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Department of Physiology and Pharmacology, Karolinska Institute, S-11486 Stockholm; National Defence Research Establishment, S-172 90 Sundbyberg; and National Institute for Working Life, S-171-84 Solna, Sweden
ABSTRACT
INTRODUCTION
METHODS
RESULTS
DISCUSSION
ACKNOWLEDGEMENTS
FOOTNOTES
REFERENCES
ABSTRACT 
Åstrand, Per-Olof, Ulf Bergh, and Åsa
Kilbom. A 33-yr follow-up of peak oxygen uptake and related
variables of former physical education students. J. Appl. Physiol. 82(6): 1844-1852, 1997.
In 1949, 27 female and 26 male physical education students were studied at a
mean age of 22 and 25 yr, respectively. They were restudied in 1970 and
1982. Measurements included oxygen uptake, heart rate, and pulmonary
ventilation during submaximal and maximal exercise on a cycle ergometer
and treadmill. After 21 yr, peak aerobic power was significantly
reduced, from 2.90 to 2.18 l/min and from 4.09 to 3.28 l/min for women
and men, respectively. After another 12 yr, the 1970 maxima were not
reduced further. From 1949 to 1982 there was a decrease in peak heart
rate from 196 to 177 beats/min in women and from 190 to 175 beats/min
in men (P < 0.05). Highest pulmonary
ventilation did not change significantly. At an oxygen uptake of 1.5 l/min, the heart rate was the same in 1949 as in 1982. In conclusion,
the physical fitness level of the subjects was well above average for
these ages. From 1970 to 1982 there was no decline in the average peak
aerobic power, a finding possibly related to increased habitual
physical activity.
submaximal and maximal exercise; heart rate; lung function; age; longitudinal study
INTRODUCTION MAXIMAL OXYGEN UPTAKE decreases at a rate of ~10%
per decade in sedentary people after the age of 25 yr (Ref. 14; for
further references, see Refs. 16, 26). On the basis of an
extensive literature review, Shvartz and Reibold (25) concluded that
between the ages of 50 and 75 yr the decline is ~15% per decade
(maximal oxygen uptake in l/min). For masters athletes a
decline of some 5% per decade in this maximum oxygen uptake has been
reported (23). Most of the data contribution to these reports are from cross-sectional studies. A common finding is that training over several
months can increase maximal oxygen uptake ~20% (however, with marked
individual differences in response; see Ref. 6, Ref. 8, chapt.
10). The adaptation of the oxygen transport system to
training is of the same relative magnitude in ages ranging from 20 to
70-80 yr, and the response is independent of gender (see Ref. 19).
A decline with age in maximal oxygen uptake and other parameters of
importance for physical fitness is an effect of apparently inevitable
changes in physiological parameters, e.g.,
1) reduced maximal heart rate (3,
22) with reduced maximal cardiac output as a consequence and
2) loss of motoneurons and thereby
of motor units and muscle mass (see Ref. 7). In addition, elderly
people usually have a more sedentary lifestyle than when they were
young. As pointed out by Jackson et al. (15, 16), the decline in maximal aerobic power over time is due not only to aging but also to
lifestyle variables such as exercise and body composition.
In 1949, 44 women and 42 men who were students in physical education
(age range from 20 to 33 yr) were subjects in a study that included
measurements of submaximal and maximal oxygen uptake and heart rate
during exercise on a cycle ergometer and/or treadmill. In
addition, static and dynamic lung volumes were measured (4). In 1970, 35 and 31 of these women and men were restudied (3). In 1982, 27 women
and 26 men, now 33 yr older and able to perform maximal exercise, were
reexamined. This reexamination offered a unique opportunity to study
the effect of interaction of habitual physical activity and aging on
maximal aerobic power.
Certainly, it is a somewhat unique group. However, one advantage is
that the former students' choice of physical education indicated that
they were very interested in personal engagement in physical
activity. The subjects' choice of physical education provides some guarantee against recall bias in subjects' estimates of
lifetime physical activity. To our knowledge, no longitudinal study
including both women and men followed for a similarly long period of
time, 33 yr, has so far been reported.
