Food Molecules Are Broken Down in Three Stages to Produce ATP
The proteins, lipids, and polysaccharides that make up most of the food we eat must be broken down into smaller molecules before our cells can use them—either as a source of energy or as building blocks for other molecules. The breakdown processes must act on food taken in from outside, but not on the macromolecules inside our own cells. Stage 1 in the enzymatic breakdown of food molecules is therefore digestion, which occurs either in our intestine outside cells, or in a specialized organelle within cells, the lysosome. (A membrane that surrounds the lysosome keeps its digestive enzymes separated from the cytosol, as described in Chapter 13.) In either case, the large polymeric molecules in food are broken down during digestion into their monomer subunits—proteins into amino acids, polysaccharides into sugars, and fats into fatty acids and glycerol—through the action of enzymes. After digestion, the small organic molecules derived from food enter the cytosol of the cell, where their gradual oxidation begins. As illustrated in Figure 2-70, oxidation occurs in two further stages of cellular catabolism: stage 2 starts in the cytosol and ends in the major energy-converting organelle, the mitochondrion; stage 3 is entirely confined to the mitochondrion.
In stage 2 a chain of reactions called glycolysis converts each molecule of glucose into two smaller molecules of pyruvate. Sugars other than glucose are similarly converted to pyruvate after their conversion to one of the sugar intermediates in this glycolytic pathway. During pyruvate formation, two types of activated carrier molecules are produced—ATP and NADH. The pyruvate then passes from the cytosol into mitochondria. There, each pyruvate molecule is converted into CO2 plus a two-carbon acetyl group—which becomes attached to coenzyme A (CoA), forming acetyl CoA, another activated carrier molecule (see Figure 2-62). Large amounts of acetyl CoA are also produced by the stepwise breakdown and oxidation of fatty acids derived from fats, which are carried in the bloodstream, imported into cells as fatty acids, and then moved into mitochondria for acetyl CoA production.
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Stage 3 of the oxidative breakdown of food molecules takes place entirely in mitochondria. The acetyl group in acetyl CoA is linked to coenzyme A through a high-energy linkage, and it is therefore easily transferable to other molecules. After its transfer to the four-carbon molecule oxaloacetate, the acetyl group enters a series of reactions called the citric acid cycle. As we discuss shortly, the acetyl group is oxidized to CO2 in these reactions, and large amounts of the electron carrier NADH are generated. Finally, the high-energy electrons from NADH are passed along an electron-transport chain within the mitochondrial inner membrane, where the energy released by their transfer is used to drive a process that produces ATP and consumes molecular oxygen (O2). It is in these final steps that most of the energy released by oxidation is harnessed to produce most of the cell’s ATP.
Because the energy to drive ATP synthesis in mitochondria ultimately derives from the oxidative breakdown of food molecules, the phosphorylation of ADP to form ATP that is driven by electron transport in the mitochondrion is known as oxidative phosphorylation. The fascinating events that occur within the mitochondrial inner membrane during oxidative phosphorylation are the major focus of Chapter 14.
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Through the production of ATP, the energy derived from the breakdown of sugars and fats is redistributed as packets of chemical energy in a form convenient for use elsewhere in the cell. Roughly 109 molecules of ATP are in solution in a typical cell at any instant, and in many cells, all this ATP is turned over (that is, used up and replaced) every 1-2 minutes.
In all, nearly half of the energy that could in theory be derived from the oxidation of glucose or fatty acids to H2O and CO2 is captured and used to drive the energetically unfavorable reaction Pi + ADP → ATP. (By contrast, a typical combustion engine, such as a car engine, can convert no more than 20% of the available energy in its fuel into useful work.) The rest of the energy is released by the cell as heat, making our bodies warm.
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