3.2.

1 GLYCOLYSIS AND FERMENTATION Glycolysis is an anaerobic metabolic pathway, found in the cytosol of all cells, which forms adenosine triphosphate ( ATP ) by degrading glucose . It also serves as a source of precursors for other pathways, and as a recipient of products of various pathways for use as metabolic fuels. Its universal and central role in metabolism suggests that glycolysis evolved early in the history of life. In the overall reaction for glycolysis, one molecule of glucose is converted to two molecules of pyruvic acid. Along the way, two molecules of adenosine diphosphate ( ADP ) are phosphorylated to ATP, and two molecules of NAD (the oxidized form of NAD, or nicotinamide adenine dinucleotide) are reduced to NADH. ATP serves as an energy carrier and can be used to power many cellular processes. The NADH carries highenergy electrons, which can be used to produce more ATP by chemiosmosis . The process of glycolysis. In glycolysis, the six-carbon glucose (shown without its hydrogens or oxygens) is first destabilized by the addition of ATP, and then split. Further transformations create NADH and new ATP, leaving a pair of threecarbon pyruvates. The pyruvic acid can be further oxidized by the Krebs cycle to yield additional ATP. The ten steps of glycolysis can be divided into two stages. The first five steps, the preparatory, or priming, phase of glycolysis, prepare the glucose by phosphorylating it twice, using two molecules of ATP as sources of phosphate. This increases the energy content of the glucose, so the preparatory phase is also sometimes called the investment stage, reflecting the need to invest two ATP molecules before a net yield of energy can be achieved. During the second five reactions, the payoff phase, the fructose-1,6-bisphosphate formed during the preparatory phase is dephosphorylated and cleaved, forming two molecules of pyruvate and four of ATP. Because two ATPs are used and four are produced during glycolysis, there is a net production of two molecules of ATP for every glucose consumed. Since glycolysis plays a central role in cellular metabolism, it has several control points. Like most pathways, it is regulated during its early steps. Hexokinase, the enzyme that catalyzes the first reaction, is inhibited by its product, glucose-6-phosphate (G-6-P). The third enzyme, phosphofructokinase (PFK), is regulated in a complex manner by several metabolites , and is also under indirect hormonal control. The last glycolytic enzyme, pyruvate kinase , is regulated by several metabolites, including ATP, which inhibits it. These control mechanisms have the effect of maintaining a constant supply of ATP for the cell, since production of ATP inhibits the process, and depletion of ATP activates it. Aerobically respiring cells will produce even more ATP through oxidative phosphorylation . However, cells that cannot respire aerobically , either because they lack the necessary metabolic pathways or because they live in anaerobic environments, cannot do this. This presents a problem since all cells must continually regenerate the NAD needed during the In the absence of oxygen the pyruvate is converted to NAD in reactions collectively referred to as fermentations. preparatory phase of glycolysis. All such cells accomplish this by converting the pyruvate to another product, oxidizing NADH to NAD in the process. These reactions are collectively referred to as fermentations. Animals, some plants, and most bacteria produce lactic acid, whereas yeast and a few bacteria produce carbon dioxide and ethanol. Rarer fermentations produce a variety of organic molecules such as other alcohols and organic acids. Fermentations are used extensively by industry to produce these compounds cheaply, as well as to produce foods such as yogurt, bread, wine, and beer. Fermentation in food processing typically is the conversion of carbohydrates toalcohols and carbon dioxide or organic acids using yeasts, bacteria, or a combination thereof, under anaerobic conditions. Fermentation in simple terms is the chemical conversion of sugars into ethanol. The science of fermentation is also known as zymology, or zymurgy. Fermentation usually implies that the action of microorganisms is desirable, and the process is used to produce alcoholic beverages such as wine, beer, and cider. Fermentation is also employed in the leavening of bread (CO2 produced by yeast activity), and for preservation techniques to produce lactic acid in sour foods such assauerkraut, dry sausages, kimchi and yogurt, or vinegar (acetic acid) for use inpickling foods. Fermentation is the process of extracting energy from the oxidation of organic compounds, such as carbohydrates, using an endogenous electron acceptor, which is usually an organic compound.[1] In

