3D intrusions transport active surface microbial assemblages to the dark ocean.

Subtropical oceans contribute significantly to global primary production, but the fate of the picophytoplankton that dominate in these low-nutrient regions is poorly understood. Working in the subtropical Mediterranean, we demonstrate that subduction of water at ocean fronts generates 3D intrusions with uncharacteristically high carbon, chlorophyll, and oxygen that extend below the sunlit photic zone into the dark ocean. These contain fresh picophytoplankton assemblages that resemble the photic-zone regions where the water originated. Intrusions propagate depth-dependent seasonal variations in microbial assemblages into the ocean interior. Strikingly, the intrusions included dominant biomass contributions from nonphotosynthetic bacteria and enrichment of enigmatic heterotrophic bacterial lineages. Thus, the intrusions not only deliver material that differs in composition and nutritional character from sinking detrital particles, but also drive shifts in bacterial community composition, organic matter processing, and interactions between surface and deep communities. Modeling efforts paired with global observations demonstrate that subduction can flux similar magnitudes of particulate organic carbon as sinking export, but is not accounted for in current export estimates and carbon cycle models. Intrusions formed by subduction are a particularly important mechanism for enhancing connectivity between surface and upper mesopelagic ecosystems in stratified subtropical ocean environments that are expanding due to the warming climate.

Biogenic carbon pool production maintains the Southern Ocean carbon sink.

Through biological activity, marine dissolved inorganic carbon (DIC) is transformed into different types of biogenic carbon available for export to the ocean interior, including particulate organic carbon (POC), dissolved organic carbon (DOC), and particulate inorganic carbon (PIC). Each biogenic carbon pool has a different export efficiency that impacts the vertical ocean carbon gradient and drives natural air-sea carbon dioxide gas (CO2) exchange. In the Southern Ocean (SO), which presently accounts for ~40% of the anthropogenic ocean carbon sink, it is unclear how the production of each biogenic carbon pool contributes to the contemporary air-sea CO2 exchange. Based on 107 independent observations of the seasonal cycle from 63 biogeochemical profiling floats, we provide the basin-scale estimate of distinct biogenic carbon pool production. We find significant meridional variability with enhanced POC production in the subantarctic and polar Antarctic sectors and enhanced DOC production in the subtropical and sea-ice-dominated sectors. PIC production peaks between 47°S and 57°S near the "great calcite belt." Relative to an abiotic SO, organic carbon production enhances CO2 uptake by 2.80 ± 0.28 Pg C y-1, while PIC production diminishes CO2 uptake by 0.27 ± 0.21 Pg C y-1. Without organic carbon production, the SO would be a CO2 source to the atmosphere. Our findings emphasize the importance of DOC and PIC production, in addition to the well-recognized role of POC production, in shaping the influence of carbon export on air-sea CO2 exchange.

δ13C of bulk organic matter and cellulose reveal post-photosynthetic fractionation during ontogeny in C4 grass leaves.

The 13C isotope composition (δ13C) of leaf dry matter is a useful tool for physiological and ecological studies. However, how post-photosynthetic fractionation associated with respiration and carbon export influences δ13C remains uncertain. We investigated the effects of post-photosynthetic fractionation on δ13C of mature leaves of Cleistogenes squarrosa, a perennial C4 grass, in controlled experiments with different levels of vapour pressure deficit and nitrogen supply. With increasing leaf age class, the 12C/13C fractionation of leaf organic matter relative to the δ13C of atmosphere CO2 (ΔDM) increased while that of cellulose (Δcel) was almost constant. The divergence between ΔDM and Δcel increased with leaf age class, with a maximum value of 1.6‰, indicating the accumulation of post-photosynthetic fractionation. Applying a new mass balance model that accounts for respiration and export of photosynthates, we found an apparent 12C/13C fractionation associated with carbon export of -0.5‰ to -1.0‰. Different ΔDM among leaves, pseudostems, daughter tillers, and roots indicate that post-photosynthetic fractionation happens at the whole-plant level. Compared with ΔDM of old leaves, ΔDM of young leaves and Δcel are more reliable proxies for predicting physiological parameters due to the lower sensitivity to post-photosynthetic fractionation and the similar sensitivity in responses to environmental changes.

Microbial dynamics of elevated carbon flux in the open ocean's abyss.