METHODS Experimental Design
Procedures
On day 1 the subjects exercised on a mechanically braked cycle ergometer (Monark) at three to four submaximal and at one maximal work rate. The time for each stage was 6-8 min, with a few minutes rest between periods of exercise. During the maximal work rate, the duration was, on average, 5.8 min (range 2.15-10.50 min) for the women and 6.0 min (3.80-8.90 min) for the men. The pedaling rate was 50 revolutions/min (rpm) for submaximal exercises and 50 or 60 rpm for maximum exercises, as decided by the subject. Within 1 min after exercise and also ~3 min after the maximal effort, blood samples for lactate assay were taken from the fingertip of a prewarmed hand. Electrocardiograms were recorded during supine rest (I, II, III, aVR, aVL, aVF, V1-V6) and during exercise (CH 1, 2, 4, 6).On day 2 the subjects walked on a horizontal motor-driven treadmill at 5 km/h. After additional two to three submaximal work rates on the cycle ergometer, depending on results of the day before and following a similar protocol, the subjects ran on the treadmill for the purpose of attaining the highest possible oxygen uptake. Because peak oxygen uptake when the subjects were cycling the day before was known, initial speed and slope were chosen that demanded ~90% of this peak value. This slope and/or speed was increased stepwise so that the subject was exhausted after ~6 min. Heart rate was measured with a pulse watch (Sporttester P-O 300, Polar Electro).
Measurements
Oxygen uptake was determined by using the Douglas bag technique, and the volume of expired air was measured in a balanced spirometer. The time for collection of expired air during maximal effort was ~0.8 min, and in some cases it was only 0.5 min. Gas samples were analyzed by a mass spectrometer (Centronics MGA 200) calibrated with gas mixtures analyzed by the Haldane technique. Blood lactate concentration was analyzed by a slight modification of Barker-Summerson's method (28). For the chemical blood profile, CALAB Medical Laboratories, Stockholm, Sweden, was responsible (a company that provides services to hospitals in the Stockholm region). Static and dynamic spirometry were performed by using a helium-dilution method with a closed spirometer system and Bernstein spirometer, respectively.Individual regression lines were calculated for data on heart rate and oxygen uptake at submaximal work rates on the cycle ergometer. Then, heart rates corresponding to oxygen uptakes of 1.5 and 2.0 l/min for all subjects, and also of 2.5 l/min for men, were interpolated.
Room temperature during exercise was kept between 18 and 20°C.
Statistics
Analysis of variance was applied to test for differences between genders and age, except for the physical activity index, for which Fisher's exact probability test was used. Difference was considered significant at P values <0.05.RESULTS
Occupation and Health Status
At the first examination in 1982, 29 women and 27 men participated. Sixteen women and 16 men still worked as physical education teachers. One man had retired, and the others had changed profession, in most cases to a sedentary job, mainly administrative. Four women and six men were cigarette smokers. All women except two were past menopause. Because of CHD, two women and one man of the examined individuals were not doing maximal exercise, and they are, therefore, not included in the following results.In general, the health status of the subjects was reasonably good (Table 1). The most common disorders were from the musculoskeletal system. In comparison with a random selection of a Stockholm population above age 45 yr, no dominance in the prevalence of subjective musculoskeletal symptoms from the neck, shoulders, or low back was found (12). No comparison can be made for leg problems because of differences in diagnostic criteria applied in the present study and that of Hagberg and Hogstedt (12). Those male subjects who still worked as physical education teachers had a tendency to report a higher prevalence of problems from the legs (Fisher's exact test, P = 0.11) and from the musculoskeletal system in general (Fisher's exact test, P = 0.10) than did subjects not active as teachers. No such trend was observed among the women or from other specific musculoskeletal locations among men.
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Body height did not change for either men or women, and for the women the mean body mass was the same in 1982 as 1949 and the range was also the same (21.3 and 20.3 kg, respectively). In 1982 the men were on average 3.8 kg heavier than 33 yr earlier (P < 0.05; range 25.3 and 20.6 kg, respectively; Table 2).
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Maximal Exercise
In 1982, 16 women and 18 men performed maximal exercise on both a treadmill and cycle ergometer. For women, the average peak oxygen uptake was 2.25 l/min when running and 2.26 l/min when cycling. For men, the peak values were 3.22 and 3.20 l/min, respectively. The same men tested in 1949 reached 4.04 and 3.98 l/min, respectively. For 13 of the women who did maximal exercise in both types of exercise, there was no significant difference in peak oxygen uptake either in 1949 or in 1982 when running and cycling were compared.Peak blood lactate concentrations are presented in Table 2. There was a slight but significant (P < 0.05) reduction from 1949 to 1982, from 11.7 to 10.1 mmol/l for women and from 12.0 to 9.9 mmol/l for men. There were no significant gender differences for any of the mean values.
