Original Research Communications-general

A reappraisal of caloric requirements

in healthy women13
Oliver E Owen, MD, Edward Kavle, BS, Rodney S Owen, BS, Marcia Polansky, ScD,
Sonia Caprio, MD, Maria A Mozzoli, BS, Zebulon V Kendrick, PhD,
Marcy C Bushman, RD, and Guenther Boden, MD

ABSTRACT The caloric expenditure of 44 healthy, lean and obese women, 8 of whom were
trained athletes, was measured by indirect calorimetry. Body composition was determined. Ages
ranged from 18-65 yr and body weights from 43-143 kg. Stepwise, multiple-regression analysis was
used to determine whether one or several variables best predicted the resting metabolic rate (RMR)
of the women. The RMR and the thermic effect of food (TEF) were measured before and after the
women consumed a mixed breakfast meal.
The results showed that the currently available tables and regression equations overestimate the
RMR of healthy women by 7-14%. Body weight was highly related to the RMR, and stepwise
inclusions of various variables did not improve predictions of RMR. The slopes of the regression
lines for nonathletes and athletes were significantly different. Regression equations for predicting
RMR of women were developed: Nonathletes RMR = 795 + 7.18 kg WT; Athletes RMR = 50.4
+ 21.1 kg WT. The range of RMR per kilogram body weight was wide for nonathletic, but narrow
for athletic women. The metabolism of some lean and obese, nonathletic women was highly efficient,
predisposing these women for developing and maintaining body fat. The TEFs were indistinguishable
between nonathletic and athletic women, and formed a continuum from the lightest to the heaviest
woman. AmJClinNutr l986;44:l-l9.

KEY WORDS Resting metabolic rate, thermic effect of food, caloric expenditure, indirect cal-
orimetry, body composition

Introduction The RMR for normal humans are based on

three large and authoritative studies done dur-
Daily caloric requirements (thermogenesis) ing the first half of this century (4-6). In 1919,
are dictated by several states of metabolism. Harris and Benedict (4) published their classic
The lowest or basal metabolic rate occurs dur- monograph on RMR of males and females.
ing the early morning hours of deep sleep (1) They developed regression formulas for pre-
and is impractical to measure. On the other
hand, the resting metabolic rate (RMR), de-
termined during the morning after an over- ‘From the Departments of Medicine and the General
night fast, can be conveniently measured. Clinical Research Center, Temple University School of
Medicine and the Departments of Statistics and HPERD,
RMR is the best predictor of overall caloric
Temple University, Philadelphia. PA.
requirements and accounts for about 65-70% 2Suppo in part by US Public Health Service Grants
of the daily energy expenditure. The post- AM 25386 and RR 00349, General Clinical Research
prandial state is associated with augmented Centers Branch, and National Institutes of Health.
energy expenditure; the thermic effects of food 3Address reprint requests to: Oliver E Owen, MD,
Temple University Hospital, 3401 N Broad Street, Phil-
(TEF) account for about 10-15% (2), and adelphia, PA 19140.
physical activity usually accounts for about Received July 10, 1985.
20-30% of the daily caloric expenditure (3). Accepted for publication November 26, 1985.

The American Journal of Clinical Nutrition 44: JULY 1986, pp 1-19. Printed in USA
© 1986 American Society for Clinical Nutrition

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 2 OWEN ET AL

dicting the RMR based on height, weight, age, TEF is dependent upon the nature (23) and
and sex. DuBois and DuBois (5) measured the the quantity (20) of the food ingested, infor-
body surface area (BSA) of humans and de- mation pertaining to the role of TEF in the
rived a formula for calculating BSA based on development or maintenance of obesity should
height and weight. Boothby and colleagues at be obtained by keeping the proportions of car-
the Mayo Foundation (6) studied normal men bohydrate, fat, and protein in a meal constant
and women to develop a standard for basal while giving adequate caloric challenges to
metabolism after having reported in 1922 that obese as well as to lean individuals by adjusting
the RMR could best be predicted from BSA the caloric value of the meal to the body
(7). Between 1930 and 1950, Robertson and weights and by determining how much of the
Reid (8) derived regression formulas for pre- food is oxidized or stored.
dicting RMR based on sex, age, and BSA. This report reappraises the caloric require-
Although additional early studies pertain to ments of healthy lean and obese women, some
RMR, values in tables widely used for pre- of whom were trained athletes, at rest and fol-
dicting RMR are smoothed means derived lowing breakfast. It shows that currently avail-
from those three large studies and are ex- able tables and formulas for healthy women
pressed as calories per square meter of BSA overestimate their RMR and that the TEF are
per hour or day for men and women of given not diminished by obesity.
ages (9).
Studies have shown that the Harris and
Materials and methods
Benedict equations overestimate the measured
RMR in morbidly obese humans (10) and in Subjects
malnourished patients (11). However, when Forty-four lean and obese healthy females were ad-
calculations were made to adjust for differ- mitted to the General Clinical Research Center (GCRC)
ences in lean body mass, the active proto- of Temple University Hospital. The nature and purpose
plasmic tissue that is thought to dictate RMR, of the study were explained to the subjects prior to ob-
taining their written consent. The study was approved by
sex, age, and nutritional status (11, 12) seemed the Institutional Review Board of Temple University
to have little direct influence on RMR. These Hospital. Eight of these 44 women were trained athletes.
conclusions were based on a reanalysis of the All subjects were eating balanced diets, had stable weights
Harris-Benedict subjects and from studies of for at least one month prior to the study, were not taking
any medications, and had no evidence of any physical
obese (10) or ill (11) humans. However, such disease. All subjects had normal body temperatures and
studies lacked their own normal control pop- were not menstruating during the study periods. They all
ulation. This becomes important because a had normal fasting blood-glucose concentrations and nor-
recent preliminary report noted discrepancies mal thyroid function.
The clinical and body compositional data for the 44
between the measured RMR and the predicted
women are presented in Table 1. Ages ranged from 18-
RMR, as calculated from the Harris-Benedict 65 yr. height ranged from 150-180 cm; weight ranged from
equation (13). 43-143 kg. Formulas used to make calculations are listed
TEF is another component of energy ex- in the appendix in numerical order. Theoretical consid-
penditure that has been frequently and more erations of indirect calorimetry, formulas for calculating
oxidation and storage rates, and regression equations are
recently studied. Some investigative groups (3,
also given in the appendix. Body mass index (BMI) (24)
14-16) have reported that the rise in metabolic and BSA (5) were calculated from equations 1 and 2, re-
rate that follows food ingestion is reduced in spectively.
obese humans, suggesting that obese people
Experimental design
have an increased metabolic efficiency that re-
sults in augmented storage of nutrients in body Forty-four subjects underwent studies to determine their
RMR. Thirty-two of these 44 subjects underwent addi-
fat. Other researchers have failed to show a tional studies to measure their TEF following a mixed
difference in the TEF between lean and obese breakfast meal. Twenty-eight of these 32 subjects under-
humans (17-19), and Hill et al (20) showed went more studies to determine their body composition.
that TEF increased as the caloric content of Body composition was determined by densitometric
the meal increased. Many of the early studies analyses. Fat mass (FATMD) (25) and fat-free mass
(FFMD) (26) based on densitometry were calculated from
used too small a caloric challenge to properly formulas 3 and 4, respectively. Fat mass (FATMSF) and
assess the differences, if any, in the TEF be- fat-free mass (FFMSF) based on skinfold thicknesses were
tween lean and obese humans (21, 22). Since calculated from formulas 5 and 6, respectively (27). Es-

