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Body composition in baboons Evaluating a morphometric method.

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American Journal of Primatology 12275-285 (1987)
Body Composition in Baboons: Evaluating A
Morphornet ric Method
GARY W. RUTENBERG, ANTHONY M. COELHO, JR.,DOUGLAS S. LEWIS,
K. DEE CAREY, AND HENRY C. McGILL, JR.
Department of Physiology and Medicine, Southwest Foundation for Biomedical Research,
San Antonio, Texas
The objective of this study was to determine whether noninvasive morphometric measurements of olive baboons (Papio cynocephalus anubis) can
reliably predict lean body mass and fat mass in this species. Crown-rump
leng$h, triceps circumference, and skinfold measures at the neck, subscapular, suprailiac, and triceps sites were obtained prior to necropsy from 21
clinically normal infant baboons at 18 weeks of age and from 22 clinically
normal adolescent baboons a t 5 years of age. At necropsy, the lean body
mass and body fat mass were measured using gravimetric methods. Pearson's correlations and regression analysis were used to test the relationship
between direct measures of lean and fat mass obtained at necropsy with
calculated estimates based on morphometric measures obtained prior to
death. Our null hypothesis was that the morphometric measures (individually or in combination) were not related to direct measures of fat mass or
lean mass. Non-linear and multiple regression estimators, by age and gender, were derived and provided increased predictability. Our results indicate
that 1) morphometric measures can accurately predict lean body mass in
male and female baboons; 2) morphometric measures used to predict lean
body mass change with age; 3) morphometric measures are strongly associated with body fat mass at 18weeks of age but are not a s strongly associated
with body fat mass in 5-year-old baboons; 4)triceps circumference provides
the best single indicator of lean body mass for both genders and age periods;
5) baboons are like humans in that adolescent females tend to accumulate
bod,y fat while males of the same age tend to develop lean mass; and 6)
combinations of these morphometric measurements explain between 70%
and 100% of the variability and can be used to estimate lean and fat mass
in baboons.
Key words: 'baboons, body composition, developmental biology, morphometry
INTRODUCTION
Recent ,studies indicate that patterns of weight gain, increase in body size, and
fat development in baboons are similar to those in humans [Lewis et al, 1983, 1984;
Coelho & Glassman, 1984; Coelho et al, 1984; Glassman et al, 1984; Coelho, 19851.
Received April 25, 1986; revision accepted October 15, 1986.
Address reprint requests to Dr. Anthony M. Coelho, Jr., Behavioral Medicine Laboratory, Department of
Physiology and Medicine, Southwest Foundation for Biomedical Research, P.O. Box 28147, San Antonio,
TX 78284.
0 1987 Alan R. Liss, Inc.
276 I Rutenberg et a1
These similarities are particularly evident during adolescence, when both baboons
and humans have a period of accelerated growth [Coelho et al, 1984; Glassman et a1
1984; Coelho, 19851.However, it is not clear whether weight in baboons is distributed
into lean and fat mass compartments in a manner similar to humans. If baboons
and humans undergo similar changes in lean mass and fat mass, the value and
attractiveness of the baboon as a n animal model is greatly enhanced. Baboons can
be used to provide experimental and prospective information on the role of nutrition
on growth, development of obesity, and human diseases associated with obesity.
One approach to determining lean body mass and fat mass is the use of direct
methods that measure tissue mass a t necropsy [Lewis et al, 1983, 1984, 1986al.
However, this method does not provide a means of studying longitudinal changes in
body composition because it requires the death of the subjects. There are a number
of indirect noninvasive methods that can provide, a t a single time point or a t
multiple time points, estimates of lean and fat mass. These include nuclear magnetic
resonance (NMR) {Lewiset al, 1986b], dilution methods t4'K assessment, desiometry,
radiometry [Forbes, 19781, and morphometry [Malina, 1969; Johnston, 19821. However, these methods must be verified by direct methods each time a new species is
studied.
There are several descriptive studies of body composition of both Old and New
World species. For example, Ausman et a1 [1982] and Russo et a1 [1980] provide
information on body composition a t necropsy in Cebus albifrons and Saimiri sciureus. In these species, water, protein, and ash were shown to be linearly related to
carcass weight, but the variability in body fat was so large that predictive equations
could not be provided for either age or weight. This variability may have been due
to limitations in sample (ie, the wide range of ages and small sample sizes a t each
age) andlor to the fact that animals were obtained opportunely as they died. This
second condition is important because it suggests that the animals were not clinically normal and that the variability may have been due to disease or illness.
