ANNALS OF HUMAN BIOLOGY, 1999, VOL. 26, NO.

1, 49± 62

Hormonal factors in the development of di€ erences in strength between

boys and girls during adolescence: a longitudinal study
J. M. R ound , D. A. Jones, J. W. Honour ² and A. M. Nevill³
The University of Birmingham, Birmingham, UK
² UCL Medical School, London, UK
³ John Moores University, Liverpool, UK

Received 24 September 1997; accepted 28 May 1998

Summary. The development of elbow ¯ exor (biceps) and knee extensor (quadriceps)
strength has been followed in a mixed longitudinal study of 50 boys and 50 girls from the
age of 8 to 17 years. Sex di€ erences in strength emerged at the time of peak height velocity
and were especially marked for the biceps. Data for individual children were aligned to the
time of peak height velocity and associations between strength, height, weight and circulat-
ing testosterone were investigated using multi-level modelling. The results show that, for
girls, quadriceps strength is proportional to height and weight while for boys there is an
additional factor which can be fully attributed to increasing levels of testosterone. Testos-
terone is important in explaining di€ erences in biceps strength between the sexes but an
additional factor is also required. It is suggested that, in addition to a direct e€ ect on muscle,
testosterone could have a second indirect action on biceps strength by promoting growth in
length of the humerus as part of the general development of the male upper limb girdle.

1. Introduction
There are well known di€ erences in muscularity between men and women, but the
reasons for this are not so clear. Di€ erences in development could be due to the
primary expression of a sex-linked genetic di€ erence in the muscle itself, to hormonal
di€ erences between the sexes which develop at puberty or to di€ erences in the
patterns of muscle use at work and play in childhood and adolescence. Of these
possibilities, the role of testosterone has received most attention and the purpose of
the present study was to examine the role of sex and growth hormones in the devel-
opment of di€ erences in strength between the sexes at puberty.
The association between testosterone and muscle growth, although widely
accepted, is based largely on anecdotal reports of drug abuse in athletes and body
builders and the observation that major di€ erences in muscular development appear
around the time of puberty. Recently it has been shown that supra-physiological
doses of testosterone, combined with speci® c training, stimulate muscle growth in
adults (Bhasin, Storer, Berman, Callegari, Clevenger, Phillips, Bunnell, Tricker,
Shirazi and Casaburi 1996) but it is not clear how this action relates to normal
patterns of growth. Testosterone is also known to restore muscle bulk in hypogo-
nadal males (Ginspoons, Corcoran, Lee, Burrows, Hubbard, Katznelson, Walsh,
Guccione, Cannan, Heller, Basgoz and Klibanski 1996) but, again, it is not clear
how this situation relates to normal muscle development.
A number of cross sectional and longitudinal studies have documented strength
changes in children and have shown that peak strength gain occurs at, or just after,
the time of peak height velocity (Jones 1949, Stoltz and Stoltz 1951, Asmussen and
Heeboll-Nielsen 1955, Malina and Johnston 1967, Faust 1977, Hughes and
Whitehouse 1981, Kemper and Verschuur 1985b, Beunen, Malina, Van’t Hof

0301-4460/99 $12. 00 Ñ 1999 Taylor & Francis Ltd.