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 3

TABLE 1
Clinical measurements and body composition

Patient
number A HT wT SSA BMI FFMD FATMD FFMSF FATMSF IBM SCM

yr cm kg m kghn’ kg kg kg kg kg kg

1 25 154 43.1 1.37 18.2 34.3 8.5 35.0 8.1 32.7 16.4
2 23 157 48.1 1.46 19.5 - - - - 35.8 18.0
3 33 156 49.9 1.47 20.5 42.4 7.2 41.0 8.9 36.0 17.9
4* 23 158 50.8 1.50 20.3 41.9 8.9 40.1 10.7 37.4 18.8
5* 31 160 51.8 1.52 20.2 44.7 5.4 45.7 6.1 37.2 18.5
6 27 158 52.2 1.51 20.9 39.3 13.3 38.5 13.7 37.8 18.9
7* 27 168 52.6 1.59 20.7 43.4 8.4 42.7 9.9 38.0 19.0
8 27 162 53.5 1.56 20.4 42.5 12.2 40.2 13.3 38.5 19.2
9 46 163 55.0 1.58 20.7 44.8 10.2 42.3 12.7 37.6 18.3
10 27 165 55.0 1.60 20.2 44.4 10.3 44.8 10.2 39.3 19.6
11 32 163 56.0 1.60 21.1 43.7 14.2 41.7 14.3 39.3 19.5
12* 25 169 57.6 1.66 20.1 48.5 9.1 46.4 11.2 40.8 20.4
13 35 153 57.8 1.55 24.7 41.0 17.2 40.6 17.2 39.9 19.8
14 35 161 59.0 1.62 22.8 47.4 11.6 45.9 13.1 40.5 20.1
15 45 173 59.0 1.70 19.7 46.3 12.8 44.0 15.0 39.5 19.3
16 25 160 59.2 1.61 23.1 - - - - 41.6 20.8
17 26 165 59.5 1.65 21.9 - - - - 41.6 20.8
18 32 168 60.0 1.68 21.3 47.3 12.8 46.6 13.4 41.3 20.5
19* 42 171 61.4 1.72 21.0 49.9 10.8 52.1 9.3 40.9 20.1
20 20 170 62.1 1.72 21.5 49.4 13.4 47.1 15.0 43.5 21.9
21* 18 166 62.3 1.69 22.6 51.2 10.7 50.2 12.1 43.8 22.1
22* 25 170 64.9 1.75 22.5 58.7 5.7 - - 44.3 22.2
23 23 164 64.9 1.71 24.1 - - - - 44.5 22.3
24 42 150 66.0 1.62 29.3 42.2 25.6 41.2 24.8 42.9 21.1
25 32 170 66.4 1.77 23.0 46.2 18.3 46.8 19.6 44.2 21.9
26 23 168 72.8 1.82 25.8 - - - - 47.8 24.0
27 59 163 74.8 1.81 28.2 - - - - 44.2 21.2
28 51 159 77.1 1.80 30.5 - - - - 46.0 22.3
29 56 169 82.0 1.93 28.7 49.0 34.7 46.9 35. 1 46.8 22.5
30 38 158 82.0 1.84 32.8 57.8 29.1 - - 49.2 24.3
31t 23 165 83.4 1.91 30.6 51.3 32.1 52.5 30.9 51.7 26.0
32 52 155 86.2 1.85 35.9 - - - - 48.5 23.5
33 55 166 88.6 1.97 32.2 52.9 35.3 49.3 39.3 48.6 23.5
34 55 158 89.3 1.90 35.8 50.1 38.1 49.3 40.0 48.8 23.5
35 65 166 91.6 1.99 33.2 - - - - 47.8 22.8
36 24 156 99.8 1.98 41.0 - - - - 55.9 28.0
37 42 160 103.0 2.04 40.2 59.8 41.5 - - 53.4 26.2
38t 35 152 104.5 1.98 45.2 47.1 57.7 51.9 52.6 54.9 27.2
39 33 180 107.7 2.27 33.2 - - - - 55.7 27.6
40 21 169 110.2 2.19 38.6 60.6 49.8 64.1 46. 1 58. 1 29.3
41 51 178 122.0 2.37 38.5 - - - - 53.3 25.8
42 39 165 123.1 2.25 45.2 - - - - 55.7 27.4
43t 39 174 125.2 2.36 41.4 74.7 50.1 68.1 57.1 55.8 27.4
44t 25 170 143.3 2.45 49.6 67.3 76.5 76.9 66.4 57.8 29.0

Mean 35 164 74.9 1.79 27.8 49.0 22.3 47.6 22.4 45.0 22.3
SD 12.2 6.8 24.6 .27 8.6 8.5 17.9 9.1 16.7 6.8 3.4

Range
Mm 18 150 43.1 1.37 18.2 34.3 5.4 35.0 6.1 32.7 16.4
Max 65 180 143.3
________________________________________________2.45 49.6 74.7 76.5 76.9 66.4 58.2 29.3
* Athlete.

t Thermogenic effect of food measured for 360 mm (6 h).

timates of lean body mass (LBM) were calculated by rear- Indirect calorimetry
ranging the expressions of Moore ci al (28) as shown in
formula 7. Estimates of body cell mass (BCM) were cal- In the morning, after a 12-h overnight fast, all 44 sub-
culated from formula 8 (29). jects were brought to the GCRC dressed in very light gowns

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 4 OWEN ET AL

(130-190 g) or clothes (430-650 g) and barefoot heights pressed in calories. The caloric values for the storage of
and weights were determined.4 They were familiarized with carbohydrate, fat, and protein were 4.1, 4.1, and 9.3
the indirect calorimetric equipment and fitted with a kcal/g, respectively (31).
mouthpiece and nose clip or an air-sealed face mask. In To assess the influence of body composition on RMR,
those subjects undergoing TEF studies, a peripheral upper TEF, and storage, the women were subdivided into lean,
extremity vein was cannulated and kept patent with a slow obese, and athletes. Obesity was defined by a BMI of greater
drip of 0.9% saline. After the physical measurements, in- than 30 kg/m2 (24). All athletes competed in strenuous
structions, fittings, and cannulations were completed, all physical events and two were Olympic participants. Their
subjects urinated and then lay in a semisupine position at maximal 02 consumption ranged from 52-62 ml#{149} kg- mm
complete bedrest for at least 30 mm before (and for 240- (n = 6). They were studied in an attempt to dissociate
360 mm throughout) the resting studies were done in a active protoplasmic tissue mass (fat-free mass, lean-body
quiet room in which the temperature was 22-24#{176}Cand mass, body-cell mass) from relatively inert triglyceride
the humidity was about 33%. stores (fat mass).
Respiratory gaseous exchange rates were measured with Ten lean and obese nonathletes underwent randomly
a Beckman Metabolic Cart (Beckman Instruments, mc, assigned studies to compare the results for measuring 02
Schiller Park, IL). The operations aspects of this machine consumption and CO2 production rates between the
have been explained by Foss et al (30). The machine was mouthpiece-nosecip technique and the ventilated-hood
calibrated before and during each study by using stan- system. Each volunteer underwent an average of two pe-
dardized gases and a 1-1 syringe. Subjects breathed through nods of 10, 1-mm measurements for both the mouthpiece-
the mouthpiece or face mask for approximately 10 mm noseclip and the ventilated hood. The mean and the stan-
for each measurement period. The first 4-5 mm were ex- dard error of the differences were computed, and a paired
pended to clear the machine of room air. The last 5-6 I test was used to determine whether the results from the
mm were used for recording and calculating 02 consump- mouthpiece-noseclip method were significantly different
tion, CO2 production, and RQ. During these periods, 02 from the results of the ventilated-hood technique.
and CO2 concentrations were analyzed continuously. Each The reproducibility of our results are of special impor-
value was derived from the mean of five or six 1-mm values. tance for this report. Therefore, the RMRs of eight lean
The built-in computer corrected 02 consumption and CO2 or obese nonathletic volunteers were determined on five
production to standard temperature, pressure, and mois- consecutive mornings after an overnight 12-h fast, Monday
ture. Two or three baseline values were determined at 15- through Friday (forty separate studies were done). Subjects
mm intervals on all subjects. Urinary-nitrogen excretion breathed through the mouthpiece for 10 mm for two or
was determined during all the RMR studies. three measurement periods. The first few minutes were
The measured RMR extrapolated to 24 h for the 44 expended to clear the machine of room air and the last
women (Table 2) were contrasted with their predicted 5-6 mm were used to measure 1-mm respiratory gaseous
RMR/24 h based on this study, and based on the Harris- exchange rates; urinary-nitrogen excretion rates were
Benedict (4), Mayo Foundation (6), Robertson-Reid (8), measured during these baseline studies.
and Cunningham (12) equations (Formulas 9-13, respec- After an overnight 12-h fast, each of eight subjects was
tively). studied for 8-9 h on a randomly selected day between
In the 32 women who underwent measurement of the Monday and Friday to evaluate possible changes in their
TEF, additional respiratory gaseous exchange rates were RMR from 0800 h to 1600-1700 h. Respiratory gaseous
made every 30 mm for 4-6 h after consuming a mixed exchange rates were determined at 15-mm intervals for
breakfast (11 kcal/kg; carbohydrate 43%, fat 42%, and the first 1#{188} h and at 30-mm intervals thereafter. Urinary-
protein 15%) eaten during a 10-mm period. In these sub- nitrogen excretion rates were determined at 2-h intervals.
jects, urinary-nitrogen and glucose-excretion rates were This was done to include a randomized control day in
determined before breakfast and at 120,240, and 360 mm which no food was given. Maintenance caloric require-
after breakfast was started. Subjects were at complete rest ments were given after this study day was completed. Two-
when not eating or voiding. Blood was drawn before and way analysis of variance (with factors for subject and time)
at the end of the study to measure plasma glucose, total was used to evaluate changes in RMR with time.
lipids, a-amino nitrogen, and urea nitrogen.
The amounts of carbohydrate, fat, and protein oxidized Data and statistical analyses
during the breakfast tests were calculated by determining
the areas under the carbohydrate, fat, and protein oxidation The relationships among RMR and WT, AGE, HT,
curves of each subject. Intracellular glucose, fat, and protein BSA, and measured or calculated body components were
(amino acids) storage was calculated by formula 14. The assessed using Pearson’s coefficient of correlation, simple
amount of glucose, fat, or protein in the extracellular space regressions, and stepwise multiple regressions (32). The
was calculated from the difference between the plasma third method was used to determine whether one or several
glucose, total lipids, a-amino nitrogen, and urea nitrogen of the variables in combination could be used for the best
concentrations at 0, 240, and 360 mm multiplied by the predictors of the RMR of our subjects.
volume of the extracellular space (29). Storage was cx- The lean, obese, and athletic women were analyzed
separately and in combinations to determine the effects
of body composition on RMR, TEF, and caloric storage.
Subjects were inconsistent about being weighed in the The derived regression lines were evaluated by the t test,
nude. Therefore, all subjects were lightly covered but cor- assuming unequal variances about the regression lines.
rections for these coverings were not made. The weight of TEF was analyzed by computing resistant-regression
the garments should have had an influence on results of lines (Minitab Statistical Package, Version 82.1.1, Penn
<1.2% of the true value. State University, State College, PA) (33) between RMR

