Binary oscillations in the Kok model of oxygen evolution in oxygenic photosynthesis

Abstract

The flash-induced kinetics of various characteristics of Photosystem II (PS II) in the thylakoids of oxygenic plants are modulated by a period of two, due to the function of a two-electron gate in the electron acceptor side, and by a period of four, due to the changes in the state of the oxygen-evolving complex. In the absence of inhibitors of PS II, the assignment of measured signal to the oxygen-evolving complex or to quinone acceptor side has frequently been done on the basis of the periodicity of its flash-induced oscillations, i.e. four or two. However, in some circumstances, the period four oscillatory processes of the donor side of PS II can generate period two oscillations. It is shown here that in the Kok model of oxygen evolution (equal misses and equal double hits), the sum of the concentrations of the S ₀ and S ₂ states (as well as the sum of concentrations of S ₁ and S ₃ states) oscillates with period of two: S ₀+S ₂→S ₁+S ₃→S ₀+S ₂→S ₁+S ₃. Moreover, in the generalized Kok model (with specific miss factors and double hits for each S-state) there always exist such ε₀, ε₁, ε₂, ε₃ that the sum ε₀[S₀] + ε₁[S₁] + ε₂[S₂] + ε₃[S₃] oscillates with period of two as a function of flash number. Any other coefficients which are linearly connected with these coefficients, % MathType!MTEF!2!1!+-% feaafiart1ev1aaatCvAUfeBSjuyZL2yd9gzLbvyNv2CaerbuLwBLn% hiov2DGi1BTfMBaeXatLxBI9gBaerbd9wDYLwzYbItLDharqqtubsr% 4rNCHbGeaGak0dh9WrFfpC0xh9vqqj-hEeeu0xXdbba9frFj0-OqFf% ea0dXdd9vqaq-JfrVkFHe9pgea0dXdar-Jb9hs0dXdbPYxe9vr0-vr% 0-vqpWqaaeaabiGaciaacaqabeaadaqaaqaaaOqaaiqbew7aLzaaja% aaaa!3917!\[\hat \varepsilon \]_i = c₁ε_i + c₂, also generate binary oscillations of this sum. Therefore, the decomposition of the flash-induced oscillations of some measured parameters into binary oscillations, depending only on the acceptor side of PS II, and quaternary oscillations, depending only on the donor side of PS II, becomes practically impossible when measured with techniques (such as fluorescence of chlorophyll a, delayed fluorescence, electrochromic shift, transmembrane electrical potential, changes of pH and others) that could not spectrally distinguish the donor and acceptor sides. This property of the Kok cycle puts limits on the simultaneous analysis of the donor and acceptor sides of the RC of PS II in vivo and suggests that binary oscillations are no longer a certain indicator of the origin of a signal in the acceptor side of PS II.