contrast, respiration is where electrons are donated to an exogenous electron acceptor, such as oxygen, via an electron transport chain. Fermentation is important in anaerobicconditions when there is no oxidative phosphorylation to maintain the production of ATP (adenosine triphosphate) by glycolysis. During fermentation, pyruvate is metabolised to various different compounds. Homolactic fermentation is the production of lactic acid from pyruvate; alcoholic fermentation is the conversion of pyruvate into ethanol and carbon dioxide; and heterolactic fermentation is the production of lactic acid as well as other acids and alcohols. Fermentation does not necessarily have to be carried out in an anaerobic environment. For example, even in the presence of abundant oxygen, yeast cells greatly prefer fermentation tooxidative phosphorylation, as long as sugars are readily available for consumption (a phenomenon known as the Crabtree effect).[2] Sugars are the most common substrate of fermentation, and typical examples of fermentation products are ethanol, lactic acid, lactose, andhydrogen. However, more exotic compounds can be produced by fermentation, such as butyric acid and acetone. Yeast carries outfermentation in the production of ethanol in beers, wines, and other alcoholic drinks, along with the production of large quantities of carbon dioxide. Fermentation occurs in mammalian muscle during periods of intense exercise where oxygen supply becomes limited, resulting in the creation of lactic acid.[3] Fermentation is one of the oldest known food preservation techniques. Along with drying and salting, fermentation was a key method of extending the life of foods, allowing them to be available, and eaten safely, in times of scarcity or seasonal nonavailability. These methods helped allow the transition from hunting and gathering to organized food cultivation and storage, which took place some ten to fifteen thousand years ago in the Middle East. Fermentation involves the action of desirable microorganisms, or their enzymes, on food ingredients to make biochemical changes, which cause significant modification to the food. Often lactic-acid bacteria convert the carbohydrate energy source of food, such as lactose in milk, to lactic acid; examples are yogurt and cheeses from milk, and pickles from fruits and vegetables. Alternatively, yeasts, often of theSaccharomyces species, may convert the glucose to ethanol and carbon dioxide in leavened breads, or the sugars in grain or fruit beverages to beers and wines. Molds also can be active in certain fermentations, such as Stilton cheese and soy sauce. It is estimated that about one-third of all the food we consume is fermented. World estimates for beer consumption are about 22 million gallons, and a total of 15 million tons of some one thousand varieties of cheese are eaten annually. 3.2.2 PYRUVIC ACID AEROBIC RESPIRATION Aerobic respiration requires oxygen in order to generate energy (ATP). Although carbohydrates,fats, and proteins can all be processed and consumed as reactant, it is the preferred method ofpyruvate breakdown in glycolysis and requires that pyruvate enter the mitochondrion in order to be fully oxidized by the Krebs cycle. The product of this process is energy in the form of ATP (Adenosine triphosphate), by substrate-level phosphorylation, NADH and FADH2 C6H12O6 (aq) + 6 O2 (g) 6 CO2 (g) + 6 H2O (l) Simplified reaction: G = -2880 kJ per mole of C6H12O6 The negative G indicates that the reaction can occur spontaneously. The reducing potential of NADH and FADH2 is converted to more ATP through an electron transport chain with oxygen as the "terminal electron acceptor". Most of the ATP produced by aerobic cellular respiration is made by oxidative phosphorylation. This works by the energy released in the consumption of pyruvate being used to create a chemiosmotic potential by pumping protons across a membrane. This potential is then used to drive ATP synthase and produce ATP from ADP and a phosphate group. Biology textbooks often state that 38 ATP molecules can be made per oxidised glucose molecule during cellular respiration (2 from glycolysis, 2 from the Krebs cycle, and about 34 from the electron transport system).[2] However, this maximum yield is never quite reached due to losses (leaky membranes) as well as the cost of moving pyruvate and ADP into the mitochondrial matrix and current estimates range around 29 to 30 ATP per glucose.[2]