In the open ocean, elevated carbon flux (ECF) events increase the delivery of particulate carbon from surface waters to the seafloor by severalfold compared to other times of year. Since microbes play central roles in primary production and sinking particle formation, they contribute greatly to carbon export to the deep sea. Few studies, however, have quantitatively linked ECF events with the specific microbial assemblages that drive them. Here, we identify key microbial taxa and functional traits on deep-sea sinking particles that correlate positively with ECF events. Microbes enriched on sinking particles in summer ECF events included symbiotic and free-living diazotrophic cyanobacteria, rhizosolenid diatoms, phototrophic and heterotrophic protists, and photoheterotrophic and copiotrophic bacteria. Particle-attached bacteria reaching the abyss during summer ECF events encoded metabolic pathways reflecting their surface water origins, including oxygenic and aerobic anoxygenic photosynthesis, nitrogen fixation, and proteorhodopsin-based photoheterotrophy. The abundances of some deep-sea bacteria also correlated positively with summer ECF events, suggesting rapid bathypelagic responses to elevated organic matter inputs. Biota enriched on sinking particles during a spring ECF event were distinct from those found in summer, and included rhizaria, copepods, fungi, and different bacterial taxa. At other times over our 3-y study, mid- and deep-water particle colonization, predation, degradation, and repackaging (by deep-sea bacteria, protists, and animals) appeared to shape the biotic composition of particles reaching the abyss. Our analyses reveal key microbial players and biological processes involved in particle formation, rapid export, and consumption, that may influence the ocean's biological pump and help sustain deep-sea ecosystems.

High Dissolved Carbon Concentration in Arid Rocky Mountain Streams.

Warming in mountains is known to intensify aridity and threaten water availability globally. Its impacts on water quality, however, have remained poorly understood. Here we collate long-term (multi-year to decadal mean), baseline stream concentrations and fluxes of dissolved organic and inorganic carbon, two essential indicators of water quality and soil carbon response to warming, across more than 100 streams in the United States Rocky Mountains. Results show a universal pattern of higher mean concentrations in more arid mountain streams with lower mean discharge, a long-term climate measure. A watershed reactor model revealed less lateral export of dissolved carbon (via less water flow) out of the watersheds in more arid sites, leading to more accumulation and higher concentrations. Lower concentrations typically occur in cold, steep, and compact mountains with higher snow fraction and lower vegetation cover, which generally have higher discharge and carbon fluxes. Inferring from a space-for-time perspective, the results indicate that as warming intensifies, lateral fluxes of dissolved carbon will decrease but concentrations will increase in these mountain streams. This indicates deteriorating water quality and potentially elevated CO2 emission directly from the land (instead of streams) in the Rockies and other mountain areas in the future climate.

Global value chains participation and trade-embodied net carbon exports in group of seven and emerging seven countries.

A vast literature has examined the empirical link between gross exports and total carbon emissions for different country groups. However, countries' increasing participation in global value chains (GVCs) challenges this traditional approach since the gross measures neglect trade-embodied carbon emissions and intermediates-driven value-added trade. Therefore, this study scrutinizes how backward participation (foreign contents in domestic exports) and forward participation (domestic contents in foreign exports) in GVCs affect per capita net exports of trade-embodied carbon dioxide emissions. The study adopts input-output accounting and value-added decomposition framework for Group of Seven (G7) and Emerging Seven (E7) countries over the 1995-2018 period. (i) Pre-estimation analyses reveal that the net carbon importer G7 group had a comparative advantage in high-tech exports and a lower export product concentration level, while the net carbon exporter E7 group had a comparative advantage in resource-intensive exports and a higher export product concentration level, albeit significant within-group heterogeneities. (ii) The augmented mean group estimates reveal that increasing backward participation raises net carbon exports for both G7 and E7. The forward participation-net carbon exports nexus is negative for G7 but positive for E7. (iii) While economic growth reduces net carbon exports in both groups, the effects of comparative advantages in resource-intensive and high-tech exports differ. Practitioners should be aware of the GVCs-driven carbon circle when assessing decarbonization performances and obligations of countries.

Manganese Limitation of Phytoplankton Physiology and Productivity in the Southern Ocean.

