The Deep Ocean's Carbon Exhaust.

The deep ocean releases large amounts of old, pre-industrial carbon dioxide (CO2) to the atmosphere through upwelling in the Southern Ocean, which counters the marine carbon uptake occurring elsewhere. This Southern Ocean CO2 release is relevant to the global climate because its changes could alter atmospheric CO2 levels on long time scales, and also affects the present-day potential of the Southern Ocean to take up anthropogenic CO2. Here, year-round profiling float measurements show that this CO2 release arises from a zonal band of upwelling waters between the Subantarctic Front and wintertime sea-ice edge. This band of high CO2 subsurface water coincides with the outcropping of the 27.8 kg m-3 isoneutral density surface that characterizes Indo-Pacific Deep Water (IPDW). It has a potential partial pressure of CO2 exceeding current atmospheric CO2 levels (∆PCO2) by 175 ± 32 µatm. Ship-based measurements reveal that IPDW exhibits a distinct ∆PCO2 maximum in the ocean, which is set by remineralization of organic carbon and originates from the northern Pacific and Indian Ocean basins. Below this IPDW layer, the carbon content increases downwards, whereas ∆PCO2 decreases. Most of this vertical ∆PCO2 decline results from decreasing temperatures and increasing alkalinity due to an increased fraction of calcium carbonate dissolution. These two factors limit the CO2 outgassing from the high-carbon content deep waters on more southerly surface outcrops. Our results imply that the response of Southern Ocean CO2 fluxes to possible future changes in upwelling are sensitive to the subsurface carbon chemistry set by the vertical remineralization and dissolution profiles.

Effects of Leg Length, Sex, Laterality, and the Intermediate Femoral Cutaneous Nerve on Infrapatellar Innervation.

Background: An iatrogenic injury to the infrapatellar branch of the saphenous nerve (IPBSN) is a common precipitant of postoperative knee pain and hypoesthesia. Purpose: To locate potential safe zones for incision by observing the patterns and pathway of the IPBSN while examining the relationship of its location to sex, laterality, and leg length. Study Design: Descriptive laboratory study. Methods: A total of 107 extended knees from 55 formalin-embalmed cadaveric specimens were dissected. The nerve was measured from palpable landmarks: the patella at the medial (point A) and lateral (point B) borders of the patellar ligament, the medial border of the patellar ligament at the patellar apex (point C) and tibial plateau (point D), the medial epicondyle (point E), and the anterior border of the medial collateral ligament at the tibial plateau (point F). The safe zone was defined as 2 SDs from the mean. Results: Findings indicated significant correlations between leg length and height (r P = 0.832; P < .001) as well as between leg length and vertical measurements (≥45°) from points A and B to the IPBSN (r P range, 0.193-0.285; P range, .004-.049). Male specimens had a more inferior maximum distance from point A to the intersection of the IPBSN and the medial border of the patellar ligament compared with female specimens (6.17 vs 5.28 cm, respectively; P = .049). Right knees had a more posterior IPBSN from point F compared with left knees (-0.98 vs-0.02 cm, respectively; P = .048). The majority of knees (62.6%; n = 67) had a nerve emerging that penetrated the sartorius muscle. Additionally, 32.7% (n = 35) had redundant innervation, and 25.2% (n = 27) had contribution from the intermediate femoral cutaneous nerve (IFCN). Conclusion: We identified no safe zone. Significant innervation redundancy with a substantial contribution to the infrapatellar area from the IFCN was noted and contributed to the expansion of the danger zone. Clinical Relevance: The location of incision and placement of arthroscopic ports might not be as crucial in postoperative pain management as an appreciation of the variance in infrapatellar innervation. The IFCN is a common contributor. Its damage could explain pain refractory to SN blocks and therefore influence anesthetic and analgesic decisions.

Observing the Global Ocean with Biogeochemical-Argo.

Biogeochemical-Argo (BGC-Argo) is a network of profiling floats carrying sensors that enable observation of as many as six essential biogeochemical and bio-optical variables: oxygen, nitrate, pH, chlorophyll a, suspended particles, and downwelling irradiance. This sensor network represents today's most promising strategy for collecting temporally and vertically resolved observations of biogeochemical properties throughout the ocean. All data are freely available within 24 hours of transmission. These data fill large gaps in ocean-observing systems and support three ambitions: gaining a better understanding of biogeochemical processes (e.g., the biological carbon pump and air-sea CO2 exchanges) and evaluating ongoing changes resulting from increasing anthropogenic pressure (e.g., acidification and deoxygenation); managing the ocean (e.g., improving the global carbon budget and developing sustainable fisheries); and carrying out exploration for potential discoveries. The BGC-Argo network has already delivered extensive high-quality global data sets that have resulted in unique scientific outcomes from regional to global scales. With the proposed expansion of BGC-Argo in the near future, this network has the potential to become a pivotal observation system that links satellite and ship-based observations in a transformative manner.