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 5

TABLE 2
Measured RMR versus predicted values (kcal/24 h)

Owen et al Harris- Robeilson-

Patient Measured (this study) Benedict Mayo Reid Cunningham
number RMR predictions predictions(4) predictions(6) predictions (8) predictions (12)

1 870 -235 -366 -305 -249 -339

2 1372 232 74 108 183 97
3* 1079 -25 -188 -183 -116 -201
4* 1146 24 -179 205 -76 -162
5* 1115 -29 -187 -191 -129 -191
6 1132 -38 -188 -166 -104 -186
7* 1161 0 -182 -201 -136 -162
8 1172 -7 -168 -163 -99 -161
9 1034 -156 -234 -295 -190 -279
10 1155 -35 -205 -214 -149 -195
11 1392 195 50 25 94 42
12* 1325 59 -77 -96 -29 -58
13 1090 -120 -237 -234 -153 -274
14 982 -237 -371 -404 -319 -395
15 1178 -41 -151 -251 -139 -178
16 1342 122 -58 -40 26 -57
17 1027 -195 -381 -389 -322 -374
18 1316 90 -74 -124 -51 -77
19* 1351 5 -11 -122 10 -35
20 1574 333 104 80 158 133
21* 1402 38 -71 -90 -15 -45
22* 1365 -55 -108 -136 -65 -92
23 1526 265 55 44 133 64
24 1268 0 -99 -112 11 -160
25 1205 -66 -250 -311 -234 -250
26 1382 65 -172 -202 -106 -153
27 1273 -59 -123 -165 -96 -184
28 1439 90 -8 -25 64 -55
29 1536 153 46 2 75 24
30 1151 -233 -403 -422 -299 -414
3lt 1248 -145 -402 -408 -308 -369
32 1466 53 -56 -40 52 -82
33 1323 -108 -229 -242 -168 -228
34 1300 -136 -244 -215 -143 -254
35 1519 66 -15 -26 36 -14
36 1639 127 -147 -78 25 -69
37 1382 -152 -357 -366 -209 -273
38t 1414 -131 -358 -281 -177 -273
39 1473 -95 -391 -468 -365 -231
40 2074 487 150 176 285 316
41 1777 106 -135 -154 -37 125
42 1640 -38 -315 -289 -132 -64
43t 1630 -64 -362 -389 -225 -75
44t 1708 -116 -515 -395 -294 -43

Mean 1340 0 -171 -172 -91 -133

SD 234 152 158 165 148 154
Range
Mm 870 -236 -515 -468 -365 -414
Max 2074 487 150 205 285 316
* Athlete.

t Thermogenic effect of food measured for 360 mm (6 h).

and Wi’ at each point in time. Similarly, resistant-regres- TEF was also determined by computing the area under
sion lines were used to study the changes in carbohydrate, the 4-h RMR response curve above baseline value for each
fat, and protein oxidation before and after the breakfast subject. The RMR measurement before the meal was used
mixed meal. as the thermic baseline for each subject. Pre- and post-

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 6 OWEN ET AL

prandial protein, carbohydrate, and fat oxidation and 0.021 to 0.061 kcal/min and the coefficient of
storage were computed from individual urinary-nitrogen
variation was<4%.
excretion, nonprotein RQ (npRQ), and change in extra-
cellularpool. Correlations,simple, multiple, and stepwise Diurnal changes in the RMR occur (1). The
regressions were used to study the relationships among lowest or basal metabolic rate develops about
TEF, fuel oxidation, and nutrient storage versus weight, 0400 h, during sleep (1). However, when we
age, BSA, and measured and calculated body components. repeatedly measured the RMR during daylight
Chemical analyses hours, there were no significant changes in our
Plasma and urine glucose were determined with a volunteers’ RMR while fasting and lying qui-
Beckman Glucose Analyzer (Beckman Instruments, Inc, etly in a comfortable position on a bed be-
Fullerton, CA). Serum urea nitrogen (34), plasma total tween 0800 h and 1600-1700 h. The coeffi-
lipids (35) and FFA (36), plasma and urine a-amino ni- cient of variation for these eight subjects stud-
trogen (37), and urine nitrogen (38) were measured by
standard techniques.
ied during 20 different time and measuring
periods over =-‘8 h was ±4.7%. Segal et al
(40) reported similar results for lean and
Results obese men.

Validation, accuracy, and reproducibility of RMR, body composition, and

indirect calorimetry anthropometric measurements
Recent refinements and automatization The measured RMRs varied from 870 to
have advanced the technique of indirect cal- 2074 kcal/24 h and were related to body size
orimetry to determine the nature and quantity (Table 3).
of fuels oxidized in a variety of states. We con- Body-composition variables reflecting active
firmed the validity of using indirect calorim- protoplasmic tissue such as WT, BSA,
etry to determine the nature and quantity of FFMSF, FFMD, LBM, and BCM were all
fuels oxidized in the postabsorptive period by highly interrelated (r> 0.80; Table 3). Also,
integrating the results obtained from cathe- it should be noted that FATMSF and FATMD
terization studies, tracer analyses, and indirect were correlated with measurements reflecting
calorimetry. These three diverse techniques active protoplasmic tissue (r> 0.67; Table 3).
showed excellent agreement for measuring the Therefore, it is extremely difficult to identify
quantities of glucose, protein, and fat produced which of these variables truly reflects active
and oxidized (39). protoplasmic tissue. Nonetheless, the variables
The mean ± SEM of the differences between most highly, but comparably, correlated with
the mouthpiece-noseclip and the ventilated- the measured RMR were FFMSF, BCM, BSA,
hood results was 0.003 ± 0.054 kcal/min. The LBM, and WT (r> 0.74; Table 3).
paired t test was not statistically significant (p Data from all 44 women were used for step-
= 0.95). Expressed as a percent of the RMR wise inclusion to determine whether some
mean, the differences between the two tech- combination of WT, HT (height), AGE, BSA,
niques is <1%. Thus, the agreement is excel- BMI, LBM, and BCM was a better predictor
lent, and the possibility that the mouthpiece- for RMR than a single variable. The combi-
noseclip technique might underestimate the nation of BSA and AGE gave the highest cor-
RMR or be different from the ventilated-hood relation (r = 0.79), but it was not statistically
technique is unfounded. Thus in our hands, different from the correlation (r = 0.74) for
the mouthpiece-noseclip technique is accurate WT alone.
for measuring respiratory gaseous exchange In addition to the above-mentioned vari-
rates in healthy volunteers. ables, FFMSF and FFMD data available for
From measuring the respiratory gaseous ex- 28 women were also variables used individu-
change rates at 1-mm intervals over several ally and in combination to predict RMR.
periods, and from repeatedly doing such stud- However, these two measured variables did
ies during five different mornings, the repro- not more accurately predict RMR. The com-
ducibility of the RMR was determined. The bination of BCM and HT was highly corre-
standard deviations of daily means for the lated with the RMR (r = 0.86), but again, it
eight volunteers that underwent repeated was comparable to the WT correlation (r
studies to measure their RMR ranged from = 0.80) for these 28 women.

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 7

TABLE 3
Resting metabolic rate, body composition, and anthropometric measurements: correlation coefficients

RMR AGE HT wT BSA BMI LBM 8CM FFMD FATMD FFMSF

A. All women (n = 44)

AGE 0.06
HT 0.41 0.00
WT 0.74 0.28 0.33
BSA 0.77 0.29 0.51 0.98
BMI 0.67 0.31 0.05 0.96 0.88
LBM 0.77 0.18 0.30 0.95 0.94 0.93
BCM 0.77 0.08 0.31 0.93 0.92 0.91 0.99
FFMD 0.71 0.07 0.53 0.84 0.90 0.72 0.83 0.83
FATMD 0.66 0.22 0.06 0.96 0.91 0.98 0.91 0.90 0.68
FFMSF 0.78 -0.05 0.52 0.92 0.94 0.82 0.87 0.88 0.95 0.82
FATMSF 0.69 0.24 0.17 0.98 0.93 0.98 0.93 0.92 0.77 0.99 0.82

B. Athletes (n = 8)

AGE -0.20
HI 0.80 0.06
WI 0.96 -0.11 0.80
BSA 0.95 -0.04 0.92 0.97
BMI 0.68 -0.26 0.20 0.76 0.58
LBM 0.96 -0.33 0.75 0.97 0.93 0.77
BCM 0.94 -0.43 0.70 0.94 0.90 0.77 0.99
FFMD 0.84 -0.14 0.71 0.95 0.90 0.77 0.93 0.91
FATMD 0.47 -0.02 0.33 0.25 0.30 0.07 0.25 0.24 -0.05
FFMSF 0.85 0.24 0.72 0.93 0.90 0.59 0.82 0.75 0.95 0.56
FAIMSF 0.62 -0.60 0.37 0.49 0.46 0.34 0.63 0.68 0.40 0.80 0.13

