PHOTOINHIBITION OF PHOTOSYNTHESIS REPRESENTS A MECHANISM FOR THE LONG-TERM REGULATION OF PHOTOSYSTEM-II

The obligate shade plant, Tradescantia albiflora Kunth grown at 50-mu-mol photons.m-2 s-1 and Pisum sativum L. acclimated to two photon fluence rates, 50 and 300-mu-mol.m-2.s-1, were exposed to photoinhibitory light conditions of 1700-mu-mol.m-2.s-1 for 4 h at 22-degrees-C. Photosynthesis was assayed by measurement of CO2- saturated O2 evolution, and photosystem II (PSII) was assayed using modulated chlorophyll fluorescence and flash-yield determinations of functional reaction centres. Tradescantia was most sensitive to photoinhibition, while pea grown at 300-mu-mol.m-2.s-1 was most resistant, with pea grown at 50-mu-mol.m-2.s-1 showing an intermediate sensitivity. A very good correlation was found between the decrease of functional PSII reaction centres and both the inhibition of photosynthesis and PSII photochemistry. Photoinhibition caused a decline in the maximum quantum yield for PSII electron transport as determined by the product of photochemical quenching (q(p)) and the yield of open PSII reaction centres as given by the steady-state fluorescence ratio, F'(v)F'(m), according to Genty et al. (1989, Biochim. Biophys. Acta 990, 81-92). The decrease in the quantum yield for PSII electron transport was fully accounted for by a decrease in F'(v)F'(m), since q(p) at a given photon fluence rate was similar for photoinhibited and noninhibited plants. Under light-saturating conditions, the quantum yield of PSII electron transport was similar in photoinhibited and noninhibited plants. The data give support for the view that photoinhibition of the reaction centres of PSII represents a stable, long-term, down-regulation of photochemistry, which occurs in plants under sustained high-light conditions, and replaces part of the regulation usually exerted by the transthylakoid DELTA-pH gradient. Furthermore, by investigating the susceptibility of differently light-acclimated sun and shade species to photoinhibition in relation to q(p), i.e. the fraction of open-to-closed PSII reaction centres, we also show that irrespective of light acclimation, plants become susceptible to photoinhibition when the majority of their PSII reaction centres are still open (i.e. primary quinone acceptor oxidized). Photoinhibition appears to be an unavoidable consequence of PSII function when light causes sustained closure of more than 40% of PSII reaction centres.