Modeling growth and maturation changes in peak oxygen
uptake in 11–13 yr olds
NEIL ARMSTRONG, JOANNE R. WELSMAN, ALAN M. NEVILL, AND BRIAN J. KIRBY
Children’s Health and Exercise Research Centre,
University of Exeter, Exeter EX1 2LU, United Kingdom
Armstrong, Neil, Joanne R. Welsman, Alan M. Nevill,
and Brian J. Kirby. Modeling growth and maturation
changes in peak oxygen uptake in 11–13 yr olds. J. Appl.
Physiol. 87(6): 2230–2236, 1999.—The influence of gender,
growth, and maturation on peak O2 consumption (V̇O2 peak) in
11–13 yr olds were examined by using multilevel regression
modeling. Subjects were 119 boys and 115 girls, aged 11.2 6
0.4 (SD) yr at the onset of the study. Sexual maturation was
classified according to Tanner’s indexes of pubic hair. V̇O2 peak
was determined annually for 3 yr. The initial model identified
body mass and stature as significant explanatory variables,
with an additional positive effect for age and incremental
effects for stage of maturation. A significant gender difference
was apparent with lower values for girls, and an age-by-
gender interaction indicated a progressive divergence in boys’
and girls’ V̇O2 peak. Subsequent incorporation of the sum of two
skinfold thicknesses into the model negated stature effects,
reduced the gender term, and explained much of the observed
maturity effects. The body mass exponent almost doubled,
but the age-by-gender interaction term was consistent with
the initial model.
aerobic fitness; growth; maturation; multilevel modeling;
gender
AEROBIC FITNESS SERVES as a functional index of the
pulmonary, cardiovascular, and hematologic compo-
nents of oxygen delivery and the oxidative mechanisms
of the exercising muscles. Peak oxygen uptake (V̇O2 peak),
the highest oxygen uptake elicited during an exercise
test to exhaustion, is recognized as the best single
indicator of young people’s aerobic fitness (4). However,
V̇O2 peak is highly correlated with body size, and, for the
effects of chronological age, maturation, and gender on
V̇O2 peak to be elucidated, the confounding influence of
body size must be accounted for. Inappropriate analy-
ses have clouded our understanding of growth and
maturational changes in V̇O2 peak (4, 5).
Although always controversial (32), the expression of
V̇O2 peak in simple ratio to body mass to account for body
size differences has been, and to a large extent remains
(20), the method of choice for normalizing V̇O2 peak data.
However, the recent literature reflects a resurgence of
interest in issues concerning the validity of ratio scal-
ing and the discussion of alternative means for control-
ling for body size differences. There is accumulating
evidence to refute the validity of conventional ratio
methods derived from both convincing theoretical and
statistical arguments and empirical evidence that dem-
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2230 8750-7587/99 $5.00 Copyright r 1999 the American Physiological Society
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onstrates the failure of per body mass ratios to produce
a size-independent exercise variable (4, 5). Scaling
techniques based on allometric principles have been
shown to produce size-free performance measures when
applied to data derived from young people, have yielded
mass exponents less than the value of 1.0 assumed by
the simple ratio method, and in cross-sectional studies
have produced results that challenge conventional inter-
pretations of the growth and maturation of V̇O2 peak
(7, 34).
The application of allometric techniques to the inter-
pretation of longitudinal data is not straightforward
and may provide an incomplete interpretation of the
performance measure under investigation. Two recent
studies (15, 27) have employed an ontogenetic allome-
tric approach (19), in which individual body mass
exponents are calculated from each subject’s longitudi-
nal body mass-V̇O2 peak regression relationship. Indi-
vidual values may be averaged subsequently to de-
scribe, for example, gender- or maturity-specific group
exponents. This approach has been criticized on the
basis that the interpretation of within-individual and
between-group responses requires a statistically ineffi-
cient two-stage process (25). Moreover, as the longitu-
dinal analysis centers on describing the body
mass-V̇O2peak relationship, limited information is gained
regarding the nature or magnitude of the pattern of
change in aerobic fitness.
