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1 Department of Kinesiology, Indiana University, Bloomington, Indiana 46405; 2 Research Laboratory for Molecular Endocrinology, Laval University, Ste-Foy, Quebec, Canada G1K 7P4; 3 School of Kinesiology and Leisure Studies, University of Minnesota, Minneapolis, Minnesota 55455; 4 Division of Biostatistics, Washington University School of Medicine, St. Louis, Missouri 63110; 5 Department of Health and Kinesiology, Texas A&M University, College Station, Texas 77843; and 6 Pennington Biomedical Research Center, Baton Rouge, Louisiana 70808
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ABSTRACT |
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Effects of age, sex, race, and initial
fitness on training responses of maximal O2 uptake
(
O2 max) are unclear. Data were
available on 435 whites and 198 blacks (287 men and 346 women), aged
17-65 yr, before and after standardized cycle ergometer training. Individual responses varied widely, but
O2 max increased significantly for all
groups. Responses by men and women and by blacks and whites of all ages
varied widely. There was no sex difference for change (
) in
O2 max
(ml · kg
1 · min
1); women
had lower initial values and greater relative (%) increases. Blacks
began with lower values but had similar responses. Older subjects had a
lower
but a similar percent change. Baseline
O2 max correlated nonsignificantly with

O2 max but significantly with
percent change. There were high, medium, and low responders in all age
groups, both sexes, both races, and all levels of initial fitness. Age,
sex, race, and initial fitness have little influence on
O2 max response to standardized training in a large heterogeneous sample of sedentary black and white
men and women.
trainability; maximal oxygen uptake
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INTRODUCTION |
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IT IS NOT CLEAR
WHETHER and how much age, sex, race, and initial fitness affect
trainability (i.e., improvement in cardiorespiratory endurance after
training), as measured by increases in maximal O2
uptake (
O2 max). A
review of the literature suggests that there is either no difference
(1, 11, 13) or a reduction (17, 18, 23) in
trainability with increasing age. A few studies (2, 16)
suggest that women are less trainable than men, but recent evidence
(1, 9) suggests little or no difference between the sexes
in trainability of
O2 max. There
appears to be little difference between blacks and whites in
O2 max before training
(5), but we failed to find any published studies directly
comparing their responses to the same training program. Relative to
initial fitness, results from earlier studies (6, 15)
showed that subjects with initially lower levels of
O2 max had larger increases in
O2 max after training than did those
with higher levels at baseline. Later studies on older black and white
men and women (9) and on a large sample of whites (3) failed to confirm these observations.
In most of the previously cited studies, the numbers of subjects who
may have completed training programs that differed in intensity,
duration, and/or frequency were small; this was true for individual
training sessions, as well as for the total program. In addition, most
of these studies focused on the contribution of only one or two of the
variables under consideration in the present report, i.e., age, sex,
race, and initial fitness, on the response of
O2 max to training. None have
looked at all four factors simultaneously. To investigate the
contribution of all four factors, it is necessary to have a large
number of sedentary black and white men and women of different ages
complete a prolonged exercise training program with the same frequency, duration, and relative intensity of exercise and then compare the
response of
O2 max; this was possible
using data from the HERITAGE Family Study.
The HERITAGE Family Study is a consortium of five laboratories that has
studied the role of the genotype in the cardiovascular and metabolic
responses to aerobic exercise training and the contribution of
inherited factors to the changes brought about by regular exercise on
aerobic fitness and on several risk factors for cardiovascular disease
and type 2 diabetes. The design of this study has been described in
detail elsewhere (4). Although it was primarily designed
to be a genetic study, the large number of subjects who performed the
same standardized training program allowed us to investigate a number
of nongenetic aspects. Thus the purpose of the present ancillary study
was to determine the contributions of sex, race, age, and initial
fitness on the response of
O2 max to a
standardized exercise training program.
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METHODS |
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Sample.
The HERITAGE subjects came from families that included the natural
mother and father (
65 yr of age) and their natural children (17-40 yr of age). The white families had at least three
offspring; the black families were smaller. Complete data were
available from 633 subjects studied at the four clinical centers
