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1. 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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2. Strong and efficient summertime carbon export driven by aggregation processes in a subarctic coastal ecosystem

   https://doi.org/10.1002/lno.12561  

   O'Daly, Stephanie H; Hennon, Gwenn_M M; Kelly, Thomas B; Strom, Suzanne L; McDonnell, Andrew_M_P (May 2024, Limnology and Oceanography)

   Sinking marine particles, one pathway of the biological carbon pump, transports carbon to the deep ocean from the surface, thereby modulating atmospheric carbon dioxide and supplying benthic food. Few in situ measurements exist of sinking particles in the Northern Gulf of Alaska; therefore, regional carbon flux prediction is poorly constrained. In this study, we (1) characterize the strength and efficiency of the biological carbon pump and (2) identify drivers of carbon flux in the Northern Gulf of Alaska. We deployed up to five inline drifting sediment traps in the upper 150 m to simultaneously collect bulk carbon and intact sinking particles in polyacrylamide gels and measured net primary productivity from deck‐board incubations during the summer of 2019. We found high carbon flux magnitude, low attenuation with depth, and high export efficiency. We quantitatively attributed carbon flux between 10 particle types, including various fecal pellet categories, dense detritus, and aggregates using polyacrylamide gels. The contribution of aggregates to total carbon flux (41–93%) and total carbon flux variability (95%) suggest that aggregation processes, not zooplankton repackaging, played a dominant role in carbon export. Furthermore, export efficiency correlated significantly with the proportion of chlorophyllain the large size fraction (> 20 μm), total aggregate carbon flux, and contribution of aggregates to total carbon flux. These results suggest that this stratified, small‐cell‐dominated ecosystem can have sufficient aggregation to allow for a strong and efficient biological carbon pump. This is the first integrative description of the biological carbon pump in this region. 

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3. Sinking carbon, nitrogen, and pigment flux within and beneath the euphotic zone in the oligotrophic, open-ocean Gulf of Mexico

   https://doi.org/10.1093/plankt/fbab001  

   Stukel, Michael R; Kelly, Thomas B; Landry, Michael R; Selph, Karen E; Swalethorp, Rasmus (February 2021, Journal of Plankton Research)

   Dolan, John (Ed.)

   Abstract During two cruises in the oligotrophic oceanic Gulf of Mexico, we deployed sediment traps at three depths: center of the euphotic zone (EZ) (60 m), base of the EZ (117–151 m), and in the twilight zone (231 m). Organic carbon export declined with depth from 6.4 to 4.6 to 2.4 mmol C m−2 d−1, suggesting that net particle production was concentrated in the upper EZ. Net primary production varied from 24 to 29 mmol C m−2 d−1, slightly more than half in the upper EZ. Export ratios varied from 11 to 25%. Trap measurements of chlorophyll and phaeopigments allowed us to quantify fluxes of fresh phytoplankton and herbivorous fecal pellets, respectively, which were both minor contributors to total flux, although their contributions varied with depth. Phytoplankton flux was more important from the upper to lower EZ; fecal pellets were more important at the EZ base and below. C:N elemental ratios and 13C and 15N isotope analyses indicated particle transformations within and beneath the EZ. 234Th-238U disequilibrium measurements varied, likely reflecting the mixing of water from multiple regions over the ~month-long time-scale of 234Th. Our results highlight the complexity of the biological carbon pump in oligotrophic regions. 

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4. The molecular-level diagenetic clock of sinking marine organic matter

   https://doi.org/10.1073/pnas.2504769122  

   Forrer, Heather J; Stukel, Michael R; McKenna, Amy M; Chen, Huan; Holt, Amy D; Kranz, Sven A; Spencer, Robert G_M (December 2025, Proceedings of the National Academy of Sciences)

   Thiemens, Mark (Ed.)

   The marine biological carbon pump is driven by sinking particulate organic matter (POM). Sinking speed and remineralization rate determine flux attenuation in the mesopelagic. Since the fate of all marine organic matter is either complete remineralization or transformation to more stable products, diagenetic modifications impact carbon dioxide sequestration time from the atmosphere. To investigate particle transformation at the molecular level, we characterize the water-extractable organic matter (WEOM) fraction of sinking particles from dominant biogeochemical environments using ultrahigh-resolution mass spectrometry. We find distinct, inverse associations in molecular-level nitrogen content and degree of transformation (i.e., “stability”) of organic matter across a productivity gradient from coastal upwelling to oligotrophic conditions. Nitrogen enrichment and low stability were observed at the coastal upwelling site and persisted to depths >400 m. Further, carbon flux is strongly correlated with the relative abundance of stable WEOM (“Island of Stability” molecular formulae) across productivity regimes and depth. This suggests emergent patterns in epi- and mesopelagic diagenesis, highlighting that the molecular composition of sinking organic matter exiting the euphotic zone varies more across regions than as a function of depth. This is attributed to highly variable sinking rates and the microbial diagenetic histories within the euphotic zone. The stability–flux relationship is considered a “diagenetic clock” relative to organic matter formation where the relative abundance of Island of Stability molecular formulae describes the degree of departure from the organic matter molecular-level composition at formation. This ubiquitous trajectory of the diagenetic clock further underpins a global ocean molecular signature of sinking POM. 

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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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