Additionally, these studies do not provide information on the relationship of body
composition to morphometric measures that could be used as noninvasive measures
of lean mass or adipose tissue.
Morphometric methods utilize a series of body size measurements, such as
skinfolds, lengths, girths, and weights. These measures are widely used in human
research because they are easy to take, inexpensive, accurate, and provide serial
(longitudinal) information on subjects. These methods have been validated for estimating lean body mass and fat mass in humans by comparison with underwater
weights and other methods [Brozek, 1963, 1966; Brozek & Henschel, 1961; Malina,
1969; Johnston, 1982; Garrow, 19821. Although these methods are used extensively
in human body composition research, there are few published studies in which
morphometry was applied to body composition in nonhuman primates. This situation is due, in part, to lack of information on the accuracy of predictions of lean body
mass and fat mass applied to nonhuman primates.
Small [1981], for example, compared skinfold measures with total body weight
and assumed a direct correlation between the skinfolds and adipose tissue. However,
there was no direct or indirect evaluation of the accuracy of skinfolds in estimating
the absolute amount of adipose tissue in the study animals. Walike et a1 [1977] and
Walker et a1 [1984] used total body water to indirectly assess total body fat. They
found strong relationships between skinfolds and estimates of total body fat. Moreover, their data suggest the potential for using morphometric variables to predict
body fat.
Body Composition in Baboons I 277
The objective of the study reported here was to evaluate the accuracy of morphometric methods in providing estimates of lean body mass and fat mass in baboons,
and is a necessary first step toward the general application of morphometric methods
to studies of body composition in nonhuman primates. The relationship between
morphometric measures and direct measures of lean body mass and fat mass were
evaluated using regression formulae. Morphometric methods were used to estimate
body composition at two different age periods: infancy and adolescence. These two
age periods correspond with the end of the neonate period (18 weeks) and the
midpoint of adolescence (5 years), when both menarche and adrenarche have been
attained and much of the pubertal growth spurt has occurred [Glassman et al, 19841.
MATERIALS AND METHODS
Subjects
The subjects were 43 clinically normal olive baboons (Papio cynocephalus anubis),born and raised at the Southwest Foundation for Biomedical Research. The first
group consisted of 21 infants (9 males, 12 females) who were 18 weeks of age a t the
time of measurement. The second group consisted of 22 adolescents (10 males, 12
females) who were 5 years of age at the time of measurement. All subjects were
assigned at birth to a n experiment whose objective was to evaluate the effects of
infant nutrition on the development of adiposity.
Dietary History
Infant baboons were fed Similac (Ross Laboratories, Columbus, OH) formulas
from birth to 18 weeks of age. The 5-year-old baboons were fed a post-weaning (> 18
weeks of age) diet based on Ralston Purina Chow (Ralston Purina, St. Louis, MO),
which contained approximately 40% calories as fat, 20% as protein, and 40% as
carbohydrate.
Morphometric Measures
Measurements were obtained of the neck, subscapular, suprailiac, and triceps
skinfolds; triceps cicumference; and the crown-rump length, as described in Coelho
[1985]. Crown-rump length was taken with a GPM anthropometer, and triceps
circumference was taken with a GPM steel tape. Each measurement was recorded
to the nearest millimeter. Skinfold measures were taken with a Holtain skinfold
caliper. Skinfold measures were taken on the left side of the animal and recorded to
the nearest 0.1 millimeter.
Measurement of Lean Body Mass and Fat Mass
Fasted baboons were anesthetized with ketamine hydrochloride (10 m g k g body
weight) and were given pentobarbital to overdose, weighed, and exsanguinated.
Lean body mass and fat mass were measured as described in detail by Lewis et a1
[l983,1986b]. At necropsy the following tissues and organs were separately analyzed
from each baboon: 1)10 adipose depots that were sufficiently distinct anatomically
to be excised [Lewis et al, 19831; 2) tissues and organs free of adipose tissue (kidneys,
hair, blood, liver, lungs, eyes, central nervous system, spleen, and adrenals); and 3)
the remaining carcass containing skin, muscle, bones, etc. Adipose-free tissue, with
the exception of blood, was minced. Both the adipose-free tissue and the tissues
containing fat cells were ground and dried to a constant weight a t 60°C in a vacuum.