Two studies have applied one or both of the theoreti-
cally proposed mass exponents of 0.67 and 0.75 (28) to
examine the longitudinal tracking of and age-related
changes in V̇O2 peak (21, 27). Although individual cross-
sectional studies in large, homogeneous subject popula-
tions have derived mass exponents approximating these
values (1, 7), there is considerable evidence to show
that exponents are highly sample specific (5), often
deviating markedly from theoretical expectations, par-
ticularly in small samples. Although producing results
that may better reflect underlying changes, the applica-
tion of a theoretical exponent may not provide an
accurate representation of true longitudinal changes
within a given subject group.
Few longitudinal studies have considered the influ-
ence of covariates other than body mass despite indica-
tions from cross-sectional studies in both adults and
young people that factors such as stature and body
fatness may be significant independent predictors (20,
24). Neither has the separate influence of age vs.
maturity on the growth of V̇O2 peak been elucidated
comprehensively, in part because of the limitations of
traditional analytic techniques.
Multilevel regression modeling (18) is a statistical
technique that enables a flexible and sensitive interpre-
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 MODELING PEAK OXYGEN UPTAKE DURING GROWTH
tation of longitudinal data while avoiding many of the
pitfalls associated with the analyses described above.
In contrast to traditional analytic approaches, multi-
level modeling not only describes the underlying popu-
lation mean response but also recognizes and describes
variation around the group mean. Furthermore, both
the number of observations per individual and the
temporal spacing of the observations may vary within a
multilevel analysis as individual growth trajectories
are modeled.
In a reanalysis of the data from a longitudinal study
of V̇O2 peak in young elite athletes (12), Nevill et al. (25)
used an allometric approach within a multilevel regres-
sion analysis to demonstrate age and, in boys, matura-
tional, influences on V̇O2 peak that were over and above
those explained by the overall increase in body size.
Unfortunately, the participation of boys and girls in
different sports precluded a detailed examination of
differential growth between the genders. The present
study sought to extend our understanding of the devel-
opment of aerobic fitness by using a multilevel regres-
sion modeling approach to interpret chronological age-,
gender-, and maturity-associated changes in V̇O2 peak in
a healthy, untrained population of subjects tested for
three consecutive years, commencing at a mean age of
11.2 yr.
METHODS
Subjects. All of the children in year 6 (age 10–11 yr) of the
15 state schools in the city of Exeter, United Kingdom, were
invited to participate in a longitudinal study of physical
activity patterns, physiological responses to exercise, and
body composition. Some 745 children (70% of those eligible)
volunteered, and written consent was obtained from both the
children and their parents and/or guardians. The project
received ethical approval from the Exeter District Health
Authority Ethical Committee. In an attempt to detect sample
bias, the stature and body mass of the volunteers were
compared with the stature and body mass of those who
declined to participate. No significant difference (P . 0.05)
was detected in either gender. Twenty-five percent of the
eligible children in each school were randomly selected from
those who volunteered. Previous publications have described
the V̇O2 peak of boys and girls who were prepubertal in the first
year of the project (1) and analyzed cross-sectional changes in
V̇O2 peak by maturity group by using data from the second year
of the project (7). The present study involves a longitudinal
analysis of the development of V̇O2 peak during the first 3 yr of
the study. The subject sample thus comprises those young
people who satisfactorily completed the V̇O2 peak test and
associated anthropometric measures and for whom maturity
assessments were available. Subject numbers for year 1 are
n 5 119 boys, n 5 115 girls; year 2, n 5 94 boys, n 5 88 girls;
and year 3, n 5 93 boys, n 5 81 girls. There were no
significant differences (P . 0.05) between those who failed to
return for a subsequent test occasion and the rest of the group
on key measures including stature, body mass, skinfold
thicknesses, hemoglobin concentration, and V̇O2 peak.
Experimental methods. Age was computed from date of
birth and date of examination. Anthropometric apparatus
was calibrated according to the manufacturers’ instructions.
