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INTRODUCTION Carbohydrates play several crucial roles in the metabolic processes of living organisms.

They serve as energy sources and as structural elements in living cells. Carbohydrate metabolism, both catabolism and anabolism, form the central core for all of metabolism; the degradation and synthesis of most of the other biological molecules feed into, and out of, this central core of reactions. Carbohydrate metabolism involves a collection of pathways. Glycolysis Hexoses 3-Carbon molecules Gluconeogenesis 3-Carbon molecules Hexoses Fermentation (anaerobic) Citric Acid Cycle (aerobic) Oxidation all the way to CO2 + H2O Pentose-Phosphate pathway Hexose Pentose Glycolysis During glycolysis, each molecule of glucose is converted into two molecules of pyruvate, generating two molecules of NADH (Nicotinamide Adenine Dinucleotide) and two molecules of ATP in the process. Gylcolysis produces ATP without molecular oxygen. It takes place in the cytosol of cells, and is the major source of energy for anaerobic microorganisms.

The glycolytic pathway can be summed up in the following equation: D-Glucose + 2ADP+ 2Pi +2 NAD+ 2 pyruvate +2 ATP+ 2 NADH + 2H+ +2H2O

Fig: Steps in Glycolysis Pathway Glycogenesis Synthesis of glycogen from glucose is called glycogenesis. Glycogen is the major storage form of glucose in our body that is reserved for future use. Glycogenesis mainly occurs in the liver and skeletal muscle. It takes part in the cytoplasm of cells, and requires energy supplied by adenosine triphosphate (ATP) and uridine triphosphate (UTP). In liver, glycogenesis occurs during periods when body has been well fed. In skeletal muscle, glycogenesis occurs during at rest.

Glycogenolysis Breakdown of glycogen to glucose (in liver) or glucose 6-phosphate (in muscle) is called glycogenolysis. Glycogenolysis occurs within the cytoplasm of cells. Glycogen is the major storage carbohydrate in our body. It stored mainly in the liver and skeletal muscle. In liver, glycogenolysis occurs during periods of fasting to maintain blood glucose level. In skeletal muscle, glycogenolysis occurs during active exercise to provide glucose 6-phosphate for glycolysis in response to need for ATP. Muscle glycogen does not directly provide free glucose due to lack of glucose 6-phosphatase enzyme in muscle.

Gluconeogenesis When dietary sources of glucose are insufficient, and glucose stores have been depleted (starch in plants and glycogen in animals), glucose is synthesized from non-carbohydrate compounds by a series of cytosolic reactions called the gluconeogenic pathway as shown in figure 10. Gluconeogenesis converts pyruvate to glucose using a set of reactions that require energy input in the form of ATP and GTP (gluconeogenesis costs 4 ATP and 2 GTP to synthesize one glucose from two pyruvate). Importantly, gluconeogenesis is not simply the reversal of glycolysis, and is in fact, a highly regulated pathway (as is glycolysis) to prevent futile cycling between glucose degradation by glycolysis and glucose synthesis by gluconeogenesis.

Fig: Steps in Gluconeogenesis

Citric Acid Cycle: The overall rate of the citric acid cycle (also known as Krebs Cycle and Tricarboxalic Acid Cycle) is controlled by the rate of conversion of pyruvate to acetyl-CoA and by the flux through citrate synthase, isocitrate dehydrogenase, and _-ketoglutarate dehydrogenase. These fluxes are largely determined by the concentrations of substrates and products: the end products ATP and NADH are inhibitory, and the substrates NAD+ and ADP are stimulatory. The production of acetyl-CoA for the citric acid cycle by the PDH complex is inhibited allosterically by metabolites that signal a sufficiency of metabolic energy (ATP, acetyl- CoA, NADH, and fatty acids) and stimulated by metabolites that indicate a reduced energy supply (AMP, NAD+, CoA).

Fig: Citric Acid Cycle/ Krebs Cycle

Pentose phosphate pathway The pentose phosphate pathway is the major source for the NADPH required for anabolic processes. There are three distinct phases each of which has a distinct outcome. Depending on the needs of the organism the metabolites of that outcome can be fed into many other pathways. Gluconeogenesis is directly connected to the pentose phosphate pathway. As the need for glucose-6-phosphate (the beginning metabolite in the pentose phosphate pathway) increases so does the activity of gluconeogenesis. Pentose Phosphate Pathways has following phases: Oxidation Phase The beginning molecule for the PPP is glucose-6-P which is the second intermediate metabolite in glycolysis. Glucose-6-P is oxidized in the presence of glucose-6-P dehydrogenase and NADP+. This step is irreversible and is highly

regulated. NADPH and fatty acyl-CoA are strong negative inhibitors to this enzyme. The purpose of this is to decrease production of NADPH when concentrations are high or the synthesis of fatty acids is no longer necessary. The metabolic product of this step is gluconolactone which is hydrolytrically unstable. Gluconolactonase causes gluconolactone to undergo a ring opening hydrolysis. The product of this reaction is the more stable sugar acid, 6-phosphoD-gluconate. + 6-phospho-D-gluconate is oxidized by NADP in the presence of 6phosphogluconate dehydrogenase which yields ribulose-5-phosphate. The oxidation phase of the PPP is solely responsible for the production of the NADPH to be used in anabolic processes. Isomerization Phase Ribulose-5-phosphate can then be isomerized by phosphopentose isomerase to produce ribose-5-phosphate. Ribose-5-phosphate is one of the main building blocks of nucleic acids and the PPP is the primary source of production of ribose5-phosphate. If production of ribose-5-phosphate exceeds the needs of required ribose-5phosphate in the organism, then phosphopentose epimerase catalyzes a chiralty rearrangement about the center carbon creating xylulose-5-phosphate. The products of these two reactions can then be rearranged to produce many different length carbon chains. These different length carbon chains have a variety of metabolic fates. Rearrangement Phase There are two main classes of enzymes responsible for the rearrangement and synthesis of the different length carbon chain molecules. These are transketolase and transaldolase. Transketolase is responsible for the cleaving of a two carbon unit from xylulose5-P and adding that two carbon unit to ribose-5-P thus resulting in glyceraldehyde-3-P and sedoheptulose-7-P. Transketolase is also responsible for the cleaving of a two carbon unit from xylulose-5-P and adding that two carbon unit to erythrose-4-P resulting in glyceraldehyde-3-P and fructose-6-P. Transaldolase is responsible for cleaving the three carbon unit from sedoheptulose-7-P and adding that three carbon unit to glyceraldehyde-3-P thus resulting in erythrose-4-P and fructose-6-P. The end results of the rearrangement phase is a variety of different length sugars which can be fed into many other metabolic processes. For example, fructose-6-P is a key intermediate of glycolysis as well as glyceraldehyde-3-P.

Galactose metabolism Galactose, which is metabolized from the milk sugar, lactose (a disaccharide of glucose and galactose), enters glycolysis by its conversion to glucose-1-phosphate (G1P). This occurs through a series of steps. First the galactose is phosphorylated by galactokinaseto yield galactose-1-phosphate. Epimerization of galactose-1 phosphate to G1P requires the transfer of UDP from uridine diphosphoglucose (UDP-glucose) catalyzed by galactose-1-phosphate uridyl transferase. This generates UDP-galactose and G1P. The UDP-galactose is epimerized to UDP glucose by UDP-galactose-4 epimerase. The UDP portion is exchanged for phosphate generating glucose-1-phosphate which then is converted to G6P by phosphoglucose mutase.

Fig: Steps in Galactose Metabolism