Aerobic metabolism is 19 times more efficient than anaerobic metabolism (which yields 2 mol ATP per 1 mol glucose). They share the initial pathway of glycolysis but aerobic metabolism continues with the Krebs cycle and oxidative phosphorylation. The post glycolytic reactions take place in the mitochondria in eukaryotic cells, and in the cytoplasm in prokaryotic cells. 3.2.3 : THE KREBS CITRIC ACID CYCLE When oxygen is present, respiration can harness more ATP from a single unit of glucose. The pyruvic acid from the glycolysis stage diffuses into a cell organelle called a mitochondrion (pl. mitochondria). These mitochondria are sausage shaped structures that host a large surface area for the respiration to occur on. The pyruvic acid is then subject to more enzymes which break it down into a 2 carbon compound, as seen below. The diagram illustrates theKreb's cycle, consisting of three main actions The carbon element is in an infinite cycle where the 2 carbon compound derived from pyruvic acid binds with the 4 carbon compound that is always present in the cycle. CO2 is released, where the oxygen that is present in aerobic respiration combines with carbon from the carbon compounds which is released as CO2. Hence the need for animals to breath out and expel this CO2. Enzymes oxidize the carbon compounds and transport the hydrogen atoms to the cytochrome system. The citric acid cycle also known as thetricarboxylic acid cycle (TCA cycle), the Krebs cycle, or the SzentGyrgyi-Krebs cycle[1][2] is a series of chemical reactions which is used by all aerobic living organisms to generate energy through the oxidization of acetate derived fromcarbohydrates, fats and proteins into carbon dioxide and water. In addition, the cycle providesprecursors for the biosynthesis of compounds including certain amino acids as well as thereducing agent NADH that is used in numerous biochemical reactions. Its central importance to many biochemical pathways suggests that it was one of the earliest established components of cellular metabolism and may have originatedabiogenically.[3] The name of this metabolic pathway is derived from citric acid (a type of tricarboxylic acid) which is first consumed and then regenerated by this sequence of reactions to complete the cycle. In addition, the cycle consumes acetate in the form of acetyl-CoA, reduces NAD+ to NADH, and produces carbon dioxide. The NADH generated by the TCA cycle is fed into the oxidative phosphorylation pathway. The net result of these two closely linked pathways is the oxidation of nutrients to produce energy in the form ofATP. In eukaryotic cells, the citric acid cycle occurs in the matrix of the mitochondrion. Bacteria also use the TCA cycle to generate energy, but since they lack mitochondria, the reaction sequence is performed in the cytosol. The components and reactions of the citric acid cycle were established in the 1930s by seminal work from the Nobel laureates Albert Szent-Gyrgyi[4] and Hans Adolf Krebs.[5] 3.2.4: OXIDATIVE PHOSPHORYLATION The NADH and FADH2 formed in glycolysis, fatty acid oxidation, and the citric acid cycle are energy-rich molecules because each contains a pair of electrons having a high transfer potential. When these electrons are used to reduce molecular oxygen to water, a large amount of free energy is liberated, which can be used to generate ATP. Oxidative phosphorylation is the process in which ATP is formed as a result of the transfer of electrons from NADH or FADH 2 to O 2 by a series of electron carriers. This process, which takes place in mitochondria, is the major source of ATP in aerobic organisms (Figure 18.1). For example, oxidative phosphorylation generates 26 of the 30 molecules of ATP that are formed when glucose is completely oxidized to CO2 and H2O. Oxidative phosphorylation is conceptually simple and mechanistically complex. Indeed, the unraveling of the mechanism of oxidative phosphorylation has been one of the most challenging problems of biochemistry. The flow of electrons from NADH or FADH2 to O2 through protein complexes located in the mitochondrial inner membrane leads to the pumping of protons out of the mitochondrial matrix. The resulting uneven distribution

