The state of plant photosystem II reaction centres affects the rate of non-photochemical quenching

Abstract Plants employ non-photochemical quenching (NPQ) to protect their photosynthetic apparatus from photodamage. The response latency of NPQ following changes in light intensity is thought to significantly decrease photosynthetic efficiency. The amount of NPQ is commonly quantified from chlorophyll-fluorescence techniques using the Stern–Volmer equation, which requires fully closed reaction centres (RCs) of photosystem II, yielding NPQ in the absence of photochemical quenching ( $${\rm{NPQ}}^{\rm{Closed}}$$ NPQ Closed ). However, in nature, NPQ and photochemical quenching are normally present simultaneously. Therefore, to obtain a full understanding of this process, NPQ should also be explored when the RCs are open. Here we developed two methodologies to obtain NPQ in the presence of photochemistry ( $${\rm{NPQ}}^{\rm{Open}}$$ NPQ Open ) using both fluorescence lifetime and fluorescence yield measurements. A detailed comparison in Arabidopsis thaliana plants reveals that the value of $${\mathrm{NPQ}}^{\mathrm{Open}}$$ NPQ Open is ~35% lower than that of $${\rm{NPQ}}^{\rm{Closed}}$$ NPQ Closed . This difference is consistently observed across all measurements and is seen both upon closing ( $${\rm{NPQ}}^{\rm{Open}}\to {\rm{NPQ}}^{\rm{Closed}}$$ NPQ Open → NPQ Closed ) and upon reopening ( $${\mathrm{NPQ}}^{\mathrm{Closed}}\to {\mathrm{NPQ}}^{\mathrm{Open}}$$ NPQ Closed → NPQ Open ) of the RCs. We show that this difference can be explained by the presence of RC-induced ‘instantaneous’ switching of the NPQ quenching rate. This means that, in plants, NPQ is much more economical than is widely believed, it is large when its presence is needed, and it decreases instantaneously when the need disappears.