Spillover is the dominant non-photochemical quenching mechanism in angiosperms

Non-photochemical quenching (NPQ) is an important photoprotective process in plants, but all molecular details of the process(es) involved are yet not understood. We have used advanced spectroscopic techniques (including simultaneous time- and spectrally-resolved room-temperature chlorophyll fluorescence analysis and spectro-kinetic deconvolution) to analyse the processes in Arabidopsis, hybrid aspen and Scots pine plants. We used four well-characterized Arabidopsis lines (npq1, npq2, npq4 and L17) affected in NPQ, together with hybrid aspen lines with corresponding modifications that we generated. The data are described best by a model for NPQ induction with up to five fluorescence components representing distinct biochemical entities. A dominant fluorescing species at the end of NPQ induction was identified as functionally detached and quenched LHCII but most importantly we believe that one represents a "PSII-PSI (Photosystem II-Photosystem I) complex" where direct energy transfer between PSII and PSI (spillover) take place. This provides strong quenching in all three plant species. We suggest a new integrated model for NPQ in higher plants where spillover is a major element and suggest roles for PsbS and zeaxanthin. Moreover, we discuss the link between NPQ and thylakoid rearrangements as thylakoid destacking facilitates direct contact between PSII and PSI; a prerequisite for spillover.