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The biological carbon pump is responsible for the transport of ∼5-20 Pg C yr-1 from the surface into the deep ocean but its variability is poorly understood due to an incomplete mechanistic understanding of the complex underlying planktonic processes. In fact, algorithms designed to estimate carbon export from satellite products incorporate fundamentally different assumptions about the relationships between plankton biomass, productivity, and export efficiency. To test the alternate formulations of export efficiency in remote-sensing algorithms formulated by Dunne et al. (2005), Laws et al. (2011), Henson et al. (2011), and Siegel et al. (2014), we have compiled in situ measurements (temperature, chlorophyll, primary production, phytoplankton biomass and size structure, grazing rates, net chlorophyll change, and carbon export) made during Lagrangian process studies on seven cruises in the California Current Ecosystem and Costa Rica Dome. A food-web based approach formulated by Siegel et al. (2014) performs as well or better than other empirical formulations, while simultaneously providing reasonable estimates of protozoan and mesozooplankton grazing rates. By tuning the Siegel et al. (2014) algorithm to match in situ grazing rates more accurately, we also obtain better in situ carbon export measurements. Adequate representations of food-web relationships and grazing dynamics are therefore crucial to improving the accuracy of export predictions made from satellite-derived products. Nevertheless, considerable unexplained variance in export remains and must be explored before we can reliably use remote sensing products to assess the impact of climate change on biologically mediated carbon sequestration.

Mesozooplankton can directly impact global biogeochemical cycles by repackaging particulate organic carbon (POC) into dense, rapidly sinking fecal pellets and by undertaking vertical migrations that transport carbon and nutrients to depth. We assessed these contributions of mesozooplankton to vertical flux in the California Current Ecosystem, a productive but spatiotemporally variable coastal upwelling system, during cruises in April 2007 and October 2008. Sediment traps and Thorium-234 (234Th) disequilibrium measurements were used to assess the total passive flux of sinking POC, while pigment analyses and microscopic enumeration of sediment trap samples provided estimates of total fecal carbon transport. Identification of mesozooplankton in paired day-night, vertically stratified plankton tows allowed calculation of the active transport of carbon by the dominant taxa of vertically migrating mesozooplankton (particularly copepods and euphausiids). Across the range of 9 ecosystem conditions encountered on the cruises, recognizable fecal pellet mass flux varied from 3.5 to 135 mg C m-2 d-1 (3 to 94% of total passive flux) at the 100 m depth horizon. The active transport of carbon by migratory mesozooplankton taxa contributed an additional 2.4 to 47.1 mg C m-2 d-1 (1.9 to 40.5% of total passive flux). Inter-cruise comparisons suggest that fecal pellets contributed a higher portion of passive export during the productive spring cruise, when fecal material may have been responsible for close to 100% of sinking material. During the fall cruise, a gradient was observed with carbon export in productive water parcels driven by a large contribution of fecal pellets. In the less productive regions, fall vertical fluxes contained a higher proportion of marine snow and unidentifiable particles. © 2013 The authors.

Despite the increasing use of linear inverse modeling techniques to elucidate fluxes in undersampled marine ecosystems, the accuracy with which they estimate food web flows has not been resolved. New Markov Chain Monte Carlo (MCMC) solution methods have also called into question the biases of the commonly used L2 minimum norm (L2MN) solution technique. Here, we test the abilities of MCMC and L2MN methods to recover field-measured ecosystem rates that are sequentially excluded from the model input. For data, we use experimental measurements from process cruises of the California Current Ecosystem (CCE-LTER) Program that include rate estimates of phytoplankton and bacterial production, micro- and mesozooplankton grazing, and carbon export from eight study sites varying from rich coastal upwelling to offshore oligotrophic conditions. Both the MCMC and L2MN methods predicted well-constrained rates of protozoan and mesozooplankton grazing with reasonable accuracy, but the MCMC method overestimated primary production. The MCMC method more accurately predicted the poorly constrained rate of vertical carbon export than the L2MN method, which consistently overestimated export. Results involving DOC and bacterial production were equivocal. Overall, when primary production is provided as model input, the MCMC method gives a robust depiction of ecosystem processes. Uncertainty in inverse ecosystem models is large and arises primarily from solution under-determinacy. We thus suggest that experimental programs focusing on food web fluxes expand the range of experimental measurements to include the nature and fate of detrital pools, which play large roles in the model. © 2011 Elsevier B.V.

