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20-September-2008 09:29:05 - Fermentation biochemistry April 2007 All or part of this article may be confusing or unclear. Please help clarify the article. Suggestions may be on the talk page. For other uses, see Fermentation. Fermentation in progress Fermentation in progress Fermentation is the process of deriving energy from the oxidation of organic compounds, such as carbohydrates, using an endogenous electron acceptor, which is usually an organic compound.1 Sugars are the common substrate of fermentation, and typical examples of fermentation products are ethanol, lactic acid, and hydrogen. However, more exotic compounds can be produced by fermentation, such as butyric acid and acetone. Yeast carries out fermentation 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.2 Contents 1 History 2 Reaction 3 Energy source in anaerobic conditions 4 Products 5 Enzymology and Zymology 6 See also 7 References 8 External links History French chemist Louis Pasteur was the first zymologist, when, in 1857, he connected yeast to fermentation. Pasteur originally defined fermentation as respiration without air. Pasteur performed careful research and concluded, I am of the opinion that alcoholic fermentation never occurs without simultaneous organization, development and multiplication of cells.... If asked, in what consists the chemical act whereby the sugar is decomposed ... I am completely ignorant of it. German chemist and zymologist, Eduard Buchner, winner of the 1907 Nobel Prize in chemistry, later determined that fermentation is actually caused by a yeast secretion that he termed zymase. Reaction See also: glycolysis The reaction of fermentation differs according to the sugar being used and the product produced. In the reaction shown below, the sugar will be glucose C6H12O6, the simplest sugar, and the product will be ethanol 2C2H5OH. This is one of the fermentation reactions carried out by yeast, and used in food production. Chemical equation C6H12O6 → 2 CH3CH2OH + 2 CO2 + 2 ATP energy released:118 kJ/mol Word equation Sugar glucose or fructose → alcohol ethanol + carbon dioxide + energy ATP The actual biochemical pathway the reaction takes varies depending on the sugars involved, but the most common involves part of the glycolysis pathway, which is shared with the early stages of aerobic respiration in most organisms. The later stages of the pathway vary considerably depending on the final product. Energy source in anaerobic conditions Fermentation is thought to have been the primary means of energy production in earlier organisms before oxygen was at high concentration in the atmosphere, and thus would represent a more ancient form of energy production in cells. Fermentation products contain chemical energy they are not fully oxidized but are considered waste products, since they cannot be metabolized further without the use of oxygen or other more highly-oxidized electron acceptors. A consequence is that the production of adenosine triphosphate ATP by fermentation is less efficient than oxidative phosphorylation, whereby pyruvate is fully oxidized to carbon dioxide. Fermentation produces 4 ATP molecules per molecule of glucose, compared to 38 by aerobic respiration: 8 are produced from FADH2, and 30 are produced from NADH, for a total of 38. Since 2 ATP molecules are used in glycolysis, the net yield for fermentation is 2 ATP versus 36 ATP from aerobic respiration. Aerobic glycolysis is a method employed by muscle cells for the production of lower-intensity energy over a longer period of time when oxygen is plentiful. Under low-oxygen conditions, however, vertebrates use the less-efficient but faster anaerobic glycolysis to produce ATP. The speed at which ATP is produced is about 100 times that of oxidative phosphorylation.citation needed While fermentation is helpful during short, intense periods of exertion, it is not sustained over extended periods in complex aerobic organisms. In humans, for example, lactic acid fermentation provides energy for a period ranging from 30 seconds to 2 minutes. The final step of fermentation, the conversion of pyruvate to fermentation end-products, does not produce energy. However, it is critical for an anaerobic cell, since it regenerates nicotinamide adenine dinucleotide NAD+, which is required for glycolysis. This is important for normal cellular function, as glycolysis is a primary source of ATP production in anaerobic conditions. Products Products produced by fermentation are actually waste products produced during the reduction of pyruvate to regenerate NAD+ in the absence of oxygen. In general, bacteria produce acids: Vinegar acetic acid is the direct result of bacterial metabolism Bacteria need oxygen to convert the alcohol to acetic acid. In milk, the acid coagulates the casein, producing curds. In pickling, the acid preserves the food from pathogenic and putrefactive bacteria. When yeast ferments, it breaks down the glucose C6H12O6 into exactly two molecules of ethanol CH3CH2OH and two molecules