50 J. M. Round et al.
1998) and, where comparisons were made between the sexes, boys have been found
to have the greater gains in strength at and after the growth spurt.
There is evidence that upper and lower body musculature show di€ erent patterns
of growth during adolescence. In a longitudinal study, Carron and Bailey (1974)
found that measures of upper body strength in boys increased 3.9 fold between the
ages of 10 and 16 years whilst for lower body strength the increase was 2.5 fold.
Tanner, Hughes and Whitehouse (1981) estimated that peak growth velocity of arm
muscles in boys was about twice that in girls while there were no sex di€ erences in the
calf. Kemper and Verschuur (1985a), in a longitudinal study measuring muscle vol-
umes, showed that sex di€ erences became apparent at puberty, with a suggestion of
greater di€ erences between boys and girls in the upper body musculature. In a cross
sectional study, Parker, Round, Sacco and Jones (1990) measured the isometric
strength of forearm ¯ exor and knee extensor muscles in boys and girls between
the ages of 8± 18 years and showed that, in the older children, whilst boys were
stronger than girls in both muscle groups, the boys had proportionally greater
strength in the upper arms.
It seems likely that these di€ erences in muscle development are due to the increase
in circulating testosterone which is a feature of maturation in normal male adoles-
cents at this time of testicular growth (e.g. Winter and Faiman 1972, Sizonenko and
Paunier 1975). However, to this study’s knowledge, a parallel longitudinal study of
changes in circulating sex hormones and increases in strength in adolescents has not
been reported.
The purpose of the present investigation was to obtain quantitative data on the
relationship between strength, height, weight and levels of circulating sex hormones
and Insulin-like Growth Factor-1 to see whether increasing testosterone levels pro-
vide a full explanation for the sex di€ erences in muscle strength which emerge at
puberty. To this end a mixed longitudinal study of boys and girls has been carried
out during the period which spans the adolescent growth spurt and sexual matura-
tion and the data have been subjected to multi-level modelling.

2. Methods
2.1. Subjects
One hundred North London school children were studied, 50 boys and 50 girls.
Three sets of measurements were made each year, once a term at roughly 4 monthly
intervals. Children were recruited in groups commencing at ages between 8 and 12
years, giving ® nal ages of 13± 17 years at the end of the study.
Permission to approach the children was obtained from the Local Education
Authority, the Head Teachers of the schools and from the committee on the
Ethics of Human Investigation at the Middlesex and University College Hospitals
(now the University College London Hospitals). In the ® rst years the younger chil-
dren were attending a junior school from which the majority of children passed on to
the local mixed comprehensive school where the older children were recruited.
Consequently in the later years of the study all the participating children were
attending the same comprehensive school. Ethnic origins of the children were
mixed and included a few children with parents of Cypriot, Pakistani and Chinese
origins.
Children volunteered for the study after being fully informed about the tests
involved and for how long they would continue in `the team’. Parental consent in
writing was then obtained for all the children who had volunteered. The parents were
 Hormones and strength in adolescent boys and girls 51
asked to supply the name of their General Practitioner who was informed in writing
about the study and given the names of the children on their list who were partici-
pating.
As in all long term studies, there was some loss of subjects during the 5 year
period. Sixty per cent of the children competed the full 5 years, with similar propor-
tions of boys and girls. The main loss occurred when children left school at 15 and 16
years of age, but most of these children had been in the study for at least 4 years by
this time. The other major loss occurred when the 11 year old children changed from
junior to senior school with some of the children continuing their education at
schools outside the area. Eighty four per cent of the girls and 82% of the boys
were studied for at least 4 years.

2.2. Muscle strength

Maximum voluntary contraction (MVC) strengths of the knee extensors and
forearm ¯ exors (hereafter referred to as quadriceps and biceps muscles respectively)
were measured using a chair described by Park, Round, Sacco and Jones (1990). The
investigators were experienced in the use of the various procedures and the main
members of the team responsible for the measurements remained the same through-
out the study.
Each child had time to become familiar with the apparatus and procedures and
then performed three MVCs, the highest of which was recorded. As found in a
previous study (Parker et al. 1990), there was no consistent trend between the
three strength measurements. On the rare occasions when there was appreciable
variation between the three e€ orts, the contractions were repeated until three
values were obtained which di€ ered by no more than 4% from the highest force.
The coe cient of variation of strength measurements of both muscle groups was
10% . The measurement of isometric strength of single muscle groups by this tech-
nique requires little or no learning and are therefore unlikely to be in¯ uenced by the
development of motor skills (Edwards, Young, Hosking and Jones 1976) and corre-
late well with measurements of muscle cross-sectional area (Chapman, Grindrod and
Jones 1984).

