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

   https://doi.org/10.1073/PNAS.2018269118  

   Poff, Kirsten E.; Leu, Andy O.; Eppley, John M.; Karl, David M.; DeLong, Edward F. (January 2021, Proceedings of the National Academy of Sciences)

   null (Ed.)

   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. 

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2. Planktonic microbial signatures of sinking particle export in the open ocean’s interior

   https://doi.org/10.1038/s41467-023-42909-9  

   Li, Fuyan; Burger, Andrew; Eppley, John M; Poff, Kirsten E; Karl, David M; DeLong, Edward F (December 2023, Nature Communications)

   Abstract A considerable amount of particulate carbon produced by oceanic photosynthesis is exported to the deep-sea by the “gravitational pump” (~6.8 to 7.7 Pg C/year), sequestering it from the atmosphere for centuries. How particulate organic carbon (POC) is transformed during export to the deep sea however is not well understood. Here, we report that dominant suspended prokaryotes also found in sinking particles serve as informative tracers of particle export processes. In a three-year time series from oceanographic campaigns in the Pacific Ocean, upper water column relative abundances of suspended prokaryotes entrained in sinking particles decreased exponentially from depths of 75 to 250 m, conforming to known depth-attenuation patterns of carbon, energy, and mass fluxes in the epipelagic zone. Below ~250 m however, the relative abundance of suspended prokaryotes entrained in sinking particles increased with depth. These results indicate that microbial entrainment, colonization, and sinking particle formation are elevated at mesopelagic and bathypelagic depths. Comparison of suspended and sinking particle-associated microbes provides information about the depth-variability of POC export and biotic processes, that is not evident from biogeochemical data alone. 

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3. Sinking particle export within and beneath the euphotic zone in the eastern Indian Ocean

   https://doi.org/10.1101/2025.08.14.670365  

   Stukel, Michael R; Biard, Tristan; Décima, Moira; Fender, Christian K; Kehinde, Opeyemi; Kelly, Thomas B; Kranz, Sven A; Laget, Manon; Landry, Michael R; Yingling, Natalia (August 2025, bioRxiv)

   Abstract The eastern Indian Ocean is substantially under sampled with respect to the biological carbon pump – the suite of processes that transport the carbon fixed by phytoplankton into the deeper ocean. Using sediment traps and other ecosystem measurements, we quantified sinking organic matter flux and investigated the characteristics of sinking particles in waters overlying the Argo Abyssal Plain directly downstream of the Indonesian Throughflow off northwest Australia. Carbon export from the euphotic zone averaged 7.0 mmol C m^-2d^-1, which equated to an average export efficiency (export / net primary production) of 0.17. Sinking particle flux within the euphotic zone (beneath the mixed layer, but above the deep chlorophyll maximum) averaged slightly higher than flux at the base of the euphotic zone, suggesting that the deep euphotic zone was a depth stratum of net particle remineralization. Carbon flux attenuation continued into the twilight zone with a transfer efficiency (export at euphotic depth + 100m / export at euphotic depth) of 0.62 and an average Martin’sb-value of 1.1. Within the euphotic zone, fresh phytoplankton (chlorophyll associated with sinking particles, possibly contained within appendicularian houses) were an important component of sinking particles, but beneath the euphotic zone the fecal pellets of herbivorous zooplankton (phaeopigments) were more important. Changes in carbon and nitrogen isotopic composition with depth further reflected remineralization processes occurring as particles sank. We show similarities with biological carbon pump functioning in a similar semi-enclosed oligotrophic marginal sea, the Gulf of Mexico, including net remineralization across the deep chlorophyll maximum. Submitted to: Deep-sea Research II HighlightsDespite low productivity, export efficiency was 17% of primary productionFlux attenuation beneath the euphotic zone (EZ) was low for a tropical regionSinking particle flux from the upper to lower EZ exceeded export from lower EZThe deep EZ was a stratum of net particle remineralization (and net heterotrophy) 