[Arizona State University (Indiana University since January 1996),
Laval University, the University of Minnesota, and the University of Texas at Austin]. The number of participants in subgroups based on
age, sex, and race is shown in Table 1.
Subjects were healthy, sedentary, and met a number of inclusion and
exclusion criteria (4). They also passed a medical
examination by a physician that included a 12-lead electrocardiogram
obtained at rest, during submaximal exercise, and during a maximal
exercise test. The study protocol was previously approved by the
committee to protect human subjects in research studies at each of the
four clinical centers. Written informed consent was obtained from each
subject.
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O2 max tests.
Two maximal exercise tests on separate days before and after the 20-wk
training program were performed using a cycle ergometer (model 800S,
SensorMedics, Yorba Linda, CA) connected to a metabolic measurement cart (model 2900, SensorMedics). An electrocardiogram machine was used to monitor heart rate (HR). During each exercise stage, gas exchange variables (O2 uptake, CO2
output, minute ventilation, and respiratory exchange ratio) were
recorded as a rolling average of three 20-s intervals. The criteria for
O2 max were as follows: respiratory
exchange ratio >1.1, plateau in O2 uptake (change of <100
ml/min in the last 3 consecutive 20-s averages), and an HR within 10 beats/min of the maximal level predicted by age. All subjects achieved
a
O2 max by at least one of these
criteria before and after training. The majority of the exercise tests
were conducted at the same time of day, with
48 h between tests.
O2 max, and 3 min at 80%
O2 max. In those cases where 50 W was >60%
O2 max, the PO associated with
60% was done first followed by that associated with 50 W. The
resistance was then increased to the highest PO attained in the first
maximal test. If subjects were able to pedal after 2 min, PO was
increased each 2 min thereafter until they reached volitional fatigue.
The reproducibility of the maximal tests was determined, and an
intraclass correlation coefficient of 0.97 and a coefficient of
variation of 5% were found. There were no differences in
reproducibility among the four clinical centers (19).
Exercise training program.
The training program was done on cycle ergometers (Universal
Aerobicycle, Cedar Rapids, IA) that were interfaced with a computer system (Universal Gym Mednet, Cedar Rapids, IA) to maintain a constant
programmed training HR by modifying PO. Participants exercised three
times per week for 20 wk, beginning at 30 min/session and progressing
to 50 min/session during the last 6 wk. Exercise intensity increased
from the HR associated with 55%
O2 max measured at baseline to that associated with 75%
O2 max during the last 8 wk of
training. All training sessions were supervised on-site. More detailed
descriptions can be found elsewhere (20).
Compliance to training.
From a total of 855 eligible participants, 742 were considered to have
complied with the study, because they had finished the training program
and had complete or nearly complete data on all tests before and after
training. These 742 participants finished
95% of the 60 required
training sessions (i.e.,
57 sessions). The 633 participants in the
present study were taken from these 742 because they had complete data
on all variables being studied. One hundred thirteen participants did
not complete the study for a variety of reasons: 6 had an injury or
illness, 7 became pregnant, 1 relocated, 22 voluntarily refused to
continue, 10 were dropped because 1 or more members of the family
dropped out or were dropped, rendering the rest of the family
ineligible by our criteria, 18 missed too many training sessions, 33 were not able to complete all the tests, and 16 did not complete the study for "other" reasons.
Body composition measurements. Body density was determined by underwater weighing (24). Reproducibility of the body density, fat mass, and pulmonary residual volume measurements was high, with intraclass correlation coefficients between 0.97 and 1.00. There were no differences in reproducibility among the four clinical centers (24).
O2 max and changes in
O2 max.
O2 max measured in ml/min is a good
indicator of the capacity of the cardiorespiratory system to transport
O2 and of the muscle system to utilize it. As such, it is a
good indicator of the power of the system. However, there can be a
problem with use of ml/min in subjects who vary markedly in body mass.
In the present study, individual values for body mass ranged from 40 to
138 kg, and values for
O2 max ranged
from 1,166 to 4,434 ml/min. A better comparison would be the relative
change (%) in
O2 max, which was the
method used in most of the studies reviewed in this report. To
partially compensate for the large variation in body mass,
O2 max can be expressed as
ml · kg
1 · min
1; this is
usually used as an index of aerobic fitness and is useful for comparing
different groups of subjects that vary in body mass. The main tissue in
the body that uses O2 during exercise is muscle, and an
estimate of muscle can be obtained from the fat-free mass (FFM).