Table 2 and Fig. 1, A and B, present data on peak oxygen uptake (cycling or running). In contrast to a comparison between the 1949 and 1970 data on oxygen uptake (l/min), there was no significant change over the next 12-yr period. Because the average body weight was approximately the same in 1982 as in 1970, there was no significant difference in maximal oxygen uptake in milliliters per kilogram per minute during this period of time. On average, the reduction in this peak value from 1949 to 1982 was 23%, with no gender difference. When expressed per kilogram of body mass, the decline was 22% for women and 27% for men.
and
, Data obtained in 1949; 
and 
, data obtained in 1970; 
and
, data obtained in 1982. B: same
data and data on peak heart rate related to line-of-identity diagrams
comparing recordings for 1949-1970, 1970-1982, and
1949-1982. , Women; , men.
The drop in peak heart rate from 1949 to 1982 was on average 19.0 beats/min for women and 15.6 beats/min for men. This decline was
significant (P < 0.05; Table 2,
Figs. 1B and
2). In this analysis three men were
excluded because they were on medication with 
-blockers.
Data on peak pulmonary ventilation, respiratory frequency, and tidal volume are presented in Table 3, and data for peak pulmonary ventilation are also presented in Fig. 3. There were no significant differences in the mean values within each function, but there was a trend for a reduction in peak pulmonary ventilation, from 92.5 to 88.5 l/min in women and from 120.9 to 114.4 l/min in men. It should be noted that the SD is large. The fraction of vital capacity utilized in peak tidal volume increased significantly, from 48 to 55% in women and from 53 to 62% in men (P < 0.05).
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Submaximal exercise. The oxygen uptake at given submaximal work rates (100 and 150 W for women and 150 W for men) was the same in 1982 as in 1949 (Table 4). In the women the heart rate at oxygen uptakes of 1.5 and 2.0 l/min was significantly higher in 1970 than in 1949 and 1982, with no difference over the 33-yr interval (Table 4, Fig. 2). Data from the men showed no significant differences in heart rate at an oxygen uptake of 1.5 l/min, but at oxygen uptakes of 2.0 and 2.5 l/min heart rate was significantly higher in 1970 than 21 yr earlier. However, heart rate stayed on the same level in the 1982 examination. When the subjects exercised at 50% of peak oxygen uptake there was, as illustrated in Table 4 and Fig. 2, a significant reduction in heart rate from ~135 to 120-115 beats/min in women. For the men there was a significant drop from 126 beats/min in 1949 to 116 beats/min in 1970. However, the additional decline to 115 beats/min in 1982 was not significant.
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1 · min1
for women (n = 27) and 14.2 ± 1.4 ml · kg1 · min1
for men (n = 20).
Static and Dynamic Spirometry
From 1970 to 1982 there was a significant (P < 0.05) reduction in vital capacity of 4% in women and 11% in men (Table 3). Total lung capacity (TLC) was 7% higher in 1982 than in 1949 for women, but for men there was no difference, with the highest values being recorded in 1970. The increase in residual volume (RV) over the 33-yr period was 52% for women and 46% for men. This increase was noted already during the measurements in 1970 (Table 3). The RV/TLC ratio increased from 0.20 to 0.29 for women and from 0.21 to 0.30 for men. The forced expiratory volume in 1 s decreased from 80% in 1970 to 75-76% in 1982 (there are no data available from 1949).Levels of Habitual Physical Activity
Data from the subjects when they were 20-30 yr of age to present time are presented in Table 5. Subjects who were still active as physical education teachers had a higher level of physical activity at work than did nonteachers. However, they did not, on the average, have a higher maximal aerobic power than did those with a sedentary profession (45.4 and 39.8 ml · kg1 · min1 for male and
female teachers, respectively, vs. 45.4 and 41.5 ml · kg1 · min1
for nonteachers). In the women there was a weak but statistically significant relationship between physical activity index and
maximal aerobic power (r = 0.60)
but not in the men (r = 0.29).
