Food

Klaas R

quotient,
H e.sterierp This composition are correlated. on have low-fat show the same studies The body ability to paper ofthe diets that

respiratory

quotient,

and

energy

balance12

ABSTRACT nutrient balance lose more weight fat but Descriptive subjects. with its elusion the fat a ready

reviews diet and total and gain energy

evidence the studies weight

that show on subjects

the that high-fat eat

macroof energy

Dietary

fat-carbohydrate balance fat is the main

ratio

maintenance

Intervention overweight

subjects Eneigi diets. Dietary relatively

determinant energy 16. energy

ofthe for dietary 16. and 37 depot in

energy

density

of has well. than (4). with surwhen

intake

as nonoverweight

our diet. The metabolizable protein. and fat is. respectively. as an minimal

carbohydrate. kJ/g (3). Fat the body The higher stores reached an energy intake energy from high in ate MJ/d). intake as

compared an important function The conFat can be stored with is that becoming overweight can be prevented by reducing density ofthe fat stores content of the diet. Studies on nutrient utilization showis the energy density of increase in carbohydrate oxidation whereas fat oxidation Thus. in most circumstances meals enriched with, in the long term, the food diets, must loss weight quotient eating the RQ mobilize in subjects (FQ) subjects respectively, respiratory carbo- the minimum quotient plus is mainly eating high- Some studies diets. is the is supported a standard intake energy they FQ. Duncan low density was cross-over twice versely. prisingly. quotient, respiratory protein. fat quotient, energy subsequent intake on (6) one consume food in the of our macronutrients is a fairly at nationwide intake 6 to changes fats of 1910 of to the in 43 diet fats 54 one Within of which wide energy studies ranges culture in food Western to en% to reason evidence ofthe the in en#{176}% . whereas or supply. countries. diet 1985(Fig for energy the for has I). from to maintain form diet. The range balance of nutrient the protein There In steadily The energy distribution between of can cultures be and intake dietary is an the shift of This correlation and the paper United increased in the fats will between maintenance balance. protein, of food and maintained. (Table The I). the fat is at changes increasing States from comhas often discuss of overFood and en-14 d mean fat, provided with in a as change in energy (6.5 design much twice there as to et al (5) kJ/g). bulk stored show a diet allowed density Each

has a limited ability oxidize carbohydrate

to oxidize fat and protein.

additional

weight. eight times (glycogen) can and energy a higher to for satiety a diet be diets.

energy

is approximately the carbohydrate energy high-fat fat. people a lower to and provided kJ/g) was change or eat from balance

Downloaded from www.ajcn.org by guest on November 18, 2011

does not change after hydrate or fat. However, (RQ) fat For is closer diets to the than it is for that on body with subjects

consuming as body that with (3 diet

for subjects high-carbohydrate than fat. This from energy a higher

density. a diet energy nearly or. conSuron energy et al a low. by for a to diet from en% fat)

high-carbohydrate

is lower body changing while for available.

subjects

indicating by data maintenance increased

5 d in a randomized

diet to a low-fat diet, even nearly 20% Direct . evidence diets is not yet

with a weekend on the high-energy little was no on the trend

in between. Subjects diet (12.5 MJ/d), diet a higher diet or high-energy different with diets. a high subjects intake diet (6.5 energy

expenditure for low-fat Nutr I 993:57(suppl):759S-65S. KEY balance, Introduction Humans ergy the tries. ratios shown carbohydrate ranges 1 1 en%. place ability contribution
en% in

..1,;i J (liii

low-energy toward

WORDS diets,

Food carbohydrate,

days on subsequent subjects medium. carbohydrate a balanced of on the trend dietary the were the like diet 8.7

the low-energy days on the with and sequence. MJ/d on the diet (45-50 over the time three one and fat.