C. Nonathletes (n = 36)

AGE 0.02
HI 0.41 0.02
WI 0.74 0.22 0.38
BSA 0.75 0.23 0.55 0.98
BMI 0.66 0.24 0.08 0.95 0.88
LBM 0.75 0.12 0.33 0.95 0.93 0.93
BCM 0.75 0.01 0.33 0.93 0.91 0.91 0.99
FFMD 0.69 0.06 0.52 0.89 0.93 0.77 0.85 0.85
FATMD 0.70 0.09 0.13 0.97 0.92 0.98 0.93 0.92 0.76
FFMSF 0.77 -0.12 0.52 0.95 0.96 0.85 0.90 0.91 0.95 0.87
FATMSF 0.71 0.16 0.21 0.98 0.94 0.98 0.94 0.92 0.82 0.99 0.87

Abbreviations RMR = resting metabolic rate; HI = height; WI = weight; BSA = body surface area; BM1 = body mass index;
LBM = lean body mass; SCM = body cell mass; FFMD = fat-free mass by densitometry; FATMD = fat mass by densitometry; FFMSF
= fat-free mass by skin folds;FAIMSF = fat mass by skin folds.

Body weight was the most accurately and for nonathletic lean and obese women were
easily determined variable and was highly cor- statistically indistinguishable. Therefore, a
related with other measurements of active single regression line with its 95%-confidence
protoplasmic tissue (r> 0.84); further, it was limits for a single prediction was developed
highly correlated with RMR (r = 0.74). for the nonathletic healthy women (RMR
Therefore, WT alone was used to derive pre- = 795 + 7.18 kg WT). Note that both non-
dictor equations for the RMR of 18-65-yr- athletic groups of women had wide 95%-con-
old, healthy women weighing 43-143 kg. fidence limits, reflecting the heterogenous na-
Figure 1 displays the relationship between ture of the RMR for lean and fat nonathietic
measured RMR and body weight. The slopes women. The regression line for the athletic
of the regression lines for the RMR and WT women (RMR = 50.4 + 21.1 kg WT) was

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 8 OWEN ET AL

2500 - 0 LEAN
#{149}
OBESE
a AThLETE

2000: #{149}

#{163}

“4 1500:

4
0
I
1000

o--------------------
500

0
I I’’ I’’ 111J1T lIIi T IT 117111 111 11IJ 111
43 53 63 73 83 93 103 113 123 133 143
WEIGHT (kg)

FIG 1. The RMRs (kcal/24 h) are contrasted against the weights (kg) of the 44 healthy lean 0 and obese S women,
8 of whom were athletes. The regression line with its wide 95%-confidence limits for a single prediction for the
nonathletes (RMR = 795 + 7.18 kg WT) is shown in black. The regression line with its narrow 95%-confidence limits
for a single prediction for the athletes (RMR = 50.4 + 21.1 kg WT) is shown in bold black.

clearly different from that for the nonathietic Mayo Foundation, Robertson-Reid, and
women (p <0.05), and the 95%-confidence Cunningham regression formulas or tables
limits for the regression line for the athletes systematically overestimated RMR by 171
were narrow. ± 158, 172 ± 165, 133 ± 154, and 91 ± 148
The regression line RMR versus WT for the kcal/24 h, respectively.
lean women alone was not significantly dif- Nature and quantity offuels oxidized
ferent from either the regression line for the and TEF
athletic or for the obese women. Therefore,
After an overnight fast in the resting state,
the lean women could conceivably have been
the nature and quantity of fuels oxidized by
grouped with either the athletic or the obese
women. However, testing for equality of vari- athletes and nonathletes were similar. Fur-
ance among these groups clearly showed the thermore, TEF was comparable in athletes and
lean and obese nonathietes. Dietary-induced
athletes to be much more predictable (less
variable) than the lean women (p < 0.05), thermogenesis was directly related to weight
while the variances for lean women alone and (r = 0.66). Adding more variables, using
stepwise-regression analysis, did not improve
obese women alone were not statistically dif-
ferent from each other. the model. With the caloric challenge and food
composition used in this study, a prediction
Our measured RMRs were contrasted
equation (TEF = -1.21 h) + 0.529 kg WT/4
against our newly developed prediction
regression formulas (see Appendix, equation for the athletic and nonathletic women was
developed.
9) for nonathletic and athletic women. The
old prediction regression formulas or tables of Figure 2 shows three-dimensional response
Harris-Benedict (41), Mayo Foundation (6), surfaces generated from resistant regression
Robertson-Reid (8), and Cunningham (12), lines from our data relating total oxidation
were also used to calculate the RMR of our rates to body weights for each time period.5
subjects. Our measured mean ± 1 SD versus 5The data from athletes were included in these resistant
predicted RMR was 0.0 ± 152 kcal/24 h. regression lines and may have caused minimal distortion
However, the use of the Harris-Benedict, of the 0 time-weight resistant regression line.

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 9

FIG 2. Three-dimensional response surface generated from resistant regression analysis of the data from 32 women
relating total metabolic rate, body weight, and time before and after breakfast.

These surfaces show a continuum from the women. Further, TEF persisted for 6 h in the
lowest to highest weights for RMR and TEF. grossly obese women (Tables 1 and 2) (data
Figure 2 shows that in a healthy woman with not shown on the 240-360 mm time period).
40 kg body weight, the first and second max- Figure 3 shows three-dimensional response
imal oxidation rates after eating a large meal surfaces generated from resistant regression
should occur at 30 and 90 mm postprandially, lines relating carbohydrate, fat, and protein
respectively. The TEF would be completed by oxidation rates to body weight for each time
210 mm postcibal. The 4-h TEF for a lean period. Before breakfast, 22 ± 13, 60 ± 8, and
individual would be 15% above RMR on av- 18 ± 1% of the RMR were derived from the
erage. As weight (and food intake) increased, oxidation of carbohydrate, lipid, and protein,
peak oxidation rate should increase and respectively. Quantities of the fuels oxidized
broaden. In addition, in the extremely obese form a continuous spectrum from lean to
140-kg healthy woman, a late-third peak obese individuals. Following breakfast, the
would develop and the TEF would last for 360 nature and quantity of fuels oxidized changed
mm postprandial (data not shown for the 240- rapidly. There was a three-fold postcibal in-
360 mm time period). The 4-h TEF for an crease on average (p <0.001) in carbohydrate-
obese individual would be 20% above RMR oxidation rate and a slight but significant in-
on average. Thus, peak oxidation rates were crease in protein oxidation (p <0.001). The
delayed, broadened, and multiple in obese increased postprandial-oxidation rates for
compared to lean women and the TEF were carbohydrate and protein were progressively
as great, if not greater, in obese as in lean delayed as body weight (and food intake) in-

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 10 OWEN ET AL

0! 30! .01 so! oI too1 ‘so sio

TIME (mm)

FIG 3. Three-dimensional response surfaces generated from resistant regression analysis of data from 32 women
relating carbohydrate, fat, and protein oxidation with body weight and time before and after breakfast.

creased. A reciprocal relationship between semiquantitative because the exact quantities

carbohydrate and lipid postprandial-oxidation of ingested calories of carbohydrate, fat, or
rates developed: simultaneously, as postcibal protein absorbed from the gut were unknown.
carbohydrate oxidation increased, lipid oxi- Further, correction for the energy cost asso-
dation decreased and vice versa. ciated with carbohydrate, fat, and protein
storage (or conversion), recycling, and synthe-
Postprandial storage of nutrients sis was not made.
Figure 4 shows caloric intake and postpran- The overnight fasting and 4-h postprandial
dial thermogenesis in kcal/4 h from which ca- plasma glucose (5.14 ± 0.08 vs 5.13 ± 0.08
loric storage was estimated. Storage is only mmol/l and 92.0 ± 2.0 vs 92.0 ± 1.0 mg/dl)

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN II

o LEAN
1650 #{149}
OBESE
1650

1500

11350 1350
0
INTAKE
I 1
900

P750

#{149} 0
600 ,-

450
METABOUSM

300: 300

150 150

0 0

43 53 63 73 83 93 103 113 123 133 143

WEIGHT (kg)

FIG 4. Caloric intake and total postprandial metabolism (RMR plus TEF kcal/4 h) or thermogenesis in 32 healthy
lean and obese females, 8 of whom were athletes. One obese subject 0 was unable to consume the entire 11 kcal/kg
meal. She ate 9.4 kcal/kg.