2.3. Height
Standing height was measured using a portable stadiometer (Holtain Ltd,
Crossways, Gwent, Wales). Children removed their shoes and stood with their
heels and back against the upright with the neck gently extended by upward
manual pressure. Sitting height was measured with the child sitting upright against
the stadiometer on a standard stool and upper body length obtained by subtracting
the stool height from the sitting height. In turn, lower body length was obtained by
subtracting upper body length from the standing height. The length of the upper arm
was measured as the distance from the acromion to the olecranon process. Height
and length were recorded to the nearest millimetre. The standard deviations of
repeated measures were, for standing height, 2.5 mm, for sitting height, 8 mm and
for upper arm length, 4.5 mm.

2.4. Weight
Weight was measured to the nearest 0.1 kg using calibrated portable scales with
the children in light indoor clothing without shoes.
52 J. M. Round et al.
2.5. Blood sampling
Ten millilitres of blood was obtained from an antecubetal vein. All blood samples
for testosterone were obtained between 9 and 10 am to minimize variation due to
diurnal changes (Wu, Brown, Butler, Stirling and Kelnar 1993). Testosterone was
measured on plasma samples using a radioimmunoassay (Holownia, Owen, Hampl,
Jacobs and Honour 1991). Oestradiol was assayed using a radioimmunoassay kit
obtained from INCSTAR (Wokingham, Berks, UK), Insulin-like Growth Factor-1
(IGF1) by the method of Morrell, Dadi, More and Taylor (1989) and Alkaline
Phosphatase by the method of Bowers and McComb (1975).

2.6. Age
Chronological age is reported in decimal years. Girls were asked about menstrual
status. Breast development in girls, and genital development in boys, was monitored
in the youngest cohort of children entering the study using the Tanner scales (Tanner
and Marshall 1969, 1970). Genital stages were assessed longitudinally by the same
male (for the boys) and female (for the girls) investigators throughout the study.

2.7. Data analysis

Growth curves were constructed for each child and height velocity calculated by
dividing the increments in height by the time between consecutive visits. Where there
was a single highest value for height velocity, time of peak height velocity (PHV) was
de® ned as the mid point between the two ages spanning PHV. Where there were two
or more equally high values, time of PHV was taken as the mid point between the
two outermost times. Identi® cation of PHV was con® rmed by measurements of
plasma alkaline phosphatase, which peaks at this time (® gure 1, Round 1973,
1977). The growth spurt in height occurred in 25 girls and 33 boys during the
time of the study. PHV is expressed as cm/year, although it should be noted that
these values relate to a rate of growth over only an approximate 4 months period.
For the boys, PHV was 11.5 cm/year, occurring at a mean age of 13.4 years while for
girls, PHV was 9.7 cm/year at the earlier age of 12.2 years. PHV occurred at genital
stage 3 in both boys and girls and, for the girls, menarche occurred at a mean age of
12.9 years. Oestrogen levels began cycling in an adult pattern by about 1 year after
PHV (® gure 2). Time of PHV was used to align the other measurements made on the
children, reporting values as years before or after PHV, with the year 0 encompass-

Figure 1. Plasma alkaline phosphatase activity as a function of years before and after peak height
velocity (PHV). Values are given as mean SD of the values for children in each year. d Boys, j Girls.
 Hormones and strength in adolescent boys and girls 53

Figure 2. Plasma oestradiol in relation to age of peak height velocity (PHV). Values are given as
mean SD of the values for girls in each year for whom the age of PHV could be clearly identi® ed.

ing 0.5 years on either side of the date identi® ed as PHV. For descriptive proposes
the data are reported as means and standard deviations but, in addition, they have
been subjected to multi-level modelling to assess the contribution that height, weight
and sex age interaction term or testosterone, make to the di€ erence in strength
seen between boys and girls.

2.8. Statistical methods

Descriptive statistics (mean SD) were performed using conventional statistical
software. Changes in strength of the quadriceps and biceps were analysed by devel-
oping the methods adopted by Baxter-Jones, Goldstein and Helms (1993) using the
Multi-level Regression Modelling Procedures (Goldstein 1986).