to a lower diet. Lissner one energy got ( 1 5-20 each increased with

density

exchanging ofcarbohydrate, differs value

All low-fat

Energy

is consumed macronutrients the There (c:p:f) by from with

between

energy 9.8 MI/d on coun- MJ/d systematic any as tation intake subjects least weight availdiet. take Animals ergy density the intake and 32 14-d to on

the medium-fat high-fat diet ofintake treatment. during

(30-35 en% fat) and en% fat). Again. there in any that ofthe there period. subjects was On the no

to I 1.4 was no during adapaverage, diet. lost high-fat ifthe en-

carbohydrate-protein-fat intake

indicating observation

3 to 82 en% country.

in energy balance low-fat diet. and the laboratory rat

on the medium-fat gained weight on compensate to achieve as controls in time

of the food reach the same

is changed hod weight

the same (7). The

energy failure

position been weight. some energy .Ini J (l/ii of

a higher

contribution increasing the diet balance.

quoted ie,

as the and

incidence

of a positive composition

reasons balance.
\zuir

macronutnient

I From the Department of Human Biology. University of Limburg. Maastricht. The Netherlands. the 2 Address reprint requests to KR Westerterp. Department of Human of Biology. University of Limburg. P0 Box 616. 6200 MD, Maastricht. The Netherlands.

1993:57(suppl):759S-65S.

Printed

in USA.

1993

American

Society

for Clinical

Nutrition

759S

760S TABLE 1 Ratio of macronutrients Country

WESTERTERP in in the diet Carbohydrate en% of people of different Protein countries* Fat etary hand. subjects 12 12 11
11

endurance of those intake. freely with

exercise studies, were

(15, mainly

16).

We using

do self

not

know to

whether measure On ordinary the intake

the diother while living The

results

report habitual under

influenced impossible choose a more their

by this to own accurate as used feasible subjects Romieu the intake. role

phenomenon. foods

it is nearly

measure alternative

Nigeria Japan India CSSR United States Greenland

*

82 77 77 59 46 3 from reference I.

6 11 12
30

conditions double-portion ofthe best with daily they came be eat to the independent

technique.

technique but is not really routines when something. conclusion of energy utilizatioui that obese Thus, that

by Jiang and Hunt ( 1 1) is one in large groups and interferes are not at home every time et ( 1 al 7) offat and intake Miller (18) may in obesity

12 43

42 54

Adapted

Substrate Knowing of humans the food may dilution Energy to compensate be a way with sucrose light humans the for for them carbohydrate polyesthers products. compensate food to compensate part ofthe items for food consumption a change to avoid or On fat has the for are of in the energy density overweight. like of do

subjects but

generally do differ

do

not from

eat

more

than

normal-weight

subjects

normal-weight

tame (8, 9) and new market for indications that not sions (10). versions. Obesiti There are quite all by increasing

high-energy-density failed substituting

subjects with respect to the fat-carbohydrate ratio of their diet, aspar- the next question is. why is an isoenergetic diet fattening when its application in the contains it relatively more fat and less carbohydrate? In terms other hand. there are ofenergy, we can make carbohydrate from protein and fat from energy dilution when carbohydrate and thus fat and carbohydrate are not essential reduced-energy ver- nutrients. However the conversion processes are very energy nonmanipulated items consuming and produce a lot ofwaste products such as ammonia analogues in energy with items high-energy intake and urea. Ideally. the of fat and carbohydrate. hydrate whereas fat her. Additionally, metabolizers (ie. of as well). body is less covers Some bulky its energy needs tissues preferentially to consume, with a mixture use carboear-

becoming

Downloaded from www.ajcn.org by guest on November 18, 2011

Subjects

increases

by covertly

as mentioned

humans part ofthe

are periodic energy intake

eaters and continuous is stored before usage

a few

studies

on

the

macronutrient Here allowed

ratio

the diet and obesity or selves to studies where

we restrict our- The storage capacities for fat and carbohydrate are very difto freely choose ferent and this may have consequences for the regulation of their own foods under ordinary living conditions. Under those body weight. Flatt ( 1 9) proposed a model with a regulated carcircumstances, overweight subjects take diets with a higher energy bohydrate (glycogen) store and with the fat store as a function density. Jiang and Hunt ( 1 1) asked 1 1 adult men to collect a of nutrient intake and nutrient utilization. An increase in the double portion of all they ate over 7 d for analysis. The energy fat content of the diet needs an increase in fat oxidation and it density of normal-weight prisingly, the the diet, including subjects to 3.9 researchers mention was subjects no alternative drinks, ranged kJ/g in overweight that the energy not higher explanation from 1 .8 subjects. density kJ/g in is hypothesized Sunthat the latter needs an expansion of the body-

