Two C 4 plants, Miscanthus × × × × giganteus and Cyperus longus L., were grown at suboptimal growth temperatures and the relationships between the quantum efficiencies of photosynthetic electron transport through photosystem II (PSII) (PSII operating efficiency; F q ′ ′ ′ ′ / F m ′ ′ ′ ′ ) and CO 2 assimilation ( φ φ φ φ CO2 ) in leaves were examined. When M. × × × × giganteus was grown at 10 °°°° C, the ratio of the PSII operating efficiency to φ φ φ φ CO2 increased relative to that found in leaves grown at 14 and 25 °°°° C. Similar increases in the F q ′ ′ ′ ′ / F m ′ ′ ′ ′ : φ φ φ φ CO2 occurred in the leaves of two C. longus ecotypes when the plants were grown at 17 °°°° C, compared to 25 °°°° C. These elevations of F q ′ ′ ′ ′ / F m ′ ′ ′ ′ : φ φ φ φ CO2 at low growth temperatures were not attributable to the development of anthocyanins, as has been suggested for maize, and were indicative of the operation of an alternative sink to CO 2 assimilation for photosynthetic reducing equivalents, possibly oxygen reduction via a Mehler reaction, which would act as a mechanism for protection of PSII from photoinactivation and damage. Furthermore, in M. × × × × giganteus grown at 10 °°°° C, further protection of PSII was effected by a 20-fold increase in zeaxanthin content in dark-adapted leaves, which was associated with much higher levels of non-photochemical quenching of excitation energy, compared to that observed in leaves grown at 14 and 25 °°°° C. These differences may explain the long growing season and remarkable productivity of this C 4 plant in cool climates, even in comparison to other C 4 species such as C. longus , which occur naturally in such climates.