Inhibition of Photosystems I and II in Chilling-Sensitive and Chilling-Tolerant Plants under Light and Low-Temperature Stress

The responses of photosystems (PS) I and II to light stress at 4 °C and 20 °C were studied in leaf discs from three chilling-sensitive plant species, Cucumis sativus, Cucurbita maxima and Nicotiana tabacum, and in the chilling-tolerant Spinacia oleracea. The chilling-sensitive plants were grown at 24 °C under 8 0 -1 2 0 j.imol photons m-2 s-1 (Cucumis and Cucurbita) or 30 [imol photons m-2 s_1 (Nicotiana). Spinacia was cultivated outdoors during winter and early spring. The P700 absorbance change around 820 nm served as a relative measure of PSI activity. The potential efficiency of PSII was determined in dark-adapted leaf discs by means of the ratio of variable to maximum chlorophyll (Chi) a fluorescence emission (Fv/ F m). In Cucurbita, Nicotiana and Spinacia, PSI was not or only slightly inhibited by 2 h illumination with 200 [imol m-2 s-1 at 4 °C or with 2000 îmol m-2 s-1 at 20 °C. In leaves of Cucurbita and Nicotiana, exposure to 2000 j.imol photons m-2 s_1 at 4 °C resulted in a decline in PSI activity and potential PSII efficiency approximately to the same extent (about 50% within 2 h). In contrast, in Cucumis, both moderate and high light at low temperature caused a PSI inhibition that proceeded considerably faster than the decline in PSII efficiency. Such preferential photoinhibition of PSI was not observed in the other three species tested. In Spinacia, a lower susceptibility of PSI and PSII to photoinhibition at 4 °C was associated with a faster de-epoxidation kinetics of violaxanthin, as compared to the three chilling-sensitive species. In addition, leaves of Spinacia were characterized by a significantly larger pool of xanthophyll-cycle pigments and a higher content of ß-carotene based on Chi a+b. When leaves of Cucurbita were preincubated with methylviologen, which catalyzes formation of superoxide anion radicals at the acceptor side of PSI, the decline in potential PSII efficiency under 2000 jimol photons m-2 s_1 at 20 °C and 4 °C was strongly enhanced, whereas the P700 signal was less affected. Our data demonstrate that in the species tested, PSI may be inhibited in vivo besides PSII under light stress, but preferential photoinhibition of PSI is not a general phenomenon in chilling-sensitive plants. At low temperatures, a reduced function of the xanthophyll cycle and of the antioxidative scavenging system might account for enhanced PSI and PSII inhibition in vivo.