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The shape of the productivity–diversity relationship (PDR) for marine phytoplankton has been suggested to be unimodal, that is, diversity peaking at intermediate levels of productivity. However, there are few observations and there has been little attempt to understand the mechanisms that would lead to such a shape for planktonic organisms. Here we use a marine ecosystem model together with the community assembly theory to explain the shape of the unimodal PDR we obtain at the global scale. The positive slope from low to intermediate productivity is due to grazer control with selective feeding, which leads to the predator-mediated coexistence of prey. The negative slope at high productivity is due to seasonal blooms of opportunist species that occur before they are regulated by grazers. The negative side is only unveiled when the temporal scale of the observation captures the transient dynamics, which are especially relevant at highly seasonal latitudes. Thus selective predation explains the positive side while transient competitive exclusion explains the negative side of the unimodal PDR curve. The phytoplankton community composition of the positive and negative sides is mostly dominated by slow-growing nutrient specialists and fast-growing nutrient opportunist species, respectively. The mechanisms that determine the relationship between diversity and productivity in marine phytoplankton remain unclear. Here, Vallina et al. show that selective predation and transient competitive exclusion determine phytoplankton community composition.

We examine the interplay between ecology and biogeochemical cycles in the context of a global three‐dimensional ocean model where self‐assembling phytoplankton communities emerge from a wide set of potentially viable cell types. We consider the complex model solutions in the light of resource competition theory. The emergent community structures and ecological regimes vary across different physical environments in the model ocean: Strongly seasonal, high‐nutrient regions are dominated by fast growing bloom specialists, while stable, low‐seasonality regions are dominated by organisms that can grow at low nutrient concentrations and are suited to oligotrophic conditions. In the latter regions, the framework of resource competition theory provides a useful qualitative and quantitative diagnostic tool with which to interpret the outcome of competition between model organisms, their regulation of the resource environment, and the sensitivity of the system to changes in key physiological characteristics of the cells.

We present a numerical model of the ocean that couples a three-stream radiative transfer component with a marine biogeochemical–ecosystem component in a dynamic three-dimensional physical framework. The radiative transfer component resolves the penetration of spectral irradiance as it is absorbed and scattered within the water column. We explicitly include the effect of several optically important water constituents (different phytoplankton functional types; detrital particles; and coloured dissolved organic matter, CDOM). The model is evaluated against in situ-observed and satellite-derived products. In particular we compare to concurrently measured biogeochemical, ecosystem, and optical data along a meridional transect of the Atlantic Ocean. The simulation captures the patterns and magnitudes of these data, and estimates surface upwelling irradiance analogous to that observed by ocean colour satellite instruments. We find that incorporating the different optically important constituents explicitly and including spectral irradiance was crucial to capture the variability in the depth of the subsurface chlorophyll a (Chl a) maximum. We conduct a series of sensitivity experiments to demonstrate, globally, the relative importance of each of the water constituents, as well as the crucial feedbacks between the light field, the relative fitness of phytoplankton types, and the biogeochemistry of the ocean. CDOM has proportionally more importance at attenuating light at short wavelengths and in more productive waters, phytoplankton absorption is relatively more important at the subsurface Chl a maximum, and water molecules have the greatest contribution when concentrations of other constituents are low, such as in the oligotrophic gyres. Scattering had less effect on attenuation, but since it is important for the amount and type of upwelling irradiance, it is crucial for setting sea surface reflectance. Strikingly, sensitivity experiments in which absorption by any of the optical constituents was increased led to a decrease in the size of the oligotrophic regions of the subtropical gyres: lateral nutrient supplies were enhanced as a result of decreasing high-latitude productivity. This new model that captures bio-optical feedbacks will be important for improving our understanding of the role of light and optical constituents on ocean biogeochemistry, especially in a changing environment. Further, resolving surface upwelling irradiance will make it easier to connect to satellite-derived products in the future.

We present a model of diverse phytoplankton and zooplankton populations embedded in a global ocean circulation model. Physiological and ecological traits of the organisms are constrained by relationships with cell size. The model qualitatively reproduces global distributions of nutrients, biomass, and primary productivity, and captures the power-law relationship between cell size and numerical density, which has realistic slopes of between −1.3 and −0.8. We use the model to explore the global structure of marine ecosystems, highlighting the importance of both nutrient and grazer controls. The model suggests that zooplankton : phytoplankton (Z : P) biomass ratios may vary from an order of 0.1 in the oligotrophic gyres to an order of 10 in upwelling and highlatitude regions. Global estimates of the strength of bottom-up and top-down controls within plankton size classes suggest that these large-scale gradients in Z : P ratios are driven by a shift from strong bottom-up, nutrient limitation in the oligotrophic gyres to the dominance of top-down, grazing controls in more productive regions.

We examine the interplay between iron supply, iron concentrations and phytoplankton communities in the Pacific Ocean. We present a theoretical framework which considers the competition for iron and nitrogen resources between phytoplankton to explain where nitrogen fixing autotrophs (diazotrophs, which require higher iron quotas, and have slower maximum growth) can co‐exist with other phytoplankton. The framework also indicates that iron and fixed nitrogen concentrations can be strongly controlled by the local phytoplankton community. Together with results from a three‐dimensional numerical model, we characterize three distinct biogeochemical provinces: 1) where iron supply is very low diazotrophs are excluded, and iron‐limited nondiazotrophic phytoplankton control the iron concentrations; 2) a transition region where nondiazotrophic phytoplankton are nitrogen limited and control the nitrogen concentrations, but the iron supply is still too low relative to nitrate to support diazotrophy; 3) where iron supplies increase further relative to the nitrogen source, diazotrophs and other phytoplankton coexist; nitrogen concentrations are controlled by nondiazotrophs and iron concentrations are controlled by diazotrophs. The boundaries of these three provinces are defined by the rate of supply of iron relative to the supply of fixed nitrogen. The numerical model and theory provide a useful tool to understand the state of, links between, and response to changes in iron supply and phytoplankton community structure that have been suggested by observations. Key Points Resource competition theory explains where diazotrophs can exist Relative iron and nitrogen supply determines community structure Fe concentrations biologically controlled over much of the low latitude Pacific Ocean