Stature was measured by using a Holtain stadiometer (Hol-
tain, Crymych, Dyfed, UK), body mass was determined by
using Avery beam balance scales (Avery, Birmingham, UK),
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2231
and skinfold thickness over the triceps and subscapular
regions was measured by using Holtain skinfold calipers
(Holtain) according to the techniques described by Weiner
and Lourie (33). Sexual maturity was visually assessed by
using Tanner’s indexes for pubic hair development (31).
Anthropometric and maturity measures were taken once on
each measurement occasion due to time and ethical restric-
tions. However, the same trained research team made the
anthropometric measures and the same trained research
nurse made the maturity assessments throughout the dura-
tion of the study. Blood hemoglobin concentration was deter-
mined from duplicate fingertip blood samples. Blood samples
were immediately assayed by using a HemoCue Photometer
(Clandon Scientific, Farnborough, UK), which was calibrated
against a control cuvette before each measurement.
The children had visited the laboratory on several occa-
sions and were habituated to both the general environment
and the experimental procedures. After a 3-min warm-up at
1.67 m/s (6 km/h), V̇O2peak was determined during an incremen-
tal treadmill running test on a motorized treadmill (Wood-
way, Cranlea Medical, Birmingham, UK). Belt speed was
increased to 1.94 m/s (7 km/h, year 1) or 2.22 m/s (8 km/h,
years 2 and 3) for the initial stage and then increased by 0.28
m/s (1 km/h) for each 3-min stage until a speed of 2.78 m/s (10
km/h) was reached. Subsequently, belt speed was held con-
stant, and the gradient was increased by 2.5% each stage for
further increments. A 1-min rest period separated the exer-
cise stages. The test was continued to voluntary exhaustion.
If the young person showed signs of intense exertion (hyper-
pnea, facial flushing, unsteady gait, sweating) and if his or
her heart rate had reached a value within 5% of age-predicted
maximum or the respiratory exchange ratio was at least 1.0,
the V̇O2 peak attained was accepted as a maximal index. All
subjects included in this paper satisfied these criteria on all
test occasions.
Throughout the test expired gases were monitored continu-
ously by using an Oxycon Sigma on-line gas-analysis system
(Cranlea Medical), which was calibrated before each test by
using gases of verified concentration. Heart rate was moni-
tored by using an electrocardiograph (Rigel, Morden, UK).
Statistical methods. Descriptive statistics (means and SDs)
for anthropometric variables, blood hemoglobin concentra-
tion, and V̇O2 peak were computed for subjects on the first test
occasion. Gender differences within each year of the study
were calculated by using analysis of variance.
Factors associated with the longitudinal development of
V̇O2 peak (gender and maturity), adjusted for differences in
anthropometric measures and age, were investigated by
using the multilevel modeling program MLwiN (17). Multi-
level modeling is an extension of multiple regression, which is
appropriate for analyzing hierarchically structured data. In
longitudinal data sets the hierarchy can be seen as the
repeated-measurement occasions (defined as level 1 units),
grouped within the individual subject (defined as the level 2
unit).
Multilevel modeling is preferable to traditional analytic
approaches (e.g., repeated-measures analysis of variance) for
longitudinal data as, in addition to describing the population
mean response, this method recognizes and describes varia-
tion around the mean at both levels. For example, at level 2,
individuals are allowed to have their own growth rates, which
vary randomly around the underlying population response
and, at level 1, each individual’s observed measurements may
vary around his or her own growth trajectory. Furthermore,
in contrast to traditional methods that require a complete
longitudinal data set, both the number of observations per
individual and the temporal spacing of the observations may
2232 MODELING PEAK OXYGEN UPTAKE DURING GROWTH

vary within a multilevel analysis as individual growth trajec-
tories can be modeled.
In this study a multiplicative, allometric approach was
adopted on the basis of the model proposed by Nevill and
associates (25) as follows
y 5 massk1 · staturek2 · exp (aj 1 bj · age 1 c · age2)eij
where all parameters were fixed, with the exception of the
constant (intercept term) and age parameters, which were
allowed to vary randomly at level 2 (between individuals), and
the multiplicative error ratio e (which varied randomly at
level 1). Subscripts i and j denote random variation at levels 1
and 2, respectively. The variable ‘‘age’’ was centered around
the group mean age of 12.0 yr.
