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Climate warming likely drives ocean deoxygenation, but models still cannot fully explain observed declines in oxygen. One unconstrained parameter is the oxygen demand per carbon respired for complete remineralization of organic matter (i.e., the total respiration quotient, rΣ‐O2:C). Here, we tested if rΣ‐O2:C declined with depth by quantifying suspended concentrations of particulate organic carbon (POC), particulate organic nitrogen (PON), particulate organic phosphorus (POP), particulate chemical oxygen demand (PCOD), and total oxygen demand (Σ‐O2 = PCOD + 2PON) down to a depth of 1,000 m in the Sargasso Sea. The respiration quotient (r‐O2:C = PCOD:POC) and total respiration quotient (rΣ‐O2:C = Σ‐O2:POC) declined with depth in the euphotic zone, but increased vertically in the disphotic zone. C:N and rΣ‐O2:N changed with depth, but surface values were similar to values at 1,000 m. C:P, N:P, and rΣ‐O2:P mostly decreased with depth. We hypothesize that rΣ‐O2:C is linked to multiple environmental factors that change with depth, such as phytoplankton community structure and the preferential production/removal of biomolecules. Using a global model, we show that the global distribution of dissolved oxygen is equally sensitive to r‐O2:C varying between surface biomes versus vertically during remineralization. Additionally, adjusting the model's r‐O2:C with depth to match our observations resulted in less dissolved oxygen throughout the upper ocean. Most of this loss occurred in the tropical Pacific thermocline, where oxygen models underestimate deoxygenation the most. This study aims to improve our understanding of biological oxygen demand as warming‐induced deoxygenation continues. Plain Language Summary: Rising ocean temperatures are likely causing the observed decline of dissolved oxygen below the ocean surface. This continued oxygen loss threatens the survival of many marine animals. Currently, global models cannot fully explain the observed rate of oxygen loss with warming. One missing component could be variance in the respiration quotient, the ratio of oxygen consumed per organic carbon respired during microbial respiration. However, the respiration quotient has yet to be estimated at different depths by directly measuring the chemical composition of organic matter. Here, we used direct measurements to find that the respiration quotient varied with depth in the Atlantic Ocean. Therefore, the respiration quotient at the surface should not represent values at deeper depths. In addition, we used a global model to find that the respiration quotient mostly affects oxygen in the tropical Pacific Ocean, where unexplained oxygen loss is the highest. Thus, more extensive data on the respiration quotient may significantly improve global models. Key Points: The respiration quotient of particulate organic matter varied with depthElemental ratios of particulate organic matter deviated from Redfield proportions at all depthsThe increase in the respiration quotient with depth may account for some previously unexplained oxygen loss [ABSTRACT FROM AUTHOR]

Are the oceans turning into deserts? Rising temperature, increasing surface stratification, and decreasing vertical inputs of nutrients are expected to cause an expansion of warm, nutrient deplete ecosystems. Such an expansion is predicted to negatively affect a trio of key ocean biogeochemical features: phytoplankton biomass, primary productivity, and carbon export. However, phytoplankton communities are complex adaptive systems with immense diversity that could render them at least partially resilient to global changes. This can be illustrated by the biology of the Prochlorococcus "collective." Adaptations to counter stress, use of alternative nutrient sources, and frugal resource allocation can allow Prochlorococcus to buffer climate‐driven changes in nutrient availability. In contrast, cell physiology is more sensitive to temperature changes. Here, we argue that biogeochemical models need to consider the adaptive potential of diverse phytoplankton communities. However, a full understanding of phytoplankton resilience to future ocean changes is hampered by a lack of global biogeographic observations to test theories. We propose that the resilience may in fact be greater in oligotrophic waters than currently considered with implications for future predictions of phytoplankton biomass, primary productivity, and carbon export. [ABSTRACT FROM AUTHOR]

In the California Current Ecosystem, El Niño acts as a natural phenomenon that is partially representative of climate change impacts on marine bacteria at timescales relevant to microbial communities. Between 2014–2016, the North Pacific warm anomaly (a.k.a., the "blob") and an El Niño event resulted in prolonged ocean warming in the Southern California Bight (SCB). To determine whether this "marine heatwave" resulted in shifts in microbial populations, we sequenced the rpoC1 gene from the biogeochemically important picocyanobacteria Prochlorococcus and Synechococcus at 434 time points from 2009–2018 in the MICRO time series at Newport Beach, CA. Across the time series, we observed an increase in the abundance of Prochlorococcus relative to Synechococcus as well as elevated frequencies of ecotypes commonly associated with low-nutrient and high-temperature conditions. The relationships between environmental and ecotype trends appeared to operate on differing temporal scales. In contrast to ecotype trends, most microdiverse populations were static and possibly reflect local habitat conditions. The only exceptions were microdiversity from Prochlorococcous HLI and Synechococcus Clade II that shifted in response to the 2015 El Niño event. Overall, Prochlorococcus and Synechococcus populations did not return to their pre-heatwave composition by the end of this study. This research demonstrates that extended warming in the SCB can result in persistent changes in key microbial populations. [ABSTRACT FROM AUTHOR]

