The recurrent assembly of C4 photosynthesis, an evolutionary tale

Today, plants using C4 photosynthesis are widespread and important components of major tropical and subtropical biomes, but the events that led to their evolution and success started billions of years ago (bya). A CO2-fixing enzyme evolved in the early Earth atmosphere with a tendency to confuse CO2 and O2 molecules. The descendants of early photosynthetic organisms coped with this property in the geological eras that followed through successive fixes, the latest of which is the addition of complex CO2-concentrating mechanisms such as C4 photosynthesis. This trait was assembled from bricks available in C3 ancestors, which were altered to fulfill their new role in C4 photosynthesis. The existence of C4-suitable bricks probably determined the lineages of plants that could make the transition to C4 photosynthesis, highlighting the power of contingency in evolution. Based on the latest findings in C4 research, we present the evolutionary tale of C4 photosynthesis, with a focus on the general evolutionary phenomena that it so wonderfully exemplifies.