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SMCH 30
SMCH 30
30
Exercise Physiology
Alon Harris, Ph.D.
Bruce Martin, Ph.D.
CHAPTER OUTLINE
RESPONSES
AGING, IMMUNE, AND PSYCHIATRIC RESPONSES
KEY CONCEPTS
xercise, or physical activity, is a ubiquitous physiological state, so common in its many forms that true physiological rest is indeed rarely achieved. Defined ultimately
in terms of skeletal muscle contraction, exercise involves
every organ system in coordinated response to increased
muscular energy demands.
551
552
Steady state
O2 uptake (L/min)
1.25
Repayment
of O2 debt
1.00
0.75
0.50
Resting level
0.25
Rest
Work
2
FIGURE 30.1
Recovery
6
8
Time (min)
10
12
TABLE 30.1
CARDIOVASCULAR RESPONSES
Local control of blood flow ensures that only working muscles with increased metabolic demands receive increased
blood and oxygen delivery. If the legs alone are active, leg
muscle blood flow should increase while arm muscle blood
flow remains unchanged or is reduced. At rest, skeletal muscle receives only a small fraction of the cardiac output. In
dynamic exercise, both total cardiac output and relative
and absolute output directed to working skeletal muscle increase dramatically (Table 30.2).
Cardiovascular control during exercise involves systemic regulation (cardiovascular centers in the brain, with
their autonomic nervous output to the heart and systemic
resistance vessels) in tandem with local control. For millennia our ancestors successfully used exercise both to escape
being eaten and to catch food; therefore, it is no surprise
that cardiovascular control in exercise is complex and
unique. Its as if a brain software program entitled Exercise
Activity
Sleeping
Sitting
Standing
Dressing, undressing
Walking (3 miles/hr)
Making a bed
Dancing
Gardening/shoveling
Climbing stairs
Crawl swimming (50 m/min)
Running (8 miles/hr)
Sedentary 22-Year-Old
Sedentary 70-Year-Old
1
2
3
3
4
5
7
8
11
16
16
6
12
19
19
25
31
44
50
69
100
100
8
17
25
25
33
42
58
67
92
CHAPTER 30
Area
Splanchnic
Renal
Brain
Coronary
Skeletal muscle
Skin
Other
Total Cardiac Output
Heavy Exercise
mL/min
mL/min
1,400
1,100
750
250
1,200
500
600
5,800
24
19
13
4
21
9
10
100
300
900
750
1,000
22,000
600
100
25,650
1
4
3
4
86
2
0.5
100
553
3
Time to exhaustion (hr)
TABLE 30.2
Exercise Physiology
1
TABLE 30.3
Rest
Strenuous Dynamic
Exercise
6
90
15
21
105
5
0
25
50
75
100
Relative aerobic exercise intensity (% maximal oxygen uptake)
554
160
Isometric
140
120
Dynamic
100
80
Rest
1 hand
1 arm
2 arms
Muscle mass
1 leg
2 legs
Effect of active muscle mass on mean arterial pressure during exercise. The highest
pressures during dynamic exercise occur when an intermediate
muscle mass is involved; pressure continues to rise in isometric
exercise as more muscle is added.
FIGURE 30.3
TABLE 30.4
Exercise Intensity
Rest
Walking
Jogging
Running fast
respiratory pump, which increases breath-by-breath oscillations in intrathoracic pressure (see Chapter 18). The
importance of these factors is clear in patients with heart
transplants who lack extrinsic cardiac innervation. Stroke
volume rises in cardiac transplant patients with increasing
exercise intensity as a result of increased venous return that
enhances cardiac preload. In addition, circulating epinephrine and norepinephrine from the adrenal medulla and norepinephrine from sympathetic nerve spillover augment
heart rate and contractility.
Maximal dynamic exercise yields a maximal heart rate:
further vagal blockade (e.g., via pharmacological means)
cannot elevate heart rate further. Stroke volume, in contrast, reaches a plateau in moderate work and is unchanged
as exercise reaches its maximum intensity (see Table 30.4).
