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Feeding Working and Sporting Dogs   331


                  Physiologic Changes Due to Exercise                 because of hydrostatic and osmotic forces. Increases in blood
        VetBooks.ir  The hallmark of exercise is increased metabolism. Many organ  pressure during exercise cause a shift of fluid from the
                                                                      intravascular space to the interstitial compartment (Harrison
                  systems increase their activity, some by several-fold, whereas
                                                                      et al, 1975). Muscle activity tends to increase intracellular
                  some systems decrease their function. The systemic changes
                  that occur during exercise seem to be driven by the muscles’  osmotic pressure, encouraging fluid movement from the
                  need for substrates and removal of metabolic waste. Working  interstitial to the intracellular spaces (Pivarnik, 1994). The
                  muscle metabolizes substrates (mostly fatty acids and glucose)  kidneys conserve plasma volume losses during exercise.
                  to release energy stored in chemical bonds for contraction.The  Decreases in plasma volume that decrease central venous
                  products of muscle metabolism are contraction, heat, CO ,  pressure cause renal vasoconstriction and diminish glomerular
                                                                 2
                           +
                  NH /NH , water, and in some cases, lactate.         filtration rate (GFR) (Pivarnik, 1994; Houpt, 1984). De-
                          4
                     3
                    Muscle metabolism can increase more than 20-fold in dogs,  creasing GFR will normally decrease urine output and thus
                  depending on the intensity of the exercise. Likewise, cardiac  diminish plasma volume losses. Increases in plasma osmotic
                  output increases proportionally with the workload. Both  concentration that occur during prolonged exercise also stim-
                  stroke volume and heart rate increase. Blood is the transport  ulate secretion of antidiuretic hormone (ADH), which con-
                  medium that carries oxygen and other substrates to the work-  serves plasma volume by stimulating production of a more
                  ing muscle and removes by-products such as heat, CO and  concentrated urine (Pivarnik, 1994; Houpt, 1984).
                                                              2
                  lactate. Increased function of the respiratory system (both  Exercise affects plasma volume and composition. Loss of
                  increased rate and depth) supplies more oxygen and disposes  fluid to the intracellular compartment increases the concen-
                  of more CO . Dogs and other mammals with contractile  trations of plasma proteins, electrolytes and all other solutes in
                            2
                  spleens can increase effective circulating blood volume and  the extravascular compartment. Other primary plasma
                  hematocrit by expelling red blood cells from the spleen before  changes that are needed to support increased muscle activity
                  or during exercise. For example, racing greyhounds increase  are a direct result of that activity. Glucose concentration may
                  blood volume as much as 24% before racing, even in the face  increase or decrease depending on the intensity and duration
                  of a 10% shift of plasma volume to other fluid compartments.  of exercise.The concentration of free fatty acids increases dur-
                  Probably as a result of both the plasma shift and splenic con-  ing prolonged exercise. At very high workloads, the partial
                  traction, hematocrit can increase as much as 29% (Toll et al,  pressure of oxygen may fall dramatically. Acidemia is also
                  1995) (Box 18-4). Plasma volume decreases during exercise  common with maximum intensity exercise because of anaero-

                     Box 18-4. Acute Physiologic Changes in Racing Greyhounds.


                     Because racing greyhounds exercise at a very high intensity, they exhibit very dramatic physical and biochemical changes during and
                     after racing. Packed cell volume (PCV) may increase to 68%, jugular venous pH values may decrease to 6.95 and plasma lactate con-
                     centrations may increase to 32 mEq/l (normal 1 to 2 mEq/l). Hyperventilation after a race can result in jugular venous pCO values as
                                                                                                            2
                     low as 14 torr (normal = 40 torr) and rectal temperatures can increase more than 1°C (1.8°F) during a 30-second race. After the race,
                     plasma sodium concentrations may reach 171 mEq/l (an 18 mEq/l increase from rest) and potassium concentrations may increase to
                     7.8 mEq/l (normal = 4 mEq/l). Plasma protein concentrations also increase after racing, implying a fluid shift out of the plasma com-
                     partment.
                      A study examined the effect of excitement before racing on these variables and quantified the effects of fluid shift on plasma volume,
                     blood volume and PCV before and after racing. Arterial blood samples were obtained at rest, just before and five minutes after a 704-
                     m race to quantify changes in hematologic variables, plasma electrolyte and protein concentrations, osmolality and acid-base variables.
                     Changes in plasma volume were estimated from the change in plasma protein concentration. Immediately before the race, plasma vol-
                     ume decreased by 10% from rest and total circulating red blood cell (RBC) volume increased by 60% attributable to increased RBC num-
                     bers rather than size. Increases in blood volume (BV) by 24% and PCV by 29% also were detected before the race. Five minutes after
                     the race, plasma volume was 21% below the resting value and total circulating RBC volume had increased 73% above the resting value,
                     resulting in a 40% increase in PCV. Contraction of the spleen appeared responsible for the increased PCV and BV before the race and
                     maintenance of BV after the race.
                      Plasma chloride concentration was the same before and after the race, meaning it decreased by the same fraction (22%) as the plas-
                                                               +
                                                                                                           +
                                      -
                     ma volume, indicating Cl loss from the plasma. Plasma Na content decreased by a smaller fraction (13%), causing Na concentra-
                                                                                   +
                     tion to increase from 151 mEq/l at rest to 167 mEq/l after the race. Assuming that Na concentration was the same throughout the
                     extracellular fluid, water likely moved into the intracellular compartment. As a consequence of these changes, the inorganic strong ion
                     difference in plasma increased by about 16 mEq/l, which tended to minimize the acid-base disturbance induced by the 33 mEq/l
                     increase in lactate concentration.
                      These results indicate that the physiologic changes taking effect during strenuous sprint exercise in racing greyhounds enhance blood
                     volume and aid in acid-base homeostasis, both of which are adaptive for this type of exercise.
                     The Bibliography for Box 18-4 can be found at www.markmorris.org.
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