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Concentrations of bioavailable inorganic nitrogen (N) and phosphorus (P) are simultaneously depleted in the (sub)tropical North Atlantic Ocean, but it remains unclear if phytoplankton growth rates are N limited or N–P co‐limited. Here we present findings from three bottle‐scale experiments using a four‐by‐four matrix of low‐level N and P additions, conducted at one site in the subtropical North Atlantic Ocean. Phytoplankton responses were assessed both in terms of bulk chlorophyll a (Chl a ) concentrations and intracellular Chl a of dominant Prochlorococcus and Synechococcus groups. Two matrix experiments suggested that N was independently limiting in situ growth, with no co‐limiting role for P, while the third showed co‐limitation by both N and P in this region. This switch from N limitation to N–P co‐limitation was attributed to an episodic wet deposition event that supplied N, thereby stimulating phytoplankton growth and consuming available P. Such rapid transitions in nutrient limitation in response to environmental forcing might be common in oceanic systems with multiple depleted nutrients.

In regions of the nitrogen limited low latitude ocean, phosphate can also be depleted to levels initiating stress responses in marine microbes. Here, we associate a broad region of phosphate depletion in the subtropical North Pacific with different levels of phosphorus stress. Nutrient and aerosol addition experiments demonstrated primary nitrogen limitation of the bulk phytoplankton community, with supply of aerosols relieving this limitation. At northern sites with depleted phosphate, alkaline phosphatase activities were enhanced, indicating elevated phosphorus stress. Analysis of satellite- and model-derived aerosol loading showed that aerosol deposition was elevated in these regions. Surface rate measurements suggested that the regional enhancement in phosphate depletion was predominantly driven by elevated nitrogen fixation, likely stimulated by the coincident supply of aerosol iron. Such observations are important for predicting future biogeochemical responses in the subtropical North Pacific to changing aerosol supply.

Atmospheric aerosol deposition into the low latitude oligotrophic ocean is an important source of new nutrients for primary production. However, the resultant phytoplankton responses to aerosol deposition events, both in magnitude and changes in community composition, are poorly constrained. Here, we investigated this with 19 d of field and satellite observations for a site in the subtropical North Atlantic. During the observation period, surface dissolved aluminum concentrations alongside satellite-derived aerosol and precipitation data demonstrated the occurrence of both a dry deposition event associated with a dust storm and a wet deposition event associated with strong rainfall. The dry deposition event did not lead to any observable phytoplankton response, whereas the wet deposition event led to an approximate doubling of chlorophyll a, with Prochlorococcus becoming more dominant at the expense of Synechococcus. Bioassay experiments showed that phytoplankton were nitrogen limited, suggesting that the wet deposition event likely provided substantial aerosol-derived nitrogen, thereby alleviating the prevalent nutrient limitation and leading to the rapid observed phytoplankton response. These findings highlight the important role of wet deposition in driving rapid responses in both ocean productivity and phytoplankton community composition.

Subtropical gyres cover 26%-29% of the world's surface ocean and are conventionally regarded as ocean deserts due to their permanent stratification, depleted surface nutrients, and low biological productivity. Despite tremendous advances over the past three decades, particularly through the Hawaii Ocean Time-series and the Bermuda Atlantic Time-series Study, which have revolutionized our understanding of the biogeochemistry in oligotrophic marine ecosystems, the gyres remain understudied. We review current understanding of upper ocean biogeochemistry in the North Pacific Subtropical Gyre, considering other subtropical gyres for comparison. We focus our synthesis on spatial variability, which shows larger than expected dynamic ranges of properties such as nutrient concentrations, rates of N-2 fixation, and biological production. This review provides new insights into how nutrient sources drive community structure and export in upper subtropical gyres. We examine the euphotic zone (EZ) in subtropical gyres as a two-layered vertically structured system: a nutrient-depleted layer above the top of the nutricline in the well-lit upper ocean and a nutrient-replete layer below in the dimly lit waters. These layers vary in nutrient supply and stoichiometries and physical forcing, promoting differences in community structure and food webs, with direct impacts on the magnitude and composition of export production. We evaluate long-term variations in key biogeochemical parameters in both of these EZ layers. Finally, we identify major knowledge gaps and research challenges in these vast and unique systems that offer opportunities for future studies. Key Points Subtropical gyres display larger spatiotemporal dynamics in biogeochemical properties than previously considered An improved two-layer framework is proposed for the study of nutrient-driven and biologically mediated carbon export in the euphotic zone Future research will benefit from high-resolution samplings, improved sen