Although iron and light are understood to regulate the Southern Ocean biological carbon pump, observations have also indicated a possible role for manganese. Low concentrations in Southern Ocean surface waters suggest manganese limitation is possible, but its spatial extent remains poorly constrained and direct manganese limitation of the marine carbon cycle has been neglected by ocean models. Here, using available observations, we develop a new global biogeochemical model and find that phytoplankton in over half of the Southern Ocean cannot attain maximal growth rates because of manganese deficiency. Manganese limitation is most extensive in austral spring and depends on phytoplankton traits related to the size of photosynthetic antennae and the inhibition of manganese uptake by high zinc concentrations in Antarctic waters. Importantly, manganese limitation expands under the increased iron supply of past glacial periods, reducing the response of the biological carbon pump. Overall, these model experiments describe a mosaic of controls on Southern Ocean productivity that emerge from the interplay of light, iron, manganese and zinc, shaping the evolution of Antarctic phytoplankton since the opening of the Drake Passage.

Interlinking diatom frustule diversity from the abyss of the central Arabian Sea to surface processes: physical forcing and oxygen minimum zone.

This study analyzed the diversity and abundance of diatom frustules including the ancillary parameters using the core top sediments from five locations (21, 19, 15, 13, and 11°N) along the central Arabian Sea (64°E), an area profoundly influenced by atmospheric forcing (monsoons) and oxygen minimum zone (OMZ) with high spatial variability. Significantly higher organic carbon (0.97 ± 0.05%) and diatom frustules (5.92 ± 0.57 × 104 valves g-1) were noticed in the north (21, 19, 15°N) where natural nutrient enrichment via open-ocean upwelling, winter convection, and lateral advection support large diatom-dominated phytoplankton blooms and intense OMZ. Conversely, the south (13, 11°N) depicted significantly lower organic carbon (0.74 ± 0.08%) as well as frustules (4.02 ± 0.87 × 104 valves g-1) as this area mostly remains nutrient-poor dominated by small-medium-sized phytoplankton. The north was dominated by large-sized diatoms like Coscinodiscus that could escape grazing and sink consequently due to higher ballasting. Furthermore, the presence of the intense OMZ in the north might reduce grazing pressure (low zooplankton stock) and mineralization speed facilitating higher phytodetritus transport. Relatively smaller chain-forming centric (Thalassiosira) and pennate diatoms (Pseudo-nitzschia, Fragilaria, Nitzschia, etc.) were found throughout the transect with higher abundance in the south. The euphotic diatom diversity from the existing literature was compared with the frustule diversity from the sediments suggesting not all diatoms make their way to the abyss. Such distinct spatial north-south variability in diatom frustule size as well as abundance could be attributed to cell size, grazing, and water column mineralization rates related to OMZ.

Using High-Sensitivity Lipidomics To Assess Microscale Heterogeneity in Oceanic Sinking Particles and Single Phytoplankton Cells.

Sinking particulate organic matter (POM) is a primary component of the ocean's biological carbon pump that is responsible for carbon export from the surface to the deep sea. Lipids derived from plankton comprise a significant fraction of sinking POM. Our understanding of planktonic lipid biosynthesis and the subsequent degradation of lipids in sinking POM is based on the analysis of bulk samples that combine many millions of plankton cells or dozens of sinking particles, which averages out natural heterogeneity. We developed and applied a nanoflow high-performance liquid-chromatography electrospray-ionization high-resolution accurate-mass mass spectrometry lipidomic method to show that two types of sinking particles─marine snow and fecal pellets─collected in the western North Atlantic Ocean have distinct lipidomes, providing new insights into their sources and degradation that would not be apparent from bulk samples. We pressed the limit of this approach by examining individual diatom cells from a single culture, finding marked lipid heterogeneity, possibly indicative of fundamental mechanisms underlying cell division. These single-cell data confirm that even cultures of phytoplankton cells should be viewed as mixtures of physiologically distinct populations. Overall, this work reveals previously hidden lipidomic heterogeneity in natural POM and phytoplankton cells, which may provide critical new insights into microscale chemical and microbial processes that control the export of sinking POM.

Major imprint of surface plankton on deep ocean prokaryotic structure and activity.