The origin of methylmercury in pelagic fish remains unclear, with many unanswered questions regarding the production and degradation of this neurotoxin in the water column. We used mercury (Hg) stable isotope ratios of marine particles and biota to elucidate the cycling of methylmercury prior to incorporation into the marine food web. The Hg isotopic composition of particles, zooplankton, and fish reveals preferential methylation of Hg within small (< 53 µm) marine particles in the upper 400 m of the North Pacific Ocean. Mass-dependent Hg isotope ratios (δ 202 Hg) recorded in small particles overlap with previously estimated δ 202 Hg values for methylmercury sources to Pacific and Atlantic Ocean food webs. Particulate compound specific isotope analysis of amino acids (CSIA-AA) yield δ 15 N values that indicate more-significant microbial decomposition in small particles compared to larger particles. CSIA-AA and Hg isotope data also suggest that large particles (> 53 µm) collected in the equatorial ocean are distinct from small particles and resemble fecal pellets. Additional evidence for Hg methylation within small particles is provided by a statistical mixing model of even mass-independent (Δ 200 Hg and Δ 204 Hg) isotope values, which demonstrates that Hg within near-surface marine organisms (0-150 m) originates from a combination of rainfall and marine particles. In contrast, in meso- and upper bathypelagic organisms (200-1,400 m), the majority of Hg originates from marine particles with little input from wet deposition. The occurrence of methylation within marine particles is supported further by a correlation between Δ 200 Hg and Δ 199 Hg values, demonstrating greater overlap in the Hg isotopic composition of marine organisms with marine particles than with total gaseous Hg or wet deposition.

Increasing inputs of organic matter (OM) are driving declining dissolved oxygen (DO) concentrations in coastal ecosystems worldwide. The quantity, source, and composition of OM transported to coastal ecosystems via stormwater runoff have been altered by land use changes associated with urbanization and subsequent hydrologic flows that accompany urban stormwater management. To elucidate the role of stormwater in the decline of coastal DO, rain event sampling of biochemical oxygen demand (BOD) in samples collected from the outfall of stormwater ponds and wetlands, as well as samples of largely untreated runoff carried by stormwater ditches, was conducted across a range of urban and suburban development densities. Sampling also included measurements of particulate and dissolved carbon and nitrogen, carbon and nitrogen stable isotopes, and chlorophyll- a . Results suggest stormwater may be a significant source of labile OM to receiving waters, especially during the first flush of runoff, even though BOD concentrations vary both among and within sites in response to rain events. BOD variability was best predicted by particulate OM (POM) and chlorophyll- a , rather than the larger pool of dissolved OM. These findings demonstrate the importance of managing episodic stormwater discharge, especially POM, from urbanized areas to mitigate DO impairment in larger downstream systems.

The short-lived radionuclide 234 Th is widely used to study particle scavenging and transport from the upper ocean to deeper waters. This manuscript optimizes, reviews and validates the collection, processing and analyses of total 234 Th in seawater and suggests areas of further improvements. The standard 234 Th protocol method consists of scavenging 234 Th from seawater via a MnO 2 precipitate, beta counting, and using chemical recoveries determined by adding 230 Th. The revised protocol decreases sample volumes to 2 L, shortens wait times between steps, and simplifies the chemical recovery process, expanding the ability to more rapidly and safely apply the 234 Th method., Competing Interests: Conflict of interestThe authors have no relevant financial or non-financial interests to disclose., (© The Author(s) 2021.)

