Chlorophyll fluorescence parameters: the definitions, photosynthetic meaning, and mutual relationships

Chlorophyll fluorescence parameters (Chl FPs) derived from the slow (long-term) induction kinetics of modulated Chl a fluorescence are reviewed and analysed with respect to their application in photosynthesis research. Only four mutually independent Chl FPs, calculated from values of five essential Chl fluorescence (ChIF) yields, are distinguished as the basic ones. These are: the maximum quantum yield of PS2 photochemistry (Phi(P0)), the photochemical quenching of variable ChlF (qp), the non-photochemical quenching of variable ChIF (q(N)), and the relative change of minimum ChIF (q(0)). Phi(P0) refers to the dark-adapted state of a thylakoid membrane, qp, qN, and q0 characterise the light-adapted state. It is demonstrated that all other Chi FPs can be determined using this quartet of parameters. Moreover, three FPs related to the non-radiative energy dissipation within thylakoid membranes are evaluated, namely: the non-photochemical ChIF quenching (NPQ), the complete non-photochemical quenching of ChIF (q(CN)), and the effective quantum yield of non-photochemical processes in PS2 (Phi(N)). New FPs, the total quenching of variable ChIF (q(TV)) and the absolute quenching of ChIF (q(A)) which allow to quantify co-action of the photochemical and non-photochemical processes during a light period are defined and analysed. The interpretation of Chi FPs and recommendations for their application in the photosynthesis research are also given. Some alternative FPs used in the laboratory practice have only an approximate character and can lead to incorrect conclusions if applied to stressed plants. They are reviewed and compared with the standard ones. All formulae and conclusions discussed herein are verified using experimental values obtained on young seedlings of the Norway spruce (Picea abies [L.] Karst.).