2.9. Modelling
Analyses were performed, using the program Multi-level Models Project ML3
(Prosser, Rashbash and Goldstein 1990), to identify factors associated with the
development of strength in the quadriceps and biceps, having adjusted for di€ erences
in body size (height and weight), age, sex and, subsequently, testosterone, as covari-
ates. The two levels of hierarchical or nested observational units were the visit
occasions at level 1 within-individuals), and the children (between-individuals) at
level 2. However, in contrast to the additive model adopted by Baxter-Jones et al.
(1993), an alternative multiplicative allometric model for strength was adopted
(Nevill 1994, Nevill and Holder 1995, Nevill, Holder, Baxter-Jones, Ropund and
Jones 1998) as follows,
strength weightk1 heightk2 exp a b age c age2 . 1
The model can be linearized with a log-transformation and a multi-level regression
analysis on loge strength can be used to estimate the unknown parameters. The
transformed log-linear multi-level regression model becomes,
loge strength k1 loge weight k2 loge height a b age c a2 . 2
The factor of sex was incorporated into subsequent analyses by introducing the
indicator variable sex (boys= 0, girls= 1), that allows the parameter in equation
(2) to vary for both boys and girls, e.g. by introducing a sex age interaction
54 J. M. Round et al.

Figure 3. Quadriceps strength in relation to PHV. Values are given as mean SD of the values for
children in each year for whom the age of PHV could be clearly identi® ed. d Boys, j Girls.

Figure 4. Biceps strength in relation to PHV. Values are given as mean SD of the values for
children in each year for whom the age of PHV could be clearly identi® ed. d Boys, j Girls.

term which allows for di€ erent values of b for boys and girls. Age is entered in the
equation as the years before or after PHV.

3. Results
Results for strength measurements of quadriceps and biceps are shown aligned for
the time of PHV in ® gures 3 and 4. Before PHV, absolute strengths were higher in
the boys compared to the girls for a given age relative to PHV as a consequence of
the later occurrence of PHV in boys. Increases in strength, however, were similar in
the two sexes until about 1 year before PHV. For both muscle groups, clear di€ er-
ences between the sexes in the rate of increase in strength were evident from 0 to 2
years after the time of PHV.
Figure 5 shows the plasma testosterone measurements centred upon the time of
PHV for boys. Circulating testosterone levels began to rise 1 year before PHV and
then increased steadily, reaching adult levels around 3 years after PHV. The increase
in testosterone coincided with the divergence of strength between boys and girls seen
in ® gures 3 and 4.
 Hormones and strength in adolescent boys and girls 55

Figure 5. Circulating testosterone in relation to PHV. Plasma testosterone in boys plotted against the
years before or after the time of peak height velocity. Values are given as mean SD of the values for
children in each year for whom the age of PHV could be clearly identi® ed.

Table 1. Parsimonious multilevel regression analysis for log-transformed strength of quadriceps.

(a) Adjusted for body size (weight and height), age (centred about time of PHV) and sex.
Random variable Constant Age
Level 1 (within individuals)
Constant 0. 009 27 0. 000 66
Level 2 (between individuals)
Constant 0. 015 9 0. 003 2
Age 0. 001 16 0. 000 8 0. 001 30 0. 000 38
Explanatory variables Estimate SEE
Constant 3.915 0. 306
Loge Weight 0.383 0. 094 34
Loge Height 1.233 0. 327 9
Sex (Female± Male) 0. 150 0. 040 3
Age 0.0533 0. 014 9
Age2 0. 0067 0. 002 5
Age Sex 0. 0441 0. 012 3
(b) Adjusted for body size (weight and height), age (centred about time of PHV), sex and testosterone.
Random variable Constant Age
Level 1 (within individuals)
Constant 0. 009 31 0. 000 66
Level 2 (between individuals)
Constant 0. 018 7 0. 003 8
Age 0. 009 26 0. 000 8 0. 001 06 0. 000 38
Explanatory variables Estimate SEE
Constant 3.677 0. 302
Loge Weight 0.368 0. 097 4
Loge Height 1.661 0. 297
Age 0. 0058 0. 010 3
Testosterone 0.0068 0. 001 44

