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FIGURE 2. Ratio of sedentary energy expenditure to basal metabolic
rate (SDE:BMR) versus age.
Infants
(ii 101)
Children
(n = 82)
Adults
(n 27)
Weight (kg) 7 ± 2 47 ± 12 64 ± 13
Height (cm) 66 ± 9 154 ± 11 164 ± 9
FFM(kg) 5±2 36±8 46±9
FM(%bodywt) 26±4 23±6 28±8
computed at 1-mm intervals from oxygen consumption andcarbon dioxide production according to Weir (7).
While in the calorimeter, subjects adhered to scheduledfeeding and sleeping times. Three meals were served withdietary energy adjusted to body weight. Subjects were asked toretire at 2200 or 2300 and were awakened at 0700, at which
time BMR was measured for 40 mm (12 h after the last meal).
To ensure that the subject remained awake, but still, heart rate
was monitored by telemetry (Dynascope 3300; Fukuda Denshi
America, Redmond, WA) and physical movement by a Dopp-
ler microwave sensor (D9/50; Microwave Sensors, Ann Arbor,MI). SDE was defined as 24-h total energy expenditure exclu-
sive of exercise.Because infants are seldom quiet while awake, SMR was
used as a proxy for BMR. We measured SMR of infants by
using a whole-body respiration chamber or a hood apparatus
for 81 ± 34 mm (8). Total energy expenditure estimated by the
doubly labeled water method was available for a subset of 40
infants at 1 and 4 mo of age (9). The total energy expenditure
of infants meets our definition of SDE because infants engagein spontaneous physical activity but not in intentional exercise.
Weights were measured on electronic balances. Heights andlengths were measured using an upright stadiometer and re-
cumbent infant board, respectively. Fat-free mass (FFM) and
fat mass (FM) were estimated from total-body electrical con-
ductivity (TOBEC, model HA-2 for adults and children and
model HP-i or HP-2 for infants; DICKEY-john, Auburn, IL).
Statistical methods
Linear regression was used with and without log transfor-
mation to determine the best model relating SDE and BMR to
weight or FFM (10). Multiple regression was used to test for
independent effects of other factors on SDE and BMR. A t test
of homogeneity was used to test for differences between
slopes.
RESULTS
SDE (kJ/d) and BMR (kJ/d) of all subjects are shown in
Figure 1. Our BMR values were 105 ± 14% of values pre-
dicted from weight and height according to Schofield (3). CVs
were 23%, 15%, and 22% for SDE, and 37%, 14%, and 17%
for BMR for infants, children, and adults, respectively. Therelatively high CVs for SDE and BMR of infants are due to the
compilation of all infants with a wide range of body weights.The CVs for BMR were 12%, 12%, 12%, 9%, and 10%,
respectively, for 3-, 6-, 9-, 12-, and 18-mo-old infants. The
ratio of SDE to BMR was 1.32 ± 0.13 for all subjects, 1.31 ±
0.14 for infants, 1.33 ± 0.12 for children, and 1.31 ± 0.14 for
Age (y)
FIGURE 1. Sedentary energy expenditure (E) and basal metabolic rate
(#{149})versus age.
adults (Figure 2). Simple correlations of SDE, BMR, and the
physical characteristics are shown in Table 2.
Before investigating the independent effects of age and bodycomposition on SDE and BMR, we investigated three models
(a constant-ratio model, a linear-regression model, and a pow-
er-function model) to remove the effect of body size on SDE
and BMR (11).
A constant-ratio model (Y1 = a X X) was applied to all
subjects (Eq 1 and Eq 2) and was found to underestimate EE in
the lower weight range (Figure 3).