The fat content of the tissues containing fat cells and the isolated fat depots were
measured by extracting the lipids with petroleum ether (15 ml/g dry weight) in a
Sohxlet (Aldrich Chemical Co., Inc., Milwaukee, WI) apparatus for 24 hours. The
2.856
2.167
2.222
1.389
8.411
32.089
1.960
0.219
1.741
0.475
0.453
0.342
0.176
0.839
1.927
0.392
0.087
0.328
SD
*
3.050
2.092
2.200
1.324
8.175
30.667
1.685
0.236
1.450
X
-
Female
(n = 12)
0.487
0.297
0.270
0.169
0.684
1.497
0.308
0.060
0.272
SD
5.010
4.050
2.990
1.690
20.780
66.000
20.288
1.245
19.043
0.781
0.470
0.348
0.242
1.678
3.037
3.287
1.332
2.395
Male
(n = 10)
X
SD
*
*
*
*
*
18 wk
.797*
.660
.501
.796*
.827*
.97 1*
5 Yr
.554
,278
.337
.371
.790*
.883*
Female
18 wk
.285
.370
.666*
,019
.887*
.897*
Lean mass
18 wk
.417
.135
.039
.344
.906*
.661
5 Yr
550
.637*
.720*
.653*
.926*
.460
Male
*Pearson’s correlation of body compartment mass to morphometric measures significant at P < .05.
Measures
Neck
Subscapular
Suprailiac
Triceps skinfold
Triceps circ
Crown-rump
Male
X
-
6.250
4.692
3.383
2.242
18.642
59.983
15.650
2.652
12.997
5 years
.791*
.489
.644*
,388
,004
,220
5 Yr
Female
Female
(n = 12)
.703*
.560
,553
,402
,597
,430
18 wk
Fat mass
TABLE 11. Univariate Correlation (r) Matrix of Body Compartment Mass to Morphometric Measures
*Student’s t-test of male-female differences significant at P < .05.
Skinfolds (mm)
Neck
Subscapular
Suprailiac
Triceps
Triceps circ (cm)
Crown-rump (cm)
Weight (kg)
Fat weight (kg)
Lean weight (kg)
X
-
Male
(n = 9)
18 weeks
TABLE I. Bodv Measurements bv Ace and Gender
.788*
.722*
.741*
.734*
.889*
.156
5Yr
2.695
1.326
0.656
0.442
1.349
2.285
3.781
2.303
1.663
SD
Body Composition in Baboons I 279
fat-free dry tissues were then weighed and the fat content was calculated. Lean
body mass was calculated as wet weight of the baboon (adjusted by subtracting gut
and bladder contents) minus the fat mass.
Statistical Analysis
The following null hypothesis was tested: morphometric measures (either individually or in combination) are not related to absolute measures of lean body mass
or fat mass in baboons. Pearson’s correlation analysis was used to compare each of
the various morphometric measures to lean body mass and fat mass. A two-tailed
alpha of 0.05 was the criterion for rejection of the null hypothesis. Non-linear and
stepwise multiple regression equations [Dixon, 19831 were calculated to predict lean
body mass and fat mass in both male and female baboons at 18 weeks and 5 years of
age. Morphometric measures were used as independent variables. An F value of
4.000 was the minimum criterion for acceptance of variables into the multiple
regression formula, and a minimum F of 3.900 was criterion for removal.
RESULTS
Body Composition and Morphometric Values
Mean values for body weight, lean body mass, and fat mass of 18-week-old
[Lewis et al, 1986al and 5-year-old [Lewis et al, 1986bl baboons are presented in
Table I. At 18 weeks of age, males had significantly more lean body mass than
females. Body weight, fat mass, and the morphometric means were not significantly
different by gender. At 5 years of age, males had greater body weight, crown-rump
length, and lean body mass than did females. Females had larger triceps skinfolds
than males. These findings are consistent with previous studies of this species
[Coelho, 1985; Coelho et al, 19841.
Correlations of Morphometric Variables to Lean Body Mass
The Pearson’s r correlations of the individual morphometric measures with lean
body mass and fat mass are presented in Table II. Among males, crown-rump length
had the highest correlation with lean body mass at both 18 weeks and 5 years of
age. Among females, crown-rump length had the highest correlation with lean body
mass at 18 weeks of age, while triceps circumference provided the highest correlation
for females at age 5 years.
