Relative activities of linear and cyclic electron flows during chloroplast CO2-fixation.

Evolution of oxygen and turnover of cytochromes b-563 and f were measured upon illumination of isolated intact spinach chloroplasts with a series of flashes. The flash yield of cytochrome f oxidation approximated the sum of the yields of cytochrome b-563 reduction and electron transfer through Photosystem II, regardless of whether HCO(-3), 3-phosphoglycerate or O2 served as the terminal electron acceptor. No absorbance contribution form cytochrome b-559 was discerned within the time range studied. Some pseudocyclic electron flow occurred when both HCO(-3) and 3-phosphoglycerate were omitted, and possibly also during induction of photosynthesis; however, the flash yield data suggest tht O2 is not reduced at a significant rate during steady state photosynthesis. The maximum rate of cytochrome f turnover (1000 microequiv./mg chlorophyll per h) was adequate to support the highest rates of photosynthesis observed in isolated chloroplasts. These results agree with the concept that cytochrome f is a component both of the linear and cyclic pathways whereas cytochrome b-563 functions only in the cyclic pathway. NH4Cl decreased the half time of cytochrome b-563 oxidation fro 11.6 to 8.2 ms and decreased the half time of cytochrome f reduction from 7.2 to 2.8 ms. The cyclic and linear pathways thus seem to be jointly regulated by a transthylakoid H+ gradient through a common control point on the reducing side of cytochrome f. Cyclic turnover also increased during the induction phase of photosynthesis, when linear throughput is limited by the rate of utilization of NADPH. The slow rise in the P-518 transient correlated with increased cyclic activity under the above conditions. It is proposed that flexibility in the utilization of linear and cyclic pathways allows the chloroplast to generate ATP and NADPH in ratios appropriate to varying needs.