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The oxidative pathways

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The oxidative pathways

A large amount of ATP is produced by the complete oxidation of carbon-containing compounds (e.g., glucose) to carbon dioxide and water in reactions of the Krebs cycle and oxidative phosphorylation. These reactions take place inside the membrane-bound components of cells called mitochondria. Carbohydrate can enter the oxidative pathways at a branch point in the glycolytic pathway. At the branch point, two kinds of reaction can occur: pyruvate, a three-carbon compound, can either react to form lactate or lose a carbon dioxide molecule to form a two-carbon unit, which then enters the Krebs cycle. If the muscle is working hard and using ATP rapidly, lactate is formed. If it is working slowly, a larger fraction of the carbon enters the oxidative pathways.

During reactions of the Krebs cycle, carbon dioxide and hydrogen atoms are removed from the compounds that enter the cycle. The carbon dioxide diffuses out of the muscle and is carried bythe blood to the lungs, where it is exhaled. The electrons from the hydrogen atoms are passed through an electron-transport chain consisting of the series of reactions involving cytochrome molecules. These events occur in the mitochondria, in which ADP is also combining with phosphate (Pi ) to form ATP. In the last step of this reaction sequence, the hydrogen atoms combine with oxygen to form water. The net reaction for each glucose molecule that enters the glycolytic pathway and proceeds through the Krebs cycle and oxidative phosphorylation is summarized below.

C6H12O6+36ADP+36Pi=6CO2+36ATP+6H2O


Although the oxidative pathways result in the formation of more ATP from each glucose molecule than does the glycolytic pathway, the complete process is much slower than glycolysis. The slowest steps are those involving the passage of the carbon-containing compounds from the sarcoplasm into the mitochondria and the delivery of oxygen, which ultimately comes from the air that is breathed. In red blood cells oxygen is combined with hemoglobin, a protein containing four identical subunits, during its transfer from the lungs to the muscles. Muscle contains myoglobin, which has a structure similar to a single subunit of hemoglobin. Myoglobin, which combines with and can store oxygen, is responsible for transporting much of the oxygen through the sarcoplasm to the mitochondria. Myoglobin is especially important in the heart and “red” muscles, which rely heavily on oxidative metabolism for the production of ATP.

During intense exercise of skeletal muscle, ATP is supplied almost entirely by carbohydrate metabolism. During rest or very light exercise, however, skeletal muscle depends largely on the oxidation of stored fats—actually of their breakdown products, fatty acids—for the production of ATP. Two-carbon units and hydrogen atoms are removed from fats in a stepwise fashion. The two-carbon units enter the Krebs cycle, as do the identical units derived from carbohydrates, as the compound acetyl co-enzyme A. The hydrogen atoms from the breakdown of the fat and from the reactions of the Krebs cycle proceed through oxidative phosphorylation, and ATP is produced.

Heart muscle, which operates constantly but at a lower level of intensity than does skeletal muscle, depends on oxidative reactions for ATP. Most of the two-carbon units are derived from either fatty acids or ketones, components of fat, brought to the heart by the blood.

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