Energy affected

Rudolph ABSTRACT
ing to

intake required to maintain by wide variation in diet

L Leibel, Diets Jules rich Hirsch, in fat may
of

body weight composition13

Gregg One C Checani

is not

Burton promote

E Appel, obesity

and by lead-

group of investigators concluded that fat intake may play in obesity that is independent of energy intake (4). do isoenergetic diets with less fat. This possibility was examined The ultimate test of the validity of such a proposition is the by a retrospective analysis of the energy needs of 16 human long-term effect on metabolic efficiency of perturbations of diet subjects (1 3 adults, 3 children) fed liquid diets ofprecisely known composition. Using the technique of caloric titrimetry (9), in
a greater deposition adipose-tissue triglycerides thana role composition with widely varied fat content, for 15-56 d (33 which 2 a formula diet ofspecified composition is fed over a period

d, i SE). Subjects lived in a metabolic ward and received fluid formulas with different fat and carbohydrate content, physical activity was kept constant, and precise data were available on energy intake and daily body weight. Isoenergetic formulas contamed various percentages ofcarbohydrate as cerelose (low, 15%; intermediate, 40% or 45%; high, 75%, 80%, or 85%), a constant
15% of energy as protein (as milk protein), and the balance

ofweeks examined changes

under circumstances ofcontrolled physical activity, we the effect on metabolic efficiency (as reflected by in body weight) ofextreme changes in the composition ofotherwise isoenergetic diets. This technique was made possible by the careful collection ofdata on the effects offormulafeeding,
in studies done over many years by EH Ahrens and 1 1). his colleagues

Downloaded from www.ajcn.org by guest on October 11, 2011

energy

as fat (as corn

oil).

Even

with

carbohydrate ratio (fat energy variedfrom intake), there was no detectable evidence in energy need as function a of percentage

extreme 0%

changes to 70%

in the of total
variation

at the of fatSubjects The

ofthe

Hospital and records

Rockefeller

of the methods

Rockefeller

University ( 10,

ofsignitlcant

fat intake.

Am

of all subjects
University

studied
Hospital

by

the

Lipid
1955

Laboratory
and 1965

Cliii

Nuir l992;55:350-5.

between

KEY WORDS diet composition, Introduction

Obesity, dietary energy requirements

fat,

dietary

carbohydrate,

who and
in the Protein

were fed liquid-formula fat composition were

research (20.9 kitchen Id/g), which of

diets ofvarious reviewed. These

the Rockefeller represented

carbohydrate diets were

University

(CHO) prepared
Hospital.

was
oil.

derived
CHO

from
(16.7

milk
kJ/g) was

protein.
supplied

Fat

a constant (37.7 kJ/g) was as cerelose,

1 5% of energy, from corn a hydrated form

the

calorimetry of these formulas are not available. calorimetry performed on eight batches ofsimilar formula physical activity (I). They found no evidence for such purely Bomb prepared in the research kitchen of the Rockefeller University endogenous obesity and also demonstrated that the long-term between April 1988 and July 1990 had a coefficient of effect of any diet onbody weight is related only to the total Hospital for Id/g of 1 .9%. Such formula shows a 0.2% corenergy content of the diet. Other features of the diet such variation as between bomb calorimetric and calculated energy carbohydrate or fat content did not, in the long run, have con- respondence content. In studies designed to examine the effects of dietary sequential effects on body weight. type and quantity on lipoprotein metabolism, the perIn recent years the adverse effects of high-fat diets on health lipid of CHO in the diet was systematically altered by isohave been emphasized, and the possibility that a high-fat intake centage may be accompanied by some special in vivo economy energy of metabolism has led to speculation on the role ofdiet composition From the Laboratory of Human Behavior and Metabolism and the in the production of obesity. Various investigators have reported Pew Center ofNutritional Excellence, Rockefeller University, New York. a positive correlation between body fat and the percentage of 2 Supported in part bygrants DK 30583 and RROO 102 from the Nafat in isoenergetic diets fed to rodents (2, 3) and humans (4, 5) tional Institutes of Health. and a lack of effect of dietary fat on respiratory quotient (RQ) 3 Address reprint requests to RL Leibel, Laboratory of Human BeI