Marine protists are integral to the structure and function of pelagic ecosystems and marine carbon cycling, with rhizarian biomass alone accounting for more than half of all mesozooplankton in the oligotrophic oceans. Yet, understanding how their environment shapes diversity within species and across taxa is limited by a paucity of observations of heritability and life history. Here, we present observations of asexual reproduction, morphologic plasticity, and ontogeny in the planktic foraminifer Neogloboquadrina pachyderma in laboratory culture. Our results demonstrate that planktic foraminifera reproduce both sexually and asexually and demonstrate extensive phenotypic plasticity in response to nonheritable factors. These two processes fundamentally explain the rapid spatial and temporal response of even imperceptibly low populations of planktic foraminifera to optimal conditions and the diversity and ubiquity of these species across the range of environmental conditions that occur in the ocean., (Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of Science. No claim to original U.S. Government Works. Distributed under a Creative Commons Attribution NonCommercial License 4.0 (CC BY-NC).)

The CARIACO (Carbon Retention in a Colored Ocean) Ocean Time-Series Program station, located at 10.50°N, 64.66°W, observed biogeochemical and ecological processes in the Cariaco Basin of the southwestern Caribbean Sea from November 1995 to January 2017. The program completed 232 monthly core cruises, 40 sediment trap deployment cruises, and 40 microbiogeochemical process cruises. Upwelling along the southern Caribbean Sea occurs from approximately November to August. High biological productivity (320-628 g C m -2 y -1 ) leads to large vertical fluxes of particulate organic matter, but only approximately 9-10 g C m -2 y -1 fall to the bottom sediments (∼1-3% of primary production). A diverse community of heterotrophic and chemoautotrophic microorganisms, viruses, and protozoa thrives within the oxic-anoxic interface. A decrease in upwelling intensity from approximately 2003 to 2013 and the simultaneous overfishing of sardines in the region led to diminished phytoplankton bloom intensities, increased phytoplankton diversity, and increased zooplankton densities. The deepest waters of the Cariaco Basin exhibited long-term positive trends in temperature, salinity, hydrogen sulfide, ammonia, phosphate, methane, and silica. Earthquakes and coastal flooding also resulted in the delivery of sediment to the seafloor. The program's legacy includes climate-quality data from suboxic and anoxic habitats and lasting relationships between international researchers.

Water column bulk Pseudo-nitzschia abundance and the dissolved and particulate domoic acid (DA) concentrations were measured in the Santa Barbara Basin (SBB), California from 2009⁻2013 and compared to bulk Pseudo-nitzschia cell abundance and DA concentrations and fluxes in sediment traps moored at 147 m and 509 m. Pseudo-nitzschia abundance throughout the study period was spatially and temporally heterogeneous (<200 cells L -1 to 3.8 × 10⁶ cells L -1 , avg. 2 × 10⁵ ± 5 × 10⁵ cells L -1 ) and did not correspond with upwelling conditions or the total DA (tDA) concentration, which was also spatially and temporally diverse (<1.3 ng L -1 to 2.2 × 10⁵ ng L -1 , avg. 7.8 × 10³ ± 2.2 × 10⁴ ng L -1 ). We hypothesize that the toxicity is likely driven in part by specific Pseudo-nitzschia species as well as bloom stage. Dissolved (dDA) and particulate (pDA) DA were significantly and positively correlated ( p < 0.01) and both comprised major components of the total DA pool (pDA = 57 ± 35%, and dDA = 42 ± 35%) with substantial water column concentrations (>1000 cells L -1 and tDA = 200 ng L -1 ) measured as deep as 150 m. Our results highlight that dDA should not be ignored when examining bloom toxicity. Although water column abundance and pDA concentrations were poorly correlated with sediment trap Pseudo-nitzschia abundance and fluxes, DA toxicity is likely associated with senescent blooms that rapidly sink to the seafloor, adding another potential source of DA to benthic organisms.