This plateau occurs in the face of ever-shortening filling
time, testimony to the increasing effectiveness of the mechanisms that enhance venous return and those that promote
cardiac contractility. Sympathetic stimulation decreases
left ventricular volume and pressure at the onset of cardiac
relaxation (as a result of increased ejection fraction), leading to more rapid ventricular filling early in diastole. This
helps maintain stroke volume as diastole shortens. Even in
untrained individuals, the ejection fraction (stroke volume
as a percentage of end-diastolic volume) reaches 80% in
strenuous exercise.
The increased blood pressure, heart rate, stroke volume,
and cardiac contractility seen in exercise all increase myocardial oxygen demands. These demands are met by a linear increase in coronary blood flow during exercise that can
reach a value 5 times the basal level. This increase in flow
is driven by local, metabolically linked factors (nitric oxide,
adenosine, and the activation of ATP-sensitive K channels) acting on coronary resistance vessels in defiance of
sympathetic vasoconstrictor tone. Coronary oxygen extraction, high at rest, increases further with exercise (up to
80% of delivered oxygen). In healthy people, there is no
evidence of myocardial ischemia under any exercise condition, and there may be a coronary vasodilator reserve in
even the most intense exercise (Clinical Focus Box 30.1).
Over longer periods of time, the heart adapts to exercise
overload much as it does to high-demand pathological
states: by increasing left ventricular volume when exercise
requires high blood flow, and by left ventricular hypertrophy when exercise creates high systemic arterial pressure
(high afterload). Consequently, the hearts of individuals
adapted to prolonged, rhythmic exercise that involves relatively low arterial pressure exhibit large left ventricular
Oxygen Uptake
(L/min)
Heart Rate
(beats/min)
Stroke Volume
(mL/beat)
Cardiac Output
(L/min)
0.25
1.0
1.8
2.5
72
110
150
190
70
90
100
100
5
10
15
19
CHAPTER 30
Exercise Physiology
555
Stress Testing
To detect coronary artery disease, physicians often record
an electrocardiogram (ECG), but at rest, many disease sufferers have a normal ECG. To increase demands on the
heart and coronary circulation, an ECG is performed while
the patient walks on a treadmill or rides a stationary bicycle. It is sometimes called a stress test.
Exercise increases the heart rate and the systemic arterial blood pressure. These changes increase cardiac work
and the demand for coronary blood flow. In many patients,
coronary blood flow is adequate at rest, but because of
coronary arterial blockage, cannot rise sufficiently to meet
the increased demands of exercise. During a stress test,
specific ECG changes can indicate that cardiac muscle is
not receiving sufficient blood flow and oxygen delivery.
As heart rate increases during exercise, the distance between any portion of the ECG (for example, the R wave) on
the ECG becomes shorter (Fig. 30.A and 30.B). In patients
suffering from ischemic heart disease, however, other
changes occur. Most common is an abnormal depression
between the S and T waves, known as ST segment depression (see Fig. 30.B). Depression of the ST segment
arises from changes in cardiac muscle electrical activity
secondary to lack of blood flow and oxygen delivery.
During the stress test, the ECG is continuously analyzed
for changes while blood pressure and arterial blood oxygen saturation are monitored. At the start of the test, the
exercise load is mild. The load is increased at regular intervals, and the test ends when the patient becomes exhausted, the heart rate safely reaches a maximum, significant pain occurs, or abnormal ECG changes are noted.
With proper supervision, the stress test is a safe method
for detecting coronary artery disease. Because the exercise load is gradually increased, the test can be stopped at
the first sign of problems.
R
T
R
T
S
2
R
T
Chronic, dynamic exercise is associated with increased circulating levels of high-density lipoproteins (HDLs) and reduced low-density lipoproteins (LDLs), such that the ratio
of HDL to total cholesterol is increased. These changes in
cholesterol fractions occur at any age if exercise is regular.
Weight loss and increased insulin sensitivity, which typically accompany increased chronic physical activity in
sedentary individuals, undoubtedly contribute to these
changes in plasma lipoproteins. Nonetheless, in people
with lipoprotein levels that place them at high risk for coronary heart disease, exercise appears to be an essential adjunct to dietary restriction and weight loss for lowering
556
LDL cholesterol levels. Because exercise acutely and chronically enhances fat metabolism and cellular metabolic capacities for -oxidation of free fatty acids, it is not surprising that regular activity increases both muscle and adipose
tissue lipoprotein lipase activity. Changes in lipoprotein lipase activity, in concert with increased lecithin-cholesterol
acyltransferase activity and apo A-I synthesis, enhance the
levels of circulating HDLs.