Sixty years ago, LH Newburgh and his colleagues examined possibility that so-called endogenous obesity might be result of special metabolic factors unrelated to energy intake

of glucose. Coefficients the final available energy the(12): protein, bomb or garding
0.92; fat,

of digestibility value ofthe

0.95; and

were used to determine constituents ofthe formula

carbohydrate, 0.98. Data re-

in

human

subjects

studied

for

9 or

24

h after

the

feeding

of in

fat havior

and

Metabolism,

Rockefeller 1990. March 1992

University,

1230

York

Avenue,

New

(6, 7). Additionally,

24-h

energy

balance

in a respiratory
or

chamber

York,

NY

10021.
October 25, for publication in USA. 6, 1991. American Society for Clinical Nutrition

is reported to be due balance and unrelated 350

almost exclusively to carbohydrate

to differences protein balance

fat Received (8). Accepted Printed

Am J C/in

Nuir l992;55:350-5.

DIET
TABLE I Anthropometry

COMPOSITION

AND

ENERGY

REQUIREMENTS

351

and

diet

characteristics

of adults Diet

ID

Sex

Age y

Mean weight kg

WT kg

INTS cm

Height

Days BMIt on diet d

Percent CHO %

Percent fat %

Energy Id

CORREC Id kJ

1 2 3 4 5 6 7 8 9 10 11 12 13

M
M M M M M M M M M F F F

56
43 55 53 41 54 49 41 57 64 49 64 57
75.28

76.01
76.10 75.42 96.21 95.95 59.42 58.81 62.26 64.02 65.93
66.11

75.59
76.19 75.41 75.68 95.46 96.21 59.03 58.19 62.31 63.87 66.25
66.13

165
-

27.9
-

24
32 43 28 23 18 39 44 15 39 34 38 25 42 22 42 52 47 25
18

45
75 45 85 15 85 15 75 15 45 45 85 45 85 45 85 45 85 15 85 45 85 15 75 15 85

40
10 40 0 70 0 70 10 70 40 40 0 40 0 40 0 40 0 70 0 40 0 70 10 70 0

10067
10067 9937 9991 13862 13862 9 100 9100 10330 10330 10201 10201 9414 9414 10590 10590 10330 9284 8498 8498 7452 6929 7322 7322 7322 7707

9071
10242 10113 10519 12016 14652 8690 8280 10330 10096 10728 10200 9765 9 121 10590 10590 9657 9577 9493 9201 6895 6929 7468 6824 7322 7707

167.5
-

26.8
-

173.5
-

32.0
-

157.5
-

24.0
-

164.5
-

23.0
-

178
-

20.8
-

63.80 64.13 70.72 71.04 61.58 61.56 70.24 70.55 45.70 46.95 52.64 52.68 49.07 48.55 of regression change

63.95 63.92 70.87 71.09 60.98 61.79 70.68 70.78 45.42 46.92 52.84 52.38 48.98 48.57 line of weight the

164
-

23.7
-

Downloaded from www.ajcn.org by guest on October 11, 2011

170
-

24.5
-

175
-

20.1
-

165
-

25.8
-

163.5
-

17.1
-

156
-

21.6
-

155.5
-

20.3
-

28 18 56 34 15 28

Time

zero

intercept

vs days. period of formula feeding.

t Body

mass t Id corrected

index, in kg/rn. for any weight

during

energetic composition

replacement were fed

with in

fat. random

These order. Center

formula Vitamin

diets and

of

varied The

daily

weights

of each
vs time. functions changes

subject
We found as the

were

examined

by regression
ofearly changes proceeded, sug-

supplements
lived living in in with the

were physical

given

daily
Research

in addition was constant

The

to 2 g NaCI.
(Rockefeller

Clinical no additional sufficient

mineral analysis ofweight All subjects in slope for these University gesting that acute

