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336       Tissues and organs



             Muscle metabolism I                              maintain glucose degradation and thus ATP
                                                              formation. If there is a lack of O 2 ,this is
             Muscle contraction is associated with a high     achieved by the formation of lactate,which
             level of ATP consumption (see p. 332). With-     is released into the blood and is resynthesized
             out constant resynthesis, the amount of ATP      into glucose in the liver (Cori cycle; see
             available in the resting state would be used up  p. 338).
             in less than 1 s of contraction.                    Muscle-specific auxiliary reactions for ATP
                                                              synthesis exist in order to provide additional
                                                              ATP in case of emergency. Creatine phosphate
             A. Energy metabolism in the white and red
                                                              (see B) acts as a buffer for the ATP level.
             muscle fibers
                                                              Another ATP-supplying reaction is catalyzed
             Muscles contain two types of fibers, the pro-    by adenylate kinase [1] (see also p. 72). This
             portions of which vary from one type of          disproportionates two molecules of ADP into
             muscle to another. Red fibers (type I fibers)    ATP and AMP. The AMP is deaminated into
             are suitable for prolonged effort. Their metab-  IMP in a subsequent reaction [2] in order to
             olism is mainly aerobic and therefore depends    shift the balance of the reversible reaction [1]
             on an adequate supply of O 2 . White fibers      in the direction of ATP formation.
             (type II fibers) are better suited for fast, strong
             contractions. These fibers are able to form
                                                              B. Creatine metabolism
             suf cient ATP even when there is little O 2
             available. With appropriate training, athletes   Creatine (N-methylguanidoacetic acid) and its
             and sports participants are able to change the   phosphorylated form creatine phosphate
             proportions of the two fiber types in the mus-   (a guanidophosphate) serve as an ATP buffer
             culature and thereby prepare themselves for      in muscle metabolism. In creatine phosphate,
             the physiological demands of their disciplines   the phosphate residue is at a similarly high
             in a targeted fashion. The expression of func-   chemical potential as in ATP and is therefore
             tional muscle proteins can also change during    easily transferred to ADP. Conversely, when
             thecourseof training.                            there is an excess of ATP, creatine phosphate
                Red fibers provide for their ATP require-     can arise from ATP and creatine. Both proce-
             ments mainly (but not exclusively) from fatty    sses are catalyzed by creatine kinase [5].
             acids,which arebrokendown via β-oxidation,          In resting muscle, creatine phosphate
             the tricarboxylic acid cycle, and the respira-   forms due to the high level of ATP. If there is
             tory chain (right part of the illustration). The  a risk of a severe drop in the ATP level during
             red color in these fibers is due to the mono-    contraction, the level can be maintained for a
             meric heme protein myoglobin,which they          short time by synthesis of ATP from creatine
             use as an O 2 reserve. Myoglobin has a much      phosphate and ADP. In a nonenzymatic reac-
             higher af nity for O 2 than hemoglobin and       tion [6], small amounts of creatine and crea-
             therefore only releases its O 2 when there is    tine phosphate cyclize constantly to form cre-
             a   severe  drop   in   O 2  partial  pressure   atinine, which can no longer be phosphory-
             (cf. p. 282).                                    lated and is therefore excreted with the urine
                At a high level of muscular effort—e. g.,     (see p. 324).
             during weightlifting or in very fast contrac-       Creatine does not derive from the muscles
             tionssuch asthose carried out by the eye         themselves, but is synthesized in two steps in
             muscles—the O 2 supply from the blood            the kidneys and liver (left part of the illustra-
             quickly becomes inadequate to maintain the       tion). Initially, the guanidino group of argi-
             aerobic metabolism. White fibers (left part of   nine is transferred to glycine in the kidneys,
             the illustration) therefore mainly obtain ATP    yielding guanidino acetate [3]. In the liver,
             from anaerobic glycolysis. They have supplies    N-methylation of guanidino acetate leads to
             of glycogen from which they can quickly re-      the formation of creatine from this [4]. The
             lease glucose-1-phosphate when needed (see       coenzyme in this reaction is S-adenosyl methi-
             p. 156). By isomerization, this gives rise to    onine (SAM; see p.110).
             glucose-6-phosphate, the substrate for glycol-
                                +
             ysis. The NADH+H formed during glycolysis
                                             +
             has to be reoxidized into NAD in order to

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