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Fates of dietary glucose

The major source of dietary carbohydrate for humans is starch from consumed plant material. This is supplemented with a small amount ofglycogen from animal tissue, disaccharides such as sucrose from products containing refined sugar and lactose in milk. Digestion in the gut converts all carbohydrate to monosaccharides which are transported to the liver and converted to glucose. The liver has a central role in the storage and distribution within the body of all fuels, including glucose. Glucose in the body undergoes one of three metabolic fates :

it is catabolised to produce ATP This occurs in all peripheral tissues, particularly in brain, muscle and kidney.

it is stored as glycogen This storage occurs in liver and muscle.

it is converted to fatty acids Once converted to fatty acids, these are stored in adipose tissue as triglycerides.

Glucose catabolism
Glucose will be oxidised by all tissues to synthesise ATP. The first pathway which begins the complete oxidation of glucose is calledglycolysis. Glycolysis This pathway cleaves the six carbon glucose molecule (C6H12O6) into two molecules of the three carbon compound pyruvate (C3H3O3-). This oxidation is coupled to the nett production of two molecules of ATP/glucose. The diagram below shows an outline of glycolysis. The full set of reactions and structures can be found in any biochemistry textbook.

as in this reaction. It uses NAD as the electron acceptor. The reduction of . This cofactor is present only in limited amounts and once reduced to NADH. It uses NAD as the electron acceptor. it must be reoxidised to NAD to permit continuation of the pathway. This re-oxidation occurs by one of two methods : Anaerobic glycolysis • pyruvate is reduced to a compound called lactate This single reaction occurs in the absence of oxygen (anaerobically) and is ideally suited to utilisation in heavily exercising muscle where oxygen supply is often insufficient to meet the demands of aerobic metabolism. One oxidation reaction occurs in the latter part of the pathway. it must be reoxidised to NAD to permit continuation of the pathway. as in this reaction. This cofactor is present only in limited amounts and once reduced to NADH.One oxidation reaction occurs in the latter part of the pathway.

Aerobic metabolism of glucose • pyruvate is transported inside mitochondria and oxidised to a compound called acetyl coenzyme A (abbreviated to "acetyl CoA"). but is now used by athletes in sprint events. Very heavily exercising muscle can use this pathway as the sole source of ATP synthesis for a short period of time. • it is converted back to glucose in the liver Gluconeogenesis The process of conversion of lactate to glucose is called gluconeogenesis. (These pathways will be discussed in more detail later. These reactions are coupled to a process known as the electron transport chain which has the role of harnessing chemical bond energy from a series of oxidation/reduction reactions to the synthesis of ATP and simultaneously re-oxidising NADH to NAD.pyruvate to lactate is coupled to the oxidation of NADH to NAD.) Fast twitch muscle fibres utilise the first of the two mechanisms described above almost exclusively. uses some of the reactions of glycolysis (but in the reverse direction) and some reactions unique to this pathway to resynthesise glucose. Accumulation of lactate (actually lactic acid) also causes a reduction in intracellular pH. acetyl CoA is oxidised ultimately to CO2. This is an oxidation reaction and uses NAD as an electron acceptor. This probably evolved in humans as a defence mechanism. This pathway requires an energy input (as ATP) but has the role of maintaining a circulating glucose concentration in the . The lactate formed is removed to other tissues and dealt with by one of two mechanisms : • it is converted back to pyruvate The pyruvate then proceeds to be further oxidised by the second mechanism described above. finally producing a large amount of ATP. The formation of lactate as an end product from glucose extracts only a relatively small amount of the bond energy contained in glucose. By a further series of reactions collectively called the citric acid cycle.

This co-operative cycle utilising both the muscle and liver tissue is called the Cori cycle. Both of these mechanisms illustrate the interdependence of tissues on each other and the co-operative activities between organs which make . Cori cycle It can be shown by a complex calculation of energy yields that this process of partially oxidising glucose to lactate in muscle. actually has a much higher energy yield than the 2 ATP/glucose produced by glycolysis alone.bloodstream (even in the absence of dietary supply) and also maintaining a glucose supply to fast twitch muscle fibres. transporting it to the liver for conversion back to glucose and then re-supplying it to muscle. The process is shown in a diagram below.

