Physiological bases for detecting and predicting photoinhibition of aquatic photosynthesis by PAR and UV radiation

Phytoplankton photosynthesis is the basis of almost all aquatic primary production in the world's oceans, estuaries and lakes. Oceanic primary production is a major portion of the global carbon budget (see other contributions this volume). Currently, we are unable to account for all the CO₂ that is leaving the atmosphere and debate continues whether the ''missing carbon'' is going into either terrestrial and oceanic sinks (7). In this context, it is important to improve our knowledge of how phytoplankton photosynthesis responds to the aquatic environment. The aquatic light environment is primary among several factors governing aquatic photosynthesis. To understand phytoplankton response to aquatic irradiance, we must consider how light propagates underwater, variations in light spectral quality as well as intensity. Also important is how these optical characteristics relate to processes of light absorption and utilization by phytoplankton cells. Considerable progress has been made on answering many of these questions (e.g. 27). One topic, phytoplankton responses to irradiance stress induced by photosynthetically available radiation (PAR2) and UJV, has become increasingly important. The primary consequence in both cases is a time-dependent loss of photosynthetic activity (photo inhibition). Concern over the effects of solar UV irradiance has recently intensified with the advent of stratospheric ozone depletion, which allows for an increase of the mid-ultraviolet (UVB 280-320 nm)irradiance, especially in the Antarctic. The sensitivity of phytoplankton photosynthesis to irradiance stress can be readily demonstrated (36), however,showing whether this stress actually occurs in the aquatic environment remains difficult. The essential problem is that phytoplankton are in suspension. Their irradiance exposure will be determined by mixing processes that transport cells over a vertical gradient in light availability. The response to irradiance stress is usually time dependent (12, 36); the light history of the cells must be known to specify the overall effect. The established method for measuring phytoplankton production, photosynthetic incorporation of ¹⁴C into organic carbon during a12-24 h bottle incubation, may seriously misrepresent irradiance regimes actually experienced by phytoplankton in situ. Further discussion of the interaction of photoinhibition and mixing can be found in (36).We propose that an integrated modeling-sampling approach is needed to define the effects of irradiance stress on productivity in situ. The model should incorporate an optical specification of the underwater irradiance environment, abiological weighting function to account for the wavelength-dependence of photoinhibition of photosynthesis, and a response function of biological action during vertical mixing to account for the differences between static incubations and natural movements of phytoplankton. Recently, excellent progress has been made toward defining the individual components of this model (See also Cullen and Neale, this volume). To verify the model, we need to detect reliably the occurrence of irradiance stress in situ. This is the sampling side of the integrated approach. In particular, we would like to differentiate between the effects of PAR and UV. We propose that such detection can be accomplished by indicator assays (or ''diagnostic markers''). Such assays would involve little or no incubation of samples, so that the measurement corresponds as closely as possible to the physiological state of the phytoplankton at time of sampling.Our objective here is to review selected aspects of irradiance stress at biophysical and molecular levels, and then proceed to examine how that information can be used to design indicator assays of irradiance stress for phytoplankton photosynthesis in situ. The effects of PAR and UV at the cellular level and the use of in vivo fluorescence and molecular probes as detection systems will be discussed