no evidence 2-wk period excretion and

in water

retention

were

Hospital);
quantities

activity
to

at
formulas

activities
were body weight

of daily not
always fed likely

occurring
the result

with

changes

in diet
exactly

composition.
2 g NaCI/d

This
in all

is most
pe-

exercise. maintain

of feeding

dietary

as riods. Therefore, all days on a given diet were included in these nearly as possible. Interim adjustments in the quantity of formula analyses. The slope of each weight-vs-time plot examined was were made to achieve this goal. Weights (to nearest 0. 1 kg) were for significant deviation from zero. If P value of this relathe obtained each morning with the subject clothed in underwear. tionship was < 0.05, the energy requirement for the individual Though stools were not collected from these subjects, similar was corrected for this intercurrent change in weight. In making subjects on similar diets did not show differences in the fraction this correction we assigned the conservative (high) value of 29.3 of energy intake excreted in the stool (EH Ahrens, personal Id/g to the weight change. The corrected daily energy requirement was calculated as: communication, 1991). The records of all patients studied from 1955 to 1965 were KJ = Id formula 29.3 searched for consecutive periods in which patients were fed for- CORREC mula diets of differing CHO-to-fat ratios. For inclusion in this x (regression slope in grams per day). analysis, each period of formula feeding had to 2 wk and be the subject had to have remained weight stable (to within 1 kg) The time zero intercepts (WT INT) of these plots are given in within that period. Reasons for rejection of records were lackTable 1. The clinical characteristics and diet specifications for of feeding of two different formulas or the absence of feeding each adult subject are given in Table 1. periods of 2 wk. A total of 47 patient records were reviewed Many ofthese subjects were fed a no-fat diet for several weeks to obtain 16 (13 adults and 3 children) fitting the above criteria. at a time. At the time these studies were done, it was uncertain
a constant
-

352

LEIBEL

ET

AL of 13 adult subjects

whether essential fatty acid (EFA) deficiency would occur adult humans. Only one subject (subject 6, Table 1) developed symptoms possibly related to EFA deficiency (ie, mild eczema relieved by the addition of fat to the diet). This study was approved by the Institutional Review Board of the Rockefeller University. Energy requirements of each subject were expressed both absolutely and corrected for significant slope ofthe weight-vs-time

TABLE 2 in Anthropornetry and energy requirements on various carbohydrate diets

Low CHO (n=6)

Intermediate

CHO (n=8)
64.4 9791

High CHO

(n=12)
3.4 356 2.3 65.7 9401 53.5 3.9 527

Weight

(kg)

65.0
9406

6.9

Energy (Id)

1004

In addition each subjects requirements were expressed per unit of body surface area (13) or per amount of body weight#{176}73. Both body surface area and weight#{176}provide good indirect estimates of metabolic mass (14). Statistical analyses were performed on a VAX PDP 1 1/780 computer runningUNIX BMDP (1 5) and UNIXSTA T(16) programs.

plots.

kJCORREC Age(y) Days on diet (d) Wt#{176}75 (kg)

Energy/SA (U/rn) Energy/wt#{176}7
(U/IgO.7i)

9217732 55.7 3.5 28.8 6.5 22.8 1.8

5519305

9611431

9485607
2.1 3.1 1.0

48.7
33.4 22.9

3.8

32.4
23.0

0.9

5611 113
17 427 8

5422
410

151
8 180

410 ICJ/SA

CORREC

Results
Adults

(kY/rn)

CORREC
(J/lgO.7S)

/07

5439

180
13

5506
418

163
13

5448
410

406

* i SE. Thirteen subjects were studied on two diets, hence a total been maintained on low CHO formula, eight The characteristics of all subjects examined CHO, and 12 on high CHO. The mean period of 26 feeding periods. or high CHO) are given. was 32 2 d (i SE). A plot of the daily weighteach diet (low, intermediate, is shown in a typical patient (subject 12) who received a 10%fat diet for 5 wk followed by a 70%-fat diet for 8 wk (Fig 1). The of means (low high) = -51.5 Id; I= -0.2236; P small corrections, calculated as described, are shown in Table [difference

Six subjects on intermediate on each diet

had

on

Downloaded from www.ajcn.org by guest on October 11, 2011

1. The

absence

of effect

ofdiet

composition

on

energy

require-

0.8305).

the no-fat diet might have induced subclinical EFA ments to maintain weight are shown in Table 2. This table pro- Because a separate analysis was done ofthe nine subjects who vides summary data on the weight-maintenance energy require- deficiency, a 0-fat diet vs the 17 dietary periods of those on 10ments of the 13 adult subjects studied on two diets of differing received 70% fat intake. Linear-regression analysis of energy and CORpercent carbohydrate. Regression plots of diet vs various body Id vs SA and O.7S was performed for these two groups, size indexes for the three diet compositions are shown in Figure REC and ANOVA of regression coefficients over groups showed no 2. This figure examines the relationship measured of energy insignificant differences between lines of regression for no-fat and take (Id) and corrected energy intake (CORREC regressed Id) fat-containing diets (Table 4). Thus, there was no aberration of on surface area (SA) and also on weight#{176}75 (%,.J1O.l5). Although energy requirement in those receiving a no-fat diet. energy requirements were highly significantly related body to size indexes, analysis of variance (ANOVA) of regression coefficients over groups did not indicate any significant used in betweenthe Children