. In both muscle and liver there is interconversion between the monomer glucose and the polymer glycogen. The control is exerted by hormones acting to control the activity of the key enzymes.either towards glycogen synthesis OR towards glycogen breakdown and mobilisation of free glucose. The activity of these enzymes is controlled such that only one is active at any one time and thus the pathway can proceed in only one direction . Glycogen stores Glycogen (g/kg tissue) 65 14 Liver Muscle Organ mass 1. The high degree of branching (about every twelve glucose residues) produces a molecule which is compact and thus can be efficiently stored in the limited space available in liver and muscle tissue. thus it is controlled to meet the body's glucose requirements at a particular time.6 kg 28 kg Total glucose ~100 g ~400 g The amount of glycogen in muscle changes substantially between the fed state and following heavy exercise. it is still a polar molecule and thus must be stored with associated water. The amount of glycogen stored in the liver is more constant and only falls substantially after prolonged starvation. There are some differences in the hormone action in liver and muscle. Glycogen and glucose interconversion Glycogen is a highly branched polymer of glucose.up the total of the body's metabolic activities. Even though the branching is designed to make the molecule compact. This has the potential to be a futile cycle wasting energy if the interconversion occurred continuously. It is stored as aggregates of glycogen molecules within cells (visible microscopically as glycogen granules) with up to 70% of the aggregate being water. Hormonal control of glycogen metabolism The control which operates is via different enzymes catalysing the synthesis and breakdown (degradation) of glycogen.

Insulin is secreted in response.this will be discussed later. excess glucose will be diverted to the synthesis of fats . The amount of glycogen which can be stored in these two tissues is limited and once the stores are saturated. Maintenance of blood glucose between meals When there is no dietary glucose intake (between meals). The pancreas secretes more glucagon and less insulin. circulating glucose concentration must be maintained. The glucagon : • • stops liver glycogen synthesis (by deactivating the synthesis enzymes) increases liver glycogen breakdown (by activating the degradation enzymes) stimulates gluconeogenesis in the liver to further increase the circulating blood glucose concentration • . Insulin : • • stimulates uptake of glucose into both muscle and liver stimulates increased glycogen synthesis in both muscle and liver This is achieved by activation of the key synthesis enzymes.HORMONE Glucagon Adrenaline Insulin Source Pancreas Adrenals Pancreas Target tissue Liver Muscle Liver and Muscle Action Stimulates glycogen breakdown Stimulates glycogen breakdown Stimulates glycogen synthesis Utilisation of glucose in the fed and fasting states Glucose utilisation after a meal A high circulating glucose concentration is present after a meal. Carbohydrate is digested and the glucose absorbed into the blood stream.

Glucose 6-phosphate is the first product in the glycolysis pathway and its formation from glucose requires the expenditure of 1 ATP molecule/glucose. This is a 50% increase in yield. glucose that is derived from glycogen and enters the glycolysis pathway (rather than starting as monomeric glucose) yields a nett production of 3 ATP/glucose rather than just 2. Liver supplies other organs with glucose so must be able to export glucose released from glycogen hydrolysis. Glucose can only be supplied to muscle cells either by utilising stored muscle glycogen or supply from the liver via the bloodstream. When glucose becomes available again after a meal glycogen stores in muscle are replenished. Supply of glucose to exercising muscle Increasing muscle activity requires adequate fuel supply for ATP synthesis by muscle. Muscle does not carry out gluconeogenesis. As glucose 6-phosphate is formed directly from glycogen hydrolysis. adrenaline secretion is switched off. Glycogen metabolism in liver and muscle Energy yield from glycogen breakdown The energy yield from the hydrolysis of stored glycogen and the subsequent oxidation of the released glucose is the same in muscle and liver. Muscle is a major consumer of glucose and thus does not export glucose. When glycogen is hydrolysed. This is easily converted to glucose 6-phosphate (these are molecules with the phosphate group attached to different carbon atoms on the glucose). When muscle activity is anticipated. . the product is glucose 1-phosphate. the adrenal glands secrete adrenaline. When muscle activity ceases. Role of glucose 6-phosphatase Muscle and liver have different metabolic needs. Adrenaline increases muscle glycogen degradation (by activating the breakdown enzymes and de-activating the synthesis enzymes).These mechanisms maintain an appropriate circulating blood glucose to supply tissues such as the brain which are major glucose consumers but do not store glycogen.

