ALBERT

Description: Marine phytoplankton are a taxonomically and functionally diverse group of organisms that are key players in the most important biogeochemical cycles. Phytoplankton taxa show different resource utilization strategies (e.g. nutrient-uptake rates and cellular allocation) and traits. Therefore, acknowledging this diversity is crucial to understanding how elemental cycles operate, including the origin and dynamics of elemental ratios. In this paper, we focus on trait-based models as tools to study the role of phytoplankton diversity in the stoichiometric phenomenology observed in the laboratory and in the open ocean. We offer a compilation of known empirical results on stoichiometry and summarize how trait-based approaches have attempted to replicate these results. By contrasting the different ecological and evolutionary approaches available in the literature, we explore the strengths and limitations of the existing models. We thus try to identify existing gaps and challenges, and point to potential new directions that can be explored to fill these gaps. We aim to highlight the potential of including diversity explicitly in our modeling approaches, which can help us gain important knowledge about changes in local and global stoichiometric patterns.

Description: Phytoplankton community composition profoundly influences patterns of nutrient cycling and the structure of marine food webs; therefore predicting present and future phytoplankton community structure is of fundamental importance to understanding how ocean ecosystems are influenced by physical forcing and nutrient limitations. In this paper, we develop a mechanistic model of phytoplankton communities that includes multiple taxonomic groups, test the model at two contrasting sites in the modern ocean, and then use the model to predict community reorganization under different global change scenarios. The model includes three phytoplankton functional groups (diatoms, coccolithophores, and prasinophytes), five nutrients (nitrate, ammonium, phosphate, silicate and iron), light, and a generalist zooplankton grazer. Each taxonomic group was parameterized based on an extensive literature survey. The model successfully predicts the general patterns of community structure and succession in contrasting parts of the world ocean, the North Atlantic (North Atlantic Bloom Experiment, NABE) and subarctic North Pacific (ocean station Papa, OSP). In the North Atlantic, the model predicts a spring diatom bloom, followed by coccolithophore and prasinophyte blooms later in the season. The diatom bloom becomes silica-limited and the coccolithophore and prasinophyte blooms are controlled by nitrogen, grazers and by deep mixing and decreasing light availability later in the season. In the North Pacific, the model reproduces the low chlorophyll community dominated by prasinophytes and coccolithophores, with low total biomass variability and high nutrient concentrations throughout the year. Sensitivity analysis revealed that the identity of the most sensitive parameters and the range of acceptable parameters differed between the two sites. Five global change scenarios are used to drive the model and examine how community dynamics might change in the future. To estimate uncertainty in our predictions, we used a Monte Carlo sampling of the parameter space where future scenarios were run using parameter combinations that produced adequate modern day outcomes. The first scenario is based on a global climate model that indicates that increased greenhouse gas concentrations will cause a later onset and extended duration of stratification and shallower mixed layer depths. Under this scenario, the North Atlantic spring diatom bloom occurs later and is of a smaller magnitude, but the average biomass of diatoms, coccolithophores and prasinophytes will likely increase. In the subarctic North Pacific, diatoms and prasinophytes will likely increase along with total chlorophyll concentration and zooplankton. In contrast, coccolithophore densities do not change at this site. Under the second scenario of decreased deep-water phosphorus concentration, coccolithophores, total chlorophyll and zooplankton decline, as well as the magnitude of the spring diatom bloom, while the average diatom and prasinophyte abundance does not change in the North Atlantic. In contrast, a decrease in phosphorus in the North Pacific is not likely to change community composition. Similarly, doubling of nitrate in deep water does not significantly affect ecosystems at either site. Under decreased iron deposition, coccolithophores are likely to increase and other phytoplankton groups and zooplankton to decrease at both sites. An increase in iron deposition is likely to increase prasinophyte and diatom abundance and decrease coccolithophore abundance at both sites, although more dramatically at the North Pacific site. Total chlorophyll and zooplankton are also likely to increase under this scenario at both sites. Based on these scenarios, our model suggests that global environmental change will inevitably alter phytoplankton community structure and potentially impact global biogeochemical cycles.

Description: We re-examine what controls the deep ocean N:P ratio in the light of recent findings that the C:N:P stoichiometry of phytoplankton varies with growth rate, nutrient and light limitation, species and phylum, and that N2-fixation may be limited by Fe or light in large parts of the world ocean. In particular, we assess whether a systematic change in phytoplankton stoichiometry can alter the deep ocean N:P ratio. To do this we adapt recent models to include non-Redfieldian stoichiometry of phytoplankton and restriction of N2-fixers to a fraction of the surface ocean. We show that a systematic change in phytoplankton C:N:P can alter the concentrations of NO3 and PO4 in the deep ocean but cannot greatly alter their ratio, unless it also alters the N:P threshold for N2-fixation. This occurs if competitive dynamics set the N:P threshold for N2-fixation, in which case it remains close to the N:P requirement of non-fixers (rather than that of N2-fixers) and consequently so does the deep ocean N:P ratio. Then, even if N2-fixers are restricted to a fraction of the surface ocean, they reach higher densities there, minimising variations in deep ocean N:P. Theoretical limits on the N:P requirements of phytoplankton suggest that since the deep ocean became well oxygenated, its N:P ratio is unlikely to have varied by more than a factor of two in either direction. Within these bounds, evolutionary changes in phytoplankton composition, and increased phosphorus weathering due to the biological colonisation of the land surface, are predicted to have driven long-term changes in ocean composition.

