The kinetics of algal photoadaptation in the context of vertical mixing

The responses of phytoplankton to turbulent motions in the surfacemixed layer can be measured to estimate the rate of verticalmixing. If the time scale for the response (photoadaptation)is shorter than that for vertical mixing, phytoplankton willexhibit a vertical gradient associated with adaptation to ambientlight, whereas if mixing occurs with a time scale shorter thanthat of photoadaptation, the surface mixed layer will be uniformwith respect to the photoadaptive parameter. To examine thephysiological bases for a model of vertical mixing and photoadaptation,we grew the marine diatom Thalassiosira pseudonana (clone 3H)at three photon flux densities and subjected the cultures toreciprocal light shifts, measuring physiological and chemicalchanges over the following 10 h. Several parameters, easilymeasured in nature and attributable primarily to phytoplankton,responded to fluctuating light on different time scales. Aftercultures were exposed to relatively bright light, both the initialslope of the photosynthesis-irradiance curve and in vivo fluorescencewere depressed on a time scale of less than an hour. Photosyntheticcapacity was also reduced transiently, but recovered over manyhours to a high level characteristic of an adapted state. First-orderkinetics (the current model of choice for describing photoadaptation)reasonably described the rapid responses of phytoplankton tobright light, but other parameters (i.e. cellular chemical compositionand photosynthetic capacity) changed as a result of unbalancedgrowth and required much longer to adapt from low to high lightas compared to from high to low light. A logistic model of thisadaptation is presented. The model suggests that hysteresisof adaptation during vertical mixing may have important consequences.The vertical distributions of photoadaptive properties in mixedlayers not only reveal the rate of vertical mixing, but showhow phytoplankton integrate environmental fluctuations.