Glacial–interglacial cycles constitute large natural variations in Earth’s climate. The Mid-Pleistocene Transition (MPT) marks a shift of the dominant periodicity of these climate cycles from ∼ 40 to ∼ 100  kyr. Recently, it has been suggested that this shift resulted from a gradual increase in the internal period (or equivalently, a decrease in the natural frequency) of the system. As a result, the system would then have locked to ever higher multiples of the external forcing period. We find that the internal period is sensitive to the strength of positive feedbacks in the climate system. Using a carbon cycle model in which feedbacks between calcifier populations and ocean alkalinity mediate atmospheric CO 2 , we simulate stepwise periodicity changes similar to the MPT through such a mechanism. Due to the internal dynamics of the system, the periodicity shift occurs up to millions of years after the change in the feedback strength is imposed. This suggests that the cause for the MPT may have occurred a significant time before the observed periodicity shift.

We employ a global three‐dimensional model to simulate diverse phytoplanktonic diazotrophs (nitrogen fixers) in the oceans. In the model, the structure of the marine phytoplankton community self‐assembles from a large number of potentially viable physiologies. Amongst them, analogs of Trichodesmium, unicellular diazotrophs and diatom‐diazotroph associations (DDA) are successful and abundant. The simulated biogeography and nitrogen fixation rates of the modeled diazotrophs compare favorably with a compilation of published observations, which includes both traditional and molecular measurements of abundance and activity of marine diazotrophs. In the model, the diazotroph analogs occupy warm subtropical and tropical waters, with higher concentrations and nitrogen fixation rates in the tropical Atlantic Ocean and the Arabian Sea/Northern Indian Ocean, and lower values in the tropical and subtropical South Pacific Ocean. The three main diazotroph types typically co‐exist in the model, although Trichodesmium analogs dominate the diazotroph population in much of the North and tropical Atlantic Ocean and the Arabian Sea, while unicellular‐diazotroph analogs dominate in the South Atlantic, Pacific and Indian oceans. This pattern reflects the relative degree of nutrient limitation by iron or phosphorus. The model suggests in addition that unicellular diazotrophs could add as much new nitrogen to the global ocean as Trichodesmium.

We interpret the environmental controls on the global ocean diazotroph biogeography in the context of a three‐dimensional global model with a self‐organizing phytoplankton community. As is observed, the model's total diazotroph population is distributed over most of the oligotrophic warm subtropical and tropical waters, with the exception of the southeastern Pacific Ocean. This biogeography broadly follows temperature and light constraints which are often used in both field‐based and model studies to explain the distribution of diazotrophs. However, the model suggests that diazotroph habitat is not directly controlled by temperature and light, but is restricted to the ocean regions with low fixed nitrogen and sufficient dissolved iron and phosphate concentrations. We interpret this regulation by iron and phosphate using resource competition theory which provides an excellent qualitative and quantitative framework. Key Points Diazotroph global habitat is not directly controlled by temperature and light It is restricted to the ocean regions with low N and sufficient Fe We interpret diazotroph regulation by Fe using resource competition theory

When aerobic microbes deplete oxygen sufficiently, anaerobic metabolisms activate, driving losses of fixed nitrogen from marine oxygen minimum zones. Biogeochemical models commonly prescribe a 1–10 μM critical oxygen concentration for this transition, a range consistent with previous empirical and recent theoretical work. However, the recently developed STOX sensor has revealed large regions with much lower oxygen concentrations, at or below its 1–10 nM detection limit. Here, we develop a simplified metabolic model of an aerobic microbe to provide a theoretical interpretation of this observed depletion. We frame the threshold as O 2 * , the subsistence oxygen concentration of an aerobic microbial metabolism, at which anaerobic metabolisms can coexist with or outcompete aerobic growth. The framework predicts that this minimum oxygen concentration varies with environmental and physiological factors and is in the nanomolar range for most marine environments, consistent with observed concentrations. Using observed grazing rates to calibrate the model, we predict a minimum oxygen concentration of order 0.1–10 nM in the core of a coastal anoxic zone. We also present an argument for why anammox may be energetically favorable at a higher oxygen concentration than denitrification, as some observations suggest. The model generates hypotheses that could be tested in the field and provides a simple, mechanistic, and dynamic parameterization of oxygen depletion for biogeochemical models, without prescription of a fixed critical oxygen concentration.

Since the mid-1980s, our understanding of nutrient limitation of oceanic primary production has radically changed. Mesoscale iron addition experiments (FeAXs) have unequivocally shown that iron supply limits production in one-third of the world ocean, where surface macronutrient concentrations are perennially high. The findings of these 12 FeAXs also reveal that iron supply exerts controls on the dynamics of plankton blooms, which in turn affect the biogeochemical cycles of carbon, nitrogen, silicon, and sulfur and ultimately influence the Earth climate system. However, extrapolation of the key results of FeAXs to regional and seasonal scales in some cases is limited because of differing modes of iron supply in FeAXs and in the modern and paleo-oceans. New research directions include quantification of the coupling of oceanic iron and carbon biogeochemistry.