Carbohydrates Digestion and Metabolism Carbohydrates are the scarcest organic molecules in life.

They have broad range of function such as providing an important fraction of the energy which is the most nutritional role in the diet of most organisms, storing energy in our body and serving as cell membrane apparatus that provoke some forms of intercellular communication. Carbohydrates also serve as a structural component of many organisms, including the cell walls of bacteria, the exoskeleton of many insects, and the fibrous cellulose of plants (Champe, Harvey & Ferrier, 2008, p.83). For the digestion of carbohydrates, it occurs mainly in the mouth and intestinal lumen. This process is fast and is usually completed by the time the stomach contents reach the connection of the duodenum and jejunum. Monosaccharide is the simplest form of carbohydrate and it does not require hydrolysis prior to absorption. However, there is just minute monosaccharide present in diets of varied animal and plant source. Other forms of carbohydrate are disaccharides, oligosaccharides and polysaccharides which are relatively larger molecules require hydrolysis prior to absorption. Therefore, enzymes are needed to hydrolyze these large molecules into smaller molecules. It involves the hydrolysis of glycosidic bonds which is catalyzed by a family glycosidases that corrupt carbohydrates into their reducing sugar components. The carbohydrates digestion begins in the mouth. The glycogen and starch which consist of amylose and amylopectin are the chief dietary polysaccharides in our food. Salivary -amylase acts momentarily on dietary starch and glycogen in an arbitrary manner, hydrolyzing some (1-4) bonds during chewing. -amylase only hydrolyze the (1-4) bond and therefore the branched amylopectin and glycogen which contain major in (1-6) bonds cannot be hydrolyzed and the digest resulting from its action contains a combination of short, branched oligosaccharides and dextrin. The digestion process arrest for the time being in the stomach because of the elevated acidity inactivates the salivary -amylase. Next, the advance process of digestion occurs in the small intestine by pancreatic enzymes, known as pancreatic amylase. According to Champe, Harvey and Ferrier (2008), the ultimate carbohydrate digestion occur at the mucosal lining of the upper jejunum, declining as they continue down the small intestine, and several disaccharides and oligosaccharides take part in the action. Then, the process of absorption takes place after the process of digestion ends.

Galactose and glucose are transported into the mucosal cells by an active, energy-

requiring process that requires a concurrent uptake of sodium ions; the transport protein is the sodium-dependent glucose cotransporter 1 (SGLT-1). Fructose uptake requires a sodium-independent monosaccharide transporter (GLUT-5) for its absorption. All three monosaccharide are transported from the intestinal mucosal cell into the portal circulation by these yet another transporter, GLUT-2. (Champe, Harvey & Ferrier, 2008, p.87) The glucose which is absorbed by the intestinal mucosal cells will undergo a process known as glycolysis. According to Benjamin III (2004), glycolysis is the foundation for all energy processes in the cell. It is utilized by all tissues in our body for the breakdown of glucose to provide energy and intermediate for other metabolic pathway. The glycolytic pathway describes the oxidation of glucose to pyruvate with the generation of ATP and NADH (Medh, n.d., p.1). Glucose can undergoes glycolysis either under aerobic or anaerobic condition. Under aerobic condition that is with presence of oxygen, the pyruvate can be further oxidized completely and produce carbon dioxide. However, under anaerobic condition, the pyruvate can be fermented to lactate and ethanol. There are several steps involve in aerobic glycolysis. The first and foremost will be the formation of glucose-6-phosphate results from the phosphorylation of glucose. The enzyme involved in the phosphorylation reaction is hexokinase. This reaction needs energy that is ATP and will lead to hydrolysis of ATP to ADP and phosphate. The second reaction will be the formation of fructose-6phosphate by isomerization of glucose-6-phosphate to fructose-6-phosphate (Medh, n.d., p.2). The enzyme known as phosphoglucomutase catalyse the reaction. Next, the third reaction will be another phosphorylation reaction. According to Medh (n.d.), it is an added kinase reaction which involve the phosphorylation of the hydroxyl group at the first carbon of fructose-6-phosphate results in formation of fructose-1, 6bisphosphate. This reaction is catalyzed by the enzyme phosphofructokinase. Since this reaction requires energy, it involves the usage of ATP and dissociates into ADP and phosphate results from hydrolysis. Next reaction is the cleaving of fructose-1, 6bisphosphate to one aldehyde and one ketone which are dihydroxyacetone phosphate and glyceraldehydes 3-phosphate respectively. The fifth reaction involves the isomerization of dihydroxyacetone phosphate. Medh (n.d.) states that,

