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  1. Theory suggests that the ocean’s biological carbon pump, the process by which organic matter is produced at the surface and transferred to the deep ocean, is sensitive to temperature because temperature controls photosynthesis and respiration rates. We applied a combined data-modeling approach to investigate carbon and nutrient recycling rates across the world ocean over the past 15 million years of global cooling. We found that the efficiency of the biological carbon pump increased with ocean cooling as the result of a temperature-dependent reduction in the rate of remineralization (degradation) of sinking organic matter. Increased food delivery at depth prompted the development of new deep-water niches, triggering deep plankton evolution and the expansion of the mesopelagic “twilight zone” ecosystem.

     
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    Abstract. During the early part of the last glacial termination (17.2–15 ka) and coincident with a ∼35 ppm rise in atmospheric CO2, a sharp 0.3‰–0.4‰ decline in atmospheric δ13CO2 occurred, potentially constraining the key processes that account for the early deglacial CO2 rise. A comparable δ13C decline has also been documented in numerous marine proxy records from surface and thermocline-dwelling planktic foraminifera. The δ13C decline recorded in planktic foraminifera has previously been attributed to the release of respired carbon from the deep ocean that was subsequently transported within the upper ocean to sites where the signal was recorded (and then ultimately transferred to the atmosphere). Benthic δ13C records from the global upper ocean, including a new record presented here from the tropical Pacific, also document this distinct early deglacial δ13C decline. Here we present modeling evidence to show that rather than respired carbon from the deep ocean propagating directly to the upper ocean prior to reaching the atmosphere, the carbon would have first upwelled to the surface in the Southern Ocean where it would have entered the atmosphere. In this way the transmission of isotopically light carbon to the global upper ocean was analogous to the ongoing ocean invasion of fossil fuel CO2. The model results suggest that thermocline waters throughout the ocean and 500–2000 m water depths were affected by this atmospheric bridge during the early deglaciation. 
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    Abstract. Temperature is a master parameter in the marine carbon cycle, exerting a critical control on the rate of biological transformation of a variety of solid and dissolved reactants and substrates. Although in the construction of numerical models of marine carbon cycling, temperature has been long recognised as a key parameter in the production and export of organic matter at the ocean surface, its role in the ocean interior is much less frequently accounted for. There, bacteria (primarily) transform sinking particulate organic matter (POM) into its dissolved constituents and consume dissolved oxygen (and/or other electron acceptors such as sulfate). The nutrients and carbon thereby released then become available for transport back to the surface, influencing biological productivity and atmospheric pCO2, respectively. Given the substantial changes in ocean temperature occurring in the past, as well as in light of current anthropogenic warming, appropriately accounting for the role of temperature in marine carbon cycling may be critical to correctly projecting changes in ocean deoxygenation and the strength of feedbacks on atmosphericpCO2. Here we extend and calibrate a temperature-dependent representation ofmarine carbon cycling in the cGENIE.muffin Earth system model, intended forboth past and future climate applications. In this, we combine atemperature-dependent remineralisation scheme for sinking organic matterwith a biological export production scheme that also includes a dependenceon ambient seawater temperature. Via a parameter ensemble, we jointlycalibrate the two parameterisations by statistically contrasting model-projected fields of nutrients, oxygen, and the stable carbon isotopicsignature (δ13C) of dissolved inorganic carbon in the oceanwith modern observations. We additionally explore the role of temperature inthe creation and recycling of dissolved organic matter (DOM) and hence itsimpact on global carbon cycle dynamics. We find that for the present day, the temperature-dependent version showsa fit to the data that is as good as or better than the existing tuned non-temperature-dependent version of the cGENIE.muffin. The main impact ofaccounting for temperature-dependent remineralisation of POM is in drivinghigher rates of remineralisation in warmer waters, in turn driving a morerapid return of nutrients to the surface and thereby stimulating organicmatter production. As a result, more POM is exported below 80 m but onaverage reaches shallower depths in middle- and low-latitude warmer waterscompared to the standard model. Conversely, at higher latitudes, colderwater temperature reduces the rate of nutrient resupply to the surface andPOM reaches greater depth on average as a result of slower subsurface ratesof remineralisation. Further adding temperature-dependent DOM processeschanges this overall picture only a little, with a slight weakening ofexport production at higher latitudes. As an illustrative application of the new model configuration andcalibration, we take the example of historical warming and briefly assessthe implications for global carbon cycling of accounting for a more completeset of temperature-dependent processes in the ocean. We find that betweenthe pre-industrial era (ca. 1700) and the present (year 2010), in response to asimulated air temperature increase of 0.9 ∘C and an associatedprojected mean ocean warming of 0.12 ∘C (0.6 ∘C insurface waters and 0.02 ∘C in deep waters), a reduction inparticulate organic carbon (POC) export at 80 m of just 0.3 % occurs (or 0.7 % including a temperature-dependent DOM response). However, due to this increased recycling nearer the surface, the efficiency of the transfer of carbon away from the surface (at 80 m) to the deep ocean (at 1040 m) is reduced by 5 %. In contrast, with no assumed temperature-dependent processes impacting production or remineralisation of either POM or DOM, global POC export at 80 m falls by 2.9 % between the pre-industrial era and the present day as a consequence of ocean stratification and reduced nutrient resupply to the surface. Our analysis suggests that increased temperature-dependent nutrient recycling in the upper ocean has offset much of the stratification-induced restriction in its physical transport. 