This model can be linearized by logarithmic transforma-
tion and multilevel regression analysis on loge(y) used to
solve for the unknown parameters. Once transformed, the
equation above becomes
loge y 5 k1 · loge mass 1 k2 · loge stature 1 aj 1 bj · age
1 c · age2 1 loge(eij)
From this baseline model additional explanatory variables
were investigated, including the sum of triceps and subscapu-
lar thicknesses, blood hemoglobin concentration, gender, and
stage of maturity (stages 1–5 for pubic hair development).
The latter two variables were incorporated into the model as
indicator variables (e.g., for gender, boys 5 0, girls 5 1), which
set the boys’ constant as the baseline from which the girls’
parameter may deviate. Similarly, the baseline model reflects
the responses of a prepubertal child from which any addi-
tional effects of later maturity may be quantified. Interaction
terms ‘‘age by gender’’ and ‘‘maturity by gender’’ were also
constructed to investigate whether age and maturational
effects on V̇O2 peak differed for boys and girls. Age was also
allowed to vary randomly at level 1 to investigate within-
individual variation around the individual growth trajectory.
The need to allow each individual his or her own mass
exponent was examined by letting body mass vary at level 2.
As demonstrated for cross-sectional data (34), this multipli-
cative, allometric modeling approach has been shown to be
theoretically and statistically superior for longitudinal analy-
ses (25) to an additive, polynomial model (12) as the former
accommodates the skewness and heteroscedasticity that of-
ten characterize size-related exercise performance data (34).
RESULTS
The subjects’ physical characteristics in each year of
the study are presented in Table 1. There were no
significant (P . 0.05) gender differences in age, body
mass, stature, or hemoglobin concentration at any
point of measurement. Boys had significantly higher
(P , 0.001) V̇O2 peak than did girls on each measurement
occasion. The sum of skinfolds for boys was signifi-
cantly lower (P , 0.05) than that for girls on the first
two measurement occasions.
The results of the initial modeling of V̇O2 peak over the
three measurement occasions are presented in Table 2.
In this model, mass and stature proved to be significant
covariates with exponents of 0.48 6 (SE) 0.03 and
0.81 6 0.12, respectively. There was a significant
positive effect for age that was larger for boys than
girls, as indicated by the significant age-by-gender
interaction term that would be deducted from the age
term for the girls. A small but significant term for age
squared was also identified for both genders, although
girls’ V̇O2 peak was shown to be significantly lower than
that for boys’ as reflected by the negative term for
gender (20.15 6 0.01). In addition to these anthropo-
metric, age, and gender effects, there was an incremen-
tal effect of maturation that was consistent for boys and
girls. The maturity-by-gender effects investigated are
not included in Table 2 as they were nonsignificant
throughout all models.
These fixed estimates describe the population mean
response, whereas the random parameters describe
variation around this response at both level 1 (within
individuals) and at level 2 (between individuals); i.e.,
these parameters reflect the variance that remains
unaccounted for by the fixed part of the model. Within
model 1 there was significant random variation at level
2 for age, reflecting differential individual growth rates
in V̇O2 peak. Allowing each individual his or her own mass
exponent proved unnecessary as there was no random
variation between individuals around the fixed (mean)
parameter.