Because men and women have differences in body mass, as well as in the
composition of that mass, another way to compensate for these
differences is to express
O2 max as
ml · kg FFM
1 · min
1. A
problem with both of these ways to express
O2 max is that changes in body mass or
FFM make it difficult to make comparisons within the same person; i.e.,
whereas training may influence the ability of the body to transport and
utilize O2 (numerator), there also may be changes in body
mass or FFM that affect the denominator. Although there are advantages
and disadvantages of each method,
O2 max will be expressed in all three ways for a more complete picture of the changes.
Statistics.
Descriptive statistics (means ± SD) of all dependent variables
were calculated for all subjects and by sex, race, age group, sex × race, and sex × race × age group. A repeated-measures
ANOVA was used to compare the changes in
O2 max with training. A one-way ANOVA
was used to examine differences in group means. A Scheffé post
hoc test was used to identify significant differences among the three
age groups. A 2 × 2 × 3 (sex × race × age
group) factorial ANOVA was used to identify significant interactions; no significant interactions were found. For the purposes of this study,
trainability was defined as the absolute (
) and relative (%) change
in
O2 max after the training program.
Because there were individual changes in body mass (22)
and in FFM, the means for
O2 max
(ml · kg
1 · min
1 and
ml · kg FFM
1 · min
1,
respectively) were adjusted by an analysis of covariance, with the
body mass or
FFM as the covariate. A correlational analysis was
used to assess relationships between baseline
O2 max (ml · kg
1 · min
1) and
changes in
O2 max
(ml · kg
1 · min
1) after
training. All family members were included in the analyses, which
brings up the issue of nonindependence of the observations. Although
nonindependence of members within families is generally considered to
inflate the effective sample size and thus increase type I error (more
false positives), extensive simulations have shown that ignoring the
within-family dependencies does not invalidate the results (M. A. Province, personal communication). The level of statistical
significance for all tests was set at P < 0.01.
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RESULTS |
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Descriptive characteristics.
Age, physical characteristics, and maximal HR of the 633 subjects with
complete data can be found in Table 2 for
the total group and for subgroups sorted by age, sex, and race. As
expected, women were shorter, weighed less, and had more body fat than
men (P < 0.01). Black women weighed more and had more
body fat than white women (P < 0.01). After training,
body mass was significantly reduced in men (0.4 kg) and in subjects
aged 30-49 yr (0.6 kg). There were no significant mean FFM changes
in any group.
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O2 max.
When expressed as ml/min,
ml · kg
1 · min
1, and
ml · kg FFM
1 · min
1,
O2 max was lower in black men and women
than in white men and women before and after training
(P < 0.01; Table 3). As
expected, women also had lower values than men before and after training (P < 0.01). In terms of
O2 max expressed as ml/min before and
after training, the youngest age group had higher (P < 0.01) values than the middle age group, which had higher values
(P < 0.01) than the oldest group. For
O2 max expressed as
ml · kg
1 · min
1 before
training, the youngest age group had higher values (P < 0.01) than the other two older age groups, but there was no difference between the two older groups. After training, the youngest group had higher values (P < 0.01) than the middle age
group, which had higher values (P < 0.01) than the
oldest group. Relative to
O2 max
expressed as ml · kg
FFM
1 · min
1 before and after
training, the youngest group had higher (P < 0.01)
values than the middle age group, which had higher values (P < 0.01) than the oldest group.
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O2 max
expressed as ml/min,
ml · kg
1 · min
1, and
ml · kg FFM
1 · min
1
increased significantly (P < 0.01) in the total group,
as well as in each subgroup classified by age, sex, or race. The mean 
O2 max was 389 ml/min for the total
group, ranging from means of 332 to 453 ml/min in the subgroups. The
mean 
O2 max was 5.4 ml · kg
1 · min
1 for the
total group, varying from means of 4.5 to 5.7 ml · kg
1 · min
1 in the
subgroups. The mean 
O2 max was 6.9 ml · kg FFM
1 · min
1 for the
total group, varying from means of 6.2 to 7.3 ml · kg FFM
1 · min
1 in the subgroups.
For the total group, the mean relative increase in
O2 max expressed in ml/min,
ml · kg
1 · min
1, and
ml · kg FFM
1 · min
1 was
17.4, 17.8, and 16.5%, respectively. These relative increases ranged
from 15.5 to 19.9% for
O2 max
expressed as ml/min, from 15.9% to 20.3% for
O2 max expressed as
ml · kg
1 · min
1, and from
14.4 to 18.8% for
O2 max expressed as
ml · kg FFM
1 · min
1 in the subgroups.
Effect of age.
There was a significant (P < 0.01) age group
difference in response to training. When expressed as ml/min,
ml · kg
1 · min
1, and
ml · kg FFM
1 · min
1,

