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In plants, photosynthesis occurs in specialized organelles called chloroplasts. The internal membranes of chloroplats are organized into sacs called thylakoids. Surrounding the thylakoid membrane system is a semiliquid substance called stroma. Photosynthetic pigments are clustered together to form photosystems. When a photon of light strikes the reaction center of photosystem 2, it excites an electron. Two water molecules bind to an enzyme at the reaction center. This enzyme splits the water and uses the electrons from the water to replace the electrons removed from the reaction center. Oxygen is produced in this process. The primary electron acceptor for the light-energized electrons leaving photosystem 2 is plastoquinone. The reduced plastquinone passes the excited electrons to a proton pump embedded in the membrane called the b6-f complex. Arrival of the energetic electrons causes the b6-f complex to pump protons from the stroma into the thylakoid space, thereby generating a proton gradient across the membrane. Because the thylakoid membrane is impermeable to protons, the protons in the stroma must pass through the channels provided by ATP synthase. As protons pass through, ADP is phosphorylated to ATP and released into the stroma. This process for making ATP is referred to as photophosphorylation. When photosystem I absorbs a photon of light, its reaction center passed high-energy electrons to ferredoxin. The enzyme NADP reductase then transfers the electrons to NADP to form NADPH. Electrons lost from photosystem I are replaced by electrons generated from photosystem 2. A small protein called plastocyanin (pC) then carries the electrons from the b6-f complex to photosystem 1.