Mesoscale eddies are common in the subtropical Northwest Pacific, however, relatively little is known about their spatial variability and temporal evolution, and how these impact upper ocean biogeochemistry. Here we investigate these using observations of a cyclonic eddy carried out along four sequential transects. Consistent with previous observations of cyclonic eddies, the eddy core had doming isopycnals, bringing elevated nutrient waters nearer to the surface. However, we also found that the upper layer of the eddy above the nutricline had significantly lower phosphate concentrations within its core relative to its edge. We attributed this to elevated N 2 fixation within the eddy core, which was likely driven by enhanced subsurface iron supply, ultimately resulting in increased phosphate consumption. Eddy‐enhanced N 2 fixation was additionally supported by the elevation of nitrate + nitrite to phosphate ratios below the euphotic zone. Moreover, we observed that while the upward displacement of isopycnals within the eddy core led to an increase in phytoplankton biomass in the lower euphotic zone, there was no significant increase in total phytoplankton biomass across the entire euphotic zone. Cyclonic eddies in the subtropical North Pacific are projected to be becoming more frequent, implying that such dynamics could become increasingly important for regulating nutrient biogeochemistry and ultimately productivity of the region. Key Points Lower phosphate concentrations were observed above the nutricline within the eddy core in comparison to the edge Enhanced N2 fixation within the eddy core is proposed to have driven increased phosphate consumption No substantial total phytoplankton biomass increase was found within the eddy core

Nutrients limiting phytoplankton growth in the ocean are a critical control on ocean productivity and can underpin predicted responses to climate change. The extensive western subtropical North Pacific is assumed to be under strong nitrogen limitation, but this is not well supported by experimental evidence. Here, we report the results of 14 factorial nitrogen–phosphorus–iron addition experiments through the Philippine Sea, which demonstrate a gradient from nitrogen limitation in the north to nitrogen–iron co-limitation in the south. While nitrogen limited sites responded weakly to nutrient supply, co-limited sites bloomed with up to ~60-fold increases in chlorophyll a biomass that was dominated by initially undetectable diatoms. The transition in limiting nutrients and phytoplankton growth capacity was driven by a gradient in deep water nutrient supply, which was undetectable in surface concentration fields. We hypothesize that this large-scale phytoplankton response gradient is both climate sensitive and potentially important for regulating the distribution of predatory fish.

Nutrients limiting phytoplankton growth in the ocean are a critical control on ocean productivity and can underpin predicted responses to climate change. The extensive western subtropical North Pacific is assumed to be under strong nitrogen limitation, but this is not well supported by experimental evidence. Here, we report the results of 14 factorial nitrogen–phosphorus–iron addition experiments through the Philippine Sea, which demonstrate a gradient from nitrogen limitation in the north to nitrogen–iron co-limitation in the south. While nitrogen limited sites responded weakly to nutrient supply, co-limited sites bloomed with up to ~60-fold increases in chlorophyll a biomass that was dominated by initially undetectable diatoms. The transition in limiting nutrients and phytoplankton growth capacity was driven by a gradient in deep water nutrient supply, which was undetectable in surface concentration fields. We hypothesize that this large-scale phytoplankton response gradient is both climate sensitive and potentially important for regulating the distribution of predatory fish.

Globally expanding hypoxia in estuaries and coastal oceans has largely been attributed to the elevated river nutrient inputs, whereas the role of river-delivered terrestrial organic matter (OMterr) in hypoxia formation has been less investigated. This study uses a coupled physical-biogeochemical model and observations to investigate how OMterr directly (via remineralization) and indirectly (via the nutrients released from OMterr remineralization) promotes hypoxia development in the coastal transition zone off the Pearl River Estuary. Results show that direct contribution of OMterr remineralization to total oxygen consumption by terrestrial and marine organic matter negatively correlates with salinity, decreasing from over 60% in the upper estuary to nearly 0% in the far reaches of the river plume, and is higher in the upstream (average 30%) than the downstream region (average 18%). Nevertheless, the nutrients released from OMterr remineralization greatly sustain an indirect contribution to oxygen depletion and hypoxia formation downstream. The increasing relative importance of indirect over the direct effect of OMterr to hypoxia along the plume path is a combined result of the wind-driven eastward shelf current and the OMterr-released nutrients being advected farther downstream than the sinking OMterr. This highlights that without including the indirect effect of OMterr may underestimate the role of OMterr in hypoxia formation in aquatic systems. Examinations of the hypoxia response to varying riverine loads further suggest that reducing the nutrient and OMterr loads is required for hypoxia mitigation in the upstream region while reducing the nutrient load alone is more effective in mitigating hypoxia in the downstream. © 2020 Association for the Sciences of Limnology and Oceanography