Correlations of Morphometric Variables to Fat Mass
Triceps circumference provided the highest correlation with fat mass tissue in
18-week-old males and 5-year-old females. Neck skinfolds were highly correlated
with fat mass in females at 18 weeks of age and in males at 5 years of age. Skinfold
measures for males a t 18 weeks were not significantly correlated to fat mass. Triceps
circumference was the single best indicator of fat mass in 5-year-old females. Males
and females exhibited significant correlations between the neck skinfold and fat
mass at 5 years of age. All skinfold thicknesses were significantly related to fat
mass in 5-year-old females.
Derivation of Regression Formulae for the Prediction of Lean Body
Mass and Fat Mass
Non-linear and stepwise multiple regression analyses were used to determine
whether combinations of the morphometric measures could provide greater accuracy
in the prediction of lean body mass and fat mass. Table III presents the amount of
variance explained (3)
and the best regression formulae for predicting lean and fat
mass in this sample.
280 I Rutenberg et a1
TABLE 111. Regression Formulae for Fat and Lean Tissues by Age and Gender
-2
Male fat
18 wk
5 Yr
Female fat
18 wk
5 Yr
Male lean
18 wk
5 Yr
Female lean
18 wk
5 Yr
- 1.883(tri circ) + .182(tri circ)' - .005(tri circI3 + 6.164
20.992tneck) - 4.792(neckI2 + .360(neckj3 - 29.428
,948
,957
.323(neck) + .183(tri circ) - 3.972
,704
.857
.666(tri circ) - 11.838
.185(neck) + .134(crn-rmp) - 3.102
.956(neck) + .62l(crn-rmp)- 26.705
,981
,869
.196(tri circ) + .lOl(crn-rmp) - 3.287
- .264(neck) + 1.552(tri circ) - 14.286
,903
,931
lmml
5
8 . 5 VLRR OLD M R L L S
4 .
3 .
1.1 uelgnl
ltg)
2 -
0 4
+
I
n r r t *tlnlold*
6
7
lmml
Fig. 1. Fat weight predictability.
The ability to predict fat mass in males a t 18 weeks and 5 years of age was not
increased by combining morphometric variables. However, non-linear (cubic)regression formulae increased the predictability of fat mass for males a t both ages (Table
111;Fig. lA,B). A bivariate exponential equation (Fig. 1C) provided the best estimate
of fat weight in 18-week-oldfemales. A univariate non-linear (exponential) equation
(Fig. 1D) provided the best predictor of fat mass for 5-year-old females. The triceps
circumference was included in the regression formulae of both males and females a t
18 weeks and in the formula of females a t 5 years of age. Conversely, the neck
skinfold was included in the female formula a t 18 weeks and was the only morphometric variable useful for 5-year-old males.
Body Composition in Baboons / 281
-------
Fig. 2. Lean weight predictability.
Lean body mass for males and females at 18 weeks was best predicted by
multiple regression estimators (Table III), including crown-rump length for both
genders, the neck skinfold for male predictor equations (Fig. 2A), and the triceps
circumference for female predictor equations (Fig. 2C). The neck skinfold was included in the formulae for both genders at 5 years, but only in that of males at 18
weeks (Fig. 2B). The neck skinfold was substituted for crown-rump length in the
female regression formula at 5 years of age (Fig. 2D).
In all but three instances (Table III), the 1.2 value of the regression equations
increased with additional morphometric variables. The neck skinfold and triceps
circumference were the most consistently used variables for predicting both lean
body mass and fat mass. As much as a 33%increase in the explanation of variability
(determined by increasing values of 1-2) was achieved through the use of non-linear
or multiple regression analysis.
Comparison of Predicted vs Observed Values of Lean Body Mass and Fat Mass
The predicted versus observed lean body mass values at both 18 weeks (Table
5 years of age (Table V) show an overall mean within animal difference
[percent difference = 1 - (predictedobserved)] of less than 3% (Table N).
These
results correspond to the high correlation coefficients obtained and verify that
regression !formulaeare appropriate for predicting lean body mass from morphometric variables.
