image
FIGURE 12-44 Linear electron flow in plants, which requires both chloroplast photosystems, PSI and PSII. Blue arrows indicate flow of electrons; red arrows indicate proton movement. LHCs are not shown. (Left) In the PSII reaction center, two sequential light-induced excitations of the same special-pair P680 chlorophyll result in a two-step reduction of the primary electron acceptor QB to QH2. On the luminal side of PSII, electrons removed from H2O by the oxygen-evolving complex are transferred to P680+, restoring the reaction-center chlorophylls to the ground state after each excitation. The oxygen-evolving complex contains a cluster of four manganese ions (Mn, violet), a Ca2+ ion (green), and a Cl ion (teal). These bound ions function in the splitting of H2O and maintain the environment essential for high rates of O2 evolution. Four sequential light-induced excitations of the P680 (double the number illustrated here) are required to oxidize two water molecules and release four protons and one molecule of molecular oxygen (O2). A tyrosine on the protein helps conducts electrons from the Mn ions to the oxidized reaction-center chlorophyll (P680+), reducing it to the ground state (P680) after excitation by each photon. (Center) The cytochrome bf complex then accepts electrons from QH2 and transports two protons into the lumen. Operation of a Q cycle in the cytochrome bf complex translocates additional protons across the membrane to the thylakoid lumen, increasing the proton-motive force. (Right) In the PSI reaction center, each electron released from light-excited P700 chlorophylls moves via a series of carriers in the reaction center to the stromal surface, where soluble ferredoxin (an Fe-S protein) transfers the electron to ferredoxin-NADP+ reductase (FNR). This enzyme uses the prosthetic group flavin adenine dinucleotide (FAD) and a proton to reduce NADP+, forming NADPH. P700+ is restored to its ground state by addition of an electron carried from PSII via the cytochrome bf complex and plastocyanin, a soluble electron carrier.