Photosynthesis of a Scots pine shoot: a comparison of two models of shoot photosynthesis in direct and diffuse radiation fields

Two models of shoot photosynthesis, the needle surface element model (SEM) and needle volume element model (VEM), were tested against empirical data obtained from measurements of the photosynthetic response of twelve Scots pine (Pinus sylvestris L.) shoots in direct and diffuse radiation. The models assume that shoot photosynthesis is obtained as the integrated response of either all needle surface area elements (SEM) or all needle volume elements (VEM) of the shoot. The models differ in that needles are treated as optically black in SEM, whereas in VEM radiation penetrates into the needle. The photosynthetic response of a surface/volume element was described as a Blackman-type curve and the distributions of irradiance on the elements were derived by computer simulation, based on a model of shoot geometry. The parameters (initial slope and maximum rate) of the Blackman-curve of an element were estimated iteratively by the method of least squares, i.e., by minimizing the residual sum of squares of simulated and measured rates of shoot photosynthesis. The parameter estimation was done separately for direct and diffuse radiation, and the models were evaluated based on the notion that, for the "ideal" model, the estimated parameter values should be the same in direct and diffuse radiation. Both models produced shoot photosynthesis curves that agreed well with measurements, but there was a discrepancy in the estimated parameter values, indicating that differences in the photosynthetic response of shoots in direct and diffuse radiation could not be explained solely on the basis of the simulated irradiance distributions. The agreement was, however, much better for the volume element model, which accounts for penetration of radiation into the needles.