being overweight. subjects were

of the

diet in overweight fat content, and

because of a higher is presented. Dreon 50

et al ( 12) related diet composition, as measured with 7-d food C records, to body composition. as measured with hydrostatic a) weighing, in 55 middle-aged 1 men. Subjects with a higher percentage body less carbohydrate. study d food positively in 244 fat male consumed Tremblay adults, also and a diet with relatively et al ( 1 3) performed measuring diet composition fat found that the en% the en% carbohydrate of the subjects. in 2 16 adult that and adiposity negatively more fat a comparable with and a) a 3-a)

40

record. They correlated

of the diet was 0 was negatively Miller 50% et alC 0 men conwith

30

correlated (14) made

with the fatness similar observations and fat

Finally, subjects, was

20

and 50% women, related with dietary dietary above and be carbohydrate ( 1 1- I 4), there

found content

positively correlated

10
C 0
0

.-

-4--

S.

consumption. In all four studies mentioned was no correlation between energy intake or obesity. that self-reported habitual intake. intake Reported tends intakes to

0 1920
I 940

indexes for overweight There have been indications an underestimate be lower than with doubly with of true energy labeled

1 960

1 980

year
of dietary energy fat (5) in the United intake between States from carbohydrate 1910 to 1985

tend to taneously and

expenditure as measured water, especially in obese intake. such as those

simulFIG I. Distribution subjects (H). protein (#{149}). and engaged (from ref 2).

subjects

a high

energy

FQ,
fat mass, Several explaining groups the studied increased the effect adiposity ofa change

RQ,
in subjects

AND

ENERGY
a test sumed

BALANCE
comparable diet. The to higher the one diet, the mentioned a mixed carbohydrate above. diet, content rate medium, or

76 1S
Subjects a high-carboofthe after the and because most of there 0.6 with initial carhigh was con-

consum-

ing high-fat diets. in nutrient be on nutrient with indirect dioxide times utilization. calorimetry Nutrient from utilization can oxygen consumption, excretion primed more be single

a low-carbohydrate

intake hydrate

production, and protein oxidation amino gas of acid: provides nutrient measuring

urinary nitrogen is measured by measuring a more utilization the effects the rapid, can ofa

3C-labeled in respiratory (2 1). studies. term with Studies

elimination split on

measured diet. the higher the carbohydrate oxidation carbon bohydrate meal: 177 5. 241 11, and 258 (20). Some- h after consumption of 500 g in the low, infusion of carbohydrate a group, respectively. Carbohydrate ofthe label higher than it was in the first experiment (22), in index of the use of short-term the carbohydrate and long- was a significant a different surplus lipid However. dietary carbohydrate. was stored as glycogen, synthesis mediumthe limited from and carbohydrate

9 g over the 14 oxidation possibly Again, although of 3.4 groups, combined

responsive meal,

usually studies. or without

covering at least one an adaptation period system lying

24-h cycle beforehand.

a fixed diet and 9.0 The indirect- respectively.

