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GLYCOGEN synthase cAMP & + glucose-6-P brancher (UDP-glucose)
Pi phosphorylase cAMP + & + AMP debrancher 2 Ca+ -calmodulin +
GLUCOSE-1-P
HMP PATHWAY
GLUCOSE-6-P
GLUCOSE
GLYCOLYSIS
Figure 17-3
Glycogen Metabolism
6-phosphate for local use through glycolysis, particularly if the muscle exerts itself so much that it restricts its blood supply and becomes anaerobic Epinephrine has the same effect on liver glycogen ( degradation); however, liver ships the glucose out for consumption by other tissues, including muscle
Integration of Energy Metabolism
Fat provides a long-term storage form for energy (ATP) Fat provides zero glucose equivalents Liver: Liver makes fat for export to other tissues Muscle: Resting muscle prefers fatty acids as an energy supply However, glucose provides short-term, high-intensity supply Adipose: Adipose tissue is the primary storage facility for fat Fat is stored in these tissues as an intracellular droplet of insoluble triglyceride A hormone-sensitive lipase mobilizes triglyceride stores by hydrolysis to free fatty acids Red blood cells: These cells can t use fat at all for energy since they have no mitochondria Brain: Brain does not burn fat as an energy source; however, after adapting to long-term starvation, brain can use ketone bodies for fuel
METABOLIC MOVEMENTS OF FAT
Feeding ( glucagon insulin): Fat synthesis Fasting ( glucagon insulin): Fat degradation Starvation ( glucagon insulin): Fat degradation Diabetes ( glucagon insulin): Fat degradation Excitement ( epinephrine): Fat degradation Fat is only an energy storage form (Fig 17-4) Fat cannot be converted to carbohydrate equivalents This is a very important point Remember it! The reason for this is a bit subtle The carbon skeleton of fatty acids is metabolized to acetyl-CoA only Glucose precursors such as oxaloacetate can be synthesized from acetyl-CoA by going around the TCA cycle However, acetyl-CoA has 2 carbon atoms Going around the TCA cycle burns off 2 carbon atoms (as CO2) The net number of carbon atoms that ends up in oxaloacetate is then zero No carbohydrate can be made from fat5
Odd-chain fatty acids are an exception While they are relatively rare in the diet, odd-chainlength fatty acids end up at propionyl-CoA (C3) Propionyl-CoA is carboxylated by propionylCoA carboxylase to give methylmalonyl-CoA Methylmalonyl-CoA is rearranged to succinyl-CoA by the enzyme methylmalonyl-CoA mutase, a vitamin-B12-requiring enzyme
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DHAP glycerol-3-P
Basic Concepts in Biochemistry
(7) malonyl-CoA ADP + Pi ATP CO2 acetyl-CoA carboxylase (biotin) Ac-CoA (1) C16-AcylCoA fatty acid synthase NADPH fatty acid triglyceride
ADIPOSE
hormone-sensitive lipase glycerol metabolized to DHAP in liver
citrate + OAA NADH AcCoA
carnitine shuttle pyruvate GLUCOSE
citrate
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+ +
citrate cAMP glucagon insulin NADP acetyl-CoA carboxylase fatty acid synthase NADPH AcCoA FAT NAD -oxidation NADH
+
hormone-sensitive lipase +
cAMP glucagon insulin malonyl-CoA
Figure 17-4
Fat Metabolism
Fat is synthesized as long-chain fatty acids and then stored by making triglyceride Most of the body s fat is stored in adipose tissue While glycogen storage is limited to about a 24-hour supply, fat can be stored in unlimited amounts Most calories in excess of those required to maintain energy demands are stored as fat In contrast to glucose metabolism, the metabolism of fat can produce ATP only when coupled to mitochondrial respiration Fat is an aerobic source of energy The formation of a triglyceride that requires the presence of glycerol introduces a carbohydrate requirement for the storage of fat When
Integration of Energy Metabolism
energy and glucose equivalents are available, adipose tissue stores fat Strangely enough, adipose tissue can synthesize the glycerol (actually the 3-phosphoglycerate) required for triglyceride synthesis However, adipose tissue cannot simply reuse the glycerol produced from triglyceride hydrolysis This glycerol is shipped to the liver, where it can be converted to glyceraldehyde 3-phosphate and either burned as fuel (during starvation) or used to make more triglyceride Many tissues (muscle, liver, renal cortex) prefer fat for an energy supply, at least in the resting state The exception is red blood cells and brain These tissues depend heavily on glycolysis for energy Red cells cannot survive without glucose (no mitochondria), but during prolonged starvation, brain can adapt to utilize fat metabolites produced by the liver (ketone bodies) The regulation of fat metabolism is relatively simple During fasting, the rising glucagon levels inactivate fatty acid synthesis at the level of acetyl-CoA carboxylase and induce the lipolysis of triglycerides in the adipose tissue by stimulation of a hormone-sensitive lipase This hormone-sensitive lipase is activated by glucagon and epinephrine (via a cAMP mechanism) This releases fatty acids into the blood These are transported to the various tissues, where they are used
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