Exercise Has a Role in Preventing and Recovering
From Several Cardiovascular Diseases
RESPIRATORY RESPONSES
CHAPTER 30
TABLE 30.5
557
The effects of training on the pulmonary system are minimal. Lung diffusing capacity, lung mechanics, and even
lung volumes change little, if at all, with training. The widespread assumption that training improves vital capacity is
false; even exercise designed specifically to increase inspiratory muscle strength elevates vital capacity by only 3%.
The demands placed on respiratory muscles increase their
endurance, an adaptation that may reduce the sensation of
dyspnea in exercise. Nonetheless, the primary respiratory
changes with training are secondary to lower lactate production that reduces ventilatory demands at previously
heavy absolute work levels.
In Lung Disease, Respiratory Limitations May Be
Evidenced by Shortness of Breath or Decreased
Oxygen Content of Arterial Blood
Ventilation
Exercise Intensity
Rest
Walking
Jogging
Running fast
Exercise Physiology
Ventilation(L/min)
Alveolar PO2
Perfusion Ratio
Arterial PO2
(mm Hg)
Arterial PCO2
(mm Hg)
Arterial pH
(mm Hg)
5
20
45
75
1
2
3
4
103
103
106
110
100
100
100
100
40
40
36
25
7.40
7.40
7.40
7.32
558
tentially dangerous systemic arterial hypoxia during exercise. The signs and symptoms of a respiratory limitation to
exercise include exercise cessation with low maximal heart
rate, oxygen desaturation of arterial blood, and severe
shortness of breath (Clinical Focus Box 30.2). The
prospects of training-based rehabilitation are modest, although locomotor muscle-based adaptations can reduce
lactate production and ventilatory demands in exercise.
Specific training of respiratory muscles to increase their
strength and endurance is of minimal benefit to patients
with compromised lung function.
Exercise causes bronchoconstriction in nearly every
asthmatic patient and is the sole provocative agent for
asthma in many people. In healthy individuals, catecholamine release from the adrenal medulla and sympathetic nerves dilates the airways during exercise. Sympathetic bronchodilation in people with asthma is
outweighed by constrictor influences, among them heat
loss from airways (cold, dry air is a potent bronchoconstrictor), release of inflammatory mediators, and increases
in airway tissue osmolality. Leukotriene-receptor antagonists block exercise-induced symptoms in most people. The
effects of exercise on airways are due to increased ventilation per se; the exercise is incidental. Individuals with exercise-induced bronchoconstriction are simply the most sensitive people along a continuum; for example, breathing
high volumes of cold, dry air provokes at least mild bronchospasm in everyone.
Events within exercising skeletal muscle are a primary factor in fatigue. These same events, when repeated during
training, lead to adaptations that increase exercise capacity
and retard fatigue during similar work. Skeletal muscle contraction also increases stresses placed on bone, leading to
specific bone adaptations.
CHAPTER 30
TABLE 30.6
Exercise Physiology
559
Effects of Training and Immobilization on the Human Biceps Brachii Muscle in a 22-Year-Old Woman
After
Strength Training
After 4 Months
Immobilization
Sedentary
After
Endurance Training
300,000
10
100
50
67
0.8
100
300,000
10
100
50
67
1.3
150
300,000
13
200
50
87
0.8
77
300,000
6
60
50
40
0.6
100
Modified from Gollnick PD, Saltin B. Skeletal muscle physiology. In: Teitz CC, ed. Scientific Foundations of Sports Medicine. Toronto: BC Decker,
1989;185242.
supported metabolically. In fact, the easy fatigability of atrophied muscle is due to the requirement that more motor
units be recruited for identical external force; the fatigability per unit cross-sectional area is normal. The magnitude
of the skeletal muscle endurance training response is limited by factors outside the muscle, since cross-innervation
or chronic stimulation of muscles in animals can produce
adaptations 5 times larger than those created by the most
intense and prolonged exercise.