SDE (kJId) = 138 X WT(kg) SEE = 1234 R2 = 0.79 (1)
BMR (kJ/d) = 106 X WT(kg) SEE = 1057 R2 = 0.72 (2)
The constant-ratio model failed to produce a variable indepen-dent of dimension, ie, the ratio of SDE to WT and of BMR to
WT were negatively correlated with weight (r = -0.92 and
-0.90, P = 0.001) (Figure 4).The linear-regression model (Y a + b X X.) of SDE or
BMR on weight (Eq 3 and Eq 4) was an improvement over the
constant-ratio model (Figure 5). Inspection of the residual
FIGURE 5. Linear-regression model of sedentary energy expenditure
(LI) and basal metabolic rate ( #{149}) versus weight.
adults. To test for the independent effects of age, height, and
body fatness (kg or % body wt) within the age-specific groups,
two approaches yielding similar results were taken: SDE orBMR was scaled by weight or FFM, and weight or FFM was
treated as a covariate. For adults, only weight and FFM con-
tributed significantly to the prediction of SDE (R2 = 0.77) and
BMR (R2 = 0.60). For children, age, weight, FFM, and FM (kgand % body wt) contributed to SDE (R2 = 0.64) and BMR(R2 0.60). For infants, age, weight, and FFM contributed tothe prediction of SDE (R2 = 0.76), and age, weight, FFM, and
FM (kg and % body wt) contributed to the prediction of BMR
(R2 0.92).
DISCUSSION
To evaluate the independent effects of variables on energyexpenditure, measurements must be scaled to a base to account
for differences in body size. In our sample, weight (R2 =
0.88-0.90) and FFM (R2 = 0.92-0.93) were interchangeably
the best predictors of SDE and BMR. Weight and FFM also
were intercorrebated with other factors of interest. To investi-
800
600 �I :1 k
400 � �
200
0
Age (y)
FIGURE 7. Power function ratios (SDE:WT�7#{176}, kJ:kg#{176}7#{176},LI: and
BMR:WT#{176}�’3, kJ:kg#{176}�3, #{149}) versus age.
gate the effect of other modulating factors on SDE or BMR, we
tested the ability of three models to remove the effect of body
size. We encountered substantial errors with the constant-ratio
and linear-regression models, as described by Tanner (12) andKatch (13). The constant-ratio model overscaled SDE andBMR, creating a negative correlation between these parameters
and body size. The power-function model fit our data well and
effectively eliminated the dimension of body size.
Application of a power-function model to energy expendituredata is not novel. In 1883 Rubner first recognized in dogs that
metabolism was a constant function of body surface area, com-
puted as the 0.67 power of body weight (14). Rubner’s student,
Voit, continued these studies across species and confirmed that
fasting metabolism was constant at = 1000 kcal . m2 � d �. Rub-ner’s body surface area law became ingrained in the field of
energy metabolism until challenged by Kleiber (15) in 1932.
Kbeiber contended that basal metabolism across species was pro-
portional to the 0.75, not the 0.67, power of body weight. Al-
though useful for cross-species comparisons, not all species con-
form to the Kleiber generalization (16).
0 10 20 30 40 50 60 70 80 90
Weight (kg)
FIGURE 6. Power function ratios (SDE:W1�7#{176}, kJ:kg#{176}7#{176},LI; and
BMR:W14163, kJ:kg#{176}�’3, #{149}) regressed against weight.
0 10 20 30 40 50
Age(y)
FIGURE 8. Power function ratios (SDE:FFM�7#{176}. kJ:kg#{176}7#{176},LI: and
BMR:WT#{176}63, kJ:kg#{176}�’3, #{149}) versus age.
Across all ages, SDE was standardized by the 0.70 power of
WT or FFM, and BMR was scaled by the 0.63 power of WT or
FFM. Inspection of Figure 8 shows that SDE and BMR per unit
body size increased during infancy, declined throughout child-
hood, and leveled off in adulthood.
Several investigators have noted that the BMR per unit bodyweight during growth differed from adult values. The greater
proportion of metabolically active organs in infants compared
with children and adults contributes to their higher metabolic
rates. The proportion of organ weight is fairly constant at 15%body weight throughout infancy, and organ metabolic rate
remains constant from infancy to maturity (17). Holliday et ab
(17) found that the best-fit exponent relating weight and BMR
of infants up to 10-12 kg was 1.0; beyond 12 kg, the exponentdecreased to 0.58. Karlberg (18) and others confirmed that
BMR of infants was more or less proportional to body weight;
the exponents of body weight reported were 0.918 (18), 1.09
(19), 1.1 1 (20), and 1.20 (21). Metabolic rate increases more
rapidly than body size, probably due to relative losses of
extraceblular fluid and increases in body cell mass (21). These
compositional changes in FFM contribute to the increased
metabolic rate of FFM seen during infancy.
