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The biological carbon pump, driven principally by the surface production of sinking organic matter and its subsequent remineralization to carbon dioxide (CO2) in the deep ocean, maintains atmospheric CO2 concentrations around 200 ppm lower than they would be if the ocean were abiotic. One important driver of the magnitude of this effect is the depth to which organic matter sinks before it is remineralised, a parameter we have limited confidence in measuring given the difficulty involved in balancing sources and sinks in the ocean's interior. One solution to this imbalance might be a temporal offset in which organic carbon accumulates in the mesopelagic zone (100–1000 m depth) early in the productive season before it is consumed later. Here, we develop a novel accounting method to address non-steady state conditions by estimating fluxes of particulate organic matter into, and accumulation within, distinct vertical layers in the mesopelagic zone using high-resolution spatiotemporal vertical profiles. We apply this approach to a time series of measurements made during the declining phase of a large diatom bloom in a low-circulation region of the Southern Ocean downstream of South Georgia. Our data show that the major export event led to a significant accumulation of organic matter in the upper mesopelagic zone (100–200 m depth) which declined over the following weeks, implying that temporal offsets need to be considered when compiling budgets. However, even when accounting for this accumulation, a mismatch in the vertically resolved organic carbon budget remained, implying that there are likely widespread processes that we do not yet understand that redistribute material vertically within the mesopelagic zone.

Through international collaborations and advances in technology, ocean observatories are increasingly capable of monitoring over long time periods. The Porcupine Abyssal Plain Sustained Observatory (PAP–SO), located at 4,850 m depth in the Northeast Atlantic, is one of a small number of oceanic sites that has achieved monitoring to full ocean depths over several decades. It has monitored seafloor ecology since 1985, water column particle flux since 1992, and surface ocean and atmosphere parameters since 2003. The observatory is serviced annually, providing the opportunity to carry out conventional ship-based observations, sensor comparison, and sampling.

The traditional separation between primary producers (autotrophs) and consumers (heterotrophs) at the base of the marine food web is being increasingly replaced by the paradigm that mixoplankton, planktonic protists with the nutritional ability to use both phago(hetero)trophy and photo(auto)trophy to access energy are widespread globally. Thus, many ‘phytoplankton’ eat, while 50% of ‘protozooplankton’ also perform photosynthesis. Mixotrophy may enhance primary production, biomass transfer to higher trophic levels and the efficiency of the biological pump to sequester atmospheric CO2 into the deep ocean. Although this view is gaining traction, science lacks a tool to quantify the relative contributions of autotrophy and heterotrophy in planktonic protists. This hinders our understanding of their impacts on carbon cycling within marine pelagic ecosystems. It has been shown that the hydrogen (H) isotopic signature of lipids is uniquely sensitive to heterotrophy relative to autotrophy in plants and bacteria. Here, we explored whether it is also sensitive to the trophic status in protists. The new understanding of H isotope signature of lipid biomarkers suggests it offers great potential as a novel tool for quantifying the prevalence of mixotrophy in diverse marine microorganisms and thus for investigating the implications of the ‘mixoplankton’ paradigm.

Our understanding of lipid biogeochemistry of the ocean’s interior is still in its infancy. Here we focus on early lipid transformation and the formation of lipid degradation products in the NE Atlantic Ocean (49.0°N, 16.5°W). We employed high resolution Fourier transform ion cyclotron resonance mass spectrometry (FT-ICR MS), a method that allows observation and elemental composition assignment of thousands of lipids in a single sample. Using these data, we infer molecular-level changes that occur during lipid transformation in the oceanic water column to shed new light on early lipid transformation processes and the formation of lipid decomposition intermediates, here termed CHO compounds (i.e., lipid-derived species that contain carbon, hydrogen and oxygen in their molecular formula). We considered the distribution of molecular rings and/or double bonds (DBE), H/C and O/C ratios, molecular diversity based on the number of mass spectral signals for monoisotopic species, and carbon number in CHO molecules. Data are elaborated for the four ocean zones, the epipelagic, mesopelagic, bathypelagic and abyssal. The highest molecular diversity characterizes CHO compounds associated with the epipelagic zone, which is explained by numerous and diverse planktonic communities inhabiting the epipelagic and by the effects of both biotic and abiotic processes on lipid transformation. Lipid transformations include crosslinking (condensation), partial degradation or fragmentation, double bond reduction, oxidation, hydrogenation, dehydrogenation and cyclization. Crosslinking likely results in a unimodal distribution of carbon number of CHO compounds, in contrast to cell lipids (referred to as Reported lipids based on the Lipid Maps Database), which have a bimodal distribution of carbon number. CHO compounds that appear to be formed by fragmentation (decrease of the number of C atoms) and ring/double bond reduction were more stable to further transformation and remained longer in the