O2 max was smaller for subjects aged
50-65 yr than for the two younger groups. When expressed in ml/min
and ml · kg
1 · min
1, there
was no significant difference in percent increase in
O2 max among the three age groups.
However, when expressed in ml · kg FFM
1 · min
1, the increase in
O2 max was larger in the 31- to 49-yr age group than in the youngest group but was not different from the
oldest group; there was no difference between the oldest and youngest
age groups. The correlation between age and change in
O2 max expressed in
ml · kg
1 · min
1 was low
(r =
0.15). Although this relationship was
statistically significant (P < 0.01), age was
associated with <1% of the rise in
O2 max.
Effect of sex.
In relative terms, women had a significantly greater mean percent rise
in
O2 max (P < 0.01)
than men when
O2 max was expressed as
ml/min, ml · kg
1 · min
1, or
ml · kg FFM
1 · min
1. When
the mean increase was expressed as

O2 max, there was no difference
between the sexes for men in terms of
ml · kg
1 · min
1 or
ml · kg FFM
1 · min
1, but
men had a larger increase (P < 0.01) when

O2 max was expressed as ml/min.
Effect of race.
There was no significant difference between blacks and whites in the
mean increase in
O2 max with training,
whether this was expressed in relative terms or when the mean

O2 max was expressed as ml/min or
ml · kg FFM
1 · min
1.
However, when the mean 
O2 max was
expressed as
ml · kg
1 · min
1, blacks had
a significantly lower rise (P < 0.01).
Effect of initial fitness.
The correlation (r = 0.08) between baseline
O2 max expressed as
ml · kg
1 · min
1 and

O2 max expressed as
ml · kg
1 · min
1 was not
statistically significant. There were high, medium, and low responders
over a wide range of baseline values (Fig.
1). When the baseline values were
correlated with relative (%) change in
O2 max, there was a significant
(P < 0.01) correlation coefficient of
0.37; i.e.,
similar
s were associated with greater relative changes for those
with lower initial values.
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DISCUSSION |
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Effect of age.
Aging has been characterized by an impaired ability to adapt to and
recover from physiological stressors (18). Training, on
the other hand, has been associated with an increased tolerance (adaptation) to the physiological stress of exercise. In a recent position stand, it was the conclusion of the American College of Sports
Medicine (1) that trainability of
O2 max is not reduced with age.
However, with the exception of the study by Kohrt et al.
(9) on 110 men and women aged 60-71 yr, the other
studies cited in the position stand had many fewer subjects. The
present study is the largest exercise training study to date (n = 633) and encompasses individuals from 17 to 65 yr
of age. Although the oldest group had a smaller