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    For most of Earth’s history, the ocean’s interior was pervasively anoxic and showed occasional shifts in ocean redox chemistry between iron-buffered and sulfide-buffered states. These redox transitions are most often explained by large changes in external inputs, such as a strongly altered delivery of iron and sulfate to the ocean, or major shifts in marine productivity. Here, we propose that redox shifts can also arise from small perturbations that are amplified by nonlinear positive feedbacks within the internal iron and sulfur cycling of the ocean. Combining observational evidence with biogeochemical modeling, we show that both sedimentary and aquatic systems display intrinsic iron–sulfur bistability, which is tightly linked to the formation of reduced iron–sulfide minerals. The possibility of tipping points in the redox state of sediments and oceans, which allow large and nonreversible geochemical shifts to arise from relatively small changes in organic carbon input, has important implications for the interpretation of the geological rock record and the causes and consequences of major evolutionary transitions in the history of Earth’s biosphere. 
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    Although the Pacific Ocean is a major reservoir of heat and CO 2 , and thus an important component of the global climate system, its circulation under different climatic conditions is poorly understood. Here, we present evidence that during the Last Glacial Maximum (LGM), the North Pacific was better ventilated at intermediate depths and had surface waters with lower nutrients, higher salinity, and warmer temperatures compared to today. Modeling shows that this pattern is well explained by enhanced Pacific meridional overturning circulation (PMOC), which brings warm, salty, and nutrient-poor subtropical waters to high latitudes. Enhanced PMOC at the LGM would have lowered atmospheric CO 2 —in part through synergy with the Southern Ocean—and supported an equable regional climate, which may have aided human habitability in Beringia, and migration from Asia to North America. 
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    The present-day marine nitrogen (N) cycle is strongly regulated by biology. Deficiencies in the availability of fixed and readily bioavailable nitrogen relative to phosphate (P) in the surface ocean are largely corrected by the activity of diazotrophs. This feedback system, termed the “nitrostat,” is thought to have provided close regulation of fixed-N speciation and inventory relative to P since the Proterozoic. In contrast, during intervals of intense deoxygenation such as Cretaceous ocean anoxic event (OAE) 2, a few regional sedimentary δ 15 N records hint at the existence of a different mode of marine N cycling in which ammonium plays a major role in regulating export production. However, the global-scale dynamics during this time remain unknown. Here, using an Earth System model and taking the example of OAE 2, we provide insights into the global marine nitrogen cycle under severe ocean deoxygenation. Specifically, we find that the ocean can exhibit fundamental transitions in the species of nitrogen dominating the fixed-N inventory––from nitrate (NO 3 − ) to ammonium (NH 4 + )––and that as this transition occurs, the inventory can partially collapse relative to P due to progressive spatial decoupling between the loci of NH 4 + oxidation, NO 3 − reduction, and nitrogen fixation. This finding is relatively independent of the specific state of ocean circulation and is consistent with nitrogen isotope and redox proxy data. The substantive reduction in the ocean fixed-N inventory at an intermediate state of deoxygenation may represent a biogeochemical vulnerability with potential implications for past and future (warmer) oceans. 
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  8. Rising oceanic and atmospheric oxygen levels through time have been crucial to enhanced habitability of surface Earth environments. Few redox proxies can track secular variations in dissolved oxygen concentrations ([O2]) around threshold levels for metazoan survival in the upper ocean. We present an extensive compilation of iodine to calcium ratios (I/Ca) in marine carbonates. Our record supports a major rise in atmospheric pO2 at ~400 million years ago (Ma), and reveals a step-change in the oxygenation of the upper ocean to relatively sustainable near-modern conditions at ~200 Ma. An Earth system model demonstrates that a shift in organic matter remineralization to greater depths, which may have been due to increasing size and biomineralization of eukaryotic plankton, likely drove the I/Ca signals at ~200 Ma 
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