and urea nitrogen (11.07 ± 0.36 vs 10.60 sented by a linear function with a positive slope
± 0.28 mmol/l and 15.5 ± 0.5 vs 14.9 ± 0.4 (G5 kcal/4 h = 3.85 + 0.739 kg WT). Lipid
mg/dl), and total lipid (4.9 ± 0.2 vs 5.0 ± 0.2 storage (L) was best represented by an in-
mg/ml) concentrations were comparable. creasing quadratic function (L kcal/4 h = 8.92
There was a small but significant increase in +0.002 kg WT2). Protein storage or repletion
plasma a-amino
nitrogen (4.78 ± 0.15 vs 5.54 (P3) was best represented by a decreasing
± 0.19 mg/dl; p <0.001 and 3.41 ± 0.11 vs square-root function (P0 kcal/4 h = -34.2
3.96 ± 0.14 mmol/l), and a large decrease in + 5.78 kg WT#{176}5).These observations were
plasma FFA (643 ± 31 vs 304 ± 25 mol/l; p corroborated by stepwise-regression analyses.
<0.001) concentrations. There was no change However, these functions are not statistically
in urinary glucose or a-amino nitrogen. better than linear functions.
The caloric equivalent changes of nutrients
in the plasma (extracellular space) before and Discussion
4 h after the mixed meal were trivial and,
therefore, were disregarded in calculating ox- This study showed that the measured RMRs
idative and nonoxidative pathways for nu- for normal women were less than their pre-
trient disposal. Caloric intake, based on body dicted RMRs based on results from the largest
weight and caloric storage, increased as weight and most authoritative reports published in
(intake) increased. However, the percentage the first half of this century (4, 6, 8). A reas-
of caloric intake that was disposed of by sessment of our measured RMR based on
mechanisms other than oxidation was pro- measured or estimated body composition did
portional to body weight. Therefore, the not change our subjects’ lower-than-predicted
heaviest individual stored the greatest number RMR. Therefore, we believe that the currently
of calories in tissue deposits. available tables (5, 8, 9) and regression equa-
Figure 5 shows carbohydrate, lipid, and tions (4, 12) overestimate the RMR of healthy
protein storage (or interconversion) in kcal/4 women. We therefore recommend that the
h following a mixed meal containing enough RMR of healthy women between the ages of
calories to be an anabolic challenge in each 18 and 65 yr be reappraised using newly de-
subject. Glucose storage (G5) was best repre- veloped prediction equations:

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 12 OWEN ET AL

0 TOTAL STORAGE
#{149}
CARBOHYDRATE STORAGE 0
o UPS STORAGE
0
S PROTEIN STORAGE

0
0
0
0

WEIGHT (kg)
FIG 5. Caloric storage of carbohydrate, fat, and protein contrasted against weight (intake). This scheme is only
semiquantitative inasmuch as the exact quantity of ingested calories in carbohydrate, fat, or protein absorption is
unknown and corrections for the energy cost associated with carbohydrate, fat, and protein storage, recycling, or
synthesis were not made.

For nonathletes, fore, if possible, the metabolic requirements

of humans should be measured rather than
RMR=795+7.l8WTkg and predicted. Recent refinements and automati-
for athletes, RMR = 50.4 + 21.1 WT kg. zation have advanced the technique of indirect
calorimetry so that measurements can be
It should be emphasized that, due to the made to determine the energy requirements
large variations in measured RMR, predicted and the nature and quantities of macronutri-
RMR may over- or underestimate the mea- ents oxidized.
sured RMR of nonathletes by 21-33%. Closer Our data did not support the frequently
estimates can be made for athletes. The pre- made claim that RMR correlates best with
dicted RMR of a well-trained, competitive LBM (12, 19, 41-46) (Table 4). In retrospect,
athlete estimates RMR within 8-10%. Athletic this is not surprising because the resting energy
women have greater increases in RMR per requirements of various aerobic tissues are
gain in body weight than nonathletic women vastly different per unit mass (47-49). In
(Fig 1). Body weight was also more highly cor- normal-weight adult women, the brain and
related with RMR (Table 3) among the ath- liver constitute about 4-5% of total body
letes (r = 0.94) than among the nonathletes (r weight and are responsible for about 40% of
= 0.74). the RMR (48, 49). Muscle comprises 35-40%
We feel that our new prediction equations of body weight but accounts for about 20% of
for the RMR of healthy women are better than the RMR (48, 49). Adipose tissue constitutes
those previously available for women. How- about 20% of body weight but accounts for
ever, the usefulness of any RMR prediction only 2-4% of RMR (48, 49). These dispro-
equation derived from a population with large portionate rates of energy requirements per
95%-confidence limits is questionable. There- mass of different tissues partly explain why the

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 13

TABLE 4
Regression analyses for the prediction of RMR
95%-Confidence 95%-Confidence
SD of RMR limit for mean limit for individuals
Predictor variable about line X RMR given X % RMR given X

A. Nonathletic women (n = 36)

WT 169.7 43 1104± 100.5 9 1104±358.9 33

77 1348± 57.7 4 1348±349.3 26
143 1823 ± 158.3 9 1823 ± 379.2 21
BSA 165.0 1.37 1047 ± 110.8 11 1047 ± 352.8 34
1.84 1363± 55.8 4 1363±339.6 25
2.45 1784 ± 140.1 8 1784 ± 363.1 20
BSAandAGE 162.6 1.37 1067±113.1 11 1067±348.9 33
1.84 1358± 55.6 4 1358±334.7 25
2.45 1841 ± 160.6 9 1841 ± 367.6 20
LBM 166.0 32.7 1004 ± 123.6 12 1004 ± 358.9 36
46.0 1357± 56.2 4 1357±341.7 25
57.8 1674± 111.5 7 1674±354.9 21
BCM 165.3 16.4 1016± 119.6 12 1016±356.2 35
23.5 1399 ± 57.0 4 1399 ± 340.4 24
29.3 1714 ± 121.4 7 1714 ± 356.9 21
BCMandHT 162.0 16.4 983± 124.9 13 983±351.8 36
23.5 1364 ± 75.3 5 1364 ± 337.4 25
29.3 1721 ± 119.4 7 1721 ± 349.9 20
FFMSF 180.4 35.0 1039 ± 137.5 13 1039 ± 401.8 39
47.1 1290± 82.9 6 1290±386.6 30
76.9 1912±251.9 13 1912 ±453.9 24

B. Athletic women (n = 8)

WT 40.6 50 1105± 54.3 5 1105± 113.2 10

58 1268 ± 35.9 3 1268 ± 105.7 8
65 1422± 64.6 5 1422± 118.5 8
BSA 44.9 1.47 1082 ± 66.8 6 1082 ± 128.6 12
1.59 1215± 39.9 3 1215±116.9 10
1.75 1403 ± 66.3 5 1403 ± 128.3 9
BSAandAGE 42.5 1.47 1068± 71.5 7 1068± 130.6 12
1.59 1218± 40.1 3 1218± 116.5 10
1.75 1410 ± 67.6 5 1410 ± 128.5 9
LBM 40.6 36.0 1093 ± 57.3 5 1093 ± 114.6 10
40.8 1282 ± 36.9 3 1282 ± 105.9 8
44.3 1421 ± 64.4 5 1421 ± 118.3 8
13CM 48.7 17.9 1094 ± 69.2 5 1094 ± 137.8 13
20.1 1258 ± 42.6 3 1258 ± 126.5 10
22.2 1416 ± 76.3 5 1416 ± 141.5 10
BCM and HT 43.3 17.9 1070 ± 74.5 7 1070 ± 133.9 13
20.4 1303 ± 50.6 4 1303 ± 122.2 9
22.2 1412 ± 71.7 5 1412 ± 132.3 9
FFMSF 75.4 41.0 1118±101.5 9 1118±210.5 19
45.7 1231 ± 69.7 6 1231 ± 197.1 16
50.2 1341 ± 105.5 8 1341 ± 212.4 16
Abbreviations: RMR = resting metabolic rate; WT = weight; BSA = body surface area; LBM = lean body mass;
BCM = body cell mass; HT = height; FFMSF = fat-free mass by skin folds.

bulk quantity of heterogeneously active pro- 19, 50), energy expenditure per kg body weight
toplasmic tissue correlates about as well with for lean and obese nonathletic and for athletic
the RMR as total body weight. These same women decreased as weight increased. Large
considerations should be extended to obese variances in body weight occur, not because
and athletic women. Although the absolute of major differences in high-energy-requiring
RMR was greater the heavier the woman (3, brain and liver masses, but because of large

