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328        Small Animal Clinical Nutrition



                                                                      periods of time. Fatty acids are stored in ample supply in adi-
                   Table 18-4. Percent fast twitch fibers (Type II) by selected   pose tissue and within muscle. They are the primary energy
        VetBooks.ir  canine breeds.*                                  source for long-lasting exercise. Although small amounts of
                   Muscle
                   Biceps femoris  Greyhound  Crossbred  Foxhound     fatty acids are stored in muscle, this source may contribute up
                                                          63.0
                                               67.2
                                   88.6
                   Semitendinosus  98.9        85.3       69.6        to 60% of the fatty acids oxidized during the first two to three
                   Triceps (long head)  94.2   77.2       64.9        hours of exercise (Weber et al, 1993).
                   Vastus lateralis  96.6      61.4       80.7          Amino acids are usually not a primary energy source for exer-
                   *Adapted from Guy PS, Snow DH. Skeletal muscle fibre compo-
                   sition in the dog and its relationship to athletic ability. Research  cise. Oxidation of amino acids may contribute up to 5 to 15%
                   in Veterinary Science 1981; 31: 244-248.           of the energy used during exercise, depending on the intensity
                                                                      and duration of the task (Young et al, 1962; Zackin, 1990;
                                                                      Hickson and Wolinsky, 1989). Most of this energy comes from
                                                                      the oxidation of the branched-chain amino acids leucine,
                   Table 18-5. Metabolism, use and storage sites of   isoleucine and valine (Miller and Massaro, 1989; Blomstrand et
                   metabolic fuels.
                                                                      al, 1988). Most amino acids are structural or functional compo-
                                                                      nents of proteins and the size of the labile amino acid pool is
                   Fuel     Metabolism  Use          Storage sites
                   ATP      Anaerobic  Primary fuel for   Muscle cells   very small, making amino acids a less significant fuel source for
                                       synthetic     (concentration is    exercise in most circumstances.
                                       processes, ion  low and highly
                                       pumps and     regulated)         The proportion of energy substrates and metabolic pathways
                                       muscle                         used during exercise depends on the intensity and duration of
                                       contraction                    the exercise. As exercise intensity increases, the power output
                   Cr-P     Anaerobic  Regenerate ATP  Muscle cells (low
                                                     concentration)   and the rate of energy metabolism must also increase. As exer-
                   Glucose  Anaerobic  Rapidly available  Muscle and liver    cise duration increases, total substrate availability and energy
                            (glycolysis)  energy source  glycogen     yield become more important (Box 18-3). High-power activi-
                            and aerobic              (1 to 2% of
                            (TCA cycle)               body weight)    ties (e.g., sprinting) rely heavily on anaerobic metabolism,
                   Fatty acids Aerobic   Long-lasting  Adipose tissue    whereas more prolonged activities require the higher energy
                            (β-oxidation  energy source  (2 to 20% of   yield provided by oxidation of glucose and fatty acids. As the
                            and TCA                  body weight)
                            cycle)                                    duration of exercise increases, oxidation of fatty acids becomes
                   Amino    Aerobic    Small         Structural       more important (Figure 18-3).
                   acids    (TCA cycle)  contribution  proteins
                                       to energy (may
                                       contribute up                  By-Products of Muscular Work
                                       to 5 to 15%)                   Heat is the primary by-product of muscle contraction; 75 to
                   Key: ATP = adenosine triphosphate, Cr-P = creatine phosphate,  80% of the energy used during muscular work is converted to
                   TCA = tricarboxylic acid.
                                                                      heat. A 10-fold increase in metabolism results in a 10-fold
                                                                      increase in heat production. Unless the animal is working in a
                                                                      very cold environment, this heat is a by-product that must be
                                                                      removed (even some sled dogs overheat). In dogs, the respira-
                                                                      tory tract is responsible for dissipating most of this heat.
                                                                      Normal body temperatures of dogs doing physical work are
                                                                      higher than their normal resting temperatures. During very
                                                                      intense exercise or exercise in hot environmental temperatures,
                                                                      heat production exceeds the ability of the respiratory tract to
                                                                      lose heat, increasing body temperature. The body temperature
                                                                      of racing greyhounds may increase more than 1°C (1.8°F) after
                                                                      a 30-second race (Rose and Bloomber, 1989). Pointing breeds
                                                                      can have normal working temperatures up to 41.1°C (106°F)
                                                                      (Gillette, 2002). Labrador retrievers can have normal working
                                                                      temperatures up to 41.6°C (107°F) (Matwichuk et al, 1999).
                  Figure 18-3. Relative contributions of the four energy-generating  Because evaporative heat loss is the primary way dogs dissipate
                  pathways, depending on exercise intensity and duration. Key: Cr-P  heat, ensuring adequate hydration is crucial for maintenance of
                  = creatine phosphate.                               normal body temperature.
                                                                        Metabolic acid is another by-product of energy metabolism
                  dation of glucose to CO and water, regenerates ATP less rap-  that must be eliminated during and after exercise. Aerobic
                                     2
                  idly (Box 18-3), but results in a much greater yield (36 ATP per  metabolism generates ATP by combustion of carbohydrates
                  molecule of glucose). Because total body glucose stores (glyco-  and fats to CO and water. Lactate is the endpoint of anaero-
                                                                                  2
                  gen) are relatively small (1 to 2% of body weight), even aerobic  bic metabolism. Either way, acid is produced that must be elim-
                  metabolism of glucose cannot sustain exercise for extended  inated in some way for exercise to continue. Muscle enzyme
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