Table 1a summarizes the results from the multi-level regression analysis of

strength in the quadriceps, adjusting for body size, age and sex. The parsimonious
solution identi® es a signi® cant increase in quadriceps strength explained by the
developmental component in body size (weight and height) plus an additional con-
tribution identi® ed by age and age2 components, that were di€ erent for boys and
girls identi® ed by the signi® cant age sex interaction.
56 J. M. Round et al.
Note that the contribution of the age term for girls is negligible, identi® ed by the
value of age sex interaction term ( 0. 0441) that is subtracted from the age par-
ameter (0.0533). When the factor testosterone is introduced into the analysis, the
parsimonious model for quadriceps, given in table 1b, only required the develop-
mental growth component of body size (height and weight) plus the additional com-
ponent testosterone.
The components of sex, age sex and age2 make no signi® cant contribution to
the prediction of strength in the quadriceps (although not signi® cant, the term age
must be retained since age is allowed to vary for each child in the random variable
part of the model). The conclusion is, therefore, that for the girls, development of
quadriceps strength is proportional to the general increase in height and weight,
while for the boys there is an additional factor than can be fully explained by the
term testosterone.
A similar result was obtained from the multi-level regression analysis of strength
of the biceps adjusting for body size, age and sex (table 2a). Note that the gap
between the boys’ and girls’ age parameter ( 0. 0717) is greater in the model for
biceps strength compared with the model for quadriceps ( 0. 0441). In addition,
the girls’ age term contribution to the regression model was negative (0. 0483±
0. 0717 0. 0234), implying that the girls became weaker with age, relative to the
changes in body size.

Table 2. Parsimonious multilevel regression analysis for log-transformed strength of biceps.

(a) Adjusted for body size (weight and height), age (centred about time of PHV) and sex.
Random variable Constant Age
Level 1 (within individuals)
Constant 0. 009 49 0. 000 66
Level 2 (between individuals)
Constant 0.015 0 0. 003 1
Age 0. 000 08 0. 000 6 0. 000 55 0. 000 23
Explanatory variables Estimate SEE
Constant 3.313 0. 298
Loge Weight 0.361 0. 091 9
Loge Height 1.062 0. 322
Sex (Female± Male) 0. 171 0. 039 4
Age 0.048 3 0. 013 8
Age2 0. 005 5 0. 002 3
Age Sex 0. 071 7 0. 009 7
(b) Adjusted for body size (weight and height), age (centred about time of PHV), sex and testosterone.
Random variable Constant Age
Level 1 (within individuals)
Constant 0. 009 66 0. 000 67
Level 2 (between individuals)
Constant 0. 015 1 0. 003 1
Age 0.000 24 0. 000 75 0. 001 13 0. 000 35
Explanatory variables Estimate SEE
Constant 3.259 0. 306
Loge Weight 0.371 0. 094 1
Loge Height 1.000 0. 332
Sex (Female± Male) 0. 143 0. 041 5
Age 0. 005 15 0. 013 8
Age2 0. 005 8 0. 002 5
Testosterone 0.005 13 0. 001 46
 Hormones and strength in adolescent boys and girls 57

Figure 6. Circulating Insulin-like Growth Factor 1 (IGF1) in relation to PHV. Plasma levels are plotted
against the years before or after the time of peak height velocity. Values are given as mean SD of the
values for children in each year for whom the age of PHV could be clearly identi® ed. d Boys, j Girls.

Figure 7. Body length in relation to PHV. A, Upper body length and B, lower body length plotted
against the years before or after the time of peak height velocity. Values are given as mean SD of
the values for children in each year for whom the age of PHV could be clearly identi® ed. d Boys, j
Girls.

When the factor testosterone was introduced into the multi-level regression analy-
sis for biceps, its inclusion was not able to explain entirely the di€ erence in biceps
strength between the boys and girls (see table 2b).
Although signi® cant, testosterone was not as successful as the age sex inter-
action term when predicting the strength of the biceps. Furthermore, unlike the
model for quadriceps, the terms of sex and age2 were retained as they still made
signi® cant contributions to the model. In summary, the analysis indicates that, whilst
testosterone explains much of the di€ erence between the strengths of boys’ and girls’
biceps, there is some additional factor making the boys disproportionately stronger
in their arms.
Growth hormone, mediated by the action of Insulin-like Growth Factor-1, is
important in growth and might be thought to play some part in sex di€ erences in
muscular development. Circulating IGF1 increased during the years before and after
PHV (® gure 6) and tended to precede the increase in sex hormones but, within the
variability of the measurements, there were no di€ erences between the sexes.
Figure 7 shows the changes in lower and upper body length in the years spanning
the time of PHV. For both upper and lower body length, boys were longer than girls
at all ages relative to the time of PHV, re¯ ecting, as with strength in ® gures 3 and 4,
58 J. M. Round et al.