dihydroxyacetone phosphate and glyceraldehydes can inter-convert to each other

gladly because; they are isomers by the deed of enzyme triose-phosphate isomerase. Thus two molecules of glyceraldehydes are formed per glucose where the dihydroxyacetone phosphate will be diminished in this reaction. Glyceraldehydes are the only substrate which can proceed for the next reaction. The subsequent reaction will be the dehydrogenation process of the glyceraldehydes to 1, 3-

bisphosphoglycerate which is catalysed by the action of enzyme glyceraldehyde 3phosphate dehydrogenase. The seventh reaction will the synthesis of 3-

phosphoglycerate. When 1, 3-bisphosphoglycerate is converted to 3-phosphoglycerate, the high-energy phosphate group of 1, 3-bisphosphoglycerate is used to synthesis ATP from ADP. The enzyme phosphoglycerate kinase catalyzes the reaction. For the eight reaction, the enzyme phosphoglycerate mutase will move the phosphate group from carbon three to carbon two of 3-phosphoglycerate to form 2-phosphoglycerate. For the ninth reaction, dehydration process is occurring. The water molecule is eradicated from 2-phosphoglycerate to form phosphoenolpyruvate and this reaction is catalysed by the enzyme enolase. The last reaction is the formation of pyruvate. Since phosphoenolpyruvate consist a lot of energy in its double bond, the hydrolysis of it will release ATP molecules. This reaction is irreversible reaction and catalysed by the enzyme pyruvate kinase. After the synthesis of pyruvate occurs from glycolysis, it enters Krebs cycle. The Krebs cycle is also known as the citric acid cycle (Krebs Cycle, n.d., p.1). It is the last pathway where most of the oxidative metabolism meets and their carbon skeletons being switch to carbon dioxide. In this cycle, the pyruvate is oxidized and decarboxylated to acetyl CoA in the mitochondria. Pyruvate, the end product of aerobic glycolysis, must be transported into the mitochondria before it can enter the TCA cycle. This is accomplished by a specific pyruvate that helps pyruvate cross the inner mitochondrial membrane. Once in the matrix, pyruvate is converted to acetyl CoA by the pyruvate dehydrogenase complex, which is a multienzyme complex (Champe, Harvey & Ferrier, 2008, p.109). Once the acetyl CoA is formed, it will condense with oxaloacetate to produce citrate which is catalyzed by citrate synthase. Next, the reaction will progress to form isocitrate. This involves the isomerization of citrate which is catalyzed by the enzyme aconitase, which is a Fe-S protein. The third step of this cycle will be the formation of -ketoglutarate. There are two event occur during this step which are oxidation and decarboxylation of the isocitrate. Isocitrate

dehydrogenase catalyzes the irreversible oxidative decarboxylation of isocitrate, yielding the first of three NADH molecules produced by the cycle, and the first release of CO2 (Champe, Harvey & Ferrier, 2008, p.112). The next step is the second oxidative decarboxylation of the -ketoglutarate. The -ketoglutarate is altered to succinyl CoA by the enzyme -ketoglutarate dehydrogenase complex. Thus the reaction discharge second carbon dioxide and generates the second NADH of the citric acid cycle. The fifth reaction will be the production of succinate. This involves the cleavage of succinyl CoA by the enzyme succinyl Coa synthetase. According to Champe, Harvey and Ferrier (2008), this reaction is accompanied with phosphorylation of guanosine diphosphate(GDP) to guanosine triphosphate(GTP). Then the succinate is oxidized to fumarate. This reaction is catalyzed by the enzyme succinate dehydrogenase and subsequently liberates a reduced coenzyme FADH2. The seventh steps will be the formation of the malate. This substrate is produced by hydrolysis of fumarate by the enzyme fumarase. The last reaction of the citric acid cycle will be the production of the oxaloacetate. This can be done by oxidizing the malate preciously to oxaloacetate by the enzyme malate dehydrogenase. From this reaction, it liberates the third and final NADH of the cycle. The last part of carbohydrate metabolisms toward energy production is the electron transport chain. The NADH and FADH2 molecules produced from the glycolysis and Krebs cycles that contain electrons will involve in the electron transport chain. According to Miles (2003), the electrons within NADH and FADH2 molecules have high transfer potential and will be transferred by a system of electron carries to oxygen molecules to form water molecules. The electron transport chain occur within the mitochondria, which it involves the inner membrane and the intermembrane space. The electron transport chain contains four complexes, which are Complex I, Complex II, Complex III and Complex IV. The processes in the electron transport chain are corresponding to the functions of each complex. First, Complex I which is also known as succinate-coenzyme Q reductase will accept electrons from NADH. It will then transfer the electrons to coenzyme Q (CoQ), which is a mobile electron carrier due to its isoprenoid tail that makes it highly hydrophobic and lipophilic. Coenzyme Q can diffuse freely in the bilipid layer of the inner mitochondrial membrane. The process of transferring electrons from NADH to CoQ by Complex I results in the net transport of protons from the matrix side of the inner