Deep ocean microbial communities rely on the organic carbon produced in the sunlit ocean, yet it remains unknown whether surface processes determine the assembly and function of bathypelagic prokaryotes to a larger extent than deep-sea physicochemical conditions. Here, we explored whether variations in surface phytoplankton assemblages across Atlantic, Pacific and Indian ocean stations can explain structural changes in bathypelagic (ca. 4,000 m) free-living and particle-attached prokaryotic communities (characterized through 16S rRNA gene sequencing), as well as changes in prokaryotic activity and dissolved organic matter (DOM) quality. We show that the spatial structuring of prokaryotic communities in the bathypelagic strongly followed variations in the abundances of surface dinoflagellates and ciliates, as well as gradients in surface primary productivity, but were less influenced by bathypelagic physicochemical conditions. Amino acid-like DOM components in the bathypelagic reflected variations of those components in surface waters, and seemed to control bathypelagic prokaryotic activity. The imprint of surface conditions was more evident in bathypelagic than in shallower mesopelagic (200-1,000 m) communities, suggesting a direct connectivity through fast-sinking particles that escape mesopelagic transformations. Finally, we identified a pool of endemic deep-sea prokaryotic taxa (including potentially chemoautotrophic groups) that appear less connected to surface processes than those bathypelagic taxa with a widespread vertical distribution. Our results suggest that surface planktonic communities shape the spatial structure of the bathypelagic microbiome to a larger extent than the local physicochemical environment, likely through determining the nature of the sinking particles and the associated prokaryotes reaching bathypelagic waters.

State of the science in reconciling top-down and bottom-up approaches for terrestrial CO2 budget.

Robust estimates of CO2 budget, CO2 exchanged between the atmosphere and terrestrial biosphere, are necessary to better understand the role of the terrestrial biosphere in mitigating anthropogenic CO2 emissions. Over the past decade, this field of research has advanced through understanding of the differences and similarities of two fundamentally different approaches: "top-down" atmospheric inversions and "bottom-up" biosphere models. Since the first studies were undertaken, these approaches have shown an increasing level of agreement, but disagreements in some regions still persist, in part because they do not estimate the same quantity of atmosphere-biosphere CO2 exchange. Here, we conducted a thorough comparison of CO2 budgets at multiple scales and from multiple methods to assess the current state of the science in estimating CO2 budgets. Our set of atmospheric inversions and biosphere models, which were adjusted for a consistent flux definition, showed a high level of agreement for global and hemispheric CO2 budgets in the 2000s. Regionally, improved agreement in CO2 budgets was notable for North America and Southeast Asia. However, large gaps between the two methods remained in East Asia and South America. In other regions, Europe, boreal Asia, Africa, South Asia, and Oceania, it was difficult to determine whether those regions act as a net sink or source because of the large spread in estimates from atmospheric inversions. These results highlight two research directions to improve the robustness of CO2 budgets: (a) to increase representation of processes in biosphere models that could contribute to fill the budget gaps, such as forest regrowth and forest degradation; and (b) to reduce sink-source compensation between regions (dipoles) in atmospheric inversion so that their estimates become more comparable. Advancements on both research areas will increase the level of agreement between the top-down and bottom-up approaches and yield more robust knowledge of regional CO2 budgets.

High-Frequency Variability of Small-Particle Carbon Export Flux in the Northeast Atlantic.

The biological carbon pump exports carbon fixed by photosynthesis out of the surface ocean and transfers it to the deep, mostly in the form of sinking particles. Despite the importance of the pump in regulating the air-sea CO2 balance, the magnitude of global carbon export remains unclear, as do its controlling mechanisms. A possible sinking flux of carbon to the mesopelagic zone may be via the mixed-layer pump: a seasonal net detrainment of particulate organic carbon (POC)-rich surface waters, caused by sequential deepening and shoaling of the mixed layer. In this study, we present a full year of daily small-particle POC concentrations derived from glider optical backscatter data, to study export variability at the Porcupine Abyssal Plain (PAP) sustained observatory in the Northeast Atlantic. We observe a strong seasonality in small-particle transfer efficiency, with a maximum in winter and early spring. By calculating daily POC export driven by mixed-layer variations, we find that the mixed-layer pump supplies an annual flux of at least 3.0 ± 0.9 g POC·m-2·year-1 to the mesopelagic zone, contributing between 5% and 25% of the total annual export flux and likely contributing to closing a gap in the mesopelagic carbon budget found by other studies. These are, to our best knowledge, the first high-frequency observations of export variability over the course of a full year. Our results support the deployment of bio-optical sensors on gliders to improve our understanding of the ocean carbon cycle on temporal scales from daily to annual.

