Multiple roles of oxygen in the photoinactivation and dynamic repair of Photosystem II in spinach leaves

Abstract

Oxygen effects have long been ambiguous: exacerbating, being indifferent to, or ameliorating the net photoinactivation of Photosystem II (PS II). We scrutinized the time course of PS II photoinactivation (characterized by rate coefficient k<inf>i</inf>) in the absence of repair, or when recovery (characterized by k<inf>r</inf>) occurred simultaneously in CO<inf>2</inf> ± O<inf>2</inf>. Oxygen exacerbated photoinactivation per se, but alleviated it by mediating the utilization of electrons. With repair permitted, the gradual net loss of functional PS II during illumination of leaves was better described phenomenologically by introducing τ, the time for an initial k<inf>r</inf> to decrease by half. At 1500 μmol photons m−2 s−1, oxygen decreased the initial k<inf>r</inf> but increased τ. Similarly, at even higher irradiance in air, there was a further decrease in the initial k<inf>r</inf> and increase in τ. These observations are consistent with an empirical model that (1) oxygen increased k<inf>i</inf> via oxidative stress but decreased it by mediating the utilization of electrons; and (2) reactive oxygen species stimulated the degradation of photodamaged D1 protein in PS II (characterized by k<inf>d</inf>), but inhibited the de novo synthesis of D1 (characterized by k<inf>s</inf>), and that the balance between these effects determines the net effect of O<inf>2</inf> on PS II functionality.