mitochondrial membrane to the inter membrane space where the H+ ions accumulate generating a proton motive force (Miles, 2003, p.6). After that, the Complex II which known as succinate-coenzyme Q reductase will receive either one or two electrons from the FADH2 molecules. Then, similar to the Complex I, the electrons received will be transfer to the conenzyme Q, that will further increase the net transport of protons to the inter membrane space. After the coenzyme Q received the electrons from either Complex I or Complex II, it will moves to the Complex III that known as the conenzyme Q reductase, where it transfers the electrons to cytochrome c. Cytochrome c is another mobile electron carrier other than coenzyme Q that can diffuse through the intermembrane space to transfer electrons from the Complex III to Complex IV (Miles, 2003).Complex IV that known as cytochrome reductase will then receive the electrons from the cytochrome c. The electrons accepted will be transferred to the oxygen molecules to form water molecules. This process will also enable the protons to be released into the intermembrane space. When the protons concentrations in the intermembrane space reached a specific concentration, a diffuse gradient will be form where the protons in the intermembrane space will diffuse towards the inner membrane through ATP synthase. This event will trigger the formation of the energy in the form of ATP (adenosine triophosphate) which is essential for the survival of the cells.

References Benjamin III, E. (2004). Review of glycolysis. USA: Morgan State University. Champe, P. C., Harvey, R. A., & Ferrier, D. R. (2008). Lippincotts illustrated reviews: Biochemistry (4th ed.). Philadelphia, USA: Lippincott Williams & Wilkins. Krebs cycle. (n.d.). Retrieved from http://www.dbbtcoc.edu.in/Krebs_Cycle.pdf Medh, J. D. (n.d.). Glycolysis. Retrieved from http://www.csun.edu/~jm77307/Glycolysis.pdf Miles, B. (2003). The electron transport chain. Retrieved from http:// www.tamu.edu/faculty/bmiles/lectures/electrontrans.pdf

Appendix

Carbohydrate Breakdown

Carbohydrates Amylase Glucose Hexokinase Glucose-6phosphate

GLCOLYSIS Fructose-6phosphate Fructose-1, 6biphosphate Aldolase Glyceraldehyde-3phosphate Triosephosphate isomerase 1, 3-bisphosphoglycerate

Phosphohexose isomerase

Phosphofructokinase -1

Dihydroxyacetone phosphate

Glyceraldehyde-3phosphate dehydrogenase

Phosphoglycerate kinase

3-phosphoglycerate Phosphoglycerate mutase Enolase Phosphoenolpyruvate Pyruvate kinase

2-phosphoglycerate

Pyruvate

Pyruvate dehydrogenase Acetyl-CoA

Oxaloacetate

Citrate

Malate dehyrogenase

Citrate synthase

Aconitase

Malate

KREBS CYCLE

Isocitrate

Fumarase

Isocitrate dehydrogenase

Fumarate

-ketoglutarate

Succinic dehydrogenase Succinate

Succinyl-CoA synthetase

-ketoglutarate dehydrogenase Succinyl-CoA

NADH and FADH2 produced from glycolysis and Krebss cycle

Electron transport chain

Complex I NADHcoenzyme Q reductase

Complex II Succinatecoenzyme Q reductase

Complex III Coenzyme Q reductase

Complex IV Cytochrome c reductase