of carbon dioxide CO2. Ethanol fermentation performed by yeast and some types of bacteria breaks the pyruvate down into ethanol and carbon dioxide. It is important in bread-making, brewing, and wine-making. Usually only one of the products is desired; in bread-making, the alcohol is baked out, and, in alcohol production, the carbon dioxide is released into the atmosphere or used for carbonating the beverage. When the ferment has a high concentration of pectin, minute quantities of methanol can be produced. Lactic acid fermentation breaks down the pyruvate into lactic acid. It occurs in the muscles of animals when they need energy faster than the blood can supply oxygen. It also occurs in some bacteria and some fungi. It is this type of bacteria that converts lactose into lactic acid in yogurt, giving it its sour taste. Hydrogen gas is produced in many types of fermentation mixed acid fermentation, butyric acid fermentation, caproate fermentation, butanol fermentation, glyoxylate fermentation, as a way to regenerate NAD+ and FAD from NADH and FADH2. Electrons are transferred to ferredoxin, which in turn is oxidized by hydrogenase, producing H2. Hydrogen gas is a substrate for methanogens and sulfate reducers, which keep the concentration of hydrogen sufficiently low to allow the production of such an energy-rich compound. 3 Enzymology and Zymology Zymology is the scientific term for the study of the chemical process of fermentation, and includes fermentation by yeast, bacteria, and other lower species. Zymology includes the study of enzymes that facilitate the chemical process of fermentation, as well as engineering issues of yeast selection, yeast physiology, and the practical issues of brewing. Enzymology, the study of enzymes, should not be confused with zymology, as the modern study of the enzyme encompasses enzymes from a multitude of sources, and studies enzymes in many terms, as well as studying enzymes from biochemical pathways including fermentation and beyond. Historically, zymologists were the first enzymologists, and since enzymology has expanded beyond the study of yeast fermentation, zymology is infrequently regarded as an archaic word for enzymology. See also Fermentation wine Fermentation food Industrial fermentation Fermentation lock Fed-batch Chemostat Ethanol fermentation References ^ Prescott, Harley, and Klein 2005 Microbiology, 6th ed., McGraw-Hill, New York,clarify This is in contrast to cellular respiration, where electrons are donated to an exogenous electron acceptor, such as oxygen, via an electron transport chain.ref/ref Fermentation does not necessarily have to be carried out in an anaerobic environment, however. For example, even in the presence of abundant oxygen, yeast cells greatly prefer fermentation to oxidative phosphorylation, as long as sugars are readily available for consumption.refDickinson, J. R. 1999. Carbon metabolism. In: The Metabolism and Molecular Physiology of Saccharomyces cerevisiae, ed. J. R. Dickinson and M. Schweizer, Philadelphia, PA: Taylor Francis./li li id=cite_note-Voet_and_Voet.2C_Biochemistry-1'''#cite_ref-Voet_and_Voet.2C_Biochemistry_1-0|^''' Voet and Voet 1995 Biochemistry, 2nd ed., John Wiley Sons, Inc., New York, NY/li li id=cite_note-Brock-2'''#cite_ref-Brock_2-0|^''' Madigan, Martinko, ''Brock Biology of Microorganisms'', 11th ed/li/ol/ref External links The chemical logic behind fermentation and respiration Inline disintegration to reduce fermentation time and improve the yield v d e Metabolism: carbohydrate metabolism Anabolism Gluconeogenesis - Glycogenesis - Photosynthesis Carbon fixation Carbohydrate catabolism Glycolysis - Glycogenolysis - Fermentation Ethanol, Lactic acid - Cellular respiration - Xylose metabolism Other Pentose phosphate pathway Retrieved from http://en..org/wiki/Fermentation_biochemistry Categories: Anaerobic digestion | Oenology | Fermented beverages | Brewing | Food science | Metabolism | Food preservation | Alchemical processes | Fermentation | Microbiology | MycologyHidden categories: articles needing clarification | Articles needing additional references from April 2007 | All articles with statements | Articles with statements since February 2007 Views Article Discussion this page History Personal tools Log in / create account Navigation Main page Contents Featured content Current events Random article Search Go Search Interaction Community portal Recent changes Contact Donate to Help Toolbox What links here Related changes Upload file Special pages Printable version Permanent link Cite this page Languages Asturianu বাংলা БългарÑ?ки Català Česky Dansk Deutsch Eesti Español Esperanto Euskara Ù?ارسی Français Gaeilge Gà idhlig í•œêµì–´ Bahasa Indonesia Italiano עברית Lietuvių Magyar Nederlands 日本語 ‪Norsk nynorsk‬ Polski Português Română Runa Simi РуÑ?Ñ?кий Shqip Sicilianu Simple English SlovenÅ¡Ä?ina СрпÑ?ки / Srpski Srpskohrvatski / СрпÑ?кохрватÑ?ки Suomi Türkçe УкраїнÑ?ька ä¸æ–‡ This page was last modified on 4 August 2008, at 13:19
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