FERMENTATION

originally, the foaming that occurs during the manufacture of wine and beer, a process at least 10,000 years old. That the frothing results from the evolution of carbon dioxide gas was not recognized until the 17th century. Louis Pasteur in the 19th century used the term fermentation in a narrow sense to describe the changes brought about by yeasts and other microorganisms growing in the absence of air (anaerobically); he also recognized that ethyl alcohol and carbon dioxide are not the only products of fermentation. In the 1920s it was discovered that extracts of muscle catalyze, in the absence of air, the formation of lactate from glucose and that the same intermediate compounds formed in the fermentation of grain are produced by muscle. An important generalization thus emerged: that fermentation reactions are not peculiar to the action of yeast but also occur in many other instances of glucose utilization. Glycolysis, which means dissolution of sugar, was originally defined around 1930 as the splitting of sugar into lactate. It can be further defined as that form of fermentation, characteristic of cells in general, in which the six-carbon sugar glucose is broken down into two molecules of the three-carbon organic acid, pyruvic acid (or its ionized form pyruvate), coupled with the transfer of chemical energy to the synthesis of adenosine triphosphate (ATP). The pyruvate may then be oxidized, in the presence of oxygen, through the tricarboxylic acid cycle, or in the absence of oxygen, be reduced to lactic acid, alcohol, or other products. The sequence from glucose to pyruvate is often called the EmbdenMeyerhof pathway after two German biochemists, who, in the late 1920s and 1930s, postulated and analyzed experimentally the critical steps in that series of reactions. The term fermentation now denotes the enzyme-catalyzed, energy-yielding pathway in cells by which fuel molecules such as glucose are broken down anaerobically. In most cells the enzymes occur in the soluble portion of the cytoplasm. The reactions leading to the formation of pyruvate thus are common to sugar transformation in muscle, yeasts, some bacteria, and plants. One product of the pathway is always the energy-rich compound adenosine triphosphate (ATP). The other product, pyruvate, can undergo various transformations, depending on the cell type and the availability of oxygen. Industrial fermentation processes begin with suitable microorganisms and specified conditions; e.g., careful adjustment of nutrient concentration. The products are of many types: alcohol, glycerol, and carbon dioxide from yeast fermentation of various sugars; butyl alcohol, acetone, lactic acid, monosodium glutamate, and acetic acid from various bacteria; citric acid, gluconic acid, and small amounts of antibiotics, vitamin B12, and riboflavin (vitamin B2) from mold fermentation.