1 .0 g in the

high-carbohydrate lipogenesis

calorimetry with subjects in on a respiration 10-20

in short-term studies or sitting. Long-term that space. allows

chamber

is a ventilated hood, the extremely high carbohydrate intake was judged to be unstudies are performed important in daily life. Finally, Flatt et al (24) studied the effects subjects to move around of the addition of fat to a standard, mixed meal. Fat and carbohydrate consumption oxidation (Table is fat begins. was not influenced 2). In the postabsorptive (2), 1with a rapid of the shift fat content by the state of increased the main when the food confat fuel

m2 offloor

Short-term
Short-term in the consumed and (22) observations measured as

studies
studies

of nutrient
of nutrient after

utilization
utilization after which an baseline are overnight usually fast. Acheson meal

of the feeding started sumed. Food is

body

to carbohydrate

independent

Downloaded from www.ajcn.org by guest on November 18, 2011

postabsorptive are the

condition continued effect of rate:

a breakfast,

measurements

Long-term

studies

of nutrient
balance

balance
should ideally cover of the several at least diurnal laboraperform the as conmeacon-

for several hours. a large carbohydrate

et al (9 Mi. Observations

of nutrient a multiple utilization

Table
lowered

2) on
the

nutrient
fat oxidation

utilization.
however.

The

high-carbohydrate
there fat. were no indications

loada 24-h
pattern

interval or of nutrient

of 24 h because (25). Nowadays.

of the conversion processing of the terval, probably only Acheson the storage sufficient 1 33 g carbohydrate et al (23)

of carbohydrate to meal in the subsequent capacity was controlled for to accommodate the

Assuming complete tories have facilities such as respiration chambers to 10-h observation inthese studies. There have been at least four studies on carbohydrate as glycogen was sequences of nutrient exchange for nutrient utilization the intake surplus because sured over 24 h (Table 3). In their diet over the next 3-6 experiment. d preceding Hunni suming Ct al (26) a mixed measured diet and. nutrient subsequently. utilization in subjects a high-carbohydrate,

oxidized.

TABLE 2 Food intake

and

nutrient

utilization Oxidation Carbohydrate Protein Percent en% Rate g/h Percent en% Rate g//z Fat Percent en%

Studs

(ref)

and

c:p:f

Intake fIji

Observation time Ii

Rate g//i

Acheson

et al (22) en% Acheson et al (23) 100:0:0 ent l00:0:0enc 100:0:0 en Flatt et al. (24) 62:27:11 en 35:l5:50enc 35:15:50 en
93:5:2
*

9.0 8.4 8.4 8.4 2.0 3.6 3.6

10 14 14 14 9 9 9

13.3 12.6 17.2 18.4 9.4 9.4 9.4

66 60 75 84 42 41 40

2.9 2.6 2.4 2.4 3.1 2.8 3.2

15 12 II 11 14 12 14

1.7 2.6 1.4 0.5 4.3 4.6 4.7

19 28 14 5 44 47 46

Carhohydrate:protein:fat ratio. 3-6 3-6 solution d preceding d preceding flavored with fruit juice. test. 14:1 1:75 en%. test. 60: 12:28 en. 80:1 1:9 en%. form of medium-chain triglycerides.

t Dextrin-maltose
:t c:p:fdiet c:p:f diet

II c:p:fdiet 3-6 #{182} Three-quarters

d preceding test. of the fat in the

762S
TABLE 3 Mean 24-h food quotient (FQ) with different carbohydrate:protein:fat Study (ref) and c:p:f Duration ofdiet d 26 44: 16:40
78:16:6 28

WESTERTERP
and respiratory ratios quotient (RQ) (c:p:f) from four was higher than fat intake, for diets synthesis or oxidation. studies Hill et al (30) measured 3 and FQ RQ and the .41J/d 7 d on a high-fat subjects three diet. were for in different Unfortunately. energy balance: assuming nutrient diets: there utilization a mixed they they did diet, not presented FQ. was in no net protein after diet, data there explacalwhether only

subjects a low-fat report

Energy balance

on diet composition was a tendency 0.4

0.5

and nutrient utilization. RQ to be lower than is the food-table difficulty nutrient

For all diets Hill et als indirect (27).

