splitting) of water a process that not only yields electrons, but is also the source of almost all the oxygen in Earth’s atmosphere ‘A molecule of P680 that has given up an energized electron to the primary electron acceptor is positively charged. This P68 ‘molecule isan oxidizing agent so strong that it pulls electrons, away from an oxygen atom that is part of a H,O molecule, Ina reaction probably catalyzed by a unique, manganese-containing enzyme, water is broken into its components: 2 electrons, pro- ‘tons, and oxygen. Bach electron is donated to a P680 molecule, ‘and the protons are released into the thylakoid lumen. Because ‘onygen does not exist in atomic form, the oxygen produced by splitting one HO molecule is written Y2 O,, Two water mol- ccules must be split to yield one molecule of oxygen. The pho tolysis of water isa remarkable reaction, but its name is some- ‘what misleading because it implies that water is broken by light. os. (Oxidation-reducton potential (vot) (lative energy level) 1s Actually, ight splits water indirectly, by causing P680 to become oxidized, Noncyclic electron transport isa continuous linear process In the presence of light, there isa continuous, one-way flow of clectrons from the ultimate electron source, HO, to the termi- nal electron acceptor, NADP*. Water undergoes enzymatically catalyzed photolysis to replace energized electrons donated to the electron transport chain by molecules of P680 in photo. system Il. These electrons travel down the electron transport chain that connects photosystem Il with photosystem I. Thus they provide a continuous supply of replacements for energized clectrons that have been given up by P700. As electrons are transferred along the electron transport chain that connects photosystem II with photosystem I, they lose energy. Some of the energy released is used to pump protons across the thylakoid membrane, from the stroma to the thyla- keoid lumen, producing a proton gradient. The energy ofthis pro- ton gradient is harnessed to produce ATP from ADP by chemi- ‘osmosis, which we will discuss shortly. ATP and NADPH, the products of the lignt-dependent reactions ae released into the stroma, where both are required in the carbon fixation reactions. ACTIVE FIGURE 8414. Noneyelie electron transport. In neneyelic electron transport, the formation of ATP is coupled to the one-way fow of energized electrons (orange arrons) from HO. (lower lft) to NADP* (midle right), Single slatrons actually pase ‘down the electron transport chain; 2 are shown inthis figure becouse 2 electrons are required to form one molecule of NADPH, @) Ele tons are supplied to the system irom the splitting of H,O by pheto- system Il, with the release of O, asa byproduct. When photosystem I 164 Chapter 8 'S activated by absorbing photons, electrons are passed along an electron transport chain and are eventually donated to photosystem | Electrons in photosyster | are "re-energized" by the absorption of additonal ight energy and are passed to NADP’, forming NADPH. BialogyZWow™ Experience the process of noneyelic electron ‘transport by clicking on ths figure on your BiologyNow CD-ROM,