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The Critical Depth Hypothesis formalized by Sverdrup in 1953 posits that vernal phytoplankton blooms occur when surface mixing shoals to a depth shallower than a critical depth horizon defining the point where phytoplankton growth exceeds losses. This hypothesis has since served as a cornerstone in plankton ecology and reflects the very common assumption that blooms are caused by enhanced growth rates in response to improved light, temperature, and stratification conditions, not simply correlated with them. Here, a nine-year satellite record of phytoplankton biomass in the subarctic Atlantic is used to reevaluate seasonal plankton dynamics. Results show that (1) bloom initiation occurs in the winter when mixed layer depths are maximum, not in the spring, (2) coupling between phytoplankton growth (μ) and losses increases during spring stratification, rather than decreases, (3) maxima in net population growth rates ( r ) are as likely to occur in midwinter as in spring, and (4) r is generally inversely related to μ. These results are incompatible with the Critical Depth Hypothesis as a functional framework for understanding bloom dynamics. In its place, a \"Dilution-Recoupling Hypothesis\" is described that focuses on the balance between phytoplankton growth and grazing, and the seasonally varying physical processes influencing this balance. This revised view derives from fundamental concepts applied during field dilution experiments, builds upon earlier modeling results, and is compatible with observed phytoplankton blooms in the absence of spring mixed layer shoaling.

Over the last ten years, satellite and geographically constrained in situ observations largely focused on the northern hemisphere have suggested that annual phytoplankton biomass cycles cannot be fully understood from environmental properties controlling phytoplankton division rates (e.g., nutrients and light), as they omit the role of ecological and environmental loss processes (e.g., grazing, viruses, sinking). Here, we use multi-year observations from a very large array of robotic drifting floats in the Southern Ocean to determine key factors governing phytoplankton biomass dynamics over the annual cycle. Our analysis reveals seasonal phytoplankton accumulation (‘blooming’) events occurring during periods of declining modeled division rates, an observation that highlights the importance of loss processes in dictating the evolution of the seasonal cycle in biomass. In the open Southern Ocean, the spring bloom magnitude is found to be greatest in areas with high dissolved iron concentrations, consistent with iron being a well-established primary limiting nutrient in this region. Under ice observations show that biomass starts increasing in early winter, well before sea ice begins to retreat. The average theoretical sensitivity of the Southern Ocean to potential changes in seasonal nutrient and light availability suggests that a 10% change in phytoplankton division rate may be associated with a 50% reduction in mean bloom magnitude and annual primary productivity, assuming simple changes in the seasonal magnitude of phytoplankton division rates. Overall, our results highlight the importance of quantifying and accounting for both division and loss processes when modeling future changes in phytoplankton biomass cycles. Phytoplankton are biogeochemically important but the drivers of their seasonal cycles in the Southern Ocean are poorly resolved. Here the authors use seven years of ARGO float data to measure bloom initiation, decline and termination throughout the Southern Ocean, finding that bloom dynamics are especially sensitive to the coupling between cell division rates and loss processes.

The Plankton, Aerosol, Cloud, ocean Ecosystem (PACE) mission represents the National Aeronautics and Space Administration’s (NASA) next investment in satellite ocean color and the study of Earth’s ocean–atmosphere system, enabling new insights into oceanographic and atmospheric responses to Earth’s changing climate. PACE objectives include extending systematic cloud, aerosol, and ocean biological and biogeochemical data records, making essential ocean color measurements to further understand marine carbon cycles, food-web processes, and ecosystem responses to a changing climate, and improving knowledge of how aerosols influence ocean ecosystems and, conversely, how ocean ecosystems and photochemical processes affect the atmosphere. PACE objectives also encompass management of fisheries, large freshwater bodies, and air and water quality and reducing uncertainties in climate and radiative forcing models of the Earth system. PACE observations will provide information on radiative properties of land surfaces and characterization of the vegetation and soils that dominate their ref lectance. The primary PACE instrument is a spectrometer that spans the ultraviolet to shortwave-infrared wavelengths, with a ground sample distance of 1 km at nadir. This payload is complemented by two multiangle polarimeters with spectral ranges that span the visible to near-infrared region. Scheduled for launch in late 2022 to early 2023, the PACE observatory will enable significant advances in the study of Earth’s biogeochemistry, carbon cycle, clouds, hydrosols, and aerosols in the ocean–atmosphere–land system. Here, we present an overview of the PACE mission, including its developmental history, science objectives, instrument payload, observatory characteristics, and data products.

