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Berg JM, Tymoczko JL, Stryer L. Biochemistry. 5th edition. New York: W H Freeman; 2002.

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Biochemistry. 5th edition.

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Chapter 21Glycogen Metabolism

Glycogen is a readily mobilized storage form of glucose. It is a very large, branched polymer of glucose residues (Figure 21.1) that can be broken down to yield glucose molecules when energy is needed. Most of the glucose residues in glycogen are linked by α-1,4-glycosidic bonds. Branches at about every tenth residue are created by α-1,6-glycosidic bonds. Recall that α-glycosidic linkages form open helical polymers, whereas β linkages produce nearly straight strands that form structural fibrils, as in cellulose (Section 11.2.3).

Figure 21.1. Glycogen Structure.

Figure 21.1

Glycogen Structure. In this structure of two outer branches of a glycogen molecule, the residues at the nonreducing ends are shown in red and residue that starts a branch is shown in green. The rest of the glycogen molecule is represented by R.

Glycogen is not as reduced as fatty acids are and consequently not as energy rich. Why do animals store any energy as glycogen? Why not convert all excess fuel into fatty acids? Glycogen is an important fuel reserve for several reasons. The controlled breakdown of glycogen and release of glucose increase the amount of glucose that is available between meals. Hence, glycogen serves as a buffer to maintain blood-glucose levels. Glycogen's role in maintaining blood-glucose levels is especially important because glucose is virtually the only fuel used by the brain, except during prolonged starvation. Moreover, the glucose from glycogen is readily mobilized and is therefore a good source of energy for sudden, strenuous activity. Unlike fatty acids, the released glucose can provide energy in the absence of oxygen and can thus supply energy for anaerobic activity.

The two major sites of glycogen storage are the liver and skeletal muscle. The concentration of glycogen is higher in the liver than in muscle (10% versus 2% by weight), but more glycogen is stored in skeletal muscle overall because of its much greater mass. Glycogen is present in the cytosol in the form of granules ranging in diameter from 10 to 40 nm (Figure 21.2). In the liver, glycogen synthesis and degradation are regulated to maintain blood-glucose levels as required to meet the needs of the organism as a whole. In contrast, in muscle, these processes are regulated to meet the energy needs of the muscle itself.

Figure 21.2. Electron Micrograph of a Liver Cell.

Figure 21.2

Electron Micrograph of a Liver Cell. The dense particles in the cytoplasm are glycogen granules. [Courtesy of Dr. George Palade.]

21.0.1. An Overview of Glycogen Metabolism:

Glycogen degradation and synthesis are relatively simple biochemical processes. Glycogen degradation consists of three steps: (1) the release of glucose 1-phosphate from glycogen, (2) the remodeling of the glycogen substrate to permit further degradation, and (3) the conversion of glucose 1-phosphate into glucose 6-phosphate for further metabolism. The glucose 6-phosphate derived from the breakdown of glycogen has three fates (Figure 21.3): (1) It is the initial substrate for glycolysis, (2) it can be processed by the pentose phosphate pathway to yield NADPH and ribose derivatives; and (3) it can be converted into free glucose for release into the bloodstream. This conversion takes place mainly in the liver and to a lesser extent in the intestines and kidneys.

Figure 21.3. Fates of Glucose 6-Phosphate.

Figure 21.3

Fates of Glucose 6-Phosphate. Glucose 6-phosphate derived from glycogen can (1) be used as a fuel for anaerobic or aerobic metabolism as in, for instance, muscle; (2) be converted into free glucose in the liver and subsequently released into the blood; (more...)

Glycogen synthesis requires an activated form of glucose, uridine diphosphate glucose (UDP-glucose), which is formed by the reaction of UTP and glucose 1-phosphate. UDP-glucose is added to the nonreducing end of glycogen molecules. As is the case for glycogen degradation, the glycogen molecule must be remodeled for continued synthesis.

The regulation of these processes is quite complex. Several enzymes taking part in glycogen metabolism allosterically respond to metabolites that signal the energy needs of the cell. These allosteric responses allow the adjustment of enzyme activity to meet the needs of the cell in which the enzymes are expressed. Glycogen metabolism is also regulated by hormonally stimulated cascades that lead to the reversible phosphorylation of enzymes, which alters their kinetic properties. Regulation by hormones allows glygogen metabolism to adjust to the needs of the entire organism. By both these mechanisms, glycogen degradation is integrated with glycogen synthesis. We will first examine the metabolism, followed by enzyme regulation and then the elaborate integration of control mechanisms.

Signal cascades lead to the mobilization of glycogen to produce glucose, an energy source for runners.

Figure

Signal cascades lead to the mobilization of glycogen to produce glucose, an energy source for runners. [(Left) Mike Powell/Allsport.]

Contents

21.1 Glycogen Breakdown Requires the Interplay of Several Enzymes

21.2 Phosphorylase Is Regulated by Allosteric Interactions and Reversible Phosphorylation

21.3 Epinephrine and Glucagon Signal the Need for Glycogen Breakdown

21.4 Glycogen Is Synthesized and Degraded by Different Pathways

21.5 Glycogen Breakdown and Synthesis Are Reciprocally Regulated

Summary

Problems

Selected Readings

By agreement with the publisher, this book is accessible by the search feature, but cannot be browsed.

Copyright © 2002, W. H. Freeman and Company.
Bookshelf ID: NBK21190

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