Improved 5-step modeling of the Photosystem II S-state mechanism in cyanobacteria

We present a model of the S-state mechanism, as well as an improved eigenvalue analysis, that integrate into a coherent ensemble several features found since the S-state model was initially developed. These features include the presence of S-1, deactivations in the dark interval between flashes, and the change in the number of active PS II centers by photoinhibition or photoactivation. A new feature is the capacity to predict the steady-state distribution of S-states under conditions of steady photoinhibition or photoactivation. The improved eigenvalue analysis allowed the calculation of the initial S-state distribution. In addition, the model resolved 'true' photochemical misses from apparent misses due to deactivations in the dark interval between flashes. The model suggested that most of the misses that are commonly reported are due to deactivations, and not to an intrinsic inefficiency of the photochemical mechanism of PS II. Because models that allow double-hits encompassing the S2 to S3 transition often predict negative initial quantities of S2 in cyanobacteria, our proposed model specifically prohibited them. The model accounts for inhomogeneous misses and a steady-state distribution of the type (S2)≈(S1)>(S3)≈(S0). This 5-step model uses only 4 probabilities, and is therefore easy to handle. The use of this model is critical for the analysis of several cyanobacterial strains, as well as for any species that show non-negligible deactivations in the dark interval between flashes.