Reassessing Southern Ocean Air-Sea CO2 Flux Estimates With the Addition of Biogeochemical Float Observations.

New estimates of pCO2 from profiling floats deployed by the Southern Ocean Carbon and Climate Observations and Modeling (SOCCOM) project have demonstrated the importance of wintertime outgassing south of the Polar Front, challenging the accepted magnitude of Southern Ocean carbon uptake (Gray et al., 2018, https://doi:10.1029/2018GL078013). Here, we put 3.5 years of SOCCOM observations into broader context with the global surface carbon dioxide database (Surface Ocean CO2 Atlas, SOCAT) by using the two interpolation methods currently used to assess the ocean models in the Global Carbon Budget (Le Quéré et al., 2018, https://doi:10.5194/essd-10-2141-2018) to create a ship-only, a float-weighted, and a combined estimate of Southern Ocean carbon fluxes (<35°S). In our ship-only estimate, we calculate a mean uptake of -1.14 ± 0.19 Pg C/yr for 2015-2017, consistent with prior studies. The float-weighted estimate yields a significantly lower Southern Ocean uptake of -0.35 ± 0.19 Pg C/yr. Subsampling of high-resolution ocean biogeochemical process models indicates that some of the differences between float and ship-only estimates of the Southern Ocean carbon flux can be explained by spatial and temporal sampling differences. The combined ship and float estimate minimizes the root-mean-square pCO2 difference between the mapped product and both data sets, giving a new Southern Ocean uptake of -0.75 ± 0.22 Pg C/yr, though with uncertainties that overlap the ship-only estimate. An atmospheric inversion reveals that a shift of this magnitude in the contemporary Southern Ocean carbon flux must be compensated for by ocean or land sinks within the Southern Hemisphere.

The integral role of iron in ocean biogeochemistry.

The micronutrient iron is now recognized to be important in regulating the magnitude and dynamics of ocean primary productivity, making it an integral component of the ocean's biogeochemical cycles. In this Review, we discuss how a recent increase in observational data for this trace metal has challenged the prevailing view of the ocean iron cycle. Instead of focusing on dust as the major iron source and emphasizing iron's tight biogeochemical coupling to major nutrients, a more complex and diverse picture of the sources of iron, its cycling processes and intricate linkages with the ocean carbon and nitrogen cycles has emerged.

Deep-Sea DuraFET: A Pressure Tolerant pH Sensor Designed for Global Sensor Networks.

Increasing atmospheric carbon dioxide is driving a long-term decrease in ocean pH which is superimposed on daily to seasonal variability. These changes impact ecosystem processes, and they serve as a record of ecosystem metabolism. However, the temporal variability in pH is observed at only a few locations in the ocean because a ship is required to support pH observations of sufficient precision and accuracy. This paper describes a pressure tolerant Ion Sensitive Field Effect Transistor pH sensor that is based on the Honeywell Durafet ISFET die. When combined with a AgCl pseudoreference sensor that is immersed directly in seawater, the system is capable of operating for years at a time on platforms that cycle from depths of several km to the surface. The paper also describes the calibration scheme developed to allow calibrated pH measurements to be derived from the activity of HCl reported by the sensor system over the range of ocean pressure and temperature. Deployments on vertical profiling platforms enable self-calibration in deep waters where pH values are stable. Measurements with the sensor indicate that it is capable of reporting pH with an accuracy of 0.01 or better on the total proton scale and a precision over multiyear periods of 0.005. This system enables a global ocean observing system for ocean pH.

Characterization of an ion sensitive field effect transistor and chloride ion selective electrodes for pH measurements in seawater.

Characterization of several potentiometric cells without a liquid junction has been carried out in universal buffer, aqueous HCl, and artificial seawater media. The electrodes studied include Ion Sensitive Field Effect Transistor (ISFET) pH electrodes, and Chloride-Ion Selective Electrodes (Cl-ISE) directly exposed to the solution. These electrodes were compared directly to the conventional hydrogen electrode and silver-silver chloride electrode in order to report the degree to which they obey ideal Nernstian laws. These data provide a foundation for operating the ISFET|Cl-ISE pair in seawater as a pH sensor. In order to obtain the highest quality pH measurements from this sensor, its response to changes in pH and salinity must be properly characterized. Our results indicate near-ideal Nernstian response for both electrodes over a wide range of pH (2-12) and Cl(-) molality (0.01-1). We conclude that the error due to sub-Nernstian response of the cell ISFET|seawater|Cl-ISE over the range of seawater pH and salinity is negligible (<0.0001 pH). The cross sensitivity of the Cl-ISE to Br(-) does not seem to be a significant source of error (<0.003 pH) in seawater media in the salinity range 20-35.