Local adaptations of skeletal muscle to endurance activity reduce reliance on carbohydrate as a fuel and allow
more metabolism of fat, prolonging endurance and decreasing lactic acid accumulation. Decreased circulating
lactate, in turn, reduces the ventilatory demands of heavier
work. Because metabolites accumulate less rapidly inside
trained muscle, there is reduced chemosensory feedback to
the central nervous system at any absolute workload. This
reduces sympathetic outflow to the heart and blood vessels,
reducing cardiac oxygen demands at a fixed exercise level.
Muscle Hypertrophies in Response to
Eccentric Contractions
companied by an acute phase reaction that includes complement activation, increases in circulating cytokines, neutrophil mobilization, and increased monocyte cell adhesion
capacity. Training adaptation to the eccentric components
of exercise is efficient; soreness after a second episode is
minimal if it occurs within two weeks of a first episode.
Eccentric contraction-induced muscle damage and its
subsequent response may be the essential stimulus for muscle hypertrophy. While standard resistance exercise involves a mixture of contraction types, careful studies show
that when one limb works purely concentrically and the
other purely eccentrically at equivalent force, only the eccentric limb hypertrophies. The immediate changes in
actin and myosin production that lead to hypertrophy are
mediated at the posttranslational level; after a week of loading, mRNA for these proteins is altered. Although its precise role remains unclear, the activity of the 70-kDa S6 protein kinase is tightly linked with long-term changes in
muscle mass. The cellular mechanisms for hypertrophy include the induction of insulin-like growth factor I, and upregulation of several members of the fibroblast growth factor family.
Exercise Plays a Role in Calcium Homeostasis
560
Cyclic
runners
180
Controls
160
Amenorrheic
runners
140
GASTROINTESTINAL, METABOLIC,
AND ENDOCRINE RESPONSES
The effects of exercise on gastrointestinal (GI) function remain poorly understood. However, chronic physical activity plays a major role in the control of obesity and type 2
diabetes mellitus.
Obesity is common in sedentary societies. Obesity increases the risk for hypertension, heart disease, and diabetes and is characterized, at a descriptive level, as an excess of caloric intake over energy expenditure. Because
exercise enhances energy expenditure, increasing physical
activity is a mainstay of treatment for obesity.
The metabolic cost of exercise averages 100 kcal/mile
walked. For exceptionally active people, exercise expenditure can exceed 3,000 kcal/day added to the basal energy
expenditure, which for a 55-kg woman averages about
1,400 kcal/day. At high levels of activity, appetite and food
intake match caloric expenditure (Fig. 30.6). The biological factors that allow this precise balance have never been
defined. In obese people, modest increases in physical activity increase energy expenditure more than food intake,
so progressive weight loss can be instituted if exercise can
be regularized (see Fig. 30.6). This method of weight control is superior to dieting alone, since substantial caloric re-
CHAPTER 30
Lean
2,500
Obese (initially
stable weight)
2,000
1,500
1,500
2,000
2,500
Caloric expenditure (kcal/day)
3,000
Caloric intake as a function of exercise-induced increases in daily caloric expenditure. For lean individuals, intake matches expenditure over a wide
range. For obese individuals during periods of weight gain or periods of stable weight, increases in expenditure are not matched by
increases in caloric intake. (Modified from Pi-Sunyer FX. Exercise
effects on calorie intake. Annals NY Acad Sci 1987;499:94103.)
FIGURE 30.6
Sedentary
150
100
50
100 g glucose
ingestion
30
After repeated
daily exercise
60
Time (min)
90
120
100 g glucose
ingestion
175
Plasma insulin (U/mL)
561
idation (thereby sparing carbohydrate stores), and oral carbohydrate intake during exercise. Frank hypoglycemia rarely occurs in healthy people during even the most prolonged or intense physical activity. When it does, it is usually in
association with the depletion of muscle and hepatic stores
and a failure to supplement carbohydrate orally.
