The global carbon cycle is affected by biological processes in the oceans, which export carbon from surface waters in form of organic matter and store it at depth; a process called the 'biological carbon pump'. Most of the exported organic carbon is processed by the water column biota, which ultimately converts it into CO2 via respiration (remineralization). Variations in the resulting decrease in organic flux with depth 9 can, according to models, lead to changes in atmospheric CO 2 of up to 200 ppm 3 , indicating a strong coupling between biological activity in the ocean interior and oceanic storage of CO 2 .A key constraint in the analysis of carbon fluxes in the twilight zone is that, at steady state, the attenuation of particulate organic carbon (POC) flux with depth should be balanced by community metabolism. Published estimates of POC flux attenuation with depth are, however, up to 2 orders of magnitude lower than corresponding estimates of heterotrophic metabolism [4][5][6][7] . This discrepancy indicates that either estimates of POC flux and/or community metabolism are unreliable, or that additional, unaccounted for, sources of organic carbon to the twilight zone exist 8 .We compiled a comprehensive carbon budget of the twilight zone based on an based on the ratio between DOC concentrations and apparent oxygen utilization 15 , and on DOC gradients coupled to turbulent diffusivity measured from previous work at the study site 16 (Methods; Extended Data Fig. 2). DOC was estimated to supply 17% of total export in agreement with previous estimates of 9-20% across the North Atlantic basin 17 . Organic matter input via lateral advection was assumed to be negligible based on analyses of back-trajectories (derived from satellite-derived near-surface velocities over 3 months) of the water masses arriving at the PAP site during the study period, which suggested that the water had not passed over the continental slope (Extended Data Fig. 1b). The final source of DOC, excretion at depth by active flux, was estimated using net samples of zooplankton biomass and allometric equations 6,18 , giving a supply of 3 mg C m -2 d -1 . Defecation and mortality at depth present further sources of organic carbon to the twilight zone, but these were excluded from the budget due to large uncertainties associated with their estimation. Finally, chemolithoautotrophy has been suggested to be a significant source of organic matter in the deep ocean 19 , but without strong evidence that this poorly understood process could provide a major contribution at our study site, we chose to exclude it from our carbon budget.The remineralization of organic carbon by zooplankton and prokaryotes was estimated from zooplankton biomass and prokaryotic activity. It is crucial to note that in a steady state system, such as we assume this to be, organic carbon is lost from the system only by export or by remineralization. We focus entirely on community respiration as a measure of remineralization, a fundamental advance over previous methods to derive...

In this paper we review the technologies available to make globally quantitative observations of particles in general-and plankton in particular-in the world oceans, and for sizes varying from sub-microns to centimeters. Some of these technologies have been available for years while others have only recently emerged. Use of these technologies is critical to improve understanding of the processes that control abundances, distributions and composition of plankton, provide data necessary to constrain and improve ecosystem and biogeochemical models, and forecast changes in marine ecosystems in light of climate change. In this paper we begin by providing the motivation for plankton observations, quantification and diversity qualification on a global scale. We then expand on the state-of-the-art, detailing a variety of relevant and (mostly) mature technologies and measurements, including bulk measurements of plankton, pigment composition, uses of genomic, optical and acoustical methods as well

Sinking organic particles transfer ∼10 gigatonnes of carbon into the deep ocean each year, keeping the atmospheric CO2 concentration significantly lower than would otherwise be the case. The exact size of this effect is strongly influenced by biological activity in the ocean's twilight zone (∼50–1,000 m beneath the surface). Recent work suggests that the resident zooplankton fragment, rather than ingest, the majority of encountered organic particles, thereby stimulating bacterial proliferation and the deep-ocean microbial food web. Here we speculate that this apparently counterintuitive behaviour is an example of ‘microbial gardening’, a strategy that exploits the enzymatic and biosynthetic capabilities of microorganisms to facilitate the ‘gardener's’ access to a suite of otherwise unavailable compounds that are essential for metazoan life. We demonstrate the potential gains that zooplankton stand to make from microbial gardening using a simple steady state model, and we suggest avenues for future research.