Photorespiratory compensation: a driver for biological diversity.

This paper reviews how terrestrial plants reduce photorespiration and thus compensate for its inhibitory effects. As shown in the equation φ = (1/Sc/o )O/C, where φ is the ratio of oxygenation to carboxylation, Sc/o is the relative specificity of Rubisco, O is stromal O2 level and C is the stromal CO2 concentration, plants can reduce photorespiration by increasing Sc/o or C, or by reducing O. By far the most effective means of reducing φ is by concentrating CO2, as occurs in C4 and CAM plants, and to a lesser extent in plants using a glycine shuttle to concentrate CO2 into the bundle sheath. Trapping and refixation of photorespired CO2 by a sheath of chloroplasts around the mesophyll cell periphery in C3 plants also enhances C, particularly at low atmospheric CO2. O2 removal is not practical because high energy and protein investment is needed to have more than a negligible effect. Sc/o enhancement provides for modest reductions in φ, but at the potential cost of limiting the kcat of Rubisco. An effective means of decreasing φ and enhancing carbon gain is to lower leaf temperature by reducing absorbance of solar radiation, or where water is abundant, opening stomata. By using a combination of mechanisms, C3 plants can achieve substantial (>30%) reductions in φ. This may have allowed many C3 species to withstand severe competition from C4 plants in low CO2 atmospheres of recent geological time, thereby preserving some of the Earth's floristic diversity that accumulated over millions of years.