The pyruvate is converted to acetyl CoA. If glucose intake continues after muscle and liver glycogen stores are saturated. which is the starting material for the synthesis of fatty acids. To leave the cell it must be converted to glucose. but rather uses it as a fuel to power muscle contraction. It is very important to be aware that muscle does not possess glucose 6-phosphatase so it does not export glucose released from its glycogen stores. Summary of carbohydrate metabolism The pathways used in carbohydrate metabolism are shown in the following diagram. glucose 6-phosphatase. The mechanism of fatty acid synthesis will be discussed under the heading of fat metabolism. It is converted to a fuel storage form which has an unlimited capacity i. triglycerides stored in adipose tissue.e. This reaction is catalysed by an enzyme. so glucose released from liver glycogen can be exported to other tissues. Glucose is converted to pyruvate by glycolysis. This synthesis occurs in the liver followed by conversion of the fatty acids to triglycerides (also in the liver) and then transport to adipose tissue for storage. Triglycerides (fat) form the major energy store in the body. Conversion of excess glucose to fat Sustained high glucose intake in the diet leads to increased fat synthesis. .Glucose 6-phosphate formed as described in the previous section is highly polar and cannot cross the cell's cytoplasmic membrane. glucose 6-phosphate → glucose + phosphate glucose 6-phosphatase Liver possesses this enzyme. the glucose is not excreted or wasted.

These calories may be used immediately for energy metabolism or may be transformed and stored as glycogen or fat to be used as an energy source as demanded. and metabolized. Carbohydrates provide roughly half of the total caloric intake of the average human diet. supplying humans or animals with energy. Plants manufacture and store carbohydrates as their main source of energy through photosynthesis. Dietary carbohydrates are comprised of a . these organic compounds can be digested.CARBOHYDRATES. Once consumed. absorbed.

The three that can be absorbed by the human body include glucose. They generally have between two to ten units. pectin. It is the repeating monosaccharide unit in starch. maltose Oligosaccharides 2–10 Includes the disaccharides Polysaccharides > 10 Glycogen. Glucose is the most abundant of the monosaccharides and the most important nutritionally. hemicellulose. consisting of glucose and galactose Maltose (from malt sugar). lignin. and cellulose. those chains containing two units. The major polysaccharides include the digestible forms (glycogen and starch) and nondigestible forms (cellulose. . galactose. For example. cellulose **A "sugar unit" is one monosaccharide—each unit is not necessarily the same monosaccharide. fructose Disaccharides 2 Sucrose. consisting of glucose and fructose Lactose (from milk sugar). Monosaccharides. glycogen. sucrose consists of one glucose uni and one fructose unit. The most common disaccharides include: Sucrose (from table. and beet sugars).wide array of compounds ranging from the simple oneor two-unit sugars to the long chain starches. cane. and polysaccharides. starch. Carbohydrate classification Classification Number of sugar units** Examples Monosaccharides 1 Glucose. galactose. lactose. and is found in all edible disaccharides. Oligosaccharides are short chains of monosaccharide units that are joined by glycosidic bonds. consisting of two glucose units Polysaccharides are long chains of monosaccharide units. di-and oligosaccharides. and gums). and fructose. are the simplest form of carbohydrates and are seldom found free in nature. glycogen and cellulose. with the disaccharides. being the most abundant. often referred to as simple sugars. Carbohydrates can be classified as monosaccharides.