Recovery of a top predator mediates negative eutrophic effects on seagrass.

A fundamental goal of the study of ecology is to determine the drivers of habitat-forming vegetation, with much emphasis given to the relative importance to vegetation of "bottom-up" forces such as the role of nutrients and "top-down" forces such as the influence of herbivores and their predators. For coastal vegetation (e.g., kelp, seagrass, marsh, and mangroves) it has been well demonstrated that alterations to bottom-up forcing can cause major disturbances leading to loss of dominant vegetation. One such process is anthropogenic nutrient loading, which can lead to major changes in the abundance and species composition of primary producers, ultimately affecting important ecosystem services. In contrast, much less is known about the relative importance of apex predators on coastal vegetated ecosystems because most top predator populations have been depleted or lost completely. Here we provide evidence that an unusual four-level trophic cascade applies in one such system, whereby a top predator mitigates the bottom-up influences of nutrient loading. In a study of seagrass beds in an estuarine ecosystem exposed to extreme nutrient loading, we use a combination of a 50-y time series analysis, spatial comparisons, and mesocosm and field experiments to demonstrate that sea otters (Enhydra lutris) promote the growth and expansion of eelgrass (Zostera marina) through a trophic cascade, counteracting the negative effects of agriculturally induced nutrient loading. Our results add to a small but growing body of literature illustrating that significant interactions between bottom-up and top-down forces occur, in this case with consequences for the conservation of valued ecosystem services provided by seagrass.

High-frequency dynamics of ocean pH: a multi-ecosystem comparison.

The effect of Ocean Acidification (OA) on marine biota is quasi-predictable at best. While perturbation studies, in the form of incubations under elevated pCO(2), reveal sensitivities and responses of individual species, one missing link in the OA story results from a chronic lack of pH data specific to a given species' natural habitat. Here, we present a compilation of continuous, high-resolution time series of upper ocean pH, collected using autonomous sensors, over a variety of ecosystems ranging from polar to tropical, open-ocean to coastal, kelp forest to coral reef. These observations reveal a continuum of month-long pH variability with standard deviations from 0.004 to 0.277 and ranges spanning 0.024 to 1.430 pH units. The nature of the observed variability was also highly site-dependent, with characteristic diel, semi-diurnal, and stochastic patterns of varying amplitudes. These biome-specific pH signatures disclose current levels of exposure to both high and low dissolved CO(2), often demonstrating that resident organisms are already experiencing pH regimes that are not predicted until 2100. Our data provide a first step toward crystallizing the biophysical link between environmental history of pH exposure and physiological resilience of marine organisms to fluctuations in seawater CO(2). Knowledge of this spatial and temporal variation in seawater chemistry allows us to improve the design of OA experiments: we can test organisms with a priori expectations of their tolerance guardrails, based on their natural range of exposure. Such hypothesis-testing will provide a deeper understanding of the effects of OA. Both intuitively simple to understand and powerfully informative, these and similar comparative time series can help guide management efforts to identify areas of marine habitat that can serve as refugia to acidification as well as areas that are particularly vulnerable to future ocean change.

Differential distributions of synechococcus subgroups across the california current system.

Synechococcus is an abundant marine cyanobacterial genus composed of different populations that vary physiologically. Synechococcus narB gene sequences (encoding for nitrate reductase in cyanobacteria) obtained previously from isolates and the environment (e.g., North Pacific Gyre Station ALOHA, Hawaii or Monterey Bay, CA, USA) were used to develop quantitative PCR (qPCR) assays. These qPCR assays were used to quantify populations from specific narB phylogenetic clades across the California Current System (CCS), a region composed of dynamic zones between a coastal-upwelling zone and the oligotrophic Pacific Ocean. Targeted populations (narB subgroups) had different biogeographic patterns across the CCS, which appear to be driven by environmental conditions. Subgroups C_C1, D_C1, and D_C2 were abundant in coastal-upwelling to coastal-transition zone waters with relatively high to intermediate ammonium, nitrate, and chl. a concentrations. Subgroups A_C1 and F_C1 were most abundant in coastal-transition zone waters with intermediate nutrient concentrations. E_O1 and G_O1 were most abundant at different depths of oligotrophic open-ocean waters (either in the upper mixed layer or just below). E_O1, A_C1, and F_C1 distributions differed from other narB subgroups and likely possess unique ecologies enabling them to be most abundant in waters between coastal and open-ocean waters. Different CCS zones possessed distinct Synechococcus communities. Core California current water possessed low numbers of narB subgroups relative to counted Synechococcus cells, and coastal-transition waters contained high abundances of Synechococcus cells and total number of narB subgroups. The presented biogeographic data provides insight on the distributions and ecologies of Synechococcus present in an eastern boundary current system.