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DISCUSSION
Did the peak oxygen uptake attained measure the subjects' real maximal
aerobic power? During exhausting cycling, the oxygen uptake for women
was on average 10.2% below the demand calculated from work rate and a
mechanical efficiency of 23%, the mean value calculated in 1949. For
men it was 5.2% below. For four subjects the measured maximum was
approximately equal to the predicted demand. All four had a peak
lactate concentration 
10.9 mmol/l. During maximal
running, the measured oxygen uptake was 9.0 and 12.3% for women and
men, respectively, which is below the demand calculated from
experimental data on the basis of measurements of oxygen uptake at a
range of speeds, slopes of the treadmill, and subject's body
mass. For the four subjects with the predicted demand
close to the measured oxygen uptake, blood lactate concentration for
one was 8 mmol/l and for the others was 10.3 mmol/l or higher. Throughout the studies the respiratory exchange ratio (RER) was on
average 1.07 or higher on all occasions (Table 2).
As mentioned in the introduction, the subjects in this study were a
somewhat select group because of their choice of education and
profession in physical education. It is a balancing act as to who, for
medical reasons, should be excluded in a study such as this one. It is
interesting to note that the average peak oxygen uptake for the three
subjects on -blockers was almost identical to the mean values for
all subjects observed in 1949, 1970, and 1982.
An evaluation of levels of habitual physical activity from questionnaires includes methodological problems, as illustrated by the following two examples. 1) How should 3 × 10 min of physical activity at a given power be compared with 1 × 30 min? 2) Vigorous training 4-6 wk just before the measurements of peak oxygen uptake can more than compensate for the effects of 12 yr of a sedentary lifestyle. Another problem is related to the effects of habitual physical activity on health and peak oxygen uptake. Therefore, Table 6 must be interpreted with some caution.
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In this study, subjects who performed maximal exercise on both the
treadmill and cycle ergometer did not differ in average values for peak
oxygen uptake between these forms of exercise. Therefore, the
discussion will concentrate on peak values attained irrespective of the
type of exercise. It is surprising that in cross-sectional studies as
well as in longitudinal studies, and also in stress tests on cardiac
patients, there is an emphasis on peak/maximal oxygen uptake related to
body weight (in
ml · kg1 · min1
or metabolic equivalents). Actually, the coefficient of
correlation between cardiac performance, i.e., cardiac output and
maximal oxygen uptake expressed in liters per minute, is high
(r = 0.90 in 54 subjects
tested in our department by application of the dye-dilution technique
with Cardio-Green for measurements of the cardiac output on the basis
of data from Refs. 5, 10, 27). None of those subjects was
overweight. Obviously, adding 5, 10, or, in some of these subjects, 15 kg of fatty tissue to the calculation will reduce the correlation
coefficient, e.g., 0.8 if cardiac output is related to peak oxygen
uptake per kilogram of body mass. The body mass-related
peak oxygen uptake may predict the person's aerobic fitness potential
to move her/his body but would give poor guidance in prediction of the
individual's cardiac performance. On average, the subjects
participating in this study maintained their gross body mass reasonably
well (Table 2), and, therefore, it is not critical whether peak oxygen
uptake is presented in liters per minute or milliliters per kilogram
per minute. However, we have presented the data both ways to emphasize
this point.
Of the original sample (1949), 35 women and 31 men were studied again 21 yr later (2). Without exception there was a decline in peak oxygen uptake, expressed in both liters per minute and milliliters per kilogram per minute (Fig. 1B). On average, the reduction was 22% for women and 20% for men (l/min). However, the data obtained after the next 12 yr showed marked individual changes, but on average there was no further reduction in peak oxygen uptake (Table 2, Fig. 1, A and B). According to questionnaires, particularly when the situation for women is focused on, there were fewer demands for child care and more time could be devoted to habitual physical activity.
As mentioned in the introduction, several studies indicate that there
is a rate of decrease in maximal aerobic power of ~10% per decade
when related to body weight. However, Jackson et al. (15) report data
from mainly cross-sectional studies including 25- to 70-yr-old healthy
men and determined that a reduction in the slope of maximal aerobic
power of 0.46 ml · kg1 · min1 · yr1
was consistent with the slope of 0.4-0.5
ml · kg1 · min1 · yr1
reported in the literature (9). In a later study, Jackson et al. (16) report data for women with an age of range 20-64 yr.
The age-related decline in peak aerobic power for that study was 0.54 ml · kg1 · min1 · yr1.
