UNIT 3: Part 1 Pathways that Harvest and Store Chemical Energy Hillis Textbook Chapter 6 Energy is stored in chemical bonds and can be released and transformed by metabolic pathways. Chemical energy available to do work is termed free energy (G). Five principles governing metabolic pathways: 1. 2. 3. 4. 5. Chemical transformations occur in a series of intermediate reactions that form a metabolic pathway. Each reaction is catalyzed by a specific enzyme. Most metabolic pathways are similar in all organisms. In eukaryotes, many metabolic pathways occur inside specific organelles. Each metabolic pathway is controlled by enzymes that can be inhibited or activated. In cells, energy-transforming reactions are often coupled: An energy-releasing (exergonic) reaction is coupled to an energy-requiring (endergonic) reaction. Adenosine triphosphate (ATP) is a kind of “energy currency” in cells. Energy released by exergonic reactions is stored in the bonds of ATP. When ATP is hydrolyzed, free energy is released to drive endergonic reactions. An exergonic reaction will release energy, allowing it to be stored in the ATP molecule! An endergonic reaction will need energy, which comes from the breaking of those bonds in ATP! The process of hydrolysis of an ATP molecule is exergonic: ATP H2O ADP Pi freeenergy ΔG is about –7.3 kcal Free energy of the bond between phosphate groups is much higher than the energy of the O—H bond that forms after hydrolysis. Phosphate groups are negatively charged, so energy is required to get them near enough to each other to make the covalent bonds in the ATP molecule. Lots of energy Not as much energy Energy can also be transferred by the transfer of electrons in oxidation–reduction, or redox reactions. Oxidation is the loss of one or more electrons. Reduction is the gain of one or more electrons. Oxidation and reduction always occur together. Transfers of hydrogen atoms involve transfers of electrons (H = H+ + e–). When a molecule loses a hydrogen atom, it becomes oxidized. The more reduced a molecule is, the more energy is stored in its bonds. Coenzyme NAD+ is a key electron carrier in redox reactions. NAD+ (oxidized form) NADH (reduced form) Reduction of NAD+ is highly endergonic: NAD H 2e NADH Oxidation of NADH is highly exergonic: NADH H 1 2 O2 NAD H 2O In cells, energy is released in catabolism (breaking bonds) by oxidation… Energy is then trapped by reduction of coenzymes such as NADH… BUT, energy for anabolic (building bonds) processes is supplied by ATP, not NADH! So, oxidative phosphorylation transfers energy from NADH to ATP. Oxidative phosphorylation couples: oxidation of NADH: NADH NAD H 2e energy with production of ATP: The coupling is called chemiosmosis—diffusion of protons across a membrane, which drives the synthesis of ATP. Chemiosmosis converts potential energy of a proton gradient across a membrane into the chemical energy in ATP. energy ADP Pi ATP ATP synthase synthesizes ATP! It is a membrane protein with two subunits: F0 is the H+ channel; potential energy of the proton gradient drives the H+ through. F1 has active sites for ATP synthesis.