Nitrate supply from deep to near-surface waters of the North Pacific subtropical gyre.

Concentrations of dissolved inorganic carbon (DIC) decrease in the surface mixed layers during spring and summer in most of the oligotrophic ocean. Mass balance calculations require that the missing DIC is converted into particulate carbon by photosynthesis. This DIC uptake represents one of the largest components of net community production in the world ocean. However, mixed-layer waters in these regions of the ocean typically contain negligible concentrations of plant nutrients such as nitrate and phosphate. Combined nutrient supply mechanisms including nitrogen fixation, diffusive transport and vertical entrainment are believed to be insufficient to supply the required nutrients for photosynthesis. The basin-scale potential for episodic nutrient transport by eddy events is unresolved. As a result, it is not understood how biologically mediated DIC uptake can be supported in the absence of nutrients. Here we report on high-resolution measurements of nitrate (NO(3)(-)) and oxygen (O(2)) concentration made over 21 months using a profiling float deployed near the Hawaii Ocean Time-series station in the North Pacific subtropical gyre. Our measurements demonstrate that as O(2) was produced and DIC was consumed over two annual cycles, a corresponding seasonal deficit in dissolved NO(3)(-) appeared in water at depths from 100 to 250 m. The deep-water deficit in NO(3)(-) was in near-stoichiometric balance with the fixed nitrogen exported to depth. Thus, when the water column from the surface to 250 m is considered as a whole, there is near equivalence between nutrient supply and demand. Short-lived transport events (<10 days) that connect deep stocks of nitrate to nutrient-poor surface waters were clearly present in 12 of the 127 vertical profiles.

Unicellular cyanobacterial distributions broaden the oceanic N2 fixation domain.

Nitrogen (N2)-fixing microorganisms (diazotrophs) are an important source of biologically available fixed N in terrestrial and aquatic ecosystems and control the productivity of oligotrophic ocean ecosystems. We found that two major groups of unicellular N2-fixing cyanobacteria (UCYN) have distinct spatial distributions that differ from those of Trichodesmium, the N2-fixing cyanobacterium previously considered to be the most important contributor to open-ocean N2 fixation. The distributions and activity of the two UCYN groups were separated as a function of depth, temperature, and water column density structure along an 8000-kilometer transect in the South Pacific Ocean. UCYN group A can be found at high abundances at substantially higher latitudes and deeper in subsurface ocean waters than Trichodesmium. These findings have implications for the geographic extent and magnitude of basin-scale oceanic N2 fixation rates.

Net production of oxygen in the subtropical ocean.

The question of whether the plankton communities in low-nutrient regions of the ocean, comprising 80% of the global ocean surface area, are net producers or consumers of oxygen and fixed carbon is a key uncertainty in the global carbon cycle. Direct measurements in bottle experiments indicate net oxygen consumption in the sunlit zone, whereas geochemical evidence suggests that the upper ocean is a net source of oxygen. One possible resolution to this conflict is that primary production in the gyres is episodic and thus difficult to observe: in this model, oligotrophic regions would be net consumers of oxygen during most of the year, but strong, brief events with high primary production rates might produce enough fixed carbon and dissolved oxygen to yield net production as an average over the annual cycle. Here we examine the balance of oxygen production over three years at sites in the North and South Pacific subtropical gyres using the new technique of oxygen sensors deployed on profiling floats. We find that mixing events during early winter homogenize the upper water column and cause low oxygen concentrations. Oxygen then increases below the mixed layer at a nearly constant rate that is similar to independent measures of net community production. This continuous oxygen increase is consistent with an ecosystem that is a net producer of fixed carbon (net autotrophic) throughout the year, with episodic events not required to sustain positive oxygen production.

Southern Ocean iron enrichment experiment: carbon cycling in high- and low-Si waters.

The availability of iron is known to exert a controlling influence on biological productivity in surface waters over large areas of the ocean and may have been an important factor in the variation of the concentration of atmospheric carbon dioxide over glacial cycles. The effect of iron in the Southern Ocean is particularly important because of its large area and abundant nitrate, yet iron-enhanced growth of phytoplankton may be differentially expressed between waters with high silicic acid in the south and low silicic acid in the north, where diatom growth may be limited by both silicic acid and iron. Two mesoscale experiments, designed to investigate the effects of iron enrichment in regions with high and low concentrations of silicic acid, were performed in the Southern Ocean. These experiments demonstrate iron's pivotal role in controlling carbon uptake and regulating atmospheric partial pressure of carbon dioxide.