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 14 OWEN ET AL

differences in the amount of moderate-energy- obese humans both have a greater predis-
requiring skeletal muscle mass in the athletes position for developing and maintaining
or because of huge differences in the amount body fat.
of low-energy-requiring adipose tissue mass in We used a powerful statistical method, re-
obese nonathletes (Table 5). Thus, a major sistant regression analysis (33), to maximize
portion of the RMR in adult women is rela- our observations. The resistant regression
tively constant because of brain and liver me- lines, calculated from data of 32 women, were
tabolism, but the increases in RMR per kg derived from measured RMR before and after
body weight depend primarily on other body eating a mixed breakfast providing a caloric
components (ie, skeletal muscle and adipose challenge proportional to body weight. The
tissue). This was recently demonstrated by Se- regression lines for RMR (cal/mm) before and
gal (40). Under unusual circumstances, in men after breakfast were related to weight (kg) and
with massively hypertrophied muscles induced time (mm) (Fig 2). There was a continuum
by power weight-lifting, RMR and TEF are from the lowest to the highest weight for RMR
augmented by lean body mass (40). and TEF: as weight increased, RMR and TEF
The first part of our studies showed that the increased. Also, as weight increased, post-
RMR of lean and obese nonathletic women prandial peak rates of metabolism were
are lower than generally recognized; our stud- broadened, delayed, and converted from bi-
ies also showed that the RMR for both lean phasic to triphasic peaks. 4-h
Further, the
and obese nonathletic women have wide 95%- postprandial thermogenic rates were persis-
confidence intervals. This latter finding sug- tently above baseline values as weight in-
gests that metabolic efficiency is not limited creased. This heightened elevation in TEF ex-
to obese women. Instead, both lean and obese hibited by the heavy women was related to the
healthy nonathletic women have wide ranges larger caloric challenges given to these women,
of RMR, and some lean as well as some obese rather than to the women’s weights (20). Data
women are very metabolically efficient while analysis shows that although there was a linear
others are not. Therefore, some lean and some increase in energy expenditure over the RMR
as the caloric intake (and weight) increased
TABLE 5 (RMR = 0.0539 kcal 5.58), the TEF#{149}
- 4 h kg
Comparison of body composition of athletes and was 0.5 ± 0.06 kcal kg#{149} mi regardless of the
nonathletes caloric content of the food ingested. Thus, if
Nona thietes Athletes
the caloric challenge of a mixed meal is large
but proportional to body weight, TEF is com-
Lean Obese
parable in lean and obese women.
WT (kg) The TEF were described about 100 years
Mean 60.3 102.3 56.4
SD 9.1 18.7 5.9
ago. They were originally attributed to the en-
FFMSF (kg) ergy requirements of urea synthesis and amino
Mean 43.0 58.9 45.5 acid degradation (51). Currently, TEF in hu-
SD 3.7 10.9 4.6 mans have been divided into obligatory energy
As%ofmeanWT 71.3 57.6 80.7
FATMSF (kg)
expenditure associated with eating, digesting,
Mean 16.1 47.5 9.7 absorbing, and storing food (52) and faculta-
SD 6.8 12.1 1.9 tive energy expenditure associated with hor-
As % of mean WT 26.7 46.4 17.2 mone secretion (53), sodium-potassium-
FFMD (kg) ATPase pump activity (54, 55), protein syn-
Mean 44.1 57.9 47.6
SD 4.1 8.9 5.7
thesis (56), and substrate recycling (57, 58).
As % of mean WT 73.1 56.6 84.4 During the past decade, about 26 studies have
FATMD (kg) been published evaluating thermogenesis in
Mean 15.4 45.6 8.3 lean and obese subjects in response to oral or
SD 7.0 14.9 2.1
parenteral macronutrients, hormones, and
As%ofmeanWT 25.5 44.3 14.7
drugs. Approximately equal numbers of these
Abbreviations: WT = weight, FFMSF = fat-free mass
articles support or deny a thermogenic defect
by skin folds; FATMSF = fat mass by skin folds; FFMD
= fat-free mass by densitometry; FATMD = fat mass by in obese subjects (personal communication
densitometry. from Eric Ravussin).

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 15

Several interrelated hormonal systems in- sumed. Before breakfast in the resting state,
fluence thermogenesis. The iodothyronines oxidation rates of carbohydrate, lipid, and
play a cardinal role in regulating the basal and protein were (mean ± 1 SD) 0.48 ± 0.40,0.59
resting metabolic rates but do not change the ± 0.20, and 0.41 ± 0.20 mg#{149} mm kg, respec-
exercise-induced or diet-induced energy ex- tively. Thirty minutes after breakfast, the
penditure (59). The central nervous system is carbohydrate-oxidation rate soared to 1.5
integrated with the sympathico-adrenome- ± 0.80 mg#{149} mm kg. Concurrently,
. the fat-ox-
dullar system, and both interrelate with the idation rate plummeted to 0.27 ± 0.27
iodothyronines. Cortisol promotes the release mg mm kg. Thus, a tight reciprocal rela-
and thermogenic response of the body to nor- tionship developed postprandially between
epinephrine and epinephrine, and the cate- carbohydrate and lipid oxidation. The post-
cholamines augment the thermogenic re- prandial increase in protein oxidation was
sponse of the body to L-triiodothyronine (60). slight, and as for carbohydrate and lipid, the
Special emphasis has been placed on the 3- change was greater and delayed in obese
adrenergic activity of norepinephrine as being women (Fig 3).
responsible for the rise in facultative energy A large proportion of the nutrients were
expenditure associated with carbohydrate me- disposed of by nonoxidative (storage) path-
tabolism (61, 62). ways. When energy requirements were met
Although insulin per se does not exert a and proteins were repleted, direct or indirect
thermogenic effect (63), insulin facilitates in- storage of excessive nutrients was the most
tracellular translocation, oxidation and storage important route of fuel disposal. In this study,
of glucose, reduces lipolysis and augments li- total caloric intake was greater per kilogram
pogenesis, and promotes amino translocation than was postprandial energy requirement per
and protein synthesis. Further, the heightened kilogram. Therefore, as weight increased, ca-
postprandial circulating-insulin concentration loric storage increased (Fig 5). Quantitatively,
could stimulate the ventromedial hypothala- the progression of calories stored as carbohy-
mus and enhance norepinephrine secretion drate, lipid, and protein varied over the 4-h
(61-63). postprandial period studied. Glucose storage
In obesity, the bioactivity of iodothyronine was linearly related with intake. Since the
hormones is considered normal (1) whereas postprandial npRQ rose only transiently to
the bioactivity of catecholamines is contro- somewhere in the range of 1.05 to 1.17 in 4
versial (15, 19, 63-65). However, insulin se- of 44 women (data not shown), only trivial
cretion does increase as weight and adiposity net quantities of carbohydrate were converted
increase (66). Nonetheless, the progressive in- into fat. The vast majority of nonoxidative
crease in the RMR and TEF among healthy disposal of glucose was via glycogen synthesis
women with weights ranging from 43-143 kg and storage. These observations from women
discredits the hypothesis suggesting a uni- are in agreement with the results from men
formly altered or defective thermogenic (68, 69) showing that de novo lipogenesis from
mechanism related to hormonal secretion, Na- glucose is negligible.
K-ATPase pumping, substrate recycling, or Lipid storage was related to body weight and
synthetic processing as causes for less or more fat ingestion in a quadratic fashion, suggesting
energy-efficient states, and, thus, secondary that when women ate mixed meals containing
leanness or obesity. substantial caloric challenges, progressively
Our data also show that TEF was directly larger quantities of the ingested fat was stored
related (r = 0.58) to carbohydrate oxidation as triglyceride. Lipid metabolism was opposite
(Fig 3), whereas no correlations existed be- to protein metabolism. Although protein con-
tween TEF and lipid or protein oxidation. tent of the breakfast meal was only 15% of the
Lusk (67) showed that thermogenesis from caloric loads, protein intake varied from 18.0
carbohydrate (glucose) oxidation was greater to 59.4 g. Thus, protein quantities in obese
than that of lipid or protein per liter of 02 women were large. In contrast to the potential
consumed and, in our study, carbohydrate massive caloric stores of lipid in adipose tissue
oxidation increased shortly after eating and and the small, yet significant storages of car-
was the dominant, postprandial fuel con- bohydrate as glycogen in liver and muscle,