In a 25-yr longitudinal study with continuously physically active men,
Kasch et al. (17) found that the decline in maximal aerobic power was
0.25 ml · kg1 · min1 · yr1.
Data from the present study show a larger reduction in peak aerobic
power over the first 21 yr of on average 0.51 ml · kg1 · min1 · yr1
in women and still greater in men, 0.65 ml · kg1 · min1 · yr1.
However, during the next 12 yr, up to the average age of 55.0 yr in
women and 58.5 yr in men, there was no further significant reduction in
maximal oxygen uptake expressed as either liters per minute or
milliliters per kilogram per minute. There are individual variations in data on peak oxygen uptake when 1970 is compared with
1982 (see Fig. 1B). One extreme
example of individual "aging" is illustrated by measurements
comparing peak aerobic power attained in 1949 with that attained in
1982. One man had a maximum of 4.0 l/min on both occasions. Physically,
he was very active, running marathon races, cross-country skiing, and
so on. Another man with the same peak aerobic power in 1949 now
attained a peak power of only 2.0 l/min. The latter is an extreme
example of the effect of regular physical activity. During his later
years, his habitual physical activity was more or less zero because he
had joint problems with pain when physically active. However, in the
cycle ergometer test he made an all-out effort, and he reached a peak
blood lactate concentration of 9.4 mmol/l and RER of 1.05.
When data from this are compared with those reported by Jackson et al.
(15), peak oxygen uptake was much higher in this study (1982), with
peak oxygen uptake on average 3.13 l/min for the men as compared with
2.60 l/min for the 160 subjects (age range 55-70 yr) of the study
of Jackson et al. When related to body weight, the values were 43.2 and
33.2 ml · kg1 · min1,
respectively. The subjects of Jackson et al. (16) were employees at
National Aeronautics and Space Administration-Johnson Space Center in
Houston, TX. In the study of Jackson et al., for women (age range
50-64 yr) the peak oxygen uptake was on average 1.63 l/min or 24.4 ml · kg1 · min1,
as compared with 1.85 l/min and 28.4 ml · kg1 · min1,
respectively, in the present study (Table 6). Rogers et al. (23) have
presented data on 14 sedentary and 15 well-trained male master
endurance athletes during a follow-up period of 8 yr. The age of the
subjects at follow-up was on average 61-62 yr. They noted a
decline in peak aerobic power of 12% per decade down to 2.45 l/min or
30.6 ml · kg1 · min1
for the sedentary subjects and a reduction of 5.5% per decade with an
end result of 3.54 l/min or 51.8 ml · kg1 · min1
for the master athletes. In absolute values the men in the present study attained a peak aerobic power between these values, but when the
values are related to body weight they are much closer to those of the
master athletes.
From an extensive literature review of studies conducted in the United States, Canada, and seven European countries, Shvartz and Reibold (25) formulated aerobic fitness norms for women and men age 6-75 yr. They presented seven fitness categories graded from 1 = excellent to 7 = very poor. An evaluation of the average subject's aerobic fitness by application of these norms to the data in the present study gives the following result. In 1949 and 1970 both women and men were in category 1 or 2, depending on whether peak oxygen uptake was expressed in liters per minute or in milliliters per kilogram per minute. In 1982 they were definitely "excellent."
In a study by Åstrand (2), peak oxygen uptake in relatively fit subjects was 1.85 l/min for 16 women (mean age 55.6 yr) and 2.54 l/min for 66 men (mean age 53.3 yr), as compared with 2.23 and 3.13 l/min, respectively, in this study. In comparison with master endurance-trained athletes, the men in the present study had lower peak aerobic power.
In the study by Heath et al. (13), 16 athletes (59 ± 6 yr of age) attained a maximum that was on average 19% higher (l/min) and, when related to body weight, 36% higher than the values in the present study. Peak pulmonary ventilation was 15 l/min higher and peak heart rate 6 beats/min lower than in the present study. The training volume in the study of Heath et al. varied from 64 to 145 km/wk. A similar difference is noted when the present data are compared with results published by Hagberg et al. (11) for eight master endurance athletes (56 ± 5 yr of age). However, when peak maximal oxygen uptake is compared in absolute values or related to body weight observed in the present study, these data are almost identical with data presented by Kavanagh and Shephard (18) on female and male master athletes competing in the 1985 World Masters' Games in Toronto, Canada. The age of the athletes was 50-59 yr.