O2 max, they also started with lower
levels, and there was no difference in response in relative terms
(i.e., percent increase). When Kohrt et al. assigned their subjects to
three age groups (60-62, 63-66, and 67-71 yr), they also
found no significant differences among the groups in the relative
increase in
O2 max with training (21, 19, and 18%, respectively).
O2 max in their group of
110 older subjects and found a nonsignificant correlation coefficient
of
0.13. This is similar to the coefficient of
0.15 found with 633 subjects in the present study. Even though the coefficient in the
present study was significantly different from zero because of the
large number of subjects, it would appear to have little practical
significance, accounting for <1% of the variance in the training
response of
O2 max.
The training program used by Kohrt et al. (9) was similar
in intensity and frequency to the program in the present study but
lasted longer (9-12 mo vs. 20 wk). The average increase in
O2 max (ml/min) found by Kohrt et al.
was 24%, but there was a wide variation in the amount of change
(0-58%). This is similar to the average increase of 17% (range
5 to 56%) found in the present study. Because of the similarity of
training programs and the results from these two large studies, data
from the 110 subjects were obtained from the study of Kohrt et al., and
the relationship between age and change in
O2 max was computed. The correlation
coefficient for the combined 743 subjects ranging in age from 17 to 71 yr was
0.08; this was not significantly different from zero. Thus,
when different age groups are compared, it appears that age has little
or no significant effect on the response of
O2 max to training.
One comment made in the position stand by the American College of
Sports Medicine (1) was that older participants may need more time to adapt to endurance training. Because the subjects in the
present study were tested only at the beginning and end of the 20-wk
training program, it is not possible to test this hypothesis. However,
according to the present data, in subjects given
20 wk to adapt, the
trainability of
O2 max in older
subjects does not differ significantly from that in younger subjects.
Effect of sex.
A position stand of the American College of Sports Medicine
(1) concluded that, "Despite many biological
differences, there appear to be no gender differences in the magnitude
of improvement in
O2 max with endurance
training." No sex-related differences in the response of
O2 max to training were seen whether these studies involved young (11), middle-aged
(12), old (9), or very old (8)
men and women. The findings of the present study support these
conclusions. Before and after training, women in the present study had
a significantly lower
O2 max (ml/min);
this was related to the fact that they weighed ~15 kg less than the
men. Thus, when a significantly smaller

O2 max (23% lower than that seen in
men) was divided by an even greater difference in baseline
O2 max (40% lower than that seen in
men), the resulting relative increase was significantly greater in
women. Women in the present study had a similar

O2 max (ml · kg
1 · min
1 and
ml · kg FFM
1 · min
1) but
began with significantly lower values, and the resulting relative
increases were significantly higher. When the individual values
for 
O2 max
(ml · kg
1 · min
1) were
plotted for men and women aged 17-65 yr (Fig.
2), there was no apparent difference
between the sexes; i.e., there were high, medium, and low responders in
both sexes at all ages. Thus it appears that sex has no significant
effect on the response of
O2 max to
endurance training.
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Effect of race.
When comparative studies have been done with various races, there tends
to be little difference in
O2 max when
such factors as age, sex, body size, and level of habitual activity are
considered. For example,
O2 max
(ml · kg
1 · min
1) was not
different between black and white men in the rural South in the United
States (14), among male white subjects, Bantus, and
Bushmen in South Africa (25), between white and Bantu male middle-distance athletes (10), among men from various
Bantu tribes (26), or between male and female Tanzanian
African and European white subjects (7). When subjects
were matched for age, body mass index, and habitual activity, Ama
(unpublished observations cited in Ref. 5) found that male
African black subjects had a lower
O2 max than white French-Canadian subjects (41.6 vs. 46.0 ml · kg
1 · min
1), but they
also had a lower maximal HR, suggesting that the African men were not
at their maximum. Even when racial differences are found, there is a
considerable overlap in aerobic performance. As a result, Boulay et al.
(5) concluded that there is no valid or reliable evidence
of clear racial differences in
O2 max. We have not found any studies comparing the trainability of
O2 max for blacks and whites.
O2 max
(ml/min, ml · kg
1 · min
1,
and ml · kg FFM
1 · min
1)
than the sedentary white men and women; the reasons for these initial
differences are not clear. Although there was no difference in