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 16 OWEN ET AL

there is no specific depot for proteins or amino 10. Feurer ID, Crosby LO, Buzby GP, Rosato EF, Mullen
JL. Resting energy expenditure in morbid obesity. Ann
acids. Therefore, it was not surprising to ob-
Surg 1983;197:17-21.
serve that protein storage (repletion) leveled, 11. Roza AM, Shizgal HM. The Harris-Benedict equation
while surplus amino acids were oxidized post- reevaluated: resting energy requirements and the body
prandially during mixed-nutrient overabun- cell mass. Am J Gin Nutr l984;40: 168-82.
12. Cunningham JJ. A reanalysis of the factors influencing
dance (Fig 3). This confirms the claim that
basal metabolic rate in normal adults. Am J Clin Nutr
excessive amino acid loads are immediately l980;33:2372-4.
and preferentially oxidized (70). 13. Heymsfield 5, Daly J, Harvey L, Nixon D. Overesti-
We concluded that 1) the RMRs of healthy mate of energy expenditure by widely used equation.
women are lower than previously recognized; Gin Res 1984;32:629A(Abst).
14. Kaplan ML, Leveille GA. Calorigenic response in
2) increased metabolic efficiency should no
obese and nonobese women. Am J Clin Nutr l976,29:
longer be considered as an important cause of 1108-13.
obesity in women; and 3) the nature and 15. Shetty PS, Jung RT, James WPT, Barrand MA, Cal-
quantities of nutrients oxidized after a mixed lingham BA. Postprandial thermogenesis in obesity.
Gin Sci 198l;60:5l9-25.
meal change dramatically with a reciprocal re-
16. Schwartz RS, Halter JB, Bierman E. Reduced thermic
lationship between carbohydrate and fat
effect of feeding in obesity: role of norepinephrine.
oxidation. II Metabolism 1983;32:l 14-7.
17. Sharief NN, Macdonald I. Differences in dietary-
The authors wish to thank Ms Rachelle Browndorf, Ms induced thermogenesis with various carbohydrates in
Jeanette Camacho, and the nurses of the General Clinical normal and overweight man. Am J Clin Nutr l982;35:
Research Center for their assistance. We would also like 267-72.
to thank the staff of the Biokinetics Research Laboratory, 18. Welle SL, Campbell RG. Normal thermic effect of
HPERD Temple University, for their work in this research glucose in obese women. Am J Clin Nutr l983;37:
project. In addition, Rodney S Owen was the recipient of 87-92.
the Harold Lamport Award for this work. 19. Felig P, Cunningham JJ, Levitt M, Hendler R, Nadel
E. Energy expenditure in obesity in fasting and post-
prandial state. Am J Physiol (Endocrin Metab 7)
References l983;244:E45-5 1.
20. Hill JO, Heymsfield SB, McMannusC III, DiGirolamo
1. Ravussin E, Burnand B, Schutz Y, Jequier E. Twenty- M. Meal size and thermic response to food in male
four hour energy expenditure and resting metabolic subjects as a function of maximum aerobic capacity.
rate in obese, moderately obese, and control subjects. Metabolism l984;33:743-9.
Am J Clin Nutr l982;35:566-73. 21. Pittet PH, Chappius PH, Acheson K, deTechterman
2. Jequier E. Long-term measurement of energy expen- F, Jequier E. Thermic effect of glucose in obese sub-
diture in man: direct or indirect calorimetry. In: jects studied by direct and indirect calorimetry. Br J
Bjorntorp P. Cairella M, eds. Recent advances in obe- Nutr l976;35:28 1-92.
sity research III. London: John Libbey, 198 1:130-5. 22. Golay A, Schutz Y, Meyer HU, et al. Glucose-induced
3. Jequier E, Schutz Y. Long-term measurements of en- thermogenesis in nondiabetic and diabetic obese sub-
ergy expenditure in humans using a respiration jects. Diabetes 1982;3l:1023-8.
chamber. Am J Clin Nutr 1983;38:989-98. 23. Horton ES, Danforth E Jr. Energy metabolism and
4. Harris JA, Benedict FG. A biometric study of basal obesity. In: Brodoff NB, Bleicher SJ, eds. Diabetes
metabolism in man. Washington, DC: The Carnegie mellitus and obesity. Baltimore, MD: Williams and
Institute, 19 19:1-266. Wilkins, 1982:261-8.
5. DuBois BS, DuBois EF. Clinical calorimetry: a for- 24. Bray GA. Obesity: benefits and risks of treatment.
mula to estimate the approximate surface area if height Drug Therapy 1984;l4:60-5.
and weight be known. Arch Intern Med 1916;17:863- 25. Goldman RF, Buskirk ER. Body volume measure-
71. ment by underwater weighing: description of a
6. Boothby WM, Berkson J, Dunn HL. Studies of the method. In: Brozek J, Henschel A, eds. Techniques
energy of metabolism of normal individuals: a stan- for measuring body composition. Washington DC:
dard for basal metabolism, with a nomogram for clin- NatlAcadSci, 1961.
ical application. Am J Physiol l936;l 16:468-84. 26. Brozek J, Grande F, Anderson R, Keys A. Densito-
7. Boothby WM, Sandiford I. Summary of the basal me- metric analysis of body composition: revision of some
tabolism data on 8,614 subjects with especial reference quantitative assumptions. NY Aced Sci l963;I:ll3-
to the normal standards for the estimation of the basal 40.
metabolism rate. J Biochem 1922;54:783-803. 27. Jackson AS, Pollock ML, Ward A. Generalized equa-
8. Robertson JD, Reid DD. Standards for the basal me- tions for predicting body density of women. Med Sci
tabolism of normal people in Britain. Lancet 1952;l: Sports Exer 1980;12(3):l75-82.
940-3. 28. Moore FD, Olesen KH, McMurrey JD, Parker
9. DuBois EF. Basal energy metabolism at various ages: HV, Ball MR. Boyden CM. The body cell mass and
man. In: Altman PL, Dittmer DS, eds. Metabolism. its supporting environment. Philadelphia, PA: WB
Bethesda, MD: FASEB, 1968:345. Saunders, 1963.

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 17

29. Albert SN. Blood volume and extracellular fluid vol- 51. Krebs HA. In: Munro HN, Allison JB, eds. Mam-
ume. Springfield, IL: Thomas, 197 1:228-32. malian protein metabolism. New York: Academic
30. Foss MC, Viachokosta V. Cunningham LN, Aoki T. Press, 1965:125.
Restoration of glucose homeostasis in insulin-depen- 52. Trayburn P, James WPT. Thermogenesis: dietary and
dent diabetic subjects. Diabetes 1982;31:46-52. nonshivering aspects. In: Cioffi LA, James WPT, Van
3 1 . Best C, Taylor NB. The physiological basis of medical Itallie TB, eds. The body weight regulatory system:
practice. 7th ed. Baltimore, MD: Williams and Wil- normal and disturbed mechanisms. New York: Raven
kins, 1961:741-68. Press, 1981:97-105.
32. Snedecor GW, Cochren WG. Statistical methods. 6th 53. Himms-Hagen J. Thermogenesis in brown adipose
ed. Ames, IA: Iowa State University Press, 1967. tissue as an energy buffer implications for obesity. N
33. Mosteller F, Tukey JW. Data analysis and regression. Engl J Med 1984;31 1:1549-58.
Reading, MA: Addison-Wesley, 1977:203-18. 54. Bray GA, York DA. Hypothalamic and genetic obesity
34. Marsh WH, Fingerhut B, Miller H. Automated and in experimental animahi an autonomic and endocrine
manual direct methods for the determination of blood hypothesis. Physiol Rev 1979;7 19-809.
urea. Gin Chem l965;l 1:624-7. 55. Guernsey DC, Stevens DD. The cell membrane so-
35. Folch J, Lees M, Stanley GH. A simple method for dium pump as a mechanism for increasing thermo-
the isolation of total lipids from animal tissues.J genesis during cold acclimation in rats. Science
Biochem 1956;226:497-509. 1977; 196:908-10.
36. Lorch E, Gey KF. Photometric titration of free fatty 56. Miller BG, Otto WR, Grimble RF, York DA, Taylor
acids with Technicon autoanalyzer. Ann Biochem TG. The relationship between protein turnover and
1966; 16:224-52. energy balance in lean and genetically obese (ob/ob)
37. Kekki M. Microdetermination of amino nitrogen as mice. BrJ Nutr 1979;4l:l85-99.
copper complexes: a modification for plasma and 57. Newsholme EA, Crabtree B. Substrate cycles in met-
urine. Scan J Clin Lab Invest 1959;1 1:311-21. abolic regulation and in heat generation. Biochem Soc
38. Koch FC, McMeekin TL. A new direct nesslerization Sympos 1976;41:61-l09.
micro Kjeldahl method and a modification of the 58. Katz J. Energy balance and futilecycling. In: Kinney
Nessler-Folin reagent for ammonia. J Am Chem Soc JM, Lense E, eds. Assessment of energy metabolism
l924;46:2066-7 1. in health and disease.Columbus, OH: Ross Labora-
39. Owen OE, Trapp YE, Reichard GA Jr,et al.Nature tories,1980:63-6.
and quantity of fuels consumed in patients with al- 59. Acheson K, Jequier E, Burger A, Danforth E Jr. Thy-
coholic cirrhosis.J Gin Invest l983;72:182l-32. roid hormones and thermogenesis: the metabolic cost
40. Segal KR, Gutin B, Nyman AM, Pi-Sunyer FX. of food and exercise. Metabolism l984;33:262-5.
Thermic effectof food at rest, during exercise,and 60. Owen OE, Reichard GA, Boden G, Patel MS. Trapp
after exercise in lean and obese men of similar body YE. Interrelationshipsamong key tissuesin the uti-
weight. J Clin Invest 1985;76:l 107-12. lization of metabolic substrates. In: Katzen HM,
41. Miller A, Blyth C. Lean body mass as a metabolic Mahler Ri, eds.Diabetes,obesity,and vasculardisease.
reference standard. J Appl Physiol 1953;5:3l 1-6. Part II:metabolic and molecular interrelationships.
42. Behnke AR. Relationship of lean body weight to me- Washington, DC: Hemisphere, 1978:517-50.
tabolism and some consequent systematizations. Ann
61. Anand BK, Chhina GS, Sharma KN, Dua 5, Singh
NY Aced Sci 1953;56:l095-l42.
B. Activity of single neurons in the hypothalamic
43. Keys A, Taylor HL, Grande F. Basal metabolism and
feeding centers: effect of glucose. Am J Physiol
age of adult man. Metabolism l973;22:579-87.
l964;207: 1146-54.
44. Tzankoff S. Norris A. Effect of muscle mass decrease
62. Young JB, Landsberg L. Catecholamines and the
on age-related BMR changes. J Appl Physiol 1977;43:
sympathoadrenal system: the regulation of metabo-
100 1-6.
lism.In:Ingbar SH, ed. Contemporary endocrinology.
45. Tzankoff 5, Norris A. Longitudinal changes in basal
Vol 1.New York: Plenum Press, 1979:24-303.
metabolism in man. J Appl Physiol 1978;45:536-9.
63. Acheson KJ, Ravussin E, Wahren J, Jequier E.
46. Bernstein RS, Thornton JC, Yang MU, et al. Predic-
Thermic effect of glucose in man: obligatory and fac-
tion of resting metabolic rate in obese patients. Am J
ultative thermogenesis. J Gin Invest 1984;74:1572-
Clin Nutr l983;37:595-601.
80.
47. Brozek J, Grande F. Body composition and basal me-
tabolism in man: correlation analysis versus physio- 64. WelleS, Campbell RG. Stimulation of thermogenesis
logical approach. Human Biol 1955;27:22-31. by carbohydrate overfeeding: evidence against sym-
48. Cahill GF, Owen OE. Some observations on carbo- pathetic nervous system mediation. J Gin Invest
hydrate metabolism in man. In: Dickens F, Randle l983;7 1:916-25.
FJ, Whelan WJ, eds. Carbohydrate metabolism and 65. Acheson K, Jequier E, Wahren J. Influence of beta-
its disorders.Vol 1. New York: Academic Press, 1968: adrenergic blockade on glucose-induced thermogenesis
497-5 18. in man. J Clin Invest l983;72:981-6.
49. Grande F, Keys A. Body weight, body composition, 66. Cahill GF. Obesity. In: Brodoff BN, Bleicher SI, eds.
and calorie status.In: Goodhart RS, Shils ME, eds. Diabetes mellitus and obesity. Baltimore, MD: Wil-
Modern nutritionin health and disease.Philadelphia, liams and 1982:215-8.
Wilkins,
PA: Lea and Febiger, 1980:3-34. 67. Lusk G. The elements of the science of nutrition.
50. James WPT, Davies HL, Bailes J, Dauncey HL. El- Philadelphia, PA: WB Saunders, 1919:61.
evated metabolic rates in obesity. Lancet 1978; 68. Acheson KJ, Schutz Y, Bessard T, Ravussin E, Jequier
1:1122-5. E, Flatt JP. Nutritional influences upon lipogenesis

