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148       Metabolism



             Fermentations                                    lated by pyruvate decarboxylase [2], which
                                                              does not occur in animal metabolism, to pro-
             As discussed on p. 146, degradation of glucose   duce acetaldehyde (ethanal). When this is re-
             to pyruvate is the only way for most organ-      duced by alcohol dehydrogenase [3], with
             isms to synthesize ATP in the absence of oxy-    NADH being consumed, ethanol [3] is formed.
                               +
             gen.The NADH+H that is also formed in this          Yeasts, unicellular fungi that belong to the
             process has to be constantly reoxidized to       eukaryotes (3), rather than bacteria, are re-
             NAD  +  in order to maintain glycolysis and      sponsible for this type of fermentation. Yeasts
             thus ATP synthesis. In the animal organism,      are also often used in baking. They produce
             this is achieved by the reduction of pyruvate    CO 2 and ethanol, which raise the dough.
             to lactate. In microorganisms, there are many    Brewers’ and bakers’ yeasts (Saccharomyces
                                 +
             other forms of NAD regeneration. Processes       cerevisiae) are usually haploid and reproduce
             of this type are referred to as fermentations.   asexually by budding (3). They can live both
             Microbial fermentation processes are often       aerobically and anaerobically. Wine is pro-
             used to produce foodstuffs and alcoholic bev-    duced by other types of yeast, some of which
             erages, or to preserve food. Features common     already live on the grapes. To promote the
             to all fermentation processes are that they      formation of ethanol, efforts are made to gen-
             start with pyruvate and only occur under         erally exclude oxygen during alcoholic fer-
             anaerobic conditions.                            mentation—for example, by covering dough
                                                              with a cloth when it is rising and by ferment-
                                                              ing liquids in barrels that exclude air.
             A. Lactic acid and propionic acid
             fermentation
                                                              C. Beer brewing
             Many milk products, such as sour milk, yo-
             gurt, and cheese are made by bacterial lactic    Barley is the traditional starting material for
             acid fermentation (1). The reaction is the same  the brewing of beer. Although cereal grains
             as in animals. Pyruvate, which is mainly de-     contain starch, they hardly have any free sug-
             rived from degradation of the disaccharide       ars. The barley grains are therefore first al-
             lactose (see p. 38), is reduced to lactate by    lowed to germinate so that starch-cleaving
             lactate dehydrogenase [1]. Lactic acid fermen-   amylases are formed. Careful warming of the
             tation also plays an important role in the       sprouting grain produces malt. This is then
             production of sauerkraut and silage. These       ground, soaked in water, and kept warm for
             products usually keep for a long time, because   a certain time. In the process, a substantial
             the pH reduction that occurs during fermen-      proportion of the starch is broken down into
             tation inhibits the growth of putrefying bac-    the disaccharide maltose (see p. 38). The
             teria.                                           product (the wort) is then boiled, yeast and
                Bacteria from the genera Lactobacillus and    hops are added, and the mixture is allowed to
             Streptococcus are involved in the first steps of  ferment for several days. The addition of hops
             dairy production (3). The raw materials pro-     makes the beer less perishable and gives it its
             duced by their effects usually only acquire      slightly bitter taste. Other substances con-
             their final properties after additional fermen-  tained in hops act as sedatives and diuretics.
             tation processes. For example, the character-
             istic taste of Swiss cheese develops during a
             subsequent propionic acid fermentation. In
             this  process,   bacteria  from   the   genus
             Propionibacterium convert pyruvate to propi-
             onate in a complex series of reactions (2).


             B. Alcoholic fermentation

             Alcoholic beverages are produced by the fer-
             mentation of plant products that have a high
             carbohydrate content. Pyruvate, which is
             formed from glucose, is initially decarboxy-


           Koolman, Color Atlas of Biochemistry, 2nd edition © 2005 Thieme
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