O2 max (ml/min or ml · kg
FFM
1 · min
1) after training, blacks
had a significantly lower 
O2 max (ml · kg
1 · min
1). When
expressed as the percent increase in
O2 max, there were no significant
differences between races in terms of ml/min,
ml · kg
1 · min
1, or
ml · kg FFM
1 · min
1. This
was also apparent when individual

O2 max
(ml · kg
1 · min
1) values
were plotted for 17- to 65-yr-old blacks and whites (Fig. 3). As was seen with men and women, there
were high, medium, and low responders in both races at all ages. Thus
our data appear to confirm previous observations that race has no
significant effect on the response of
O2 max to endurance training.
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Effect of initial fitness.
Although a few studies (6, 15) have suggested that initial
fitness level can affect the response to training, other studies (3, 9, 21) have not reached the same conclusion. Thomas et
al. (21) found a nonsignificant correlation
(r = 0.20) between initial
O2 max and the change in
O2 max in older men after a 12-mo
training program of moderate intensity. Kohrt et al. (9)
found nonsignificant correlations in their older men (r = 0.04) and women (r =
0.23). They also found no
significant difference in improvement in
O2 max between those men whose initial
O2 max was in the lowest quartile
(20%) and those in the highest quartile (19%). For women, the
respective values for the lowest and highest quartiles of initial
O2 max was 19 and 14%
(P < 0.09). On the basis of data on 481 sedentary whites from the HERITAGE Family Study, Bouchard et al. (3) reported nonsignificant correlation coefficients ranging from 0.03 to
0.16 computed separately for fathers, mothers, sons, and daughters.
The present study, which includes data from blacks in the HERITAGE
Family Study, also found a nonsignificant correlation coefficient
(r = 0.08) between initial
O2 max
(ml · kg
1 · min
1) and

O2 max
(ml · kg
1 · min
1) for all
sedentary subjects. However, when the relative (%) changes in
O2 max were correlated with initial
levels, the relationship was significant (r =
0.38).
From the data in Fig. 1, it is apparent that there are high, medium,
and low responders to training at all levels of initial
O2 max. Thus it appears that initial fitness level has no significant effect on

O2 max after training but that
similar 
O2 max values represent a higher percentage of the lower initial values.
O2 max to a standardized 20-wk endurance exercise training program. There appear to be high, medium, and low responders to training at all ages,
in both sexes, in both races, and at all levels of initial fitness
studied. Genetics (13) may help explain the large
variation in training response in these subjects.
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ACKNOWLEDGEMENTS |
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The authors thank all the coprincipal investigators, investigators, coinvestigators, local project coordinators, research assistants, laboratory technicians, and secretaries who have contributed to this study. The HERITAGE consortium is very thankful to those hard-working families whose participation made these data possible.
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FOOTNOTES |
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The HERITAGE Family Study is supported by the National Heart, Lung, and Blood Institute through the following grants: HL-45670 (to C. Bouchard, Principal Investigator), HL-47323 (to A. S. Leon, Principal Investigator), HL-47317 (to D. C. Rao, Principal Investigator), HL-47327 (to J. S. Skinner, Principal Investigator), and HL-47321 (J. H. Wilmore, Principal Investigator). A. S. Leon is partially supported by the Henry L. Taylor Endowed Professorship in Exercise Science and Health Enhancement. C. Bouchard is partially supported by the George Bray Chair in Nutrition.
Address for reprint requests and other correspondence: J. S. Skinner, Dept. of Kinesiology, Indiana University, 11001 North 26th St., Phoenix, AZ 85028 (E-mail: jskinner{at}iupui.edu).
The costs of publication of this article were defrayed in part by the payment of page charges. The article must therefore be hereby marked "advertisement" in accordance with 18 U.S.C. Section 1734 solely to indicate this fact.
Received 5 July 2000; accepted in final form 1 December 2000.
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