Can Rates of Ocean Primary Production and Biological Carbon Export Be Related Through Their Probability Distributions?

We describe the basis of a theory for interpreting measurements of two key biogeochemical fluxes-primary production by phytoplankton (p, µg C · L-1 · day-1) and biological carbon export from the surface ocean by sinking particles (f, mg C · m-2 · day-1)-in terms of their probability distributions. Given that p and f are mechanistically linked but variable and effectively measured on different scales, we hypothesize that a quantitative relationship emerges between collections of the two measurements. Motivated by the many subprocesses driving production and export, we take as a null model that large-scale distributions of p and f are lognormal. We then show that compilations of p and f measurements are consistent with this hypothesis. The compilation of p measurements is extensive enough to subregion by biome, basin, depth, or season; these subsets are also well described by lognormals, whose log-moments sort predictably. Informed by the lognormality of both p and f we infer a statistical scaling relationship between the two quantities and derive a linear relationship between the log-moments of their distributions. We find agreement between two independent estimates of the slope and intercept of this line and show that the distribution of f measurements is consistent with predictions made from the moments of the p distribution. These results illustrate the utility of a distributional approach to biogeochemical fluxes. We close by describing potential uses and challenges for the further development of such an approach.

Blue carbon: past, present and future, with emphasis on macroalgae.

Blue carbon did not originally include macroalgal ecosystems; however evidence is mounting that macroalgal ecosystems function in marine carbon sequestration. The great majority of present day marine macroalgal net primary productivity (NPP) involves haptophytic algae on eroding shores. For these organisms the long-term storage of particulate organic carbon involves export from the site of production of biomass that has evaded parasites and grazers, and that some of the exported biomass is sedimented and stored rather than being mineralized en route by detritivores (microbes and fauna). Export from eroding shores, and subsequent storage, of haptophytic marine macroalgal particulate organic carbon could have started by 1.6 Ga. Storage on depositing shores close to the site of NPP by rhizophytic macroalgae and then by rhizophytic coastal seagrasses, tidal marshes and mangroves began not less than 209 Ma ago. Future increases in surface ocean temperatures may bring tropical marine macroalgae to their upper temperature limit, while temperate marine macroalgae will migrate poleward, in both cases assuming that temperature increases faster than genetic adaptation to higher temperature. Increased CO2 in the surface ocean will generally favour uncalcified over calcified marine macroalgae. This results in decreased CO2 release from decreased calcification, as well as decreased ballasting by CaCO3 of exported particulate organic carbon resulting in decreasing sedimentation. While much more work is needed, the available information suggests that macroalgae play a significant role in marine organic carbon storage.

Spring-summer net community production, new production, particle export and related water column biogeochemical processes in the marginal sea ice zone of the Western Antarctic Peninsula 2012-2014.

New production (New P, the rate of net primary production (NPP) supported by exogenously supplied limiting nutrients) and net community production (NCP, gross primary production not consumed by community respiration) are closely related but mechanistically distinct processes. They set the carbon balance in the upper ocean and define an upper limit for export from the system. The relationships, relative magnitudes and variability of New P (from 15NO3- uptake), O2 : argon-based NCP and sinking particle export (based on the 238U : 234Th disequilibrium) are increasingly well documented but still not clearly understood. This is especially true in remote regions such as polar marginal ice zones. Here we present a 3-year dataset of simultaneous measurements made at approximately 50 stations along the Western Antarctic Peninsula (WAP) continental shelf in midsummer (January) 2012-2014. Net seasonal-scale changes in water column inventories (0-150 m) of nitrate and iodide were also estimated at the same stations. The average daily rates based on inventory changes exceeded the shorter-term rate measurements. A major uncertainty in the relative magnitude of the inventory estimates is specifying the start of the growing season following sea-ice retreat. New P and NCP(O2) did not differ significantly. New P and NCP(O2) were significantly greater than sinking particle export from thorium-234. We suggest this is a persistent and systematic imbalance and that other processes such as vertical mixing and advection of suspended particles are important export pathways.This article is part of the theme issue 'The marine system of the west Antarctic Peninsula: status and strategy for progress in a region of rapid change'.