en% en% en% en% en% en% en% en% en% eri% en% en%

7 7 0 0 5-43 6-32 3 7 3 7 3 7 different from

1.4

0.85
0.95

0.80 0.0 1
0.88 0.01*

-1.9

nation orimetry

for

the discrepancy combined with

in using analysis

45:15:40
82:15:3 29 43:15:42

-0.1 -0.1

0.85 0.96

0.82 0.01 0.87 0.01 0.84 0.01 0.91 0.88

0.75 0.01 0.75 0.01 0.79 0.01

Nutrient
Dallosso

addition
and lames

and

nutrient
(3 1 ) increased

balance
energy after intake a l-wk for 1 wk by

65:15:20 30 20:20:60 20:20:60 35:20:45 35:20:45 60:20:20 60:20:20

-0.3 0.3 0.83 -0.1 0.2

-

0.77 0.77 0.83

50% with fat, mainly as double 0.011 on a maintenance diet with were observed on days when on days when they and protein balances the judging

cream.

observation Subjects level and to fat,

a c:p:f of 57: 1 3:30 en%. they had a low activity activity affected balance +151 respectively.

had a high were not fat -14 high of days energy and change in fat activity.

level. The carbohydrate by the addition of fat was from from on 33 mainly g/d The these high-active stored on mean figures days. days net Schutz as + 14 25 to +163 with

0.83 0.92
0.92
<

0.78 0.00 0.86 0.02 0.86 0.01

<

diet.
27 g/d

The from and in

5-MI the from and

supplement 19 to calculated 5. 1 Ml

Downloaded from www.ajcn.org by guest on November 18, 2011

*t Significantly

FQ: P *

0.05,

tP

0.001.

low storage Mi

activity on

fat

terms

is 5.5

low-active

et al (32) did a comparable experiment in which they measured low-fat diet. Both diets were consumed for 7 d, with a 2-wk energy expenditure and substrate utilization over 2-d intervals washout period in between, and measurements of nutrient uti- subjects in getting a maintenance diet on the first day (c:p:f 50: lization took place on the last day of each 7-d period. Unfor- 1 5:35 en%) and the diet with the same amounts of carbohydrate tunately. subjects were in negative energy balance: intake was and protein but twice the amount of fat on the second day. On on average 14% lower than expenditure during the 24-h respi- the first day, the mean difference between intake and expenditure ration-chamber ance, intake, within hydrate diet flexibility Lean energetic consumption sured fat diet. penditure fasting Mean plus with carbohydrate even the compared ofthe and James of and, 5%, exchange on error the range more with body the (28) measurements. and protein diet. than of the mixed Despite oxidations However, methods on diet. metabolic one meal. on than 24-h was 24-h the applied doubled carbohydrate the over evening after based higher the negative were these lower results (27). illustrating effect interval diet fasting mean Subjects The energy than are bal-was their The 0.1 24-h 0.2 FQ
MI.

indicating mean day changed RQ that

subjects values subjects the energy by the

were were

in 0.87

energy and

balance. 0.85 0.01,

and

still respectively. carbo- well. On utilization

indicating the second were not

were in nutrient balance as expenditure and substrate fat supplement. The energy extra intake as observed I wk with

oxidation

high-carbohydrate the metabolism. ofan after were

low-carbohydrate measured

to use of nutrients

for energy

balance became +4. of4. I 0. 1 Ml, that before (3 )1 Acheson . iso-

1 0.3 Mi, compared with the is, the fat supplement was stored et al (33) overfed subjects for

a standard subsequently, intake, was 1 .7% energy individual

a low-fat

carbohydrate after a 3-d interval with a restricted diet to deplete the stores. The energy-restricted diet was 6.7 Ml/d. mea-the glycogen c:p:fof 10:15:75 en%. whereas energy expenditure on the and a high-with third day was 9.6 Ml. Subsequently. subjects got a diet with a energy exenergy were

c:p:fof86: 1 1:3 en%, increasing energy intake over 7 d from I 5.5 exto 2 1 .0 MI/d. On the first day ofoverfeeding. the energy surplus stored as glycogen: then, de novo lipogenesis started in energy balance). Nutrient utilization was closer to nutrient was fully making up all of the energy surplus from the fifth day of overintake for the high-fat diet than for the low-fat diet. comparing feeding onwards. At the end of the overfeeding period. there the calculated FQ with the presented RQ value (Table 2). gain of 0.7 kg and a fat gain . 1of kg, 1 together Abbott et al (29) measured energy metabolism in subjects was a glycogen representing 75% of the energy consumed in excess of mainteafter 5-43 d on high-fat and low-fat diets in a metabolic ward. nance requirements. penditure, differences
< all 10%

