AbstractWe used extensive ecological and biogeochemical measurements obtained from quasi-Lagrangian experiments during two California Current Ecosystem Long-Term Ecosystem Research cruises to analyze carbon fluxes between the epipelagic and mesopelagic zones using a linear inverse ecosystem model (LIEM). Measurement constraints on the model include 14C primary productivity, dilution-based microzooplankton grazing rates, gut pigment-based mesozooplankton grazing rates (on multiple zooplankton size classes), 234Th:238U disequilibrium and sediment trap measured carbon export, and metabolic requirements of micronekton, zooplankton, and bacteria. A likelihood approach (Markov Chain Monte Carlo) was used to estimate the resulting flow uncertainties from a sample of potential flux networks. Results highlight the importance of mesozooplankton active transport (i.e., diel vertical migration) for supplying the carbon demand of mesopelagic organisms and sequestering carbon dioxide from the atmosphere. In nine water parcels ranging from a coastal bloom to offshore oligotrophic conditions, mesozooplankton active transport accounted for 18% - 84% (median: 42%) of the total carbon supply to the mesopelagic, with gravitational settling of POC (12% - 55%; median: 37%) and subduction (2% - 32%; median: 14%) providing the majority of the remainder. Vertically migrating zooplankton contributed to downward carbon flux through respiration and excretion at depth and via consumption loses to predatory zooplankton and mesopelagic fish (e.g. myctophids and gonostomatids). Sensitivity analyses showed that the results of the LIEM were robust to changes in nekton metabolic demands, rates of bacterial production, and mesozooplankton gross growth efficiency. This analysis suggests that prior estimates of zooplankton active transport based on conservative estimates of standard (rather than active) metabolism should be revisited.Contribution to the FieldUnderstanding the flows of carbon within the ocean is important for predicting how global climate will shift; yet even after decades of research, the magnitude with which the ocean sequesters carbon is highly uncertain. One reason behind this uncertainty is that a variety of mechanisms control the balance between carbon input and carbon output within the ocean. The topic of this work is to inspect the role of biological organisms in physically transferring organic carbon from the surface to the deep ocean. As opposed to other mechanisms—such as sinking particles, the biological transfer of carbon is difficult to measure directly and is often quite variable, leading to large uncertainties. Here we use an extensive set of in situ observations off the coast of southern California to model the flow of carbon through the ecosystem. The model determined that in our study area nearly half of the total transfer of carbon from the surface ocean to deep was carried out by zooplankton that swim up to the surface each night to feed. This finding has direct implications for global carbon budgets, which often underestimate this transfer of carbon.