Table 5 shows data for the three children studied. Although need for weight maintenance was 30-40% higher (ku both energy (1 5%) and very-high-CHO (75-85%) diets were kg#{176}) than that for adults, no significant effect of diet composition on energy requirements seen in children, is as is the case separately examined. Paired t tests of the energy requirements with adults. (CORREC Id) of these subjects on the two diets indicated no difference due to diet composition [difference of means (low high) = -334.7 Id; t = -0.558; P = 0.6 1]. Similarly, a comDiscussion parison (paired t test) ofenergy requirements in the seven subjects who received both moderate-CHO (45%) and very-high-CHO The energy content ofoodstuffs f is conventionally determined by measuring heat released by the complete combustion of the (75, 85%) diets indicated no difference due to diet composition group
abscissa very-low-CHO

difference, regardless of the body (Table 3). Five of the subjects

size index

who

had

received

Days

1O%Fat

70% FatS

FIG 1. A l3-wk study ofsubject 12, first 10% (75% on CHO) ofenergy intake as fat and then on 70% (15% CHO) ofenergy intake as fat. During both periods, 7322 U per day fed. Actual Id combusted were were corrected, described as in Subjects and Methods, to 7468 U and 6824 Id per day, respectively. It is notable that there was only 1 kg of weight variation during the entire study.

DIET

COMPOSITION

AND

ENERGY

REQUIREMENTS

353

14000

16000
0

12000
-)

14000
-)

12000 10000

c
8000

DO 0

10000 0
U

8000

6000

6000 1 .4 1 .6 1 .8 2.0 2.2 2.4

1 .4
1 .6 1

.8

2.0

2.2

2.4

SA

SA

14000

16000
0

12000 ,
.

14000
C
C.)

12000

10000

Downloaded from www.ajcn.org by guest on October 11, 2011

10000

0
8000 .00 6000 15 20 25 30 0.75
&

8000
6000

c0

35

15

20

25

30

35

wr

.
0

LOWCHO=15% INTERMEDIATE
HIGH CHO = 75%,

CHO = 40% or 45% 80% or 85% or weight#{176} (WT#{176}75) for intermediate, and

FIG 2. Regression liquid-formula diets

CHO

diets.

analyses of 24-hr of different CHO All subjects are adults.

energy content.n

intakes vs either body surface area (SA) body 6, 8, and 12, respectively, for low-CHO,

high-

specific

components

of the

diet.

A recent

analysis

of the

in foodstuffs combustion (corrected for

showed that of 37.690-40.091 urea) of

conventional Id/g. conventional

food fats have The metabolizable food protein (17).

energy value offoodstuffs as used to calculate total energy energy ofthe by indirect calorimetry may be in error 5% of by a heat expenditure of energy the true value even under the most careful circumstances (17, from 1 8). There could in the metabolic bution energy to systemic as the and actual release be even handling fuel measured of greater error because if, of different foods, oxidation in vitro. is not Specifically,

varies

of differences
the directly actual predicted the ATP to contriby retheir of could terms of

I 7.43 1 to 2 . 11 50 Id/g. CHOs vary glucose, 1.560 Id/g; 5 and sucrose,

less:
16.481

starches,
kJ/g

17.48 1 Id/g;
An analysis

TABLE
Regression

3 equations Plot (y= rnx

+

b) for the plots Equation

in Figure r

2 P(cx)

quired for incorporation glucose reduce to the

activation into proteins from glycogen energy

amino acids and the ready via of these

preparatory interconversion steps in

energy-requiring substrates

value

A Low CHO Intermediate HighCHO CHO y = 9699x y = 699 1x y=9025x-6146 y y

= = =
-

6950 2393

0.92 0.88 0.93 0.94 0.83 0.96 0.92 0.79 0.92 0.93 0.72 0.95

0.009 0.005 0.000 0.004 0.0 16 0.000 0.010 0.020 0.000 0.008 0.050 0.000

TABLE

4
regressions for each of the for differences in regression

B
Low CHO Intermediate High CHO CHO 7243x 7979x 9443x
-

2992 4297 8962

For each of the four plots, the separate three levels of CHO feeding examined coefficients by ANOVA* Plot A: Id vs SA