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 18 OWEN ET AL

and thermogenesis after a carbohydrate meal. Am J 8) Body Cell Mass equation:

Physiol (Endocrinol Metab) 1984;246:E62-70.
BCM (kg) = 0.493 kg WT - 0.00184 kg WT2
69. Acheson KJ, Raft JP, Jequier E. Glycogen synthesis
versus lipogenesis after a 500-g carbohydrate meal. - 0.00 144 kg WT X yr age + 0.00000221 kg WT2
Metabolism 1982; 1234-40.
70. Krebs HA. Some aspects of the regulation of fuel sup- X yr age + 0.00000102 yr age2 X kg WT (29)
ply in omnivorous animals. Adv Enzyme Regal
1972; 10:397-420. 9) Owen et al (this study) regression equation:
71. Cathcart EP, Cuthbertson DP. The composition and Athletes: RMR (kcal/24 h) = 50.4 + 21.1 kg WT
distribution of the fatty substances of human subjects.
J Physiol l931;72:349-60. Nonathietes: RMR (kcal/24 h) = 795 + 7.18 kg Wi’

10) Harris and Benedict equation:

Appendix
RMR (kcal/24 h) = 655.0955 + 1.8496 cm HT
Formulas, methods, and theory
+ 9.5634 kg Wi’ - 4.6756 yr age (4)
1) Body Mass Index equation:
11) Mayo Foundation equation:
BMI (kg/m2) = kg WT/m HT2 (24)

2) Body Surface Area equation: RMR (kcal/24 h)

BSA (m2) = 0.007184 kg WT#{176}425

X cm HT#{176}725
(5) = BSA (m2) X tabulated value X 24 (6)

Underwater weighing for determining densitometric 12) Robertson and Reid equation:
analyses was conducted in a 5 X S X 5#{189}ft tank in which RMR (kcal/24 h)
a light metal chair with a 10-lb weighted belt was suspended
from a Chatillion 15-kg scale following the method out- = BSA (m2) X tabulated value X 24(8)
lined by Goldman and Buskirk (25). The underwater-
weighing procedure was repeated six or more times until 13) Cunningham equation:
three similar readings nearest to 25 g were obtained. Re- RMR (kcal/24 h) = 501.6 + 21.6 kg LBM (12)
sidual lung volume was determined by the nitrogen wash- 14) Storage (g/4 h) = intake - (oxidation + excretion
out technique (26).
3) FATMD (kg) = kg WT (4.570/density - 4.142) (25) + quantity of fuel left in extracellular space)

Density = kg WT in air/ Extracellular space (1) = blood volume (1)

([(kg WT in air - kg WT underwater)/water density] X 2.6 blood volume from nomogram (29)

- residual lung volume - 0.10) Theory of indirect calorimetry

4) FFMD (kg) = kg WT - FATMD (kg) (26) The energy expenditure of humans can be determined
Skinfold thicknesses were measured at the chest, axilla, accurately by measuring respiratory gaseous exchange rates
triceps, subscapular, abdomen, suprailium, thigh, and knee coupled with urinary nitrogen excretion; furthermore, the
with a Lange caliper (Cambridge Scientific, Cambridge, nature and quantity of fuels oxidized can be determined.
MD). The caliperfor measuring skinfold thickness had a Protein oxidation can be calculated from the quantity of
constant tension of 10 g/mm2 and measurements were nitrogen excreted in the urine. Carbohydrate and lipid ox-
taken on the right side of the body. Skinfold thicknesses idation rates can be determined from the nonprotein re-
were measured to the nearest 0.5 mm. Published recom- spiratory quotient (npRQ) tables of Lusk as modified by
mendations were followed to obtain body density data Cathcart and Cuthbertson (71), which are based on npRQ
(27). The three skinfolds used for calculations were triceps, of 0.707 for 100% fat oxidation and 1.00 for 100% car-
suprailium, and subscapular (27). bohydrate oxidation.
5) FATMSF (kg) = (4.95/body density - 4.5) kg WT
(27) Measured parameters for indirect calorimetry

Density = 1.0094921 I. 02 consumption (ml/min) at STP (Metabolic Cart)

II. CO2 production (ml/min) at STP (Metabolic Cart)
- (9.929 X l0 X sum of three skinfolds) III. RQ (Metabolic Cart)
IV. Urine sample: measure total volume (ml) and
+ (2.3 X l06 x sum of three skinfolds squared)
elapsed time (mm)
-(l.392X l0Xyrage) V. Urine total nitrogen (mg/ml and mg/mm)

6) FFMSF (kg) = kg WT - FATMSF (kg) Cakulated parameters for indirect calorimetry

7) Lean Body Mass equation: I. Factor for calories/liter 02: Table IV4.7 from ref-
LBM (kg) erence (31)
II. Percent calories of the npRMR derived from fat
= (69.8 - 0.26 kg Wi’ - 0.12 yr age)kg WT/73.2 (28) and carbohydrates: Table IV4.7 from reference (31)

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018
 CALORIC REQUIREMENTS OF WOMEN 19

III. npRQ: VIII. Fat (cal/mm) = RMR (cal/mm) X % fatcalories.

a. Urinary-nitrogen excretion, 1 mg = 4.75 ml CO2 IX. Carbohydrate (cal/mm) = RMR (cal/mm) X % car-
production and 6.02 ml 02 consumption. bohydrate calories.
b. Protein CO2 production (ml/min) = N2(mg/min) fat calories
X 4.75. X. Fat % calories = total RMR X 100.
c. Protein 02 consumption (ml/min) = N2 (mg/ carbohydrate calories
mm) X 6.02. XI. Carbohydrate % calories =
total RMR
d. Nonprotein CO2 production = total CO2 (ml/
X 100.
mm) - protein CO2 (ml/min).
protein calories
e. Nonprotein 02 consumption = total 02 (mI/mm) XII. Protein % calones = X 100.
- protein 02 (mI/mm). total RMR
XIII. Total postprandial metabolism (kcal/4 h) = area
f n ‘ RQ = Nonprotein
Nonprotein CO2
02 (ml/min)
(ml/min) under RMR vs time curve
-OR-
IV. npRMR(ca1/min) =02 (cal/mI) X np O2(ml/min). total postprandial metabolism (kcal/4 h)
V. Protein (cal/mm) = nitrogen (mg/mm) X 26.4.
VI. RMR (cal/mm) = npRMR (cal/mm) + protein (cal/
interval
mm). = 2 x(Y1+Y+2XY1). i-2
VII. RMR (kcal/24 h) = npRMR (kcal/24 h) + protein XIV. Thermic effect of food = total postprandial metab-
(kcal/24 h). olism (kcal/4 h) - baseline RMR (kcal/4 h).

Downloaded from https://academic.oup.com/ajcn/article-abstract/44/1/1/4691946

by guest
on 23 February 2018