Figure 8. Upper arm in relation to PHV. Upper arm length plotted against the years before or after the
time of peak height velocity. Values are given as mean SD of the values for children in each year for
whom the age of PHV could be clearly identi® ed. d Boys, j Girls.

the later growth spurt in boys compared to girls. Lower body length (® gure 7a)
showed a fairly steady increase up to 1 year after the time of PHV when growth
markedly slowed; the pattern of growth being very similar in boys and girls. The
upper body showed an acceleration in growth in the 2 years spanning the time of
PHV (® gure 7b) with evidence of continuing, albeit slower, growth in both sexes until
3 years after PHV.
Figure 8 shows data for upper arm length. Two and 3 years before the time of
PHV, boys and girls had upper arms of the same length, but whilst for the girls there
was little change in the next 6 years, in the boys there was marked increased in length
in the 2 years spanning the time of PHV. The patterns of growth of upper and lower
body length (® gure 7) were similar for boys and girls but those for the upper arm
(® gure 8) were qualitatively, as well as quantitatively, di€ erent between the sexes.

4. Discussion
In the mixed longitudinal study presented here the plasma testosterone has been
measured and multi-level modelling has been used which makes it possible to esti-
mate the extent to which testosterone can explain the sex di€ erences in strength. In
part, the results con® rm what is probably a widespread belief that changes in tes-
tosterone are a su cient explanation of the di€ erence in strength between boys and
girls for the quadriceps. For the biceps, however, the results are surprising in that
they show an additional factor(s) is needed to explain the di€ erence in strength
between the sexes.
The present cohort of children was very similar in absolute strength and pattern of
growth to that studied in the cross sectional survey of Parker et al. (1990) which
implies that the increases in strength seen in the longitudinal study were unlikely to
have been the consequence of learning the technique by repeating the tests over the 5
year period.
This study’s data, when aligned to the time of PHV, clearly show sex di€ erences in
strength becoming apparent at about the time of the boys’ growth spurt (® gures 3
and 4). Figure 5 shows that in the year of PHV, plasma testosterone had risen to a
mean value of around one third of the adult level. The visual impression from these
® gures is that the time course of the rise in testosterone is very similar to that of the
development of sex di€ erences in strength for both muscle groups.
 Hormones and strength in adolescent boys and girls 59
Multi-level modelling of mixed longitudinal data e€ ectively allows individuals to
have their own growth curves as well as ® tting separate growth curves for speci® c
groups within the sample (e.g. boys and girls). Thus group e€ ects (i.e. comparing the
boys’ and girls’ growth parameters) that are larger than the within-individual vari-
ation can be identi® ed. The technique that has been used is very similar to that used
by Baxter-Jones et al. (1993) except that this study has used a multiplicative allo-
metric model rather than additive. The multiplicative model is intuitively superior
since, when extrapolating to zero weight or height, the predicted strength will also be
zero as opposed to a ® nite value for the intercept given by an additive approach.
The multi-level modelling analysis for the quadriceps indicates that, for the girls
during puberty, strength increases in proportion to the size of the body and that no
other age-related factor is required to explain the increase in strength. For the boys,
however, height and weight alone are not su cient to explain the increase in strength
and an additional term is required which is provided by circulating testosterone. The
results of the modelling show that testosterone fully explains the di€ erences between
the sexes which remain after making allowance for body size (height and weight).
For the girls’ biceps, a negative age term is required in addition to height and
weight which indicates that the girls’ biceps did not grow in strength in proportion to
body height and weight. In addition, the di€ erences between boys and girls is not
fully explained by testosterone, implying that some additional factor promotes
biceps growth in boys during puberty.
This additional factor is unlikely to be growth hormone since the time course of
the rise in IGF1 (® gure 6) is consistent with growth hormone and IGF being
involved in the adolescent growth spurt but not directly contributing to the di€ er-
ential muscle development of boys and girls.
Body weight is an important predictor of strength for both quadriceps and biceps
(tables 1 and 2) but since muscle makes a major contribution to body mass, ® nding
that one is related to the other gives very little indication of a causal relationship. It
has been previously pointed out (Parker et al. 1990) that, from dimensional argu-