The independent effect of introducing the sum of two
skinfold thicknesses into the model was examined, with
the results summarized in Table 2, model 2. The
addition of skinfolds to the baseline model rendered the
stature and age squared terms nonsignificant and re-
duced the magnitude of the gender term, whereas the
mass exponent was almost doubled to a value of 0.86 6
0.03. Inclusion of skinfolds also explained much of the
observed maturity effects, with the parameter estimates
for maturity stages 3–5 almost halved. Furthermore,

Table 1. Physical characteristics and peak V̇O2 peak data for subjects in each year of the study
Boys, Year 1 Girls, Year 1 Boys, Year 2 Girls, Year 2 Boys, Year 3 Girls, Year 3
(n 5 119) (n 5 115) (n 5 94) (n 5 88) (n 5 93) (n 5 81)

Age, yr 11.2 6 0.4 11.2 6 0.4 12.1 6 0.4 12.2 6 0.4 13.1 6 0.04 13.1 6 0.04
Stature, m 1.46 6 0.06 1.46 6 0.08 1.51 6 0.08 1.51 6 0.08 1.57 6 0.09 1.57 6 0.08
Mass, kg 36.7 6 6.5 38.1 6 7.8 41.1 6 7.4 43.2 6 8.3 46.4 6 9.1 48.4 6 8.8
Skinfolds, mm 19.9 6 10.5 22.8 6 8.6* 19.7 6 8.8 22.6 6 8.8* 21.3 6 10.7 23.5 6 8.1
Hemoglobin concentration, g/l 13.5 6 0.6 13.6 6 0.6 13.5 6 0.7 13.4 6 0.7 13.7 6 0.07 13.6 6 0.07
V̇O2 peak , l/min 1.81 6 0.26 1.63 6 0.28† 2.11 6 0.34 1.89 6 0.29† 2.39 6 0.44 2.10 6 0.30†
Heart rate at V̇O2 peak , beats/min 200 6 7 202 6 7 201 6 8 203 6 7 200 6 7 202 6 7
RER at V̇O2 peak 1.06 6 0.07 1.05 6 0.06 1.04 6 0.05 1.05 6 0.05 1.03 6 0.06 1.05 6 0.05
Values are means 6 SD. n, No. of subjects; V̇O2 peak , peak O2 uptake; RER, respiratory exchange ratio. * P , 0.05, † P , 0.001 (boys vs. girls
within yr group).

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 MODELING PEAK OXYGEN UPTAKE DURING GROWTH 2233

Table 2. Multilevel regression analysis for V̇O2 peak ing V̇O2 peak has therefore been the focus of much debate
in exercise science (5), but few studies have considered
Parameter Model 1 SEE Model 2 SEE the likelihood that the value of the mass exponent is
Fixed dependent on not only sample size (4, 5) and homogene-
Constant 21.2526 6 0.0978 21.8735 6 0.0945 ity (20) but also the effect of other confounding covari-
Loge mass 0.4765 6 0.0320 0.8629 6 0.0317 ates. For example, with stature included as an addi-
Loge stature 0.8105 6 0.1172 NS
Loge skinfolds 20.1704 6 0.0134 tional covariate the value of the mass exponent has
Age 0.0428 6 0.0116 0.0450 6 0.0108 been reduced in several data sets (20, 34). Conversely,
Age2 20.0073 6 0.0035 NS inclusion of a measure of body fatness tends to raise the
Gender 20.1495 6 0.0094 20.1340 6 0.0084 value of the mass exponent (21). The present study
Age 3 gender 20.0177 6 0.0068 20.0177 6 0.0065
Maturity level 2 0.0382 6 0.0090 0.0301 6 0.0086
demonstrates these effects clearly. When, for illustra-
Maturity level 3 0.0548 6 0.0106 0.0372 6 0.0105 tive purposes, the data in the present study were
Maturity level 4 0.0902 6 0.0140 0.0571 6 0.0138 analyzed with mass as the sole body size variable, a
Maturity level 5 0.0892 6 0.0221 0.0435 6 0.0212 mass exponent of 0.67 6 (SE) 0.03 was obtained,
Random
Level 2
reflecting the values reported in cross-sectional investi-
Constant 0.0042 6 0.0005 0.0029 6 0.0004 gations of these children at prepuberty (1) and 12 yr of
Age 0.0008 6 0.0003 0.0006 6 0.0003 age (7). The value of the mass exponent decreased to
Covariance 0.0004 6 0.0003 0.0003 6 0.0002 0.48 6 0.03 with the addition of stature (model 1) and
Level 1 increased to 0.86 6 0.03 when a measure of body
Constant 0.0024 6 0.0003 0.0025 6 0.0003
Age 0.0004 6 0.0005 0.0004 6 0.0005 fatness was incorporated (model 2), illustrating the
Covariance 20.0006 6 0.0002 20.0007 6 0.0002 interdependence of covariates and supporting the view
22 3 log(likelihood) 21359.6270 21432.6730 that a lack of statistical control over known covariates
Values are means 6 SE of the estimate (SEE). NS, not significant. is a factor underlying the variability in reported mass
Age was centered around group mean of 12.0 yr. exponents (20). These findings also illustrate that it is
inappropriate to assume that scaling to either the
traditional per body mass ratio or to one of the theoreti-
the addition of this measure of body fatness explained a cal mass exponents (i.e., 0.67 or 0.75) will control
considerable proportion of the remaining level 2 (be- adequately for body size differences in children and
tween-individuals) variance associated with the con- adolescents.