of protons generates a pH gradient and a transmembrane electrical potential that creates a proton-motive force.ATP is synthesized when protons flow back to the mitochondrial matrix through an enzyme complex. Thus, the oxidation of fuels and the phosphorylation of ADP are coupled by a proton gradient across the inner mitochondrial membrane (Figure 18.2). Oxidative phosphorylation is the culmination of a series of energy transformations that are called cellular respiration or simply respiration in their entirety. First, carbon fuels are oxidized in the citric acid cycle to yield electrons with high transfer potential. Then, this electron-motive force is converted into a proton-motive force and, finally, the proton-motive force is converted into phosphoryl transfer potential. The conversion of electron-motive force into proton-motive force is carried out by three electron-driven proton pumpsNADHQ oxidoreductase, Q-cytochrome c oxidoreductase, and cytochrome c oxidase. These large transmembrane complexes contain multiple oxidation-reduction centers, including quinones, flavins, iron-sulfur clusters, hemes, and copper ions. The final phase of oxidative phosphorylation is carried out by ATP synthase, an ATP-synthesizing assembly that is driven by the flow of protons back into the mitochondrial matrix. Components of this remarkable enzyme rotate as part of its catalytic mechanism. Oxidative phosphorylation vividly shows thatproton gradients are an interconvertible currency of free energy in biological systems. Oxidative phosphorylation (or OXPHOS in short) is a metabolic pathway that uses energy released by the oxidation of nutrients to produce adenosine triphosphate (ATP). Although the many forms of life on earth use a range of different nutrients, almost all aerobic organisms carry out oxidative phosphorylation to produce ATP, the molecule that supplies energy to metabolism. This pathway is probably so pervasive because it is a highly efficient way of releasing energy, compared to alternative fermentation processes such as anaerobic glycolysis. During oxidative phosphorylation, electrons are transferred fromelectron donors to electron acceptors such as oxygen, in redox reactions. These redox reactions release energy, which is used to form ATP. In eukaryotes, these redox reactions are carried out by a series of protein complexes within mitochondria, whereas, inprokaryotes, these proteins are located in the cells' inner membranes. These linked sets of proteins are called electron transport chains. In eukaryotes, five main protein complexes are involved, whereas in prokaryotes many different enzymes are present, using a variety of electron donors and acceptors. The energy released by electrons flowing through this electron transport chain is used to transport protons across the inner mitochondrial membrane, in a process called chemiosmosis. This generates potential energy in the form of a pH gradient and anelectrical potential across this membrane. This store of energy is tapped by allowing protons to flow back across the membrane and down this gradient, through a large enzyme called ATP synthase. This enzyme uses this energy to generate ATP from adenosine diphosphate(ADP), in a phosphorylation reaction. This reaction is driven by the proton flow, which forces the rotation of a part of the enzyme; the ATP synthase is a rotary mechanical motor. Although oxidative phosphorylation is a vital part of metabolism, it produces reactive oxygen species such as superoxide and hydrogen peroxide, which lead to propagation of free radicals, damaging cells and contributing to disease and, possibly, aging (senescence). The enzymes carrying out this metabolic pathway are also the target of many drugs and poisons that inhibit their activities.
SOURCE: Read more: Glycolysis and Fermentation - Biology Encyclopedia - cells, body, process, cycle, life, used, molecules, energy http://www.biologyreference.com/FoGr/Glycolysis-and-Fermentation.html#ixzz1dlJRZ2mW http://en.wikipedia.org/wiki/Glycolysis http://en.wikipedia.org/wiki/Citric_acid_cycle http://en.wikipedia.org/wiki/Oxidative_phosphorylation http://www.ncbi.nlm.nih.gov/books/NBK21208/