The Costa Rica Dome is a picophytoplankton-dominated, open-ocean upwelling system in the Eastern Tropical Pacific that overlies the ocean's largest oxygen minimum zone. To investigate the efficiency of the biological pump in this unique area, we used shallow (90-150 m) drifting sediment traps and 234 Th: 238 U deficiency measurements to determine export fluxes of carbon, nitrogen and phosphorus in sinking particles. Simultaneous measurements of nitrate uptake and shallow water nitrification allowed us to assess the equilibrium balance of new and export production over a monthly timescale. While f -ratios (new:total production) were reasonably high (0.36 ± 0.12, mean ± standard deviation), export efficiencies were considerably lower. Sediment traps suggested e- ratios (export/ 14 C-primary production) at 90-100 m ranging from 0.053 to 0.067. ThE-ratios ( 234 Th disequilibrium-derived export) ranged from 0.038 to 0.088. C:N and N:P stoichiometries of sinking material were both greater than canonical (Redfield) ratios or measured C:N of suspended particulates, and they increased with depth, suggesting that both nitrogen and phosphorus were preferentially remineralized from sinking particles. Our results are consistent with an ecosystem in which mesozooplankton play a major role in energy transfer to higher trophic levels but are relatively inefficient in mediating vertical carbon flux to depth, leading to an imbalance between new production and sinking flux.

Near-surface waters ranging from the Pacific subarctic (58°N) to the Southern Ocean (66°S) contain the neurotoxin domoic acid (DA), associated with the diatom Pseudo-nitzschia. Of the 35 stations sampled, including ones from historic iron fertilization experiments (SOFeX, IronEx II), we found Pseudo-nitzschia at 34 stations and DA measurable at 14 of the 26 stations analyzed for DA. Toxin ranged from 0.3 fg·cell(-1) to 2 pg·cell(-1), comparable with levels found in similar-sized cells from coastal waters. In the western subarctic, descent of intact Pseudo-nitzschia likely delivered significant amounts of toxin (up to 4 μg of DA·m(-2)·d(-1)) to underlying mesopelagic waters (150-500 m). By reexamining phytoplankton samples from SOFeX and IronEx II, we found substantial amounts of DA associated with Pseudo-nitzschia. Indeed, at SOFeX in the Antarctic Pacific, DA reached 220 ng·L(-1), levels at which animal mortalities have occurred on continental shelves. Iron ocean fertilization also occurs naturally and may have promoted blooms of these ubiquitous algae over previous glacial cycles during deposition of iron-rich aerosols. Thus, the neurotoxin DA occurs both in coastal and oceanic waters, and its concentration, associated with changes in Pseudo-nitzschia abundance, likely varies naturally with climate cycles, as well as with artificial iron fertilization. Given that iron fertilization in iron-depleted regions of the sea has been proposed to enhance phytoplankton growth and, thereby, both reduce atmospheric CO(2) and moderate ocean acidification in surface waters, consideration of the potentially serious ecosystem impacts associated with DA is prudent.

Mesoscale eddies may play a critical role in ocean biogeochemistry by increasing nutrient supply, primary production, and efficiency of the biological pump, that is, the ratio of carbon export to primary production in otherwise nutrient-deficient waters. We examined a diatom bloom within a cold-core cyclonic eddy off Hawaii. Eddy primary production, community biomass, and size composition were markedly enhanced but had little effect on the carbon export ratio. Instead, the system functioned as a selective silica pump. Strong trophic coupling and inefficient organic export may be general characteristics of community perturbation responses in the warm waters of the Pacific Ocean.

We have developed a new method for the collection, purification, and measurement of natural levels of (32)P and (33)P in rain, marine particulates, and dissolved constituents of seawater. (32)P and (33)P activities were measured using a recently developed ultra-low-level liquid scintillation counter. Measurement by liquid scintillation counting allows, for the first time, simultaneous measurement of both (32)P and (33)P. Furthermore, (33)P activities are measured with high efficiency (>50%), regardless of the amount of stable phosphorus in the sample. Liquid scintillation also produces energy specific β spectra which has enabled us to identify previously unrecognized β-emitting contaminants in natural samples. In order to remove these contaminants, new methods of purification have been developed which utilize a series of precipitations and anion and cation exchange columns. Rainwater and dissolved seawater samples were extracted from large volumes of rain- and seawater, 5-20 and >5000 L, respectively, using iron-impregnated polypropylene filters. On these filters, it was possible to load between 25 and 30% Fe(OH)(3) by weight, over twice that loaded on previously utilized materials. Using our collection, purification, and liquid scintillation counting techniques, it was possible to obtain specific (32)P and (33)P activities with less than 10% error (2σ) in rainwater and 20% error (2σ) in seawater.