In 1949, the values on peak oxygen uptake for the women averaged 71% of the men's values, in 1970 66%, and in 1982 again 71% (l/min). When related to body weight the values are 82, 84, and 89%, respectively. It is interesting to note that in data on peak aerobic power related to body mass and measured in 1949, there was little overlap between data on women and men. This is in contrast to the situation in 1982 (Fig. 1A).
Peak oxygen pulse (see Table 2), reflecting stroke volume and
arteriovenous oxygen difference, was 14% lower in 1982 as compared with 1949 for women and 16% lower for men. The decline in peak aerobic
power was 23% for both groups. One interpretation of this finding is
that there was an ~15% reduction in stroke volume (because the
hemoglobin concentration was normal on both occasions). However, Rivera
et al. (20) reported that male master runners [66 ± 8 (SD)
yr] had a significantly smaller arteriovenous oxygen difference (11%) than did younger runners [32 ± 5 (SD) yr]. If
this finding applies to the present study, then a 9.6% reduction in
peak heart rate in women and 8.2% decrease in men could explain the
downward slope in peak aerobic power. However, then it
remains to be explained why the heart rate at a submaximal work rate
demanding an oxygen uptake of 1.5 l/min was not different when the data
obtained in 1949 are compared with those obtained in 1982. For the
women the result was the same at an oxygen uptake of 2.0 l/min, but in
the men heart rate was significantly higher at this oxygen uptake as
well as when the demand was 2.5 l/min. Possibly, these findings may
indicate that stroke volume may remain at the same level with aging for
submaximal exercise and may only decline during heavier exertion.
Rodeheffer et al. (22) studied 61 subjects with an age range from 25 to
79 yr. They concluded that during vigorous exercise the cardiac output
was maintained because of age-related increases in end-diastolic volume
and stroke volume, which compensate for an age-related decrease in
heart rate. The decrease in heart rate they explained by a hypothesis
that the effectiveness of -adrenergic modulation of myocardial
contractility, heart rate, and vascular tone declines with advancing
adult age. Unfortunately, peak oxygen uptake was not measured. They
also noticed a reduction in maximal work rate achieved with age. It is
surprising that in the 25-yr-old group the maximal work rate was on
average 150 W, with an oxygen demand of 2.1 l/min, which is a very low
aerobic power at this age.
Saltin (24) reports an oxygen pulse of 0.17 ml · kg1 · min1 · beat1
in sedentary men but 0.34 ml · kg1 · min1 · beat1 in elite athletes of
the same age. Similar values are reported by Hagberg et
al. (11). In the present study the mean value is 0.25 ml · kg1 · min1 · beat1
for men (0.22 ml · kg1 · min1 · beat1
for women).
Pulmonary function was well maintained. Peak pulmonary ventilation during maximal exercise did not decline significantly (Fig. 3), nor did tidal volume. However, there was a significant reduction in vital capacity of 5% in women and 12% in men. The percentage of vital capacity utilized as tidal volume increased from 48 to 55% in women and from 53 to 62% in men. In the women there was a significant 6.5% increase in TLC, but no change was recorded in the men. In a follow-up study of close to 40 yr that included 6 women and 19 men, all former physical education students, Asmussen et al. (1) did not observe a change in TLC. In both our study and that of Asmussen et al., RV was close to 20% of TLC when the subjects were in their 20s. In the present follow-up study, RV reached 29% of TLC in women and 30% in men. The first data of Asmussen et al. were at a similar level, but 40 yr later the women's RV took 36% of TLC and the men's RV took 33%.
In conclusion, the subjects in this study had a physical fitness level, if related to peak oxygen uptake, that was well above data reported for nonathletes of the same ages. It was interesting to note that over a 12-yr period, up to an average age of 55.0 yr for women and 58.5 yr for men, there was no significant reduction in mean peak oxygen uptake, but large individual variations were observed.
ACKNOWLEDGEMENTS
The authors are grateful to C. Käll for technical assistance and U. Siltberg for help with the organization of the study.
FOOTNOTES
Address for reprint requests: P.-O. Åstrand, Dept. of Physiology and Pharmacology, Karolinska Institute, Box 5626, S-114 86 Stockholm, Sweden.
Received 30 September 1996; accepted in final form 6 February 1997.
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