The predicted versus observed fat mass values indicate greater variability than
that observed for lean body mass. However, much of the variability can be explained
by using non-linear regression formulae. Female baboons show a mean difference of
6% when predicted and observed values are compared at 5 years of age. Adolescent
male baboons show considerably more deviation of predicted values from observed
values than do male infants; almost 11% deviation at 5 years of age versus 1%
deviation a.t 18 weeks of age. Fat in females at both ages tends to increase exponen-
IV)and
282 I Rutenberg et a1
TABLE IV. Observed and Predicted Lean Body Mass and Fat Mass in 18-week-old Baboons
Animal
Males
M1
M2
M3
M4
M5
M6
M7
M8
M9
Mean percent
difference
SD
Females
F1
F2
F3
F4
F5
F6
F7
F8
F9
F10
F11
F12
Mean percent
difference
SD
'Percent difference
Lean mass (kg)
Percent
Predicted
Observed differencea
1.98
1.77
1.90
2.08
1.21
1.41
2.14
1.48
1.58
2.02
1.82
1.92
2.05
1.25
1.45
2.17
1.40
1.59
01.9
03.2
00.7
-01.4
03.5
02.7
01.5
-06.1
-00.7
00.8
Fat mass (kg)
Predicted
Observed
0.28
0.38
0.21
0.26
0.14
0.14
0.26
0.15
0.15
0.29
0.38
0.20
0.30
0.15
0.16
0.24
0.15
0.12
1.79
1.94
1.62
1.22
1.18
1.19
1.20
1.30
1.35
1.26
1.70
1.66
06.5
04.7
-01.2
05.1
-05.4
11.0
16.4
-06.0
-08.9
05.1
02.1
05.2
02.9
0.24
0.32
0.26
0.28
0.20
0.17
0.16
0.22
0.21
0.23
0.26
0.23
07.3
=
01.4
-00.8
-02.0
11.1
05.4
13.3
-11.4
-04.1
-22.9
-01.1
11.2
03.0
1.67
1.85
1.64
1.21
1.24
1.06
1.00
1.38
1.47
1.19
1.66
1.57
Percent
differencea
0.25
0.32
0.33
0.34
0.19
0.18
0.18
0.21
0.18
0.19
0.23
0.22
03.9
-00.9
19.8
17.6
-05.8
08.8
10.5
-04.8
-18.4
-21.2
-11.2
-04.9
-00.6
13.1
1 - (predicted value/observed value) x 100.
tially relative to the morphometric measures. Conversely, fat in males at 5 years of
age shows a linear trend for the smaller fat weights but is affected by a single
outlier. It cannot be demonstrated from these data whether the outlier is a deviant
value, or if the outlier truly represents the overall trend of the relationships between
male fat weight and morphometric measures at 5 years of age. However, polynomial
regression at both 18 weeks and 5 years of age substantially improves the predictability of male fat weight over simple linear or multiple regression estimators (including the outlier a t 5 years of age) and explains much of the variance attributable to
other extraneous variables.
DISCUSSION
Evaluated in this study was the ability of a select number of morphometric
measurements to predict lean body mass and fat mass in nonhuman primates at two
distinct age periods, infancy and adolescence. Selection of these age periods was
based on previous findings by Coelho et a1 [1984],which documented three patterns
of growth in the baboon: 1)a period of continuous growth to the end of puberty for
measures of body size; 2) continuous growth to the onset of puberty, followed by a
Body Composition in Baboons I 283
TABLE V. Observed and Predicted Lean Body Mass and Fat Mass in 5-year-old Baboons
Animal
Males
M10
M11
M12
M13
M14
M15
M16
M17
M18
M19
Mean percent
difference
SD
Fema1es
F13
F14
F15
F16
F17
F18
F19
F20
F2 1
F22
F23
F24
Mean percent
difference
SD
Lean mass (kg)
Percent
Predicted Observed difference*
22.16
17.28
17.49
16.07
17.81
16.86
19.31
21.47
20.38
21.62
21.62
18.12
16.18
17.46
17.65
15.95
18.61
22.08
20.53
22.24
Fat mass (kg)
Predicted
04.6
-03.8
-08.1
-05.7
08.0
-02.5
00.7
-00.9
02.8
02.8
-00.2
0.76
0.88
0.78
0.86
0.86
0.74
0.76
0.89
0.97
4.95
Percent
Observed-differencea
0.96
0.45
0.60
0.85
1.42
0.64
0.57
0.80
1.23
4.94
05.0
10.54
12.12
12.18
13.65
13.37
10.24
11.86
14.05
13.90
15.17
15.01
13.85
10.16
12.22
11.75
13.05
14.08
10.87
11.78
14.43
13.58
15.23
15.33
13.48
-03.7
00.9
-03.7
-04.6
05.0
05.8
-00.7
02.6
-02.4
00.4
02.1
-02.7
-00.1
20.7
-97.5
-29.7
-00.8
39.3
- 15.3
-33.7
-11.1
20.8
-00.2
- 10.8
38.2
0.56
1.16
0.89
1.98
1.62
0.43
0.95
3.16
2.77
4.41
8.04
4.41
03.5
0.39
1.31
0.61
1.81
0.98
0.76
0.97
4.68
5.01
4.44
7.55
3.32
-45.1
11.2
-46.5
-09.4
-65.8
43.4
01.4
32.4
44.8
00.6
-06.5
-33.0
-06.0
36.2
aPercent difference = 1 - (predicted value/observed value) x 100.
