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  1. Abstract

    A quantitative understanding of the mesopelagic zooplankton food web is key to development of accurate carbon budgets and geochemical models in marine systems. Here we use compound specific nitrogen stable isotope analysis of amino acids to quantify the trophic structure of the microzooplankton and mesozooplankton community during summer in the subarctic northeast Pacific Ocean during the EXport Processes in the Ocean from Remote Sensing (EXPORTS) field campaign. Source amino acid values in particles and zooplankton provide strong evidence that basal resources for the mesopelagic zooplankton food web were primarily small (), suspended or slow‐sinking particles, but that surface organic matter delivered by vertically migrating zooplankton may have also been important. Comparisons of values of source and trophic amino acids provide estimates of food web length, which decrease significantly with depth and suggest that protistan microzooplankton are key components of the food web from the surface to at least 500. These results emphasize the importance of small particles as a source of carbon and nitrogen to mesopelagic communities in this region, support observations of an inverse relationship between zooplankton vertical migration and small particles as sources of carbon to deep‐sea food webs in low productivity environments, and document the role of heterotrophic protists as key trophic intermediaries in the mesopelagic zone at this location.

     
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  3. Abstract

    Particulate organic matter settling out of the euphotic zone is a major sink for atmospheric carbon dioxide and serves as a primary food source to mesopelagic food webs. Degradation of this organic matter encompasses a suite of mechanisms that attenuate flux, including heterotrophic metabolic processes of microbes and metazoans. The relative contributions of microbial and metazoan heterotrophy to flux attenuation, however, have been difficult to determine. We present results of compound specific nitrogen isotope analysis of amino acids of sinking particles from sediment traps and size‐fractionated particles from in situ filtration between the surface and 500 m at Ocean Station Papa, collected during NASA EXPORTS (EXport Processes in the Ocean from RemoTe Sensing). With increasing depth, we observe: (1) that, based on theδ15N values of threonine, fecal pellets dominate the sinking particle flux and that attenuation of downward particle flux occurs largely via disaggregation in the upper mesopelagic; (2) an increasing trophic position of particles in the upper water column, reflecting increasing heterotrophic contributions to the nitrogen pool and the loss of particles via remineralization; and (3) increasingδ15N values of source amino acids in submicron and small (1–6μm) particles, reflecting microbial particle solubilization. We further employ a Bayesian mixing model to estimate the relative proportions of fecal pellets, phytodetritus, and microbially degraded material in particles and compare these results and our interpretations of flux attenuation mechanisms to other, independent methods used during EXPORTS.

     
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  4. The fate of organic matter (OM) in the deep ocean remains enigmatic, with little understood regarding the flux and its utilization by deep food webs. We used compound-specific nitrogen stable isotope ratios of source amino acids measured in particle size classes and deep zooplankton (700–1500 m) to determine the contribution of small (0.7–53 μm) vs. large particles (> 53 μm) to their diet at four sites in the North Pacific. Our results show that small particles constitute between 9% and 98% of zooplankton diets, being the contribution higher at sites with lower flux regimes. The contribution of small particles to the diet of deep zooplankton was also higher when biomass of vertical migrators, and therefore actively transported OM, was lower. Climate-driven changes in primary production and export are expected to shift particle fluxes to smaller size classes, and thus their importance in midwater food webs may become more widespread. 
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  5. Abstract

    Zooplankton contribute a major component of the vertical flux of particulate organic matter to the ocean interior by packaging consumed food and waste into large, dense fecal pellets that sink quickly. Existing methods for quantifying the contribution of fecal pellets to particulate organic matter use either visual identification or lipid biomarkers, but these methods may exclude fecal material that is not morphologically distinct, or may include zooplankton carcasses in addition to fecal pellets. Based on results from seven pairs of wild‐caught zooplankton and their fecal pellets, we assess the ability of compound‐specific isotope analysis of amino acids (CSIA‐AA) to chemically distinguish fecal pellets as an end‐member material within particulate organic matter. Nitrogen CSIA‐AA is an improvement on previous uses of bulk stable isotope ratios, which cannot distinguish between differences in baseline isotope ratios and fractionation due to metabolic processing. We suggest that the relative trophic position of zooplankton and their fecal pellets, as calculated using CSIA‐AA, can provide a metric for estimating the dietary absorption efficiency of zooplankton. Using this metric, the zooplankton examined here had widely ranging dietary absorption efficiencies, where lower dietary absorption may equate to higher proportions of fecal packaging of undigested material. The nitrogen isotope ratios of threonine and alanine statistically distinguished the zooplankton fecal pellets from literature‐derived examples of phytoplankton, zooplankton biomass, and microbially degraded organic matter. We suggest that δ15N values of threonine and alanine could be used in mixing models to quantify the contribution of fecal pellets to particulate organic matter.

     
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  6. The goal of the EXport Processes in the Ocean from RemoTe Sensing (EXPORTS) field campaign is to develop a predictive understanding of the export, fate, and carbon cycle impacts of global ocean net primary production. To accomplish this goal, observations of export flux pathways, plankton community composition, food web processes, and optical, physical, and biogeochemical (BGC) properties are needed over a range of ecosystem states. Here we introduce the first EXPORTS field deployment to Ocean Station Papa in the Northeast Pacific Ocean during summer of 2018, providing context for other papers in this special collection. The experiment was conducted with two ships: a Process Ship, focused on ecological rates, BGC fluxes, temporal changes in food web, and BGC and optical properties, that followed an instrumented Lagrangian float; and a Survey Ship that sampled BGC and optical properties in spatial patterns around the Process Ship. An array of autonomous underwater assets provided measurements over a range of spatial and temporal scales, and partnering programs and remote sensing observations provided additional observational context. The oceanographic setting was typical of late-summer conditions at Ocean Station Papa: a shallow mixed layer, strong vertical and weak horizontal gradients in hydrographic properties, sluggish sub-inertial currents, elevated macronutrient concentrations and low phytoplankton abundances. Although nutrient concentrations were consistent with previous observations, mixed layer chlorophyll was lower than typically observed, resulting in a deeper euphotic zone. Analyses of surface layer temperature and salinity found three distinct surface water types, allowing for diagnosis of whether observed changes were spatial or temporal. The 2018 EXPORTS field deployment is among the most comprehensive biological pump studies ever conducted. A second deployment to the North Atlantic Ocean occurred in spring 2021, which will be followed by focused work on data synthesis and modeling using the entire EXPORTS data set. 
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