It is made up of repeating glucose units and is highly branched. it also encases some of the starch. Insoluble fiber. excess calories from fat. indigestible. and lignin (a noncarbohydrate component of the cell wall that is often included as dietary fiber). found primarily in the liver and skeletal muscle. absorbs water. animal products do not contain large amounts of glycogen because it is depleted at the time of slaughter due to stress hormones. These bonds are resistant to mammalian digestive enzymes rendering cellulose.4. cellulose is not considered to be a significant source of energy for the body. However. When energy intake exceeds energy expenditure. This increases the time the food is in the small intestine. protein. which makes up the greatest percent of the total starch content.4 glycosidic bonds. unbranched molecule that is bound solely by a-1. Although found in animal tissue. preventing constipation. It helps to speed the movement through the intestinal tract. allowing the starch to be digested. The increased nutrition for the bacteria can increase microbial growth. and other substances containing these bonds. and carbohydrate can be used to form glycogen. Bacteria in the bowel can use fiber as a food source. which can then lead to increased stool bulk. Since cellulose is a major part of the plant cell wall. it is important for intestinal bacteria. these chains can be broken down to single glucose units and used as an energy source for the body. These bacteria can degrade the fiber and release some components that can then be absorbed and used by the body. is branched with a-1. which includes pectin and gums. including cellulose. dissolves in water to form a gel in the digestive tract. Cooking causes the granules to swell and also softens and ruptures the cellulose wall. thereby increasing the bulk and volume of the stool. Just as starch and glycogen. Cellulose is the major component of cell walls in plants. Soluble fiber. it too is made up of repeating glucose molecules. During times of fasting or in between meals. Thus. with little of the fiber actually found in the stool. Amylopectin. It can be found in two forms—amylose and amylopectin.6 bonds at the branch points. However. preventing the digestive enzymes from reaching it and decreasing the digestibility of some raw foods such as potatoes and grains. It is believed that soluble fiber plays a role in lowering blood LDL cholesterol. thus increasing the chance of nutrients being absorbed. and is prescribed in the treatment of . the glycosidic bonds connecting the units are b-1. hemicellulose. This could be due to the binding and increased excretion of fat and bile acid (a derivative of cholesterol) or other mechanisms not yet understood.Starch is the most common digestible polysaccharide found in plants. Dietary Fiber Fiber can be classified as soluble and insoluble. Amylose is a linear. Glycogen is the major storage form of carbohydrates in animals. as a fiber.

Absorption. as well as to receive other added health benefits. The digestive process begins in the mouth with salivary a-amylase that partially breaks down starch by hydrolyzing some of the a-1. Digestion. and phytochemicals (chemicals found in plants believed to contain protective properties) that contribute to the health benefits of whole grain foods. and Transportation In order for carbohydrates to be absorbed by the intestinal mucosal cells. However. helping to decrease cancer risk.irritable bowel syndrome.4 bonds. they must first be converted into monosaccharides. Examples of carbohydrate food sources Monosaccharides Glucose Fructose Galactose Fruit High-fructose corn syrup Milk Vegetables Honey Milk products Honey Fruit Disaccharides Sucrose Lactose Maltose Table sugar Milk Beer . but along with it many of the vitamins. although this may differ depending on chewing time. Refined and processed foods have not only most of the fiber removed. the digestion that takes place here is of little significance since food remains in the mouth for only a brief period. The enzyme continues to work for a short time in the stomach until the pH is lowered due to hydrochloric acid that inhibits the enzyme. minerals. It has also been shown that insoluble fibers bind fat-soluble carcinogens and remove them from the gastrointestinal tract. The federal government's Dietary Guidelines for Americans encourage individuals to include whole grain foods in their diet to ensure adequate fiber to promote proper bowel function.

bread) Dietary Fiber Soluble Pectin Gums Fruits (apples.Examples of carbohydrate food sources Monosaccharides Glucose Fructose Galactose Maple sugar Milk products Malt liquor Fruit Vegetables Honey Polysaccharides Starch (rye. legumes. cereals. potatoes. oats. barley Jams and jellies (additive) Ice cream (additive) Legumes Insoluble Cellulose Hemicellulose Lignin Whole wheat foods Whole grains Fruit Bran Seeds . rice. berries) Oats. wheat.

