Variations in Chlorophyll Fluorescence Yields in Phytoplankton in the World Oceans

Abstract

The ocean is optically thin and lends itself to large-scale measurements of in vivo chlorophyll fluorescence. In the open ocean, however, phytoplankton chlorophyll concentrations average only 0.2 μg L^-1, and hence high sensitivity is required for precise measurements of the fluorescence yields. Over the past decade, we have developed two approaches to achieve the required sensitivity; these are the pump- and probe-technique and a fast repetition rate (FRR) method. Both methods have been adapted for in situ studies and are used to rapidly measure the maximum change in the quantum yield (ΔØ_max) of photosystem II (PSII), as well as the effective absorption cross-section of PSII (σ_PSII). Sections of variable fluorescence across the Pacific and Atlantic Oceans reveal the influence of geophysical processes in controlling the quantum yields of phytoplankton photosynthesis. Areas of upwelling, such as off the coast of north-westem Africa, have F_v/F_m values of 0.65, which are close to the maximum achievable values in nutrient-replete cultures. Throughout most of the nutrient-deficient central ocean basins, this quantum efficiency is reduced by more than 50%. In high-nutrient, low- chlorophyll regions of the eastern Equatorial Pacific, the deliberate, large-scale addition of nanomolar iron directly to the ocean leads to a rapid increase in quantum efficiency of the natural phytoplankton community, thereby revealing that in these regions phytoplankton photosynthetic energy conversion efficiency is iron limited. Diel patterns of variation in the upper ocean display midday, intensity- dependent reductions in both upsII and AØmax. We interpret the former as indicative of non- photochemical quenching in the antenna, while the latter is a consequence of both rapidly reversible and slowly reversible damage to reaction centres. From knowledge of the incident spectral irradiance, ΔØ_max, σ_PSII, and photochemical quenching, the absolute photosynthetic electron transport rate can be derived in real-time. Using unattended, moored continuous measurements of in vivo fluorescence parameters, the derived in situ electron transport rates can be related to satellite observations of the global ocean with basin-scale, seasonal estimates of phytoplankton carbon fixation. Thus, unlike any other photosynthetic parameter, chlorophyll fluorescence can be used to bridge the scales of biophysical responses to ecosystem dynamics.