The decline in SDE and BMR relative to weight or FFM seenduring childhood is due to the slower growth of organs with
high metabolic rates (eg, brain, liver, heart, and kidney) relative
to those with lower metabolic rates (eg, muscle, bone, and fat).
Organ weight is 8% of body weight in adolescents and 6% inadults (22). Muscle and skeletal mass increase steadily
throughout childhood. The percentage of body fat declines in
the midchildhood years, and then increases during the adobes-
cent growth spurt in girls (23).
Factors underlying the variability in SDE and BMR havebeen investigated in several studies of adults. In 177 adults,
Ravussin et al (4) found that FFM was the single best predictor
of SDE; weight, height, % FM, but not age and sex, were alsoindependent predictors of SDE. BMR was related to FFM, but
not to FM. The ratio of SDE to BMR was 1.27 ± 0.12, similar
to the mean value and variance observed in our room calorim-
eters. In 37 men and women, Webb and Sangal (5) reported
that the best prediction equation for SDE combined FFM and
body mass index (R2 = 0.92). FM also improved the prediction
of SDE in this study.In a compilation of adult studies, FFM was the best predictor
of resting metabolic rate (RMR), explaining 53-88% of itsvariability (24). FM made a significant contribution to the
prediction of RMR in obese individuals. The contribution of
FM to RMR was not supported for the general population
because of the relative constancy of FM in nonobese individ-
uabs. It is of interest that FM as a predictor of BMR and SDE
emerged strongest in our data set of children. These girls were
not obese, but were entering or were in their adolescent growth
spurt, a time of considerable fat deposition.
Weinsier et al (22) compiled 3 1 data sets (ii 1 1 1 1) com-
prising a wide range of ages from infants to adults and found
the relation between RMR and FFM to be nonlinear. RMR felldisproportionately to the rise in FFM. The slopes relating RMRand FFM were significantly higher for infants than for adoles-
cents and adults due to the increasing proportion of less met-
abolicalby active muscle mass.
In children aged 6-10 y, FFM was identified as the strongest
independent predictor of RMR (25). FFM predicted 74% of thevariability of TEE and 88% of the variability in BMR in young
children (26, 27). In nonobese and obese adolescents aged
12-18 y, FFM accounted for 85% ofthe variability in TEE and
87% of the variability in BMR (28). Heretofore, SDE values
for children had not been published.The standardized measurement of basal metabolism was
developed in an effort to compare metabolic rates rather than to
establish minimal metabolism within and between species.
Standardization ensured comparability, but also diminishedobservable biological variability between individuals. In themeasurement of SDE, factors known to influence metabolismsuch as cold- and food-induced thermogenesis, muscular
movement, and sleep are included, which therefore makes the
measurement more encompassing of individual variability. In
our study, the ratio of SDE to BMR averaged 1.3 from infancyto adulthood. We found that factors affecting BMR similarlyinfluenced SDE. FFM and weight were the strongest determi-nants of SDE and BMR. However, the relation between SDE or
BMR and weight or FFM was a function of age. Other mod-
ulating factors were age-specific. In adults, age, height, and
body fatness did not contribute independently to the prediction
of SDE or BMR. In infants and children, age and FM were
identified as independent predictors of SDE and BMR. Be-
cause the relation between SDE or BMR and weight or FFM
was found to be a function of age, separate predictive equations
for SDE and BMR should be developed for infants, children,
and adults. U
We thank A Adolph, M Puyau, and F Vohra for performing the energy
expenditure measurements; L Clarke and M Llaurador for performing the
mass spectrometric measurements; and C Heinz, K Wallace, and J Stuff for
technical assistance.
REFERENCES
1. FAO/WHO/UNU. Energy and protein requirements: report of a joint
FAO/WHO/UNU expert consultation. World Health Organ Tech Rep
Ser l985;724.2. National Research Council. Recommended dietary allowances. 10th
ed. Washington, DC: National Academy Press, 1989.
3. Schofield WN. Predicting basal metabolic rate, new standards and
review of previous work. Human Nutr Clin Nutr 1985;39C:5-41.
4. Ravussin E, Lilloja 5, Anderson TE, Christin L, Bogardus C. Deter-