Diatoms are the most recent major algal lineage added to the geological record, appearing more than 200 million years ago. They are stramenopile protists resulting from a secondary endosymbiotic event that yielded the only photosynthetic protistan lineage expressing external siliceous cell wall structures called frustules. Many diatoms also have large internal vacuoles, and a common assumption in the literature is that success of the diatoms is largely attributable to these two morphological inventions: the frustule for defense and vacuole for luxury nutrient uptake. Here, we revisit the evolution of these inventions, propose sequential steps in frustule development, replace luxury nutrient uptake with predator defense and buoyancy control as the driver of vacuole expansion, and suggest that perhaps the greatest significance of the frustule for diatom evolution is the secondary consequence of enhancing sexual reproduction. In this synthesis, we emphasize a distinction between the \"general\" success of diatoms and the success of \"bloom-forming\" species, as the physiological and morphological drivers of these successes differ. Importantly, the bloom-forming species are responsible for the major role of diatoms in aquatic biogeochemical cycles. The bloom-forming habit we ascribe to specific physiological attributes that, at their core, revolve around influencing the balance between diatom growth and losses to predators. We propose that these physiological adaptations are linked to size-dependent maximum division rates in bloom-forming diatoms, because of size scaling of predator–prey interactions. The existence of these bloom-forming species yields an apparent allometric relationship that has previously been interpreted in terms of nutrient acquisition. Our analysis yields insights into species successions during blooms, considers the fundamental benefit of blooming (and subsequent sinking) from a reproductive standpoint, and provides some reinterpretation of diatoms success over geologic time and in the modern ocean.

Global satellite observations document expansions of the low-chlorophyll central ocean gyres and an overall inverse relationship between anomalies in sea surface temperature and phytoplankton chlorophyll concentrations. These findings can provide an invaluable glimpse into potential future ocean changes, but only if the story they tell is accurately interpreted. Chlorophyll is not simply a measure of phytoplankton biomass, but also registers changes in intracellular pigmentation arising from light-driven (photoacclimation) and nutrient-driven physiological responses. Here, we show that the photoacclimation response is an important component of temporal chlorophyll variability across the global ocean. This attribution implies that contemporary relationships between chlorophyll changes and ocean warming are not indicative of proportional changes in productivity, as light-driven decreases in chlorophyll can be associated with constant or even increased photosynthesis. Extension of these results to future change, however, requires further evaluation of how the multifaceted stressors of a warmer, higher-CO 2 world will impact plankton communities. Satellite measurements of chlorophyll are used to infer phytoplankton biomass changes and the relationship to sea surface temperature. This study shows that chlorophyll changes can be light-driven so the temperature–biomass relationship may not hold in the future.

Polar plankton communities are among the most productive, seasonally dynamic and rapidly changing ecosystems in the global ocean. However, persistent cloud cover, periods of constant night and prevailing low solar elevations in polar regions severely limit traditional passive satellite ocean colour measurements and leave vast areas unobserved for many consecutive months each year. Consequently, our understanding of the annual cycles of polar plankton and their interannual variations is incomplete. Here we use space-borne lidar observations to overcome the limitations of historical passive sensors and report a decade of uninterrupted polar phytoplankton biomass cycles. We find that polar phytoplankton dynamics are categorized by ‘boom–bust’ cycles resulting from slight imbalances in plankton predator–prey equilibria. The observed seasonal-to-interannual variations in biomass are predicted by mathematically modelled rates of change in phytoplankton division. Furthermore, we find that changes in ice cover dominated variability in Antarctic phytoplankton stocks over the past decade, whereas ecological processes were the predominant drivers of change in the Arctic. We conclude that subtle and environmentally driven imbalances in polar food webs underlie annual phytoplankton boom–bust cycles, which vary interannually at each pole. Phytoplankton productivity is high in the polar oceans. Lidar observations from 2006–2015 reveal that phytoplankton biomass was characterized by annual cycles influenced by sea-ice extent in the Antarctic and ecological processes in the Arctic.

