Inorganic carbon concentrating mechanisms (CCMs) catalyse the accumulation of CO2 around rubisco in all cyanobacteria, most algae and aquatic plants and in C4 and crassulacean acid metabolism (CAM) vascular plants. CCMs are polyphyletic (more than one evolutionary origin) and involve active transport of HCOK3 , CO2 and/or HC, or an energized biochemical mechanism as in C4 and CAM plants. While theCCMin almost all C4 plants and manyCAM plants is constitutive, many CCMs show acclimatory responses to variations in the supply of not only CO2 but also photosynthetically active radiation, nitrogen, phosphorus and iron. The evolution of CCMs is generally considered in the context of decreased CO2 availability, with only a secondary role for increasing O2. However, the earliest CCMs may have evolved in oxygenic cyanobacteria before the atmosphere became oxygenated in stromatolites with diffusion barriers around the cells related to UV screening. This would decrease CO2 availability to cells and increase the O2 concentration within them, inhibiting rubisco and generating reactive oxygen species, including O3.