Fast cyclic electron transport around photosystem I in leaves under far-red light: a proton-uncoupled pathway?

Fast cyclic electron transport (CET) around photosystem I (PS I) was observed in sunflower (Helianthus annuus L.) leaves under intense far-red light (FRL) of up to 200 mumol quanta m(-2) s(-1). The electron transport rate (ETR) through PS I was found from the FRL-dark transmittance change at 810 and 950 nm, which was deconvoluted into redox states and pool sizes of P700, plastocyanin (PC) and cytochrome f (Cyt f). PC and P700 were in redox equilibrium with K(e) = 35 (ΔE(m) = 90 mV). PS II ETR was based on O(2) evolution. CET [(PS I ETR) - (PS II ETR)] increased to 50-70 mumol e(-) m(-2) s(-1) when linear electron transport (LET) under FRL was limited to 5 mumol e(-) m(-2) s(-1) in a gas phase containing 20-40 mumol CO(2) mol(-1) and 20 mumol O(2) mol(-1). Under these conditions, pulse-saturated fluorescence yield F(m) was non-photochemically quenched; however, F(m) was similarly quenched when LET was driven by low green or white light, which energetically precluded the possibility for active CET. We suggest that under FRL, CET is rather not coupled to transmembrane proton translocation than the CET-coupled protons are short-circuited via proton channels regulated to open at high ΔpH. A kinetic analysis of CET electron donors and acceptors suggests the CET pathway is that of the reversed Q-cycle: Fd -> (FNR) -> Cyt c(n) -> Cyt b(h) -> Cyt b(l) -> Rieske FeS -> Cyt f -> PC -> P700 ->-> Fd. CET is activated when PQH(2) oxidation is opposed by high ΔpH, and ferredoxin (Fd) is reduced due to low availability of e(-) acceptors. The physiological significance of CET may be photoprotective, as CET may be regarded as a mechanism of energy dissipation under stress conditions.