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4. Particle cycling rates at Station P as estimated from the inversion of POC concentration data

   https://doi.org/10.1525/elementa.2021.00018  

   Amaral, Vinícius J.; Lam, Phoebe J.; Marchal, Olivier; Roca-Martí, Montserrat; Fox, James; Nelson, Norman B. (January 2022, Elementa: Science of the Anthropocene)

   Particle cycling rates in marine systems are difficult to measure directly, but of great interest in understanding how carbon and other elements are distributed throughout the ocean. Here, rates of particle production, aggregation, disaggregation, sinking, remineralization, and transport mediated by zooplankton diel vertical migration were estimated from size-fractionated measurements of particulate organic carbon (POC) concentration collected during the NASA EXport Processes in the Ocean from RemoTe Sensing (EXPORTS) cruise at Station P in summer 2018. POC data were combined with a particle cycling model using an inverse method. Our estimates of the total POC settling flux throughout the water column are consistent with those derived from thorium-234 disequilibrium and sediment traps. A budget for POC in two size fractions, small (1–51 µm) and large (> 51 µm), was produced for both the euphotic zone (0–100 m) and the upper mesopelagic zone (100–500 m). We estimated that POC export at the base of the euphotic zone was 2.2 ± 0.8 mmol m−2 d−1, and that both small and large particles contributed considerably to the total export flux along the water column. The model results indicated that throughout the upper 500 m, remineralization leads to a larger loss of small POC than does aggregation, whereas disaggregation results in a larger loss of large POC than does remineralization. Of the processes explicitly represented in the model, zooplankton diel vertical migration is a larger source of large POC to the upper mesopelagic zone than the convergence of large POC due to particle sinking. Positive model residuals reveal an even larger unidentified source of large POC in the upper mesopelagic zone. Overall, our posterior estimates of particle cycling rate constants do not deviate much from values reported in the literature, i.e., size-fractionated POC concentration data collected at Station P are largely consistent with prior estimates given their uncertainties. Our budget estimates should provide a useful framework for the interpretation of process-specific observations obtained by various research groups in EXPORTS. Applying our inverse method to other systems could provide insight into how different biogeochemical processes affect the cycling of POC in the upper water column. 

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5. Variable particle size distributions reduce the sensitivity of global export flux to climate change

   https://doi.org/10.5194/bg-18-229-2021  

   Leung, Shirley W.; Weber, Thomas; Cram, Jacob A.; Deutsch, Curtis (January 2021, Biogeosciences)

   Abstract. Recent earth system models predict a 10 %–20 % decrease in particulate organic carbon export from the surface ocean by the end of the21st century due to global climate change. This decline is mainly caused by increased stratification of the upper ocean, resulting in reducedshallow subsurface nutrient concentrations and a slower supply of nutrients to the surface euphotic zone in low latitudes. These predictions,however, do not typically account for associated changes in remineralization depths driven by sinking-particle size. Here we combinesatellite-derived export and particle size maps with a simple 3-D global biogeochemical model that resolves dynamic particle size distributions toinvestigate how shifts in particle size may buffer or amplify predicted changes in surface nutrient supply and therefore export production. We showthat higher export rates are empirically correlated with larger sinking particles and presumably larger phytoplankton, particularly in tropical andsubtropical regions. Incorporating these empirical relationships into our global model shows that as circulation slows, a decrease in export isassociated with a shift towards smaller particles, which sink more slowly and are thus remineralized shallower. This shift towards shallowerremineralization in turn leads to greater recycling of nutrients in the upper water column and thus faster nutrient recirculation into the euphoticzone. The end result is a boost in productivity and export that counteracts the initial circulation-driven decreases. This negative feedbackmechanism (termed the particle-size–remineralization feedback) slows export decline over the next century by ∼ 14 % globally (from −0.29to −0.25 GtC yr−1) and by ∼ 20 % in the tropical and subtropical oceans, where export decreases are currently predicted tobe greatest. Our findings suggest that to more accurately predict changes in biological pump strength under a warming climate, earth system modelsshould include dynamic particle-size-dependent remineralization depths. 

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