e-photosynthesis: a comprehensive dynamic mechanistic model of C3 photosynthesis from light capture to sucrose synthesis
Photosynthesis is arguably the most researched of all plant processes. A dynamic model of leaf photosynthesis that includes each discrete process from light capture to carbohydrate synthesis, e-photosynthesis, is described. It was developed by linking and extending our previous models of photosystem II (PSII) energy transfer and photosynthetic C3 carbon metabolism to include electron transfer processes around photosystem I (PSI), ion transfer between the lumen and stroma, ATP synthesis and NADP reduction to provide a complete representation. Different regulatory processes linking the light and dark reactions are also included: Rubisco activation via Rubisco activase, pH and xanthophyll cycle-dependent non-photochemical quenching mechanisms, as well as the regulation of enzyme activities via the ferredoxin-theoredoxin system. Although many further feedback and feedforward controls undoubtedly exist, it is shown that e-photosynthesis effectively mimics the typical kinetics of leaf CO2 uptake, O2 evolution, chlorophyll fluorescence emission, lumen and stromal pH, and membrane potential following perturbations in light, [CO2] and [O2] observed in intact C3 leaves. The model provides a framework for guiding engineering of improved photosynthetic efficiency, for evaluating multiple non-invasive measures used in emerging phenomics facilities, and for quantitative assessment of strengths and weaknesses within the understanding of photosynthesis as an integrated process.