the laboratory and determined life-history traits for each species, important to categorize its competition strategy. Then, using monocultures we determined maximum biomass density under pulsed resource supply. These same conditions of resource supply were used with polycultures comprised of combinations of the isolated species. Our focal species were consistent of either r- or K-strategies and the biomass production achieved in monocultures depended on their efficiency to convert resources to biomass. For these species, the K-strategists were less efficient resource users. This affected biomass production in polycultures, which were characteristic of underyielding. In polycultures, K-strategists sequestered more resources than the r-strategists. This likely occurred because the intermittent periods of nutrient limitation that would have occurred just prior to the next nutrient supply pulse would have favored the K-strategists, leading to overall less efficient use of resources by the polyculture. This study provides evidence that fluctuation in resource concentrations resulting from pulsed resource supplies in aquatic ecosystems can result in phytoplankton assemblages' underyielding.

The anticipated long‐term rise in temperature can have numerous effects on freshwater phytoplankton, however, there is a great deal of uncertainty about how biogeochemical processes modify community functioning under a warming climate. Previous works regarding a large shallow lake and a mesocosm study showed a positive relationship between the temporal variability of phytoplankton composition and mean temperature, which might be a sign of warming‐stimulated instability in ecosystem processes. To elucidate whether these observations are part of a more general phenomenon, we combined a numerical model describing the dependence of algal growth on temperature and nutrient concentration with a multispecies, multi‐nutrient model. We ran simulations representative of changes in phytoplankton communities with randomized initial species composition under different temperature scenarios (current annual temperature pattern increased with 1°C, 2°C, or 3°C), various levels of nitrogen and phosphorus supply and different degrees of temporal variability in those supplies. The model outputs showed that seasonal variability in composition can be enhanced by warming through the mechanistic links between nutrient availability, temperature, and species‐specific growth rates, but the rate of increase is also dependent on nutrient ratios and the size of the species pool. Moreover, temporal variability in nutrient loads also enhanced compositional variability, but the rate of increase also depended on the level of nutrient supply. The results of our study hint that the impact of climate change and eutrophication on phytoplankton dynamics can be more intricate than spatiotemporally limited field or experimental observations would suggest. [ABSTRACT FROM AUTHOR]

Climate change‐related heatwaves are major threats to biodiversity and ecosystem functioning. However, our current understanding of the mechanisms governing community resistance to and recovery from extreme temperature events is still rudimentary. The spatial insurance hypothesis postulates that diverse regional species pools can buffer ecosystem functioning against local disturbances through the immigration of better‐adapted taxa. Yet, experimental evidence for such predictions from multi‐trophic communities and pulse‐type disturbances, like heatwaves, is largely missing. We performed an experimental mesocosm study to test whether species dispersal from natural lakes prior to a simulated heatwave could increase the resistance and recovery of plankton communities. As the buffering effect of dispersal may differ among trophic groups, we independently manipulated the dispersal of organisms from lower (phytoplankton) and higher (zooplankton) trophic levels. The experimental heatwave suppressed total community biomass by having a strong negative effect on zooplankton biomass, probably due to a heat‐induced increase in metabolic costs, resulting in weaker top‐down control on phytoplankton. While zooplankton dispersal did not alleviate the negative heatwave effects on zooplankton biomass, phytoplankton dispersal enhanced biomass recovery at the level of primary producers, providing partial evidence for spatial insurance. The differential responses to dispersal may be linked to the much larger regional species pool of phytoplankton than of zooplankton. Our results suggest high recovery capacity of community biomass independent of dispersal. However, community composition and trophic structure remained altered due to the heatwave, implying longer‐lasting changes in ecosystem functioning. [ABSTRACT FROM AUTHOR]

River phytoplankton has been studied to understand its occurrence and composition since the end of the nineteenth century. Later, pioneers addressed mechanisms that affected river phytoplankton by "origin of plankton", "turbulent mixing", "flow heterogeneity", "paradox of potamoplankton maintenance" and "dead zones" as keywords along the twentieth century. A major shift came with the recognition that characteristic units in phytoplankton compositions could be linked to specific set of environmental conditions, known as the "Phytoplankton Functional Group concept" sensu Reynolds. The FG concept could successfully be applied to river phytoplankton due to its close resemblance to shallow lakes phytoplankton. The FG approach enables one to separate the effects of "natural constraints" and "human impacts" on river phytoplankton and to evaluate the ecological status of rivers. The FG classification has mainly been advocated in the context of how the environment shaped the functional composition of phytoplankton. It may be further developed in the future by a trait-based mechanistic classification of taxa into FGs, and by the exact quantification of FGs on ecosystem functioning. These improvements will help quantify how global warming and human impacts affect river phytoplankton and corresponding alterations in ecosystem functioning. [ABSTRACT FROM AUTHOR]