ANOVA

P (a)
0.761 0.351 0.522 0.328 the occasional small

C
LowCHO Intermediate High CHO CHO y=5l5x-2330 y 310x y = 473x
-

2720 1477 590 1883 3343

D
Low CHO Intermediate High CHO

B: CORREC C: Id vs O.75 D: CORREC

*

U vs SA Id vsO.73
was
=

F14,01 = 0.464 F14201 = 1. 1 77 F14,,1 = 0.830

F14201 =
1.236

y
CHO y y

= = =

38 1 x 339x 556x

There

no significant
body

correlation stores
(r =

between

shifts in apparent (r = -0.027, P

energy 0.89) or O.75

(Id - CORREC 0.15, P = 0.94).

U) and either

SA

354
TABLE 5 Anthropometry and diet characteristics of children

LEIBEL

ET

AL

Diet Mean weight kg 19.91 20.77 20.80 21.44 26.19 25.88 Days on diet d 15.0
-

ID

Sex y

Age

WT kg 19.66 20.37 20.59 21.18 26.10 25.91

INT cm 115
-

Height

BMI

Percent CHO % 40 80 40 80 40 80

Percent fat % 40 0 40 0 40 0

Energy k.J 6146 6146 5623 5623 6590 6590

CORREC kJ

14 15 16

M M F

9 10 8

115.5
-

15.7
-

137
-

20.0
-

36 41 40 30 42 23

5766 5590 5330 5125 6590 6590

net the

energy presumably

provision lower offree

for

in vivo rate and acids and

chemical energy might

respiration. cost result of in greater

Conversely, hydrolysis efficiency relative in the

Although and each

the 33

number

ofsubjects

studied

was

relatively

small,

individual

was

studied

for extended

periods

of time

(15no

reesterification ofhandling in net in effect of

fatty

ofdietary fats vivo generation feeding when cause of foods

hence of ATP. are

an apparent Differences compared with

56 d, i increase difference thermic content or fat extended design in tions: our

d) on two in energy were conducted

diets. Comparable requirements on for a maximum

studies concluding diets of differing of ( 1 9-22). 6 d The

Downloaded from www.ajcn.org by guest on October 11, 2011

CHO

CHOs for in

protein

period
of this assignment

of our
study, as ofa

diets
value

adds
of29.3 by

considerable
by the kJ/g using the to any

power
following

to
calcula-

the

is yet an additional the energy value measures Short-term stantial promote that frame adult who this ( (4l00 the of heat

potential systematic vivo as compared h) in humans

differences with laboratory that

demonstrated

intercurrent

of combustion. (24-48 suggest does immediately not (7). Our results in a more storage extended of dietary

weight
a subbeing

changes
I 3- 1 7 Id/g.

is extremely
Nonetheless, interval g/d) and the days 95%

conservative,

a more
figure

likely
slope by

figure
Id!

studies

of 29.3

Id) dietary fat supplement use offat as a metabolic fuel does to This that produce point human the not operate preferential

g, the confidence suggest vs days (8.2 timesubject fat in energy minus Id/32

(level 0.95) around the the total energy ingested confidence on diets equals limits ofdiffering 2.7% (range)

of weight each adult differences equals

phenomenon 8 wk)

ofthe composition

requirements which

humans. emphasized

was in fact made subjects achieve operation studying

by Schutz et al (7), 7853 weight equilibrium gested.

of total

kilojoules

in-

Thus,

despite

the

relatively

small

number

of

subjects

on high-fat diets despite nomena. Other investigators,

phe-studied, when the large number of total kilojoules fed to each time subject during each diet period are taken into account, it is unthat we failed to detect a5% difference in energy effiperiods than we used here, reported no effect on various aspects likely ofenergy expenditure (24-h expenditure, resting metabolic rate) ciency between diets. However, very small differences in the of energy metabolism integrated over prolonged peof highvs low-CHO diets (19-23). On the other hand, Prewitt efficiency oftime can theoretically lead to clinically significant shifts et al (24) recently compared the effects on energy need and body riods composition of switching 18 outpatient female subjects (BMI in body composition. For example, 2% greater metabolic effi18-44) from a 37% fat diet (1 4 wk) to a 21.4% fat diet (1 ciency 20 of one isoenergetic diet vs another would (assuming

of these short-term subjects over shorter

wk).
required ofbody weight.

