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Chapter 8 Summary Metabolic pathways consist of many interrelated, enzyme-catalyzed chemical reactions. These pathways can be categorized as either catabolic or anabolic. Anabolic pathways promote the synthesis of new compounds and energy storage, whereas catabolic pathways promote the mobilization of stored energy and the breakdown of energy-yielding nutrients. Chemical reactions are catalyzed by enzymes, some requiring cofactors or coenzymes to function. A cofactor is an inorganic substance that is part of the enzyme’s structure. Coenzymes are organic molecules that assist enzymes in redox reactions by transferring hydrogen ions (H+) and electrons (e-) to and from molecules. Coenzymes have oxidized (NAD+, FAD, and NADP+) and reduced (NADH + H+, FADH2, and NADPH + H+) forms. Enzymes involved in metabolic pathways are regulated primarily by hormones. The hormone insulin promotes energy storage, whereas the hormone glucagon promotes energy mobilization. The hormones cortisol and epinephrine also promote catabolic pathways, being released in response to stress. Cells rely on the energy contained in the chemical bonds of ATP. Some ATP is generated by substrate phosphorylation, a process that adds a phosphate group (P i) directly to ADP. However, most ATP is synthesized by oxidative phosphorylation, which involves a series of chemical reactions that make up the electron transport chain. When NADH + H+ and FADH2 enter the electron transport chain, their electrons and hydrogen ions are removed. The electrons are passed along protein complexes, and the energy released is used to pump the hydrogen ions (H+) out of the mitochondrial matrix. The movement of hydrogen ions back into the mitochondrial matrix releases energy that is used by the enzyme ATP synthase to attach a phosphate group to ADP, generating ATP. Last, iron-containing protein complexes called cytochromes reunite the electrons (e-) and hydrogen ions (H+), which in turn combine with oxygen to form water. Liver and muscle cells break down glycogen into glucose by a process called glycogenolysis. Glucose catabolism begins with glycolysis, an anaerobic pathway that converts glucose to pyruvate. Oxygen availability determines if pyruvate is converted to acetyl-CoA or lactate. If oxygen is available, acetylCoA is formed and enters the citric acid cycle, resulting in the formation of NADH + H + and FADH2. These coenzymes can enter the electron transport chain and drive ATP formation via oxidative phosphorylation. Protein is broken down to amino acids by proteolysis. For amino acids to be used as an energy source, the nitrogen-containing amino group is removed via transamination and deamination. The remaining structure (-ketoacid) can enter the citric acid cycle, ultimately generating ATP via the electron transport chain. Lipid catabolism begins with lipolysis, releasing fatty acids and glycerol. Fatty acids are oxidized via -oxidation and the citric acid cycle to form NADH + H+ and FADH2, which then enter the electron transport chain to generate additional ATP. Anabolic pathways play important roles in storing excess energy and in synthesizing energy-yielding molecules when glucose availability is limited. The hormone insulin stimulates liver and muscle tissues to store excess glucose as glycogen (glycogenesis). Insulin also promotes the conversion of glucose and amino acids to fatty acids and the subsequent production of triglyceride in adipose and liver tissues (lipogenesis). Thus anabolic pathways are important during periods of excess energy availability. During starvation, anabolic pathways are used to synthesize glucose from noncarbohydrate sources (gluconeogenesis). Substances used for gluconeogenesis include glucogenic amino acids, lactate, and glycerol. However, high rates of gluconeogenesis deplete the amount of oxaloacetate. When this occurs, acetyl-CoA cannot participate in the citric acid cycle. Rather, acetyl-CoA is diverted to ketone production. Some amino acids are converted to ketones as well. Tissues such as muscle, brain, and kidney have enzymes that allow them to use ketones for ATP production. Energy metabolism pathways are responsive to intermittent states of feeding and fasting, called the absorptive state, postabsorptive state, acute starvation, and prolonged starvation. Most of the major anabolic pathways operate during the absorptive state, including those promoting protein, triglyceride, and glycogen synthesis. During this time, glucose is the major source of energy for all tissues. The postabsorptive state, which is the period 4 to 24 hours after the last intake of food, relies heavily on energy supplied by the breakdown of stored energy reserves, especially glycogen. As the body enters acute starvation, defined as the first five days after the postabsorptive state, the body begins to rely more on the mobilization of adipose reserves for ATP production, while producing sufficient glucose via gluconeogenesis. The body enters the nutritional state of prolonged starvation when a person is deprived of food for more than one week. During this time, the body must preserve lean tissue from its gluconeogenic fate by increasing the use of ketones as an energy source. This is accomplished via ketogenic pathways. However, prolonged starvation can eventually result in the extensive use of muscle for glucose and, ultimately ATP production. If refeeding is not resumed, the consequences of starvation can be severe.