Observing climate change trends in ocean biogeochemistry: when and where.

Understanding the influence of anthropogenic forcing on the marine biosphere is a high priority. Climate change-driven trends need to be accurately assessed and detected in a timely manner. As part of the effort towards detection of long-term trends, a network of ocean observatories and time series stations provide high quality data for a number of key parameters, such as pH, oxygen concentration or primary production (PP). Here, we use an ensemble of global coupled climate models to assess the temporal and spatial scales over which observations of eight biogeochemically relevant variables must be made to robustly detect a long-term trend. We find that, as a global average, continuous time series are required for between 14 (pH) and 32 (PP) years to distinguish a climate change trend from natural variability. Regional differences are extensive, with low latitudes and the Arctic generally needing shorter time series (<~30 years) to detect trends than other areas. In addition, we quantify the 'footprint' of existing and planned time series stations, that is the area over which a station is representative of a broader region. Footprints are generally largest for pH and sea surface temperature, but nevertheless the existing network of observatories only represents 9-15% of the global ocean surface. Our results present a quantitative framework for assessing the adequacy of current and future ocean observing networks for detection and monitoring of climate change-driven responses in the marine ecosystem.

UAV-Based Estimation of Carbon Exports from Heterogeneous Soil Landscapes--A Case Study from the CarboZALF Experimental Area.

The advantages of remote sensing using Unmanned Aerial Vehicles (UAVs) are a high spatial resolution of images, temporal flexibility and narrow-band spectral data from different wavelengths domains. This enables the detection of spatio-temporal dynamics of environmental variables, like plant-related carbon dynamics in agricultural landscapes. In this paper, we quantify spatial patterns of fresh phytomass and related carbon (C) export using imagery captured by a 12-band multispectral camera mounted on the fixed wing UAV Carolo P360. The study was performed in 2014 at the experimental area CarboZALF-D in NE Germany. From radiometrically corrected and calibrated images of lucerne (Medicago sativa), the performance of four commonly used vegetation indices (VIs) was tested using band combinations of six near-infrared bands. The highest correlation between ground-based measurements of fresh phytomass of lucerne and VIs was obtained for the Enhanced Vegetation Index (EVI) using near-infrared band b899. The resulting map was transformed into dry phytomass and finally upscaled to total C export by harvest. The observed spatial variability at field- and plot-scale could be attributed to small-scale soil heterogeneity in part.

Modeling the Vertical Flux of Organic Carbon in the Global Ocean.

The oceans play a fundamental role in the global carbon cycle, providing a sink for atmospheric carbon. Key to this role is the vertical transport of organic carbon from the surface to the deep ocean. This transport is a product of a diverse range of physical and biogeochemical processes that determine the formation and fate of this material, and in particular how much carbon is sequestered in the deep ocean. Models can be used to both diagnose biogeochemical processes and predict how the various processes will change in the future. Global biogeochemical models use simplified representations of food webs and processes but are converging on values for the export of organic carbon from the surface ocean. Other models concentrate on understanding specific processes and can be used to develop parameterizations for global models. Model development is continuing by adding representations and parameterizations of higher trophic levels and mesopelagic processes, and these are expected to improve model performance.

Phaeocystis: A Global Enigma.

The genus Phaeocystis is globally distributed, with blooms commonly occurring on continental shelves. This unusual phytoplankter has two major morphologies: solitary cells and cells embedded in a gelatinous matrix. Only colonies form blooms. Their large size (commonly 2 mm but up to 3 cm) and mucilaginous envelope allow the colonies to escape predation, but data are inconsistent as to whether colonies are grazed. Cultured Phaeocystis can also inhibit the growth of co-occurring phytoplankton or the feeding of potential grazers. Colonies and solitary cells use nitrate as a nitrogen source, although solitary cells can also grow on ammonium. Phaeocystis colonies might be a major contributor to carbon flux to depth, but in most cases, colonies are rapidly remineralized in the upper 300 m. The occurrence of large Phaeocystis blooms is often associated with environments with low and highly variable light and high nitrate levels, with Phaeocystis antarctica blooms being linked additionally to high iron availability. Emerging results indicate that different clones of Phaeocystis have substantial genetic plasticity, which may explain its appearance in a variety of environments. Given the evidence of Phaeocystis appearing in new systems, this trend will likely continue in the near future.