Description: We re-examine what controls the deep ocean N:P ratio in the light of recent findings that the C:N:P stoichiometry of phytoplankton varies with growth rate, nutrient and light limitation, species and phylum, and that N2-fixation may be limited by Fe, temperature and/or light in large parts of the world ocean. In particular, we assess whether a systematic change in phytoplankton stoichiometry can alter the deep ocean N:P ratio. To do this we adapt recent models to include non-Redfieldian stoichiometry of phytoplankton and restriction of N2-fixers to a fraction of the surface ocean. We show that a systematic change in phytoplankton C:N:P can alter the concentrations of NO3 and PO4 in the deep ocean but cannot greatly alter their ratio, unless it also alters the N:P threshold for N2-fixation. This occurs if competitive dynamics set the N:P threshold for N2-fixation, in which case it remains close to the N:P requirement of non-fixers (rather than that of N2-fixers) and consequently so does the deep ocean N:P ratio. Then, even if N2-fixers are restricted to a fraction of the surface ocean, they reach higher densities there, minimising variations in deep ocean N:P. Theoretical limits on the N:P requirements of phytoplankton suggest that whilst the deep ocean has been well oxygenated (i.e. since the Neoproterozoic, with the possible exception of Oceanic Anoxic Events), its N:P ratio is unlikely to have varied by more than a factor of two in either direction. Within these bounds, evolutionary changes in phytoplankton composition, and increased phosphorus weathering due to the biological colonisation of the land surface, are predicted to have driven long-term changes in ocean composition.

Description: There is a long-established, remarkable correspondence between the nitrogen-to-phosphorus ratio N:P~15 of deep ocean water and the ''Redfield ratio'' of N:P~16 required by phytoplankton. Redfield and subsequent workers have suggested that it is due to N-fixing organisms being selected when N:P16. Models have shown this mechanism can work, but recent observations reveal that the real system is more complex. First, the C:N:P stoichiometry of phytoplankton varies with growth rate, nutrient and light limitation, species and phylum. Second, although N-fixation is sometimes P-limited and suppressed by N-addition, there is also evidence for Fe-limitation, light-limitation and P and Fe co-limitation of N-fixers. Here we adapt recent models to include non-Redfieldian stoichiometry of phytoplankton and limitation of N-fixers by resources other than P. We show that the deep ocean N:P is set by the N:P threshold that triggers N-fixation, and is not directly related to the N:P ratio of sinking material. However, assuming competitive dynamics set the N:P threshold for N-fixation, it will be close to the N:P requirement of non-fixers (rather than that of N-fixers) and consequently so will the deep ocean N:P ratio. Theoretical limits on the N:P requirements of phytoplankton suggest that since the deep ocean became well oxygenated, its N:P has remained within the range 7.7–32.3. Decreases in phytoplankton C:P and N:P ratios over the past ~1 Gyr are predicted to have driven a decrease in deep ocean N:P, probably via increasing PO4. Even if Fe or light limitation restrict N-fixers to a fraction of the surface ocean, they reach higher densities there, minimising variations in deep ocean N:P. Thus Redfield's mechanism is robust and we expand it to suggest that phytoplankton C:N:P and deep ocean N:P have co-evolved.

Description: Phytoplankton community composition profoundly affects patterns of nutrient cycling and the dynamics of marine food webs; therefore predicting present and future phytoplankton community structure is crucial to understand how ocean ecosystems respond to physical forcing and nutrient limitations. We develop a mechanistic model of phytoplankton communities that includes multiple taxonomic groups (diatoms, coccolithophores and prasinophytes), nutrients (nitrate, ammonium, phosphate, silicate and iron), light, and a generalist zooplankton grazer. Each taxonomic group was parameterized based on an extensive literature survey. We test the model at two contrasting sites in the modern ocean, the North Atlantic (North Atlantic Bloom Experiment, NABE) and subarctic North Pacific (ocean station Papa, OSP). The model successfully predicts general patterns of community composition and succession at both sites: In the North Atlantic, the model predicts a spring diatom bloom, followed by coccolithophore and prasinophyte blooms later in the season. In the North Pacific, the model reproduces the low chlorophyll community dominated by prasinophytes and coccolithophores, with low total biomass variability and high nutrient concentrations throughout the year. Sensitivity analysis revealed that the identity of the most sensitive parameters and the range of acceptable parameters differed between the two sites. We then use the model to predict community reorganization under different global change scenarios: a later onset and extended duration of stratification, with shallower mixed layer depths due to increased greenhouse gas concentrations; increase in deep water nitrogen; decrease in deep water phosphorus and increase or decrease in iron concentration. To estimate uncertainty in our predictions, we used a Monte Carlo sampling of the parameter space where future scenarios were run using parameter combinations that produced acceptable modern day outcomes and the robustness of the predictions was determined. Change in the onset and duration of stratification altered the timing and the magnitude of the spring diatom bloom in the North Atlantic and increased total phytoplankton and zooplankton biomass in the North Pacific. Changes in nutrient concentrations in some cases changed dominance patterns of major groups, as well as total chlorophyll and zooplankton biomass. Based on these scenarios, our model suggests that global environmental change will inevitably alter phytoplankton community structure and potentially impact global biogeochemical cycles.