Marine phytoplankton stoichiometry links nutrient supply to marine carbon export. Deviations of phytoplankton stoichiometry from Redfield proportions (106C : 1P) could therefore have a significant impact on carbon cycling, and understanding which environmental factors drive these deviations may reveal new mechanisms regulating the carbon cycle. To explore the links between environmental conditions, stoichiometry, and carbon cycling, we compared four different models of phytoplankton C: P: a fixed Redfield model, a model with C: P given as a function of surface phosphorus concentration (P), a model with C: P given as a function of temperature, and a new multi-environmental model that predicts C: P as a function of light, temperature, and P. These stoichiometric models were embedded into a fivebox ocean circulation model, which resolves the three major ocean biomes (high-latitude, subtropical gyres, and tropical upwelling regions). Contrary to the expectation of a monotonic relationship between surface nutrient drawdown and carbon export, we found that lateral nutrient transport from lower C: P tropical waters to high C: P subtropical waters could cause carbon export to decrease with increased tropical nutrient utilization. It has been hypothesized that a positive feedback between temperature and pCO2,atm will play an important role in anthropogenic climate change, with changes in the biological pump playing at most a secondary role. Here we show that environmentally driven shifts in stoichiometry make the biological pump more influential, and may reverse the expected positive relationship between temperature and pCO2,atm. In the temperature-only model, changes in tropical temperature have more impact on the Δ pCO2,atm (~41 ppm) compared to subtropical temperature changes (~4.5 ppm). Our multi-environmental model predicted a decline in pCO2,atm of ~46 ppm when temperature spanned a change of 10 °C. Thus, we find that variation in marine phytoplankton stoichiometry and its environmental controlling factors can lead to nonlinear controls on pCO2,atm, suggesting the need for further studies of ocean C: P and the impact on ocean carbon cycling. [ABSTRACT FROM AUTHOR]

Marine phytoplankton stoichiometry links nutrient supply to marine carbon export. Deviations of phytoplankton stoichiometry from Redfield proportions (106C : 1P) could therefore have a significant impact on carbon cycling, and understanding which environmental factors drive these deviations may reveal new mechanisms that regulate the carbon cycle. To explore the links between environmental conditions, stoichiometry, and carbon cycling, we compared four different models for variations in phytoplankton C : P: a fixed Redfield model, a model with C : P given as a function of surface phosphorus concentration ([P]), a model with C : P given as a function of temperature, and a new multi-environmental model that predicts C : P as a function of light, temperature, and [P]. These stoichiometric models were embedded into a box model of the ocean circulation, which resolves the three major ocean biomes (high-latitude, subtropical gyres, and iron-limited tropical upwelling regions). Contrary to the expectation of a monotonic relationship between surface nutrient drawdown and carbon export, we found that lateral nutrient transport from lower C : P tropical waters to high C : P subtropical waters could cause carbon-export to decrease with increased tropical nutrient utilization. Temperature is thought to be one of the primary drivers of changes in atmospheric pCO2 (pCO2,atm) across glacial/interglacial periods, and it has been hypothesized that a positive feedback between temperature and pCO2,atm will play an important role in anthropogenic climate change, with changes in the biological pump playing at most a secondary role. Here we show that environmentally driven shifts in stoichiometry make the biological pump more influential, and may reverse the expected negative relationship between temperature and pCO2,atm. In the temperature-only model changes in tropical temperature have more impact on the Δ pCO2,atm (~ 41 ppm) compared to subtropical temperature (~ 4.5 ppm). Our multi-environmental model produced a decline in pCO2,atm of ~ 46 when temperature spanned a change of 10 °C. Thus, we find that variation in marine phytoplankton stoichiometry and its environmental controlling factor can lead to counterintuitive controls on pCO2,atm, suggesting the need for further studies of ocean C : P and the impact on ocean carbon cycling. [ABSTRACT FROM AUTHOR]