© The Author(s), 2021. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Cael, B. B., Bisson, K., Conte, M., Duret, M. T., Follett, C. L., Henson, S. A., Honda, M. C., Iversen, M. H., Karl, D. M., Lampitt, R. S., Mouw, C. B., Muller-Karger, F., Pebody, C. A., Smith, K. L., & Talmy, D. Open ocean particle flux variability from surface to seafloor. Geophysical Research Letters, 48(9), (2021): e2021GL092895, https://doi.org/10.1029/2021GL092895., The sinking of carbon fixed via net primary production (NPP) into the ocean interior is an important part of marine biogeochemical cycles. NPP measurements follow a log-normal probability distribution, meaning NPP variations can be simply described by two parameters despite NPP's complexity. By analyzing a global database of open ocean particle fluxes, we show that this log-normal probability distribution propagates into the variations of near-seafloor fluxes of particulate organic carbon (POC), calcium carbonate, and opal. Deep-sea particle fluxes at subtropical and temperate time-series sites follow the same log-normal probability distribution, strongly suggesting the log-normal description is robust and applies on multiple scales. This log-normality implies that 29% of the highest measurements are responsible for 71% of the total near-seafloor POC flux. We discuss possible causes for the dampening of variability from NPP to deep-sea POC flux, and present an updated relationship predicting POC flux from mineral flux and depth., B. B. Cael and S. A. Henson acknowledge support from the National Environmental Research Council (NE/R015953/1) and the Horizon 2020 Framework Programme (820989, project COMFORT). The work reflects only the authors' views; the European Commission and their executive agency are not responsible for any use that may be made of the information the work contains. S. A. Henson also acknowledges support from a European Research Council Consolidator grant (GOCART, agreement number 724416). C. L. Follett acknowledges support from the Simons Foundation (grants #827829 and #553242). M. H. Iversen acknowledges support from the DFG-Research Center/Cluster of Excellence “The Ocean Floor – Earth's Uncharted Interface”: EXC-2077-390741603 and the HGF Young Investigator Group SeaPump “Seasonal and regional food web interactions with the biological pump”: VH-NG-1000. M. C. Honda acknowledges financial support from the Ministry of Education, Culture, Sports, Science, and Technology – Japan (grants #: KAKENHI JP18H04144 and JP19H05667). M. Conte acknowledges support from the US National Science Foundation, Division of Ocean Sciences for support for the Oceanic Flux Program time-series since inception, most recently by NSF OCE grant 1829885. D. M. Karl acknowledges support from the Gordon and Betty Moore Foundation (#3794) and the Simons Foundation (SCOPE #329108).

The Joint Global Ocean Flux Study (JGOFS) started in 1987 and stimulated massive integration of efforts to measure and understand the processes of downward flux in the ocean. A small number of sustained observatories have persisted from this time in the belief that a key to gaining this understanding is by prolonged time-series observations. The sustained observatory over the Porcupine Abyssal Plain (PAP-SO) in the Northeast Atlantic is one such site which has provided nearly continuous measurements of downward flux at a depth of 3000m since 1989. This was using sediment traps but, in order to understand the data, we have exploited a wide range of other approaches such as optical methods, drifting sediment traps in the upper 1000m, chemical variables near the surface and computational modelling. Insights are also gained from the Continuous Plankton Recorder (CPR), satellite observations and various climatic indices. This presentation draws together these measurements to quantify and understand flux at 3000m. Seasonal and interannual variability is large but after 30 years of observation, explanations are now possible. In addition, some of the conclusions identify major and surprising features about the ways ecosystems in one year may be influenced by their structure and function in previous years. At the same time as these time-series observations have been in progress, major new developments have taken place globally and at PAP-SO to provide additional and alternative means to asses flux. The sediment trap has significant advantages as well as uncertainties which have been described previously. The new era of approaches using, for instance, BGC Argo, the Carbon Flux Explorer and a variety of other optical techniques offer what may be a quantum leap in our understanding of downward particle flux now and how it is changing in response to changes in the global climate. This presentation will give a personal and optimistic view of the opportunities which are now develo

The sinking of carbon fixed via net primary production (NPP) into the ocean interior is an important part of marine biogeochemical cycles. NPP measurements follow a log‐normal probability distribution, meaning NPP variations can be simply described by two parameters despite NPP’s complexity. By analyzing a global database of open ocean particle fluxes, we show that this log‐normal probability distribution propagates into the variations of near‐seafloor fluxes of particulate organic carbon (POC), calcium carbonate, and opal. Deep‐sea particle fluxes at subtropical and temperate time‐series sites follow the same log‐normal probability distribution, strongly suggesting the log‐normal description is robust and applies on multiple scales. This log‐normality implies that 29% of the highest measurements are responsible for 71% of the total near‐seafloor POC flux. We discuss possible causes for the dampening of variability from NPP to deep‐sea POC flux, and present an updated relationship predicting POC flux from mineral flux and depth.

