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. Chl a / b are essential pigments in plants for harvesting light used in photosynthesis to produce energy, and carotenoids are essential for photoprotection of chloroplast photosystems via nonphotochemical quenching ( Bartley and Scolnik, 1995

harvesters of photons. These energized compounds and the processes they drive (photosynthesis, nonphotochemical quenching) affect metabolism and therefore develop metabolic product states that are sensed and used as information within the cell. Retrograde

nonphotochemical quenching (NPQ) of chlorophyll fluorescence], which possibly limits the formation of chlorophyll triplets and prevents ROS from being produced ( Demmig-Adams and Adams, 1992 ). Lutein and β-carotene are also key components of the light

photosynthetic system II (PSII) ( Huner et al., 1998 ; Lichtenthaler et al., 2005 ): the maximal quantum yield ( F v / F m ) = ( F m − F o )/ F m ; the effective quantum yield (Φ PSII ) = ( F m ′ − F s )/ F m ′; nonphotochemical quenching (qN) = ( F m − F m

quenching (qP), and nonphotochemical quenching (NPQ), were measured using a chlorophyll fluorescence imager (CF Imager; Techologica, Essex, UK). All fluorescence measurements were performed five times following the first NPs treatment, once every 10 d. The