• We developed a simplified mechanistic model of Pseudo-Nitzschia to disentangle the environmental factors leading to the production of the neurotoxin domoic acid. • Silicic acid limitation shows the strongest impact on domoic acid production. • Our simplified model can be embedded in complex ecosystem models to expand Pseudo-Nitzschia and domoic acid predictability across the U.S. Western Coast. Pseudo-nitzschia species are one of the leading causes of harmful algal blooms (HABs) along the western coast of the United States. Approximately half of known Pseudo-nitzschia strains can produce domoic acid (DA), a neurotoxin that can negatively impact wildlife and fisheries and put human life at risk through amnesic shellfish poisoning. Production and accumulation of DA, a secondary metabolite synthesized during periods of low primary metabolism, is triggered by environmental stressors such as nutrient limitation. To quantify and estimate the feedbacks between DA production and environmental conditions, we designed a simple mechanistic model of Pseudo-nitzschia and domoic acid dynamics, which we validate against batch and chemostat experiments. Our results suggest that, as nutrients other than nitrogen (i.e., silicon, phosphorus, and potentially iron) become limiting, DA production increases. Under Si limitation, we found an approximate doubling in DA production relative to N limitation. Additionally, our model indicates a positive relationship between light and DA production. These results support the idea that the relationship with nutrient limitation and light is based on direct impacts on Pseudo-nitzschia biosynthesis and biomass accumulation. Because it can easily be embedded within existing coupled physical-ecosystem models, our model represents a step forward toward modeling the occurrence of Pseudo-nitzschia HABs and DA across the U.S. West Coast. [ABSTRACT FROM AUTHOR]

Although iron is the fourth most abundant element in the Earth's crust, bioavailable iron limits marine primary production in about one third of the ocean. This lack of iron availability has implications in climate change because the removal of carbon dioxide from the atmosphere by phytoplankton requires iron. Using literature values for global fish biomass estimates, and elemental composition data we estimate that fish biota store between 0.7–7×1011 g of iron. Additionally, the global fish population recycles through excretion between 0.4–1.5×1012 g of iron per year, which is of a similar magnitude as major recognized sources of iron (e.g. dust, sediments, ice sheet melting). In terms of biological impact this iron could be superior to dust inputs due to the distributed deposition and to the greater solubility of fecal pellets compared to inorganic minerals. To estimate a loss term due to anthropogenic activity the total commercial catch for 1950 to 2010 was obtained from the Food and Agriculture Organization of the United Nations. Marine catch data were separated by taxa. High and low end values for elemental composition were obtained for each taxonomic category from the literature and used to calculate iron per mass of total harvest over time. The marine commercial catch is estimated to have removed 1–6×109 g of iron in 1950, the lowest values on record. There is an annual increase to 0.7–3×1010 g in 1996, which declines to 0.6–2×1010 g in 2010. While small compared to the total iron terms in the cycle, these could have compounding effects on distribution and concentration patterns globally over time. These storage, recycling, and export terms of biotic iron are not currently included in ocean iron mass balance calculations. These data suggest that fish and anthropogenic activity should be included in global oceanic iron cycles. [ABSTRACT FROM AUTHOR]

• We analyze potential drivers of domoic acid HABs in California waters from 20 years of data. • Silicic acid deficit correlates with increased domoic acid HABs ocurrence in California. • Upwelling drives Chl-a concentrations and more intense domoic acid HABs. • Climate modes affect subregional domoic acid HABs frequencies differently. • Riverine and wastewater discharges are positively associated with domoic acid HABs in the south. Pseudo-nitzschia species with the ability to produce the neurotoxin domoic acid (DA) are the main cause of harmful algal blooms (HABs) along the U.S. West Coast, with major impacts on ecosystems, fisheries, and human health. While most Pseudo-nitzschia (PN) HAB studies to date have focused on their characteristics at specific sites, few cross-regional comparisons exist, and mechanistic understanding of large-scale HAB drivers remains incomplete. To close these gaps, we compiled a nearly 20-year time series of in situ particulate DA and environmental observations to characterize similarities and differences in PN HAB drivers along the California coast. We focus on three DA hotspots with the greatest data density: Monterey Bay, the Santa Barbara Channel, and the San Pedro Channel. Coastwise, DA outbreaks are strongly correlated with upwelling, chlorophyll-a, and silicic acid limitation relative to other nutrients. Clear differences also exist across the three regions, with contrasting responses to climate regimes across a north to south gradient. In Monterey Bay, PN HAB frequency and intensity increase under relatively nutrient-poor conditions during anomalously low upwelling intensities. In contrast, in the Santa Barbara and San Pedro Channels, PN HABs are favored under cold, nitrogen-rich conditions during more intense upwelling. These emerging patterns provide insights on ecological drivers of PN HABs that are consistent across regions and support the development of predictive capabilities for DA outbreaks along the California coast and beyond. [Display omitted] [ABSTRACT FROM AUTHOR]