The energy flux theory 35 years later: formulations and applications

Several models have been proposed for the energetic behavior of the photosynthetic apparatus and a variety of experimental techniques are nowadays available to determine parameters that can quantify this behavior. The Energy Flux Theory (EFT) developed by Strasser 35 years ago provides a straightforward way to formulate any possible energetic communication between any complex arrangement of interconnected pigment systems and any energy transduction by these systems. We here revisit the EFT, starting from the basic general definitions and equations and presenting applications in formulating the energy distribution in photosystem (PS) II units with variable connectivity, as originally derived, where certain simplifications were adopted. We then proceed to the derivation of equations for a PSII model of higher complexity, which corresponds, from the formalistic point of view, to the later formulated and now broadly accepted exciton-radical-pair model. We also compare the formulations derived with the EFT with those obtained, by different approaches, in the classic papers on energetic connectivity. Moreover, we apply the EFT for the evaluation of the excitation energy distribution between PSII and PSI and the distinction between state transitions and PSII to PSI excitation energy migration. Our analysis demonstrates that the EFT is a powerful approach for the formulation of any possible model, at any complexity level, even of models that may be proposed in the future, with the advantage that any possible energetic communication or energy transduction can be easily formulated mathematically by trivial algebraic equations. Moreover, the biophysical parameters introduced by the EFT and applicable for any possible model can be linked with obtainable experimental signals, provided that the theoretical resolution of the model does not go beyond the experimental resolution.