stant in model 1; i.e., the estimate was reduced from Welsman et al. (34) used log-linear analysis of covari-
0.0042 6 0.0005 to 0.0029 6 0.0004. ance to partition out body size from V̇O2 peak in groups of
Blood hemoglobin concentration was introduced as preteen and teenage boys and girls and adult men and
an additional explanatory variable, but a nonsignifi- women. These data challenged the conventional inter-
cant parameter estimate was obtained. pretation of V̇O2 peak during growth by demonstrating
that V̇O2 peak increases progressively in boys from prepu-
DISCUSSION
berty through puberty into adulthood, whereas in girls
The V̇O2 peak data in liters per minute are in accor- increases are observed from prepuberty into puberty
dance with the extant literature (4). The present data, with no significant decline into adulthood (34). Subse-
however, offer further insights into the growth of quently, the same group examined the relationship
V̇O2 peak, with body size appropriately accounted for. The between maturation and V̇O2 peak, with body mass con-
conventional interpretation of V̇O2 peak ‘‘corrected’’ for trolled by using allometry (7). Twelve-year-old boys and
body mass (ml · kg21 · min21 ) is that, during the teen girls were classified into maturity stages 1–4 (31), and
years, boys’ values remain remarkably consistent, the results demonstrated significant increases in V̇O2peak
whereas girls’ values progressively decline (4). How- across maturity stages that were over and above the
ever, although it has been argued that there is no need changes attributable to increased body mass alone (7).
to abandon the use of the above-mentioned unit of These maturational effects had been masked in previ-
measure to normalize V̇O2 peak for body size (29), the ous cross-sectional studies by the use of the ratio
recent literature reflects a growing awareness of the standard (ml · kg21 · min21 ) to partition out body-size
theoretical and statistical limitations of this approach effects (10, 17). The present study has sought to further
(4, 20). clarify age, growth, and maturity effects on V̇O2 peak in
There is increasing empirical evidence to refute boys and girls studied longitudinally by using appropri-
the assumption that scaling V̇O2 peak to a mass exponent ate multilevel regression modeling techniques.
of 1.0 produces a size-free variable (1, 7, 21, 34), and Most longitudinal studies have analyzed age-,
theoretical arguments based on considerations of geo- growth-, or maturity-associated changes in V̇O2 peak by
metric similarity or surface law predict that V̇O2 peak correcting data for a single body-size indicator within
should scale to mass raised to the power 0.67 (28). each analysis, usually body mass (15, 27) or anthropo-
Nevertheless, when V̇O2 peak is modeled in subject groups metrically predicted lean body mass (21, 27). However,
heterogeneous for factors such as body size and age, a more comprehensive understanding of developmental
several studies have reported mass exponents close to changes in V̇O2 peak should ideally investigate simulta-
the power 0.75 (20). The numeric value of an alterna- neously the influence of other known covariates. For
tive ‘‘universal’’ or ‘‘true’’ mass exponent for normaliz- example, despite valid concerns regarding issues of

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2234 MODELING PEAK OXYGEN UPTAKE DURING GROWTH
collinearity among covariates (11), stature has been
shown to be a significant, independent predictor of
V̇O2 peak in both adults and young people when incorpo-
rated alongside mass in an allometric analysis (20, 25,
34). This was confirmed in the baseline model pre-
sented here (Table 2, model 1) with a significant
exponent for stature of 0.78 6 0.12.