growth spurt and a plateau of growth at the end of puberty for the neck and
suprailiac skinfolds; and 3) rapid growth to the beginning of adolescence, followed
by a slowed growth rate through adolescence, including puberty, for the triceps and
subscapular skinfolds. Both of the age periods chosen for this study contained
individuals exhibiting a large amount of variability in relative and absolute fat and
lean mass. Large variability in these measurements provided a major challenge for
accurate prediction with indirect methods.
This study indicates that morphometric measures are viable estimators of lean
mass in nonhuman primates. However, the relationship between the lean and fat
body compartments and morphometric measures is complicated and cannot be accurately predicted by simple linear regression formulae. When non-linear and multiple
regression estimators are derived, the accuracy of the predicted values for estimating lean body mass and fat mass increases considerably. Combinations of these
measurements a t weaning and at puberty explain between 70% and almost 100%of
any variability in lean body mass and fat mass for this sample. The skinfold
284 I Rutenberg et al
measures of subcutaneous fat, while consistent for most of the older animals, vary
occasionally and render the adult predictor equations for fat mass less reliable than
those for lean mass.
Moreover, our results demonstrate that these morphometric predictors change
as the animals develop. The amount of subcutaneous fat, indicated by skinfold
thickness, in young baboons is positively and linearly related to other measures of
lean body mass. However, at 5 years of age this relationship is very different. There
are several reasons for the changing relationship of morphometric measures to lean
body mass. The 5-year-old baboon is undergoing or has passed through the pubertal
growth spurt. As baboons reach puberty, they are like humans in that females tend
to accumulate body fat while males tend to develop muscle mass.
In human studies, morphometry is a n important method of estimating body
composition because it provides a n easy, inexpensive, noninvasive method of doing
serial evaluations of the development of body composition. For example, Jellife and
Jellife [1969] have demonstrated that triceps circumference can be used as a rapid
means of identifying malnutrition in infants. The circumference is a reflection of
lean and fat mass development. In nonhuman primates, morphometric measures
are important for the same reasons. Further, they provide for noninvasive serial
evaluations of the development of body composition in subjects that can be exposed
to a variety of experimental treatments. Morphometric techniques for estimating
fat and lean body mass in live baboons are important because of the use of baboons
as animal models for humans in studies of obesity and risk factors for diseases. The
morphometric methods used in this study to estimate body composition may be used
for other nonhuman primate species; however, the potential of species differences in
fat deposition necessitates validation of these measures in each species.
CONCLUSIONS
1)Morphometric measures can accurately predict lean body mass in a sample of
male and female baboons.
2) Morphometric measures used to predict lean body mass change with age.
3) Morphometric measures are strongly associated with body fat mass at 18
weeks of age, but these predictors are not as strongly associated with body fat mass
in 5-year-old baboons.
4) Triceps circumference provides the best single indicator of lean body mass for
both genders and age periods.
5) Baboons are like humans in that adolescent females tend to accumulate body
fat, while males of the same age tend to develop lean mass.
6) Combinations of these morphometric measurements explain between 70%
and 100% of the variability and may be used to estimate lean and fat mass in
baboons.
ACKNOWLEDGMENTS
This research was supported by National Institutes of Health, National Heart,
Lung and Blood Institutes, and grants HL28728, HL28972, and HL19362, and a n
Andrew G. Cowles Research Fellowship.
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