C4 photosynthesis evolution: the conditional Mt. Fuji

C4 photosynthesis implements a biochemical carbon pump to suppress photorespiration. While this mechanism allows for increased photosynthetic efficiency, it requires the ancestral C3 state to be modified in terms of leaf anatomy, expression of metabolic genes, and enzyme kinetics. Despite the complexity of the C4 syndrome, it evolved in more than 60 independent lineages. Because the phylogenetic distribution of these origins appears to be non-random, enabling factors that are initially unrelated to C4 photosynthesis are assumed to be acting in certain C3 lineages. In recent years, substantial progress has been made on firstly, the nature of enabling events and finally, quantitative models of C4 evolution that are based on C3-C4 intermediate species. I discuss the synthesis of these approaches as a consensus trajectory towards C4 photosynthesis and hypothesize on the effect of enabling factors on the fitness landscape of C4 evolution. A complete understanding of these mechanisms will require both further experimental studies and improved quantitative models of leaf physiology.