On average,
an average fat) despite Differences

while

ingesting the lower fat diet, these subjects 10 460 Id/d intake and 29.3 Id/g weight gain) lead to a 3.2 kg gain in 1 y. of 10.5% more calories and lost 2.8% (1 1.3%weight efforts to feed sufficient calories to maintain A formal analysis ofthese data can be made. However, such in physical activity and noncompliance on analysis is very sensitive to sample size and cannot readily be

to account for the special power of this study in the large the lower-fat diet may account for some of the observed differ- made ences. number ofdays for which each subject was maintained on each Forbes (25) recently reviewed a series of studies in which hudiet. With this caveat in mind, the /3 (type II) error can be esman subjects were overfed for 2- 1 2 wk and found that although timated in two ways. 1) The variance about the slope of the obese individuals appear to deposit a larger proportion of excess regression equation relating CORREC Id to surface area (plot
energy intake as fat, the energy cost or obese subjects was not significantly position of the excess energy be fed. our data and those rodents are growing The lack in rodents ously, of agreement (2, 3) may between because ofweight gain influenced in either by the leanB, Fig 2) may com- ferences ofany

on

formulas

be used to estimate the likelihood desired magnitude between energy of different composition. Assuming,
in energy efficiency (CORREC (75% or 85%) formula (n = 12 adult

of missing difrequirements for example,

Id) conferred studies) and by

obtained a 10% decrease continu- the high-CHO

introducing

a possible

to diet composition. lack of diet-composition children in this study libitum intake (permitting was designed

proportional effect ofgrowth itselfon the response variance probability ofa /3 error Against this explanation is our finding of effect on energy requirements in three P = 0.1 1 (26). 2) A direct ofCORREC Id for the (Table 5). These rodents were also fed ad and 75% weight gain), whereas our human subjects (intermediate) = -0.2236, P(a) = 0.8304, to maintain body weight.

to

that

for

the

high-CHO
null

formula,
hypothesis)

the
is

(acceptance

of a false

comparison seven adult

or 85% and SD

between the distributions subjects studied on both (high) CHO gives paired
ofdifferences
=

45%

607

Id.