The idea that 'everything is everywhere, but the environment selects' has been seminal in microbial biogeography, and marine phytoplankton is one of the prototypical groups used to illustrate this. The typical argument has been that phytoplankton is ubiquitous, but that distinct assemblages form under environmental selection. It is well established that phytoplankton assemblages vary considerably between coastal ecosystems. However, the relative roles of compartmentalization of regional seas and site-specific environmental conditions in shaping assemblage structures have not been specifically examined. We collected data from coastal embayments that fall within two different water compartments within the same regional sea and are characterized by highly localized environmental pressures. We used principal coordinates of neighbour matrices (PCNM) and asymmetric eigenvector maps (AEM) models to partition the effects that spatial structures, environmental conditions and their overlap had on the variation in assemblage composition. Our models explained a high percentage of variation in assemblage composition (59–65%) and showed that spatial structure consistent with marine compartmentalization played a more important role than local environmental conditions. At least during the study period, surface currents connecting sites within the two compartments failed to generate sufficient dispersal to offset the impact of differences due to compartmentalization. In other words, our findings suggest that, even for a prototypical cosmopolitan group, everything is not everywhere. [ABSTRACT FROM AUTHOR]

Evidence from terrestrial ecosystems indicates that biodiversity relates to ecosystem functions (BEF), but this relationship varies in its strength, in part, as a function of habitat connectivity and fragmentation. In primary producers, common proxies of ecosystem function include productivity and resource use efficiency. In aquatic primary producers, macroecological studies have observed BEF variance, where ecosystems with lower richness show stronger BEF relationships. However, aquatic ecosystems are less affected by habitat fragmentation than terrestrial systems and the mechanism underlying this BEF variance has been largely overlooked. Here, we provide a mechanistic explanation of BEF variance using a trait-based, numerical model parameterized for phytoplankton. Resource supply in our model fluctuates recurrently, similar to many coastal systems. Our findings show that following an extinction event, the BEF relationship can be driven by the species that are the most efficient resource users. Specifically, in species-rich assemblages, increased redundancy of efficient resource users minimizes the risk of losing function following an extinction event. On the other hand, in species-poor assemblages, low redundancy of efficient resource users increases the risk of losing ecosystem function following extinctions. Furthermore, we corroborate our findings with what has been observed from large-scale field studies on phytoplankton. [ABSTRACT FROM AUTHOR]

Fluctuations in nutrient ratios over seasonal scales in aquatic ecosystems can result in overyielding, a condition arising when complementary life-history traits of coexisting phytoplankton species enables more complete use of resources. However, when nutrient concentrations fluctuate under short-period pulsed resource supply, the role of complementarity is less understood. We explore this using the framework of Resource Saturation Limitation Theory (r-strategists vs. K-strategists) to interpret findings from laboratory experiments. For these experiments, we isolated dominant species from a natural assemblage, stabilized to a state of coexistence in the laboratory and determined life-history traits for each species, important to categorize its competition strategy. Then, using monocultures we determined maximum biomass density under pulsed resource supply. These same conditions of resource supply were used with polycultures comprised of combinations of the isolated species. Our focal species were consistent of either r- or K-strategies and the biomass production achieved in monocultures depended on their efficiency to convert resources to biomass. For these species, the K-strategists were less efficient resource users. This affected biomass production in polycultures, which were characteristic of underyielding. In polycultures, K-strategists sequestered more resources than the r-strategists. This likely occurred because the intermittent periods of nutrient limitation that would have occurred just prior to the next nutrient supply pulse would have favored the K-strategists, leading to overall less efficient use of resources by the polyculture. This study provides evidence that fluctuation in resource concentrations resulting from pulsed resource supplies in aquatic ecosystems can result in phytoplankton assemblages’ underyielding. [ABSTRACT FROM AUTHOR]

Dispersal in heterogeneous ecosystems, such as coastal metacommunities, is a major driver of diversity and productivity. According to theory, both species richness and spatial averaging shape a unimodal relationship of productivity with dispersal. We experimentally tested the hypothesis that disturbances acting on local patches would buffer the loss of productivity at high dispersal by preventing synchronized species oscillations. To simulate these disturbances, our experimental assemblages involved species that self-organized in isolation under three inflow pulsing frequencies, where hydraulic displacement and nutrient loading affected assemblage diversity and composition. At steady-state, the emerging isolated assemblages were connected at three levels of dispersal creating three metacommunities of different connectivity. Consistent with theory, as dispersal increased, species richness in the metacommunity declined; productivity however remained high. This occurred because the most productive species in our study (which dominated the isolated patch of intermediate inflow pulsing frequency) dominated all three patches (low, intermediate and high inflow pulsing frequencies) after dispersal commenced in our metacommunities. This experimental result provides empirical support for the mechanism of spatial averaging. Furthermore, disturbances, in the form of localized pulsed inflows, prevented population oscillation synchrony caused by homogenization. Overall, our observations suggest that localized environmental fluctuations and the identity of species seem to be more influential than dispersal in shaping the diversity and composition of phytoplankton assemblages and stabilizing productivity. [ABSTRACT FROM AUTHOR]