For individuals left untreated. At this time the glycogen is broken down into individual glucose units. Liver and skeletal muscle can convert excess glucose to glycogen through a pathway known as glycogenesis. which will lead to serious consequences if steps are not taken to correct it. where they are taken up. Specific disaccharidases located on the intestinal mucosal cells help to further break down the carbohydrates into the monosaccharides: glucose. Once transported through the intestinal wall. Although glucose is metabolized extensively in the liver. a process known as . fructose. facilitated diffusion requires a specific carrier. Glucose and galactose enter by active transport. resulting in hyperglycemia. but instead of needing energy.Examples of carbohydrate food sources Monosaccharides Glucose Fructose Galactose Leafy vegetables Bran. whereas the brain and liver do not. The glycogen is stored after meals to be used as an energy source when energy demands are higher than intake. wheat Vegetables The bulk of carbohydrate digestion occurs in the small intestine by pancreatic a-amylase. Fructose is absorbed by facilitated diffusion. Metabolism of Carbohydrates The liver is the major site of galactose and fructose metabolism. Glucose can be metabolized through the glycolysis pathway to pyruvate where it is either converted to lactate or completely oxidized to CO2. allowing the enzyme activity to occur. dietary carbohydrates cause glucose levels to rise. the monosaccharides enter the blood through the capillaries and are carried to the portal circulation and then to the liver. which requires energy as well as specific receptors and carriers. and energy. The pH of the small intestines is increased due to the addition of bicarbonate and bile. This dependence on insulin becomes a problem for diabetics who either cannot make insulin (IDDM) or are resistent to insulin (NIDDM). it relies on the low levels of fructose inside the cell to "pull" the fructose inside. H2O. the monosaccharides can be absorbed by the mucosal cells. converted to glucose derivatives. and either stored as liver glycogen or used for energy immediately when needed. Like active transport. and galactose. Once in the tissues. unlike galactose and fructose. Tissues like skeletal muscle and adipose tissue depend on insulin for glucose uptake. the fate of glucose depends on the energy demands of the body. it is also passed into the blood supply to be used by other tissues. Once the carbohydrates have been broken down.

Johnson STARCH Starch from plants makes up about half of our dietary carbohydrates. BIBLIOGRAPHY Ettinger. "Types of Carbohydrates in an Ordinary Diet Affect Insulin Action and Muscle Substrates in Humans. Because of these characteristics. they do differ by characteristics such as solubility. Guthrie. flavor. Susan. 14–18 April 1997. starch is often removed from the source to use commercially. corn. Starch molecules can aggregate to form granules that differ by size and shape depending on the source of the starch. Saunders. "Carbohydrate as a Nutrient in Adults: Range of Acceptable Intakes. help to maintain adequate blood glucose. Debra Coward McKenzie Rachel K. through leaching and drying. Rome. and E. Food and Agriculture Organization of the United Nations."European Journal of Clinical Nutrition. Carbohydrates are an essential part of a healthy diet. Fiber. Richter. Table 2 contains examples of foods that contain the various types of carbohydrates. 1998.. and manioc. 51. and the glucose can be metabolized further. Rome: World Health Organization. 10th ed. "Food Sources of Added Sweetners in the Diets of Americans. The potato starch is often used as a thickener or instead of cornstarch in recipes. They provide an easily available energy source." American Journal of Clinical Nutrition. Excess carbohydrates also can be used as a substrate for fat synthesis. Proteins and Lipids. 63 (1996): 47–53. Mahan and Sylvia Escott-Stump. "Macronutrients: Carbohydrates. or in the case of manioc. A. B. and Diet Therapy. Dietary. Joanne. are an important vehicle for micronutrients and phytochemicals. potato. 2000. Macdonald. and are important in maintaining the integrity and function of the gastrointestinal tract." In Krause's Food. B. while manioc is best known as tapioca .glycogenolysis.: W. Philadelphia. A. Carbohydrates in Human Nutrition: Report of a Joint FAO/WHO Expert Consultation. 53 (1999): S101–S106. Pa. for example. Nutrition. Kiens. and Joan Morton. edited Kathleen L. Starch. FAO/WHO. Although there is no difference in the nutritional value between the starches since all cooked starches are broken down in the body into glucose molecules. For example. I. See also Digestion. and thickening power." Journal of the American Dietetic Association 100 (2000): 43–48. the starch can be removed from tubers such as potatoes and manioc (also known as cassava) through a wet milling process.