THE LIGHT-DEPENDENT REACTIONS Learning Objectives [B] Describe the flow of electrons through photosystems and In the noneyelc electron transport pathway; contrast this wth eyclic electron transport. Explain how a proton (H*) gradient is established across the thylakoid membrane and how thi oradiont functions in ATP synthesis. In the light-dependent reactions, the radiant energy from sun- light phosphorylates ADP, producing ATP, and reduces NADP*, forming NADPH. The light energy that chlorophyll captures is temporarily stored in these two compounds. The light-dependent reactions are summarized as follows: Light Chorepbril 60, + 12 NADPH+ 18 ATP 12 HO + 12 NADF* + 18 ADP+ 18 Py Photosystems | and II each consist of a reaction center and multiple antenna complexes ‘The light-dependent reactions of photosynthesis begin when. chlorophyll « and/or accessory pigments absorb light. Accord- ing to the currently accepted model, chlorophylls @and band accessory pigment molecules are organized with pigment-bind- ing proteins inthe thylakoid membrane into units called antenna complexes. The pigments and associated proteins are arranged as highly ordered groups of about 250 chlorophyll molecules, associated with specific enzymes and other proteins. Each antenna complex absorbs light energy and transfers it to the reaction center, which consists of chlorophyll molecules and proteins, including electron transfer components, that partic- pate directly in photosynthesis (Fig. 10). Light energy is con- verted to chemical energy in the reaction centers by a series of clectron transfer reactions “Two types of photosynthetic units, designated photosystem I and photosystem I, are involved in photosynthesis. Their eac- tion centers are distinguishable because they are associated with proteins in a way that causes a slight shift in their absorption spectra. Ordinary chlorophyll a has a strong absorption peak at about 660 nm. In contrast, the chlorophyll molecule that, rakes up the reaction center associated with photosystem T hasan absorption peak at 700 nm and is referred to as P700. ‘The reaction center of photosystem II is made up of a chloro- Phyil a molecule with an absorption peak of about 680 nm and is referred to as P680. ‘When a pigment molecule absorbs light energy, that energy is passed from one pigment molecule to another until it reaches the reaction center. When the energy reaches a molecule of 700 (in a photosystem I reaction center) or P680 (in a photo- system II reaction center), an electron is then raised to a higher nergy level. As we explain in the next section, this energized électron can be donated to an electron acceptor that becomes reduced in the proces | ke TRGURESA0 | A photosystem. CChlorapinyll molecules and accessory pigments are arranged in light- harvesting array, or antenna complexes. When a molecule in an antenna complex absorés a photon, the photon’s energy is fun- reled into the reaction centr. When this energy reaches the P700 (or P60) chlorophyll molecule in the reaction center, an electron [becomes energized and is accepted by a primary electron acceptor. Noncyclic electron transport produces ATP and NADPH Let’ begin our discussion of noncyelic electron transport with, the events associated with photosystem I (Fig. 8-11). pigment ‘molecule in an antenna complex associated with photosystem I absorbs a photon of light. The absorbed energy is transferred to the reaction center, where it excites an electron in a molecule ‘of P700, This energized electron is transferred to a primary elec- tron acceptor, which isthe fist of several electron acceptors in a series. (Uncertainty exists regarding the exact chemical nature ‘of the primary electron acceptor for photosystem L.) The ener~ gized electron is passed along an electron transport chain from, one electron acceptor to another, until it is passed to ferredoxin, an iron-containing protein. Ferredoxin transfers the electron to, NADP* in the presence of the enzyme ferredoxin-NADP* re ductase, When NADP * accepts 2 electrons, they unite with a proton (H”); thus the reduced form of NADP* is NADPH, which is released into the stroma, P700 becomes positively charged when it gives up an electron to the primary electron ‘acceptor; the missing electron is replaced by one donated by photosystem I. Like photosystem I, photosystem Ili activated when a pig ‘ment molecule in an antenna complex absorbs a photon of ight, energy. The energy is transferred to the reaction center, where it causes an electron in a molecule of P680 to move toa higher energy level. This energized electron is accepted by a primary electron acceptor (a highly modified chlorophyll molecule known as pheophytin) and then passes along an electron trans- port chain until it is donated to P700 in photosystem I. ‘How is the electron that has been donated to the electron ‘transport chain replaced? This occurs through photolysis (light Photosynthesis: Capturing Energy | 163