(ie, subjects

Part
first in

of the

subjects

ate

the

high-fat

diet

on

the

first

admission

and

the and energy

others second

started the

with

the was

low-fat more difference 5%) was were intake

diet. than within

The between the

interval Subjects energy error on

between wereDiscussion intake

admission mean (<

4 wk.

and

conclusion

balance: expenditure

and

energy

both measurements. fat diet, indicating were atically the same. lower On than

FQ and RQ that nutrient the FQ < 0.00 (P

not different and nutrient diet, RQ that

high-carbohydrate 1), indicating

range of Based on the literature referred to. there is evidence that a the high-change to a fattier diet leads to an increase in body weight. Comutilization bining this with the fact that obese people tend to eat more fat was systemleads to the conclusion that becoming overweight can be prevented by reducing the fat content of the diet. Secondly. there fat oxidation

FQ,
is evidence compared that with the the body ability has a limited

RQ. AND
to and

ENERGY fat References

BALANCE

763S

ability

oxidize protein. are

to oxidize

carbohydrate

pattern and nutritional adaptations. Prague: AcainI . Fabry P. Feeding demia. and London: Butterworths. 1969. Research Council (US). Committee on Diet and Health. more likely to become obese consuming a high-fat diet than are 2. National Diet and health: implications for reducing chronic disease risk. others. On the other hand. intervention studies do not support Washington. DC: National Academy Press. 1989. unequivocally the bodys limitations to burn fat. 3. Atwater WO. Bryant AP. The availability and food values of food Short-term studies, measuring substrate utilization over up to materials. In: The I 2th annual report ofthe Storrs. CT. Agricultural 14 h after a meal, show how the carbohydrate oxidation is inExperimental Station. Storrs, CT: Storrs. CT. Agricultural Expericreased after a high-carbohydrate meal whereas addition of fat ment Station, 1900:73-I 10. to a meal does not influence fat oxidation (Table 2). In the long 4. Bray GA. Bethune JE. eds. Treatment and management of obesity. term. measuring substrate utilization over 24 h, there are inHagerstown. MD: Harper & Row, 1974. dications of the opposite. At diets higher in carbohydrate there 5. Duncan KH. Bacon IA. Weinsier RL. The effects of high and low is a bigger discrepancy between FQ and RQ such that RQ is energy density diets on satiety. energy intake. and eating time of subjects. Am I Clin Nutr 1983:37:763-7. lower than FQ (Table 3). This should mean that substrate uti- obese and nonobese L, Levitsky DA. Strupp BJ. Kalkwarf HI. Roe DA. Dietary lization is closer to substrate intake for diets higher in fat than 6. Lissner of energy intake in human subjects. Am J for diets higher in carbohydrate. a phenomenon that cannot be fat and the regulation Clin Nutr 1987:46:886-92. readily explained. It is often suggested that this obesity-susceptible individuals limitation (34). Thus is more pronounced some individuals Under must ance, near1 not use energy equal all conditions RQ. energy protein Under in the and The ofperfect conditions adult form carbohydrate stores energy and ofenergy or mobilizes, fat. store Then. for in the reserves of body nutrient or balance. nutrient in the the energy long body storage FQ imbalterm. does of liver or 9.
8.

7.

Adolph EF. Urges 1 10-25.

to eat and

drink

in rats.

Am

I Physiol

1947:151:

a normal-weight

Porikos KP. Booth G. Van Itallie TB. Effect ofcovert nutritive dilution on the spontaneous food intake of obese individuals: a pilot study. Am I Clin Nutr 1977:30:1638-44.

