When leaves are exposed to intensities of light in excess of those that can be used with maximum quantum efficiency, the excess energy is dissipated nonphotochemically; this is detected as the quenching of Chl fluorescence and the process is therefore referred to as nonphotochemical quenching, qN (for reviews, see Demmig-Adams and Adams, 1992;Horton and Ruban, 1992;. The heterogeneous nature of qN has been revealed by the study of its relaxation in darkness or after application of inhibitors (Horton and Hague, 1988;Walters and Horton, 1991). A rapidly relaxing component, which in isolated chloroplast relaxes as the pH gradient decays, has been referred to as energy-dependent quenching, qE (Briantais et al., 1979). Relaxation of qE is followed by a number of slowly recovering phases of qN, which include the state transition and processes collectively grouped together under the term qI. This latter quenching process is found especially after exposure to either extreme or prolonged light stress, and it can be essentially irreversible (Demmig and Bjorkman, 1987;Demmig and Winter, 1988).