ments, the strength of a muscle would be expected to be proportional to its cross-
sectional area and thus the square of linear body dimensions. The results of this
previous study con® rmed that, for pre-pubertal children, the strength of the biceps
was proportional to the square of body height but, for the quadriceps the relation-
ship involved the cube of height. It was suggested that the stimulus for growth in
both cases was growth of the skeleton, accounting for the square relationship, but
that with the quadriceps there was an additional stimulus due to loading with body
weight, accounting for the cube relationship with height. The analysis in that paper
was restricted to pre-pubertal children because the relationships clearly altered (at
least for the boys) with the onset of puberty. With the present longitudinal data, and
using multi-level modelling the e€ ect of testosterone can be taken into account and
the analysis extended to the full age range. From the information in tables 1 and 2, it
is possible to translate the exponent for weight into a term for height by multiplying
by three (since weight is proportional to height3 ) and adding this to the existing
exponent for height. The combined height exponent becomes 2.77 for the quadriceps
(table 1b), while for the biceps (table 2b) the exponent is 2.11. These exponents are
similar to the values of 3 and 2 suggested by Parker et al. (1990).
It is interesting to speculate on the nature of the additional factor, that is required
to explain the di€ erence in biceps strength between boys and girls. The di€ erence
might re¯ ect cultural attitudes with boys, at around the time of puberty, taking up
60 J. M. Round et al.
sports and activities which train the upper body musculature and Verschur and
Kemper (1985) have shown that boys of this age spend signi® cantly more time
than girls undertaking `heavy’ activities. This possibility is di cult to refute but, if
it were true, one would expect to see an overlap in strength between the sexes with
some of the more aesthetic boys in the cohort avoiding such training whilst some
active girls de® ed the social norms for their sex. The results in ® gure 4 show a very
clear separation between the sexes with no suggestion of the increased variance,
which might be expected in the older age groups if habitual activity was a major
determinant of strength. Of all the 16 year old children, only one boy had a biceps
strength below 200 N, while no girl had a biceps strength greater than 199 N. For the
quadriceps strength there was more overlap between the largest and strongest girls
and the smaller and weaker of the boys.
Another possibility for the additional factor required to explain the large sex
di€ erences in the strength of the biceps is that the standing height measurement,
and possibly the height component implicit in the body weight factor, used to predict
strength in tables 1 and 2 may not be a true re¯ ection of the length signal `seen’ by
the developing biceps. It is notable that the two components of standing height,
upper and lower body length, show very similar patterns of change between the
sexes during puberty (® gure 7) and consequently are unlikely to account for sex
di€ erences in the biceps. In contrast, the upper arm development, which determines
the length of the biceps, shows a very di€ erent pattern of change between the sexes
(® gure 8). it is well known that there are di€ erences in skeletal development which
become apparent at puberty with men having a greater development of the upper
limb girdle, presumably under the in¯ uence of testosterone, which will be re¯ ected in
di€ erences in growth of the upper arm.
It is important to understand how the statistical associations found between
strength, height, weight, and circulating testosterone relate to the physiological sti-
muli for muscle growth and increase in strength. The statistical associations are
consistent with the notion that increasing bone length is an important factor pro-
moting the growth of muscle during childhood and adolescence and it is reasonable
to assume that, for the quadriceps, the relevant bone is the femur while, for the
biceps, the humerus or upper arm length is the critical measurement. Body mass also
plays a role determining the strength of the quadriceps (since the composite height
exponent is greater than 2; see above) and, teleologically, this is understandable since
the quadriceps are major weight bearing muscles. Testosterone probably causes an
increase in strength by directly stimulating the growth of contractile tissue in both
muscle groups but it also has an additional indirect action on the biceps as a conse-
quence of the hormonal stimulation of skeletal development in the upper limb girdle
of adolescent boys. The large upper limb girdle and the resultant disproportionate
strength of the biceps in males (in relation to body size) may be seen, therefore, as
another example of a secondary sexual characteristic.