Phytoplankton growth and productivity relies on light, multiple nutrients and temperature. These combined factors constitute the 'integrated growth environment'. Since their emergence in the Archaean ocean, phytoplankton have experienced dramatic shifts in their integrated growth environment and, in response, evolved diverse mechanisms to maximize growth by optimizing the allocation of photosynthetic resources (ATP and NADPH) among all cellular processes. Consequently, co-limitation has become an omnipresent condition in the global ocean. Here we focus on evolved phytoplankton populations of the contemporary ocean and the varied energetic pathways they employ to solve the optimization problem of resource supply and demand. Central to this discussion is the allocation of reductant formed through photosynthesis, which we propose has the following three primary fates: carbon fixation, direct use and ATP generation. Investment of reductant among these three sinks is tied to cell cycle events, differentially influenced by specific forms of nutrient stress, and a strong determinant of relationships between light-harvesting (pigment), photosynthetic electron transport and carbon fixation. Global implications of optimization are illustrated by deconvolving trends in the 10-year global satellite chlorophyll record into contributions from biomass and physiology, thereby providing a unique perspective on the dynamic nature of surface phytoplankton populations and their link to climate.

Under most natural marine conditions, phytoplankton cells suspended in the water column are too distantly spaced for direct competition for resources (i.e., overlapping cell boundary layers) to be a routine occurrence. Accordingly, resource-based competitive exclusion should be rare. In contrast, contemporary ecosystem models typically predict an exclusion of larger phytoplankton size classes under low-nutrient conditions, an outcome interpreted as reflecting the competitive advantage of small cells having much higher nutrient ‘affinities’ than larger cells. Here, we develop mechanistically-focused expressions for steady-state, nutrient-limited phytoplankton growth that are consistent with the discrete, distantly-spaced cells of natural populations. These expressions, when encompassed in a phytoplankton-zooplankton model, yield sustained diversity across all size classes over the full range in nutrient concentrations observed in the ocean. In other words, our model does not exhibit resource-based competitive exclusion between size classes previously associated with size-dependent differences in nutrient ‘affinities’.

Four North Atlantic Aerosol and Marine Ecosystems Study (NAAMES) field campaigns from winter 2015 through spring 2018 sampled an extensive set of oceanographic and atmospheric parameters during the annual phytoplankton bloom cycle. This unique dataset provides four seasons of open-ocean observations of wind speed, sea surface temperature (SST), seawater particle attenuation at 660 nm (c p,660, a measure of ocean particulate organic carbon), bacterial production rates, and sea-spray aerosol size distributions and number concentrations (N SSA). The NAAMES measurements show moderate to strong correlations (0.56 < R < 0.70) between N SSA and local wind speeds in the marine boundary layer on hourly timescales, but this relationship weakens in the campaign averages that represent each season, in part because of the reduction in range of wind speed by multiday averaging. N SSA correlates weakly with seawater cp,660 (R = 0.36, P << 0.01), but the correlation with cp,660, is improved (R = 0.51, P < 0.05) for periods of low wind speeds. In addition, NAAMES measurements provide observational dependence of SSA mode diameter (d m) on SST, with d m increasing to larger sizes at higher SST (R = 0.60, P << 0.01) on hourly timescales. These results imply that climate models using bimodal SSA parameterizations to wind speed rather than a single SSA mode that varies with SST may overestimate SSA number concentrations (hence cloud condensation nuclei) by a factor of 4 to 7 and may underestimate SSA scattering (hence direct radiative effects) by a factor of 2 to 5, in addition to overpredicting variability in SSA scattering from wind speed by a factor of 5.

Ocean ecosystems play a critical role in the Earth’s carbon cycle and the quantification of their impacts for both present conditions and for predictions into the future remains one of the greatest challenges in oceanography. The goal of the EXport Processes in the Ocean from Remote Sensing (EXPORTS) Science Plan is to develop a predictive understanding of the export and fate of global ocean net primary production (NPP) and its implications for present and future climates. The achievement of this goal requires a quantification of the mechanisms that control the export of carbon from the euphotic zone as well as its fate in the underlying “twilight zone” where some fraction of exported carbon will be sequestered in the ocean’s interior on time scales of months to millennia. Here we present a measurement / synthesis / modeling framework aimed at quantifying the fates of upper ocean NPP and its impacts on the global carbon cycle based upon the EXPORTS Science Plan. The proposed approach will diagnose relationships among the ecological, biogeochemical and physical oceanographic processes that control carbon cycling across a range of ecosystem and carbon cycling states leading to advances in satellite diagnostic and numerical prognostic models. To collect these data, a combination of ship and robotic field sampling, satellite remote sensing and numerical modeling is proposed which enables the sampling of the many pathways of NPP export and fates. This coordinated, process-oriented approach has the potential to foster new insights on ocean carbon cycling that maximizes its societal relevance through the achievement of research goals of many international research agencies and will be a key step towards our understanding of the Earth as an integrated system.