Deep maxima of phytoplankton biomass, primary production and bacterial production in the Mediterranean Sea

The deep chlorophyll maximum (DCM) is a ubiquitous feature of phytoplankton vertical distribution in stratified waters that is relevant to our understanding of the mechanisms that underpin the variability in photoautotroph ecophysiology across environmental gradients and has implications for remote sensing of aquatic productivity. During the PEACETIME (Process studies at the air-sea interface after dust deposition in the Mediterranean Sea) cruise, carried out from 10 May to 11 June 2017, we obtained 23 concurrent vertical profiles of phytoplankton chlorophyll a, carbon biomass and primary production, as well as heterotrophic prokaryotic production, in the western and central Mediterranean basins. Our main aims were to quantify the relative role of photoacclimation and enhanced growth as underlying mechanisms of the DCM and to assess the trophic coupling between phytoplankton and heterotrophic prokaryotic production. We found that the DCM coincided with a maximum in both the biomass and primary production but not in the growth rate of phytoplankton, which averaged 0.3 d−1 and was relatively constant across the euphotic layer. Photoacclimation explained most of the increased chlorophyll a at the DCM, as the ratio of carbon to chlorophyll a (C:Chl a) decreased from ca. 90–100 (g:g) at the surface to 20–30 at the base of the euphotic layer, while phytoplankton carbon biomass increased from ca. 6 mg C m−3 at the surface to 10–15 mg C m−3 at the DCM. As a result of photoacclimation, there was an uncoupling between chlorophyll a-specific and carbon-specific productivity across the euphotic layer. The ratio of fucoxanthin to total chlorophyll a increased markedly with depth, suggesting an increased contribution of diatoms at the DCM. The increased biomass and carbon fixation at the base of the euphotic zone was associated with enhanced rates of heterotrophic prokaryotic activity, which also showed a surface peak linked with warmer temperatures. Considering the phytoplankton