The addition of the sum of two skinfolds (Table 2,
model 2) to the baseline model made the terms for both
stature and age squared redundant. Incorporating this
measure of body fatness also explained a large propor-
tion of the observed maturity effects, with the magni-
tude of the effect increasing with progression through
maturational stages. A similar effect was observed
when the development of mean power obtained during
a Wingate Anaerobic Test in 12–13 yr olds from the
present population (9) was modeled. In the present
study, differences in skinfold thickness also explained
part of the gender difference observed in addition to a
considerable proportion of the residual between-
individual variance (level 2).
Studies investigating age, growth, and maturational
changes in the body mass-V̇O2 peak relationship by using
ontogenetic allometry (15, 27) have noted extreme
variability in individual mass exponents with, for ex-
ample, values ranging from 0.18 to 1.74 in one study
(27). The flexibility of the multilevel modeling proce-
dure used in the present study enables the underlying
mean response to be described while concurrently
allowing individuals to have their own mass exponent.
The need for this was examined by fitting a random
component for mass at level 2 of the analysis (i.e.,
between individuals). When this was done, however,
the model failed to converge, indicating that the fixed
(mean) parameter adequately described this popula-
tion. Individual variation in overall growth rates was
evident, as indicated by the significant random variance
at level 2 associated with the age term. The variability
in overall rates of change in V̇O2 peak during the 3 yr of
this study is clearly evident from the individual growth
trajectories illustrated in Fig. 1.
Fig. 1. Examples of individual predicted changes in peak O2 uptake
(V̇O2 peak) adjusted for body mass skinfold thicknesses and maturity
(see Table 2, model 2).
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The data from the first year of the study support
evidence of gender differences in V̇O2 peak at age 11 yr
(4). In fact, a significant gender difference in V̇O2 peak
was demonstrated even when prepubescent girls (n 5
53) and boys (n 5 111) were compared, and with this
subsample there was no significant gender difference in
skinfold thicknesses (1). Explanations for girls’ lower
V̇O2 peak at a young age are speculative but the lower
values may be due to a lower exercise stroke volume
than in boys. Boys have consistently demonstrated
higher stroke volumes than girls at the same submaxi-
mal oxygen uptake or exercise intensity, although
differences have been small and in some cases statisti-
cally insignificant (26). Comparative data at V̇O2 peak
appear to be limited to one published study (23), which,
by using carbon dioxide rebreathing, reported boys’
stroke index to be 15% higher than those in girls at
9–10 yr of age and 5% higher at 11–12 yr of age.
However, recent work by Rowland (26), using Doppler
echocardiography, provides supportive data, with 12-yr-
old boys reported to have 13% higher maximal stroke
indexes relative to lean body mass than similarly aged
girls.
The results of the multilevel regression models exam-
ined here confirm previous cross-sectional indications
(34) in showing increases in size-related V̇ O2 peak
that contrast with conventional interpretation of V̇O2peak
during growth (4). Furthermore, the significant age-by-
gender interaction term in the present data set indi-
cates a progressive divergence in boys’ and girls’ values
over the age range examined. The increasing gender
difference in V̇O2 peak during childhood and adolescence
has been attributed to the greater accumulation of body
fat in relation to body mass in girls, boys’ higher
hemoglobin concentration, and girls’ lower levels of
habitual physical activity (4).
The addition of the sum of skinfold thicknesses to the
baseline model reduced the magnitude of the gender
term. This may be reflective of boys’ relatively greater
increase in muscle mass, which would not only facili-
tate the use of oxygen during exercise but may also
supplement the venous return to the heart and there-
fore augment stroke volume, through the peripheral
muscle pump (6, 26).
During puberty there is a marked increase in hemo-
globin concentration and hence oxygen-carrying capac-
ity in boys, whereas girls’ values plateau (16). It might
therefore be expected that differences in hemoglobin
levels between boys and girls would be a contributory
factor to the observed gender difference in V̇O2 peak, and
this has been demonstrated with 14 and 15 yr olds (10).