For

DIET Pu.
=

COMPOSITION

AND

ENERGY

REQUIREMENTS

355

WGH, Howard BV, Christin L, et al. Short-term energy between-diet difference is 912 Id (one tail). That 8. Abbott balance: relationship with protein, carbohydrate, and fat balances. of a difference between the two diets as large Am J Physiol l988;255:E332-7. as 9 12 kJ/d escaping detection is 0.05. This difference represents 9. Leibel RL, Hirsch J. Reduced energy requirements in reduced-obese 9.5% of total energy intake per day. For reasons discussed patients. Metabolism l984;33:164-70. above, such calculations overstate the probable error in this study. 10. Ahrens EH, Insull W, Hirsch J, et al. The effect on human serumSimilar results might not have been obtained in a group of lipids of a dietary fat, highly saturated, but poor in essential fatty obese individuals or lean individuals susceptible to obesity. There acids. Lancet l959;l:l15-9. may be an interaction of diet composition and predisposition 1 1. Ahrens EH. The use ofliquid formula diets in metabolic studies: 15 to obesity, with higher dietary fat content facilitating the expresyears experience. Adv Metab Disord l970;4:297-332. sion ofa more efficient metabolic phenotype (27). In this regard 12. Watt BK, Merrill AL. Composition of Foods: raw, processed, preit is of interest to note that the three subjects (subjects 1-3 in pared. Agriculture handbook no. 8. Washington, DC: US GovernTable 1 with highest BMIs had lower CORREC ) Id on the higherment Printing Office, 1963. 13. DuBois D, DuBois EF. A formula to estimate the approximate surface fat diets, whereas all but one of the other adult subjects had area ifheight and weight be known. Arch Intern Med l9l6;17:863equal or higher CORREC U on the higher-fat diets. It is also 71. possible that fats putative enhanced efficiency as a metabolic M. The fire of life. An introduction to animal energetics. substrate is seen mainly in circumstances ofpositive energy bal-14. Kleiber Malabar, FL: Robert E Krieger, 1975. ance. 15. Dixon WJ, ed. BMDP statistical software manual. Berkeley, CA: Finally, the physical activity of some of our subjects was University of California Press, 1988. somewhat diminished by their restriction to a metabolic ward. 16. Perlrnan G. Unixstat. Data analysis programs for UNIX. Tyngsboro, There may be interactions between physical activity and diet MA: Wang Institute ofGraduate Studies, 1986. composition that predispose to fat accumulation (28); such in17. Livesey G, Elia M. Estimation of energy expenditure, net carboteractions were not examined in this study. hydrate utilization, and net fat oxidation and synthesis by indirect It has been noted that obese individuals have a preference for calorimetry: evaluation of errors with special reference to the detailed high fat-foods (29). This preference, based on palatability, may composition of fuels. AmJ Clin Nutr l988;47:608-28. favor obesity by increasing total caloric intake. However, from 18. Elia M, Livesey G. Theory and validity ofindirect calorimetry during Am J Clin Nutr 1988;47:59 1-607. our results here and with the caveats mentioned above, such fat net lipid synthesis. WPT. Metabolic effects of isoenergetic nutrient preference probably does not provide obese individuals with an19. Lean MEJ, James exchange over 24 hours in relation to obesity in women. Int J Obes intrinsically more-efficient fuel source. Variations in fat intake l988;12: 15-27. from 0% to 70% oftotal energy under conditions ofequal energy 20. Hurni M, Burnand B, Pittet P, Jequier E. Metabolic effects of a intake produced no significant changes in body weight over pemixed and a high-carbohydrate diet in man, measured over 24 hours nods of observation averaging 33 d. 0 in a respiration chamber. Br J Nutr 1982;47:33-43. These studies could not have been performed without the generous 21. McNeill G, Bruce AC, Ralph A, James WPT. Inter-individual difhelp and advice of Dr EH Ahrens. Two anonymous reviewers provided ferences in fasting nutrient oxidation and the influence ofdiet cornhelpful comments. position. Int J Obes l988;12:455-63. 22. Abbott WGH, Howard BV, Ruotolo G, Ravussin E. Energy expenReferences diture in humans: effects of dietary fat and carbohydrate. Am J Physiol 1990;258:E347-5 1. 1 . Newburgh LH, Johnston MW. Endogenous obesity-a misconcep23. Hill JO, Peters JC, ReedGW, Schlundt DG, Sharp T, Green HL. tion. JAMA 1930;3:8l5-25. Nutrient balance in humans: effects ofdiet composition. Am J Clin 2. Schemmel R, Mickelsen 0, Motavi K. Conversion of dietary to Nutr l991;54:lO-7. body energy in rats as affected by strain, sex and ration. Nutr J 24. Prewitt TE, Schmeisser D, Bowen PE, et al. Changes bodyin weight, l972;l02:1 187-97. body composition, and energy intake in women fed high- and low3. Salmon DMW, Flatt JP. Effect ofdietary fat content on the incidence fat diets. Am J Clin Nutr 199 1;54:304-lO. ofobesity among ad libitum fed mice. Int J Obes l985;9:443-9. gain weight more easily than non4. Romieu I, Willett W, Stampfer MJ, et al. Energy intakes and other 25. Forbes GB. Doobese individuals obese individuals. Am J Clin Nutr l990;52:224-7. determinants of relative weight. Am Clin Nutr 1988;47:406-l J 2. RR, Rohlf FJ. Introduction to biostatistics. San Francisco: 5. Dreon DM, Frey-Heywitt B, Ellsworth N, Williams PT, Terry RB, 26. Sokal WH Freeman, 1973:116-27. Woud PD. Dietary fat:carbohydrate ratio and obesity in middle27. Sims EAH Jr. Expenditure and storage of energy in man. J Clin aged women. Am J Clin Nutr 1988;47:995-l000. Invest l987;79: 10 19-25. 6. Flatt JP, Ravussin E, Acheson KL, Jequier E. Effects ofdietary fat FX. Exercise and obesity. Med Clin North Am on postprandial substrate oxidation and on carbohydrate and fat28. Segal KR, Pi-Sunyer 1989;73:2 17-36. balances. J Clin Invest 1985:76:1019-24. 7. Schutz Y, Flatt JP, Jequier E. Failure ofdietary fat intake to promote 29. Drewnowski A, Brunzell JD, Sande K, Iverius PH. Sweet tooth refat oxidation: a factor favoring the development of obesity. Am J considered: taste responsiveness in human obesity. Physiol Behav Clin Nutr l989;50:307-l4. l985;35:6 17-22. is, the 0.05, the probability

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