Until the 1980s, the deep sea was generally considered to be a particularly stable environment, free from major temporal variations (Sanders, 1968). Studies in the abyssal northeast Atlantic by Billett et al. (1983), and subsequently Lampitt (1985) discovered seasonal pulses of surface primary production-derived particulate organic matter (phytodetritus), and hence carbon, at abyssal depths. These early observations were subsequently extended to the central oceanic region of the NE Atlantic (Pfannkuche, 1993; Thiel et al., 1989), and prompted the establishment of more concerted time series studies in the Porcupine Abyssal Plain area. Today, the Porcupine Abyssal Plain Sustained Observatory (PAP–SO) is a multidisciplinary open-ocean time series site in the NE Atlantic (48°50′N 16°30′W, 4850 m water depth; Fig. 1), focused on the study of connections between the surface and deep ocean. In situ measurements of climatically and environmentally relevant variables have been made for more than 30 years. This represents an exceptionally long time series - a recent compilation of biological time series data, across terrestrial, freshwater, and marine realms, indicates an average duration of only 13-years (Dornelas et al., 2018). Long-term time series in the deep sea are rare, particularly those collecting data from surface to seabed. The PAP-SO is one of two abyssal long-term time series sites globally (Smith et al. 2015), the other being a thirty-year time series at Station M in the northeastern Pacific Ocean (34°50′N, 123°00′W, ~4000 m water depth), maintained by the Monterey Bay Aquarium Research Institute (Smith et al., 2020). This ‘sibling’ abyssal time series site also aims to understand the connections between the surface ocean and the seabed, using many similar techniques (Smith et al., 2017), facilitating comparisons between the two sites (e.g. Durden et al., 2019; Durden et al., 2020a; Laguionie-Marchais et al., 2013; Smith et al., 2009). Another source of extended

The sinking of carbon fixed via net primary production (NPP) into the ocean interior is an important part of marine biogeochemical cycles. NPP measurements follow a log-normal probability distribution, meaning NPP variations can be simply described by two parameters despite NPP's complexity. By analyzing a global database of open ocean particle fluxes, we show that this log-normal probability distribution propagates into the variations of near-seafloor fluxes of particulate organic carbon (POC), calcium carbonate, and opal. Deep-sea particle fluxes at subtropical and temperate time-series sites follow the same log-normal probability distribution, strongly suggesting the log-normal description is robust and applies on multiple scales. This log-normality implies that 29% of the highest measurements are responsible for 71% of the total near-seafloor POC flux. We discuss possible causes for the dampening of variability from NPP to deep-sea POC flux, and present an updated relationship predicting POC flux from mineral flux and depth.

The sinking of carbon fixed via net primary production (NPP) into the ocean interior is an important part of marine biogeochemical cycles. NPP measurements follow a log-normal probability distribution, meaning NPP variations can be simply described by two parameters despite NPP's complexity. By analyzing a global database of open ocean particle fluxes, we show that this log-normal probability distribution propagates into the variations of near-seafloor fluxes of particulate organic carbon (POC), calcium carbonate, and opal. Deep-sea particle fluxes at subtropical and temperate time-series sites follow the same log-normal probability distribution, strongly suggesting the log-normal description is robust and applies on multiple scales. This log-normality implies that 29% of the highest measurements are responsible for 71% of the total near-seafloor POC flux. We discuss possible causes for the dampening of variability from NPP to deep-sea POC flux, and present an updated relationship predicting POC flux from mineral flux and depth.

The North Atlantic Ocean is a unique area that has regular occurrences of storm events and mesoscale structures (eddies, fronts). These physical forcings are known to promote episodic fluxes of macronutrients from the deep waters to the oligotrophic photic zone. This fertilizes the surface waters and promotes higher primary production, which in turn stimulates the trophic food web and biogeochemical fluxes. However, the character and magnitude of the biological response vary according to the physics of the mesoscale structure (front, eddy type, wind stress, duration). Here we present an interdisciplinary data set (concentrations of nutrients and chlorophyll a, zooplankton biomass and community structure, aggregate abundance and sinking velocity) that was collected in a cyclonic eddy, which was further impacted by a strong storm event at the Porcupine Abyssal Plain (PAP). We observed two nutrient intrusion events with a 2-fold increase in the surface ocean nutrient concentrations. This increase in nutrients had biological responses ranging from increased chlorophyll to elevated aggregate abundance and export out of the photic zone. At the same time, we also observed changes in the zooplankton community. We will discuss the importance of the mesoscale structures on the biological carbon pump in the open ocean.