食品生物化学Chapter 21 photosynthesis.pptVIP

21.7 Light-Driven ATP Synthesis—Photophosphorylation The Mechanism of Photophosphorylation Is Chemiosmotic 化学渗透 Experiments with Isolated Chloroplasts Provided the First Direct Evidence for the Chemiosmotic Hypothesis 20.8 The Thermodynamic View of Chemiosmotic Coupling 化学渗透假说 [kemi?zm?tik] Mitchell first described his chemiosmotic hypothesis in 1961. The proton and electrochemical gradients across the inner mitochondrial membrane generated by electron transport represents an enormous source of potential energy. This potential energy is used to synthesize ATP as protons flow back into the matrix. Peter Mitchell was awarded the Nobel Prize in chemistry in 1978 for this discovery. Racker and Stoeckenius Confirmed the Mitchell Model in a Reconstitution 重构Experiment--1974 In this experiment, the bovine mitochondrial ATP synthase was reconstituted in simple lipid vesicles with bacteriorhodopsin, 噬菌调理素 a light-driven proton pump from Halobacterium halobium. Upon illumination, bacteriorhodopsin pumped protons into these vesicles, and the resulting proton gradient was sufficient to drive ATP synthesis by the ATP synthase. Because the only two kinds of proteins present were one that produced a proton gradient and one that used such a gradient to make ATP, this experiment essentially verified Mitchell’s chemiosmotic hypothesis. The mechanism of photophosphorylation is chemiosmotic. In 1966, Jagendorf and Uribe experimentally demonstrated for the first time that establishment of an electrochemical gradient across the membrane of an energy-transducing organelle could lead to ATP synthesis. They equilibrated isolated chloroplasts for 60 seconds in a pH 4 bath, adjusted the pH to 8 in the presence of ADP and Pi, and allowed phosphorylation to proceed for 15 seconds. The entire experiment was carried out in the dark. Collapse of the gradient: the chemical potential of a concentration difference is transduced into synthesis of ATP. CF1CF0 ATP Synthase Is the Chloroplast Equiva