Exercise suppresses insulin secretion by increasing sympathetic tone at the pancreatic islets. Despite acutely
falling levels of circulating insulin, both non-insulin-dependent and insulin-dependent muscle glucose uptake increase during exercise. Exercise recruits glucose transporters from their intracellular storage sites to the plasma
membrane of active skeletal muscle cells. Because exercise
increases insulin sensitivity, patients with type 1 diabetes
(insulin-dependent) require less insulin when activity increases. However, this positive result can be treacherous
because exercise can accelerate hypoglycemia and increase
the risk of insulin coma in these individuals. Chronic exercise, through its reduction of insulin requirements, up-regulates insulin receptors. This effect appears to be due less to
training than simply to a repeated acute stimulus; the effect
is full-blown after 2 to 3 days of regular physical activity
and can be lost as quickly. Consequently, healthy active
people show strikingly greater insulin sensitivity than do
their sedentary counterparts (Fig. 30.7). In addition, up-
Obese (during
weight gain)
3,000
Exercise Physiology
Sedentary
140
105
70
After repeated
daily exercise
35
30
60
Time (min)
90
120
Repeated daily exercise and the blood glucose and insulin response to glucose ingestion. Both responses are blunted by repeated exercise, demonstrating increased insulin sensitivity.
FIGURE 30.7
562
Endurance-trained
4
3
2
Sedentary
1
0
10
20
30
40
Age (yr)
50
60
70
Maximal oxygen uptake, endurance training, and age. Endurance-trained subjects possess greater maximal oxygen uptake than sedentary subjects, regardless of age.
FIGURE 30.8
In healthy people, prolonged exercise increases subsequent deep sleep, defined as stages 3 and 4 of slow-wave
sleep (see Chapter 7). This effect is apparently mediated
entirely through the thermal effects of exercise, since
equivalent passive heating produces the same result.
Whether or not exercise can improve sleep in patients
with insomnia is not known.
Clinical depression is characterized by sleep and appetite dysfunction and profound changes in mood.
Whether acute or chronic exercise can help relieve depression remains unproven. The two most prominent biological
theories of depressionthe dysregulation of central
monoamine activity and dysfunction of the hypothalamicpituitary-adrenal axishave received almost no study with
regard to the impact of exercise.
Panic disorder patients, often characterized by agoraphobia, have reduced exercise capacity. Although sodium
lactate infusion does provoke panic in these patients, the
anxiety mediator appears to be hypernatremia, not lactate;
even strenuous exercise with substantial lactic acidosis will
not trigger panic attacks in these individuals.
CHAPTER 30
Exercise Physiology
563
REVIEW QUESTIONS
5.
6.
7.
8.
564
PART VIII
CHAPTER 30
2. If we assume that the patients total body water was 36 L
when he came for treatment, it can be shown that giving the
patient 3 L of water without salt (by mouth and/or as an intravenous infusion of glucose in water) would reduce serum
[Na] to 144 mmol/L. Such treatment would improve the patients condition considerably. How might the medical officer argue the case for giving 2 L of normal saline?
3. What other (and relatively unusual) condition could produce
the patients symptoms? Did the medical officer rule this
possibility out by appropriate means?
Answers to Case Study Questions for Chapter 29
1. The patients nausea is probably a result of constriction of
the splanchnic vascular beds, which is part of the homeostatic cardiovascular response that helps maintain cardiac
output and blood pressure when central blood volume is reduced. Central blood volume, in turn, was reduced by the
loss of body water and pooling of blood in the peripheral
vascular beds. This homeostatic response also includes
constriction of the renal vascular beds, which, in turn, contributes (along with the release of vasopressin and activation of the renin-angiotensin system) to scanty urine production.
2. Because the weather was cool back home, the patient probably was probably not acclimatized to heat and was not
conserving salt in his sweat. He was probably secreting
large amounts of sweat, and losing correspondingly large
amounts of salt because of the weather and the activity involved in the exercise. If the patient returns to training the
next morning without correcting the salt deficit, he is likely
to have further difficulties in the heat. Even if the medical
officer has guessed incorrectly about the patients salt balance, a patient with normal renal function and adequate
fluid intake should be able to excrete any excess salt resulting from the treatment.
3. Hyponatremia can produce symptoms similar to the patients symptoms. However, the medical officer was able to
exclude hyponatremia (although not necessarily some degree of salt deficit) on the basis of elevated serum [Na].
Giving a hyponatremic patient large volumes of fluid without an equivalent of salt (which would have been a reasonable alternative treatment for the patient in this example)
would worsen the hyponatremia, perhaps to a dangerous
degree.
Reference
Knochel JP. Clinical complications of body fluid and electrolyte balance. In: Buskirk ER, Puhl SM, eds. Body Fluid Balance: Exercise and Sport. Boca Raton, FL: CRC Press,
1996;297317.
Exercise Physiology
565