Fluorescence of chlorophyll a (Chia) is a noninvasive and very sensitive intrinsic probe of photosynthesis. It monitors the composition and orgnization of the photosystems, the exciton energy tfer, the photochemistry, [2] where kp is the bimolecular rate constant for PSII photochemistry.In PSII, light initiates the electron transfer from the primary Chla donor (P680) to the primary electron acceptor pheophytin (Pheo), forming P680+ and Pheo-(5). The latter transfers its electron to QA, which, when reduced, transfers it to the secondary quinone acceptor (QB). Unlike QA, which is a one-electron carrier, QB is a two-electron acceptor (see ref. 6). Plastoquinol (QBH2), generated after a double turnover ofPSII, then exchanges with an oxidized molecule ofthe plastoquinone pool (see review in ref. 7). This two-step process, also called the two-electron gate, leads to a period 2 oscillation in Chla fluorescence yield because electron flow from Qi to QB is faster than that from Q; to Qi (8). On the other hand, P680+, produced in the photochemical reaction ofPSII, transfers its positive charge to a manganese complex (its redox state being labeled as S), via an intermediate, Yz (a tyrosine moiety). Here, however, four positive charges must accumulate before water is oxidized to molecular 02. This leads to a periodicity of 4 in the 02 evolution when measured as a function of flash number (e.g., ref. 9; reviews in refs. 10 and 11).An increase in the Chla fluorescence yield after short flashes in the nanosecond to submicrosecond time range (12) suggested that P680+ is also a quencher of Chla fluorescence (13). To accommodate the quenching characterofboth P680+ and QA, Sonneveld et al. (14,15) assumed that the most fluorescent state of PSII is PQ4 (where P represents P680).With such an assumption, it was easy to explain the dependence ofthe rate ofChla fluorescence-yield changes upon the redox state of not only the acceptor side, but also the donor side, ofP51 (16). Depending on the experimental conditions, one can observe, when plants are exposed to a series of flashes, a periodicity of4 in the fast (microsecond) (17,18) or in the slow (millisecond and second) components of Chla fluorescence yield (18,19) as affected by the four-step charge-accumulating process in the oxygen-evolving complex. The flash number-dependent period 4 oscillation in the Chla fluorescence yield is explained by the different rate of Abbreviations: Chl, chlorophyll; PSII, photosystem II; P680 (or P), electron donor of PSII; QA, primary one-electron quinone acceptor of PSII; QB, secondary two-electron quinone acceptor; Sk, chargeaccumulating state on the donor side of PSI1 (where k is 0, 1, 2, 3, or 4); Y., a tyrosine of PSII that is the electron donor to P680+. tOn leave