Observations of the vertical structure of density, concentrations of chlorophyll a and nitrate, and turbulent dissipation rates were made over a period of 25 h in a well-stratified shelf region in the Western English Channel, between neap and spring tides. Maximum turbulent dissipation at the base of the thermocline occurred almost 5 h after maximum tidal currents. This turbulence aids phytoplankton growth by supplying bottom-layer nutrients into the subsurface chlorophyll maximum but reduces phytoplankton concentrations in the thermocline by mixing cells from the base of the subsurface maximum into the bottom mixed layer. The turbulent dissipation observations were used to estimate an average nitrate flux into the thermocline of 2.0 (0.8-3.2, 95% confidence interval) mmol m, which is estimated to have been capable of supporting new phytoplankton growth at a rate of 160 (64-256) mg C m Ϫ2 d Ϫ1 . Turbulent entrainment of carbon from the base of the subsurface biomass maximum into the bottom mixed layer was observed to be 290 (120-480) mg C m Ϫ2 d Ϫ1 . This apparent excess export from the chlorophyll maximum is suggested to be a feature of the spring-neap cycle, with export dominating as the tidal turbulence increases toward spring tides and erodes the base of the thermocline. The observed rate of carbon export into the bottom mixed layer could account for as much as 25% of the gross annual primary production in stratifying shelf seas. Such turbulent losses, combined with grazing losses and low light levels, suggest that phytoplankton need to be highly adapted to environmental conditions within the thermocline in order to survive.The seasonal thermocline is an important physical barrier in the ocean, separating the surface mixed layer from the deeper water. The stability of the vertical temperature (density) gradient inhibits diapycnal transfer of properties (e.g., momentum, heat, nutrients, algal cells, and oxygen). As the transition region between the nutrient-poor, well-lit surface layer and the darker, nutrient-rich deeper water, the thermocline plays a role in determining the biological properties of the water column. Since the development of continuous fluorescence measurements as a technique for observing the vertical structure of algae (Lorenzen 1966), the thermocline has been observed to be a region of enhanced chlorophyll concentration (e.g., Anderson 1969;Cullen and Eppley 1981;Holligan et al. 1984). Observed concentrations of subsurface chlorophyll range between 0.5 mg mϪ3 in the open ocean up to 100 mg m Ϫ3 in shelf seas. Although in some cases this higher chlorophyll concentration has been shown to be the result of a lower carbon : chlorophyll ratio (Steele 1 Corresponding author (j.sharples@soc.soton.ac.uk).
AcknowledgmentsRay Wilton (University of Wales, Bangor) provided invaluable technical support for FLY. Our thanks to the crew and RVS support staff on RRS Challenger during cruise CH145.