Hemoglobin levels were determined routinely in the
present study at each laboratory visit, but, when inves-
tigated as an additional explanatory variable, a nonsig-
nificant parameter estimate was obtained with these
11–13 yr olds. This is not an unexpected finding given
the minimal change in hemoglobin concentration across
the 3 yr of observation in both boys and girls (see
Table 1).
Boys have been consistently demonstrated to have
higher levels of habitual physical activity than girls,
 MODELING PEAK OXYGEN UPTAKE DURING GROWTH
but the evidence relating habitual physical activity to
V̇O2 peak is conflicting (5). In parallel studies the physical
activity patterns of the present children were moni-
tored over 3 days, using continuous heart rate monitor-
ing, and no significant relationships were detected
between V̇O2 peak and heart rate indicators of level of
physical activity in either the first (3) or second year (8)
of the study. In a multilevel regression analysis of the
3-yr heart rate data, V̇O2 peak was examined as an addi-
tional explanatory variable, but a nonsignificant param-
eter estimate was obtained (2). Habitual physical activ-
ity is unlikely to influence children’s and adolescents’
V̇O2 peak because such activity typically lacks the inten-
sity and duration sufficient to improve aerobic fitness (5).
In the exercise sciences, maturity is usually assessed
by using indicators of skeletal, somatic, or sexual
maturity. Although no single assessment gives a com-
plete description of the tempo of maturation, there is a
high concordance among the aforementioned indicators
(13). Previous studies have revealed that skeletal age
adds little to the description of physiological variables
yielded by chronological age and body size (30). Beunen
and Malina (14) reviewed the literature concerning
V̇O2 peak and the adolescent growth spurt, commented
that the available data should be interpreted with
caution, and concluded that the evidence suggests a
spurt in V̇O2 peak in boys that reaches a maximum gain
at the time of peak height velocity, but secure data are
insufficient to offer any generalization for girls. Few
longitudinal studies have investigated the influence of
maturity, independent of body size, on V̇O2 peak. Baxter-
Jones et al. (12) used an additive polynominal model
within a multilevel regression structure. They assessed
maturity by using secondary sexual characteristics (31)
and reported that, in athletic boys, there was a signifi-
cant increase in V̇O2 peak toward the end of puberty that
was in direct contrast to the nonsignificant increase in
V̇O2 peak found in athletic girls during this time. The
authors attributed this additional increase in V̇O2 peak
partially to the boys’ trained status. Using stature
velocity as an indicator of maturity status and analysis
of covariance to control for body mass, Malina et al. (22)
observed similar results with children from a sports
school. In the present study of untrained boys and girls,
maturity positively affected V̇O2 peak in both genders,
and an additional effect of chronological age indicated
the importance of incorporating both age and maturity
into analyses of V̇O2 peak during growth and maturation.
In summary, the multilevel modeling approach has
revealed age, gender, and maturity effects, independent
of body size, on the V̇O2 peak of untrained boys and girls.
These effects may have been masked in previous stud-
ies by the inappropriate use of V̇O2 peak in ratio to body
mass and/or the failure to consider the impact of
covariates other than body mass. Physiological explana-
tions for these effects cannot be established fully by the
present data set, but it appears that gender, age, and
maturity differences in the increase in fat-free mass
relative to body mass are the predominant influences
on the differential growth of boys’ and girls’ V̇O2 peak in
11–13 yr olds.
Downloaded from journals.physiology.org/journal/jappl (188.025.174.033) on December 27, 2023.
2235
We gratefully acknowledge the technical assistance of Jenny
Frost, Alison Husband, and Sue Vooght.
The work was supported by the British Heart Foundation and the
Healthy Heart Research Trust. Alan Nevill is with the School of
Human Sciences, Liverpool John Moores University, Liverpool, UK.
Address for reprint requests and other correspondence: N. Arm-
strong, Children’s Health and Exercise Research Centre, Univ. of
Exeter, Heavitree Rd., Exeter, EX1 2LU, UK (E-mail: N.
Armstrong@exeter.ac.uk).
Received 27 April 1999; accepted in final form 10 August 1999.
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