Acknowledgements
We are grateful to Action Research for their generous ® nancial support of this
work, to Mr P. Hudson, and the sta€ and pupils of Southgate and DeBohun Schools
for their enthusiastic help and participation. Ms Alison Gerharts and Dr M. Mills
provided excellent assistance during this long study and we thank Dr P. Holownia
for measuring the sex steroids.
 Hormones and strength in adolescent boys and girls 61
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Address for correspondence: Dr J. M. Round, School of Sport and Exercise Sciences, The University
of Birmingham, Birmingham B15 2TT, UK.

Zusammenfassung. Im Rahmen einer gemischten Longitudinalstudie an 50 Jungen und 50 MaÈ dchen im

Alter von 8 bis 17 Jahren wurde die Entwicklung der Kraft des Ellenbogenbeugers (Biceps) und des
Kniestreckmuskels (Quadriceps) untersucht. Die Geschlechtsunterschiede in der Kraft entwickelten sich
zum Zeitpunkt des Auftretens der maximalen Wachstumsgeschwindigkeit der KoÈ rperhoÈ he, sie waren fuÈ r
den Biceps besonders ausgepraÈ gt. Die Individualdaten wurden am Zeitpunkt des Auftretens der maxima-
len Wachstumsgeschwindigkeit der KoÈ rperhoÈ he ausgerichtet, und die ZusammenhaÈ nge zwischen der
Kraft, der KoÈ rperhoÈ he, dem KoÈ rpergewicht und der Konzentration des zirkulierenden Testosterons wur-
den mittels Multilevel-Modellierung analysiert. Die Ergebnisse zeigen, dass fuÈ r MaÈ dchen die Kraft des
Quadriceps proportional ist zur KoÈ rperhoÈ he und zum KoÈ rpergewicht waÈ hrend bei Jungen ein zusaÈ tzlicher
Faktor hinzukommt, der uneingeschraÈ nkt auf die ansteigende Testoteronkonzentration zuruÈ ckgefuÈ hrt
werden kann. Es wird vermutet, dass Testosteron zusaÈ tzlich zu seinem direkten Ein¯ uû auf den Muskel
durch seine wachstumsfoÈ rdernde Wirkung auf den Humerus als Teil der Wirkung auf die generelle
Entwicklung des SchulterguÈ rtels im maÈ nnlichen Geschlecht einen zweiten indirekten Ein¯ uû auf die
Kraft des Biceps ausuÈ ben koÈ nnte.

Re sume . On a observe le de  veloppement de la force de ¯ exion du coude (biceps) et de l’extension du

genou (quadriceps) au cours d’une Âetude longitudinale mixte de 50 garcËons et 50 ® lles aÃge s de 8 aÁ 17 ans.
Les di€  erences de force lie es au sexe apparaissent au moment du pic de vitesse de croissance de la stature et
sont particulieÁrement marque es pour le biceps. Les donne es individuelles ont Âete pour chaque enfant
aligne es sur le moment du pic de vitesse de croissance de la stature et les associations entre force, taille,
poids et testoste rone circulante, ont  ete recherche es au moyen de modeÁles de niveaux multiples. Les
re sultats montrent que la force du quadriceps est proportionnelle aÁ la stature et au poids chez les ® lles,
tandis qu’un facteur supple mentaire attribuable aÁ l’accroissement des niveaux de testoste rone, intervient
chez les garcËons. La testoste rone est importante pour expliquer les di€  erences de force du biceps entre les
sexes, mais un facteur supple mentaire est aussi ne cessaire. On suggeÁre qu’en plus de son action directe sur
le muscle, la testoste rone pourrait avoir une seconde action indirecte sur la force du biceps en entraõà nant la
croissance de l’humerus, qui est elle meÃme une partie du de veloppement ge ne ral de la ceinture des membres
supe rieurs.