Downloaded from www.ajcn.org by guest on November 18, 2011

Porikos KP. Control offood intake in man: response to covert caloric dilution of a conventional and palatable diet. In: Cioffi LA, James and muscle glycogen fluctuates between 250 and 500 g, or 4 and WPT, Van Itallie TB, eds. The body weight regulatory system: normal 8 Ml. Reference man has a muscle mass of 30 kg with 7.5 kg and disturbed mechanisms. New York: Raven Press, 198 1:83-7. protein, or an energy equivalent of 120 Mi. but changes in mus- 10. Foltin RW. Fischman MW, Emurian CS, Rachlinski. Compensation cle mass are insignificant in terms of energy compared with for caloric dilution in humans given unrestricted access to food in laboratory. Appetite I 988:10:13-24. changes in fat mass, unless body fat is nearly depleted. Thus, in a residential I 1. Jiang C-L. Hunt IN. The relation between freely chosen meals and the long term, an RQ higher than the FQ implicates conversion body habitus. Am I Clin Nutr 1983:38:32-40. of carbohydrate or protein to body fat. and an RQ lower than mobilization. carbohydrate form the for FQ, a mobilization ofenergy observation diet with from body fat. RQ measured
12.

aged men. Am I Clin Nutr 1988:47:995-1000. I 3. Trembla A. Plourde G. Despres I-P. Bouchard C. Impact of dietary over a long-term interval can only be lower than the FQ by fat content and fat oxidation on energy intake in humans. Am I mobilizing body fat leads to the conclusion that high-carbohyClin Nutr 1989:49:799-805. drate diets induce body fat loss. Apart from studies referred earlier 14. Miller WC. Lindeman AK. Wallace I. Niederpruem M. Diet com(5. 6), a recent study suggests that the macronutrient composition position, energy intake. and exercise in relation to body fat in men of the diet plays a role in the energy requirement for weight and women. Am I Clin Nutr 1990:52:426-30. maintenance. Prewitt et al (35) reported data on 18 women con- I 5. Schoeller DA. How accurate is self-reported dietary energy intake? suming a standard diet for 4 wk (c:p:f44:19:37 en%), followed Nutr Rev 1990:10:373-9. by 20 wk of low-fat diet (c:p:f 60: 19:2 1 en%). Energy intake of Westerterp 16. KR. Verboeket-van de Venne WPHG. Meijer GAL. Hoor the subjects was adjusted to maintain body weight throughout F ten. Self-reported intake as a measure for energy intake. a validation against doubly labelled water. In: Ailhaud G. Guy-Grand B. Lafontan the study period. that is. intake was increased or decreased when M. Riequier D. eds. Obesity in Europe . London: 91 John Libbey. body weight decreased or increased > by 1 kg, respectively. Comparing
1992:17-22. diet with I. Willett WC. Stampfer MI, et al. Energy intake and other intake increased I 7. Romieu determinants of relative weight. Am I Clin Nutr I 988:47:406-12. with 2 kg. Thus. energy intake, and nutritional status a high-carbohydrate diet resulted in a significant reduction in 18. Miller WC. Diet composition. in relation to obesity in men and women. Med Sci Sports Exerc body weight despite a substantial increase in energy intake aimed I991:23:280-4. at weight maintenance. 19. Flatt IP. The difference in the storage capacities for carbohydrate This review leads to the suggestion that energy expenditure is and for fat. and its implications in the regulation of body weight. higher for low-fat, high-carbohydrate diets than for high-fat, lowAnn NY Acad Sci 1987:499:104-23. carbohydrate diets. Convincing evidence for this at this point is 20. Frayn KN. Calculation of substrate oxidation rates in vivo from not yet available. Current studies include measurement of energy gaseous exchange. I AppI Physiol 1983:55:628-34. expenditure over short time intervals, up to 24 h in a respiration 2 1. Garlick P1, McNurlan MA. McHardy KC. Factors controlling the chamber, and calculation of energy expenditure from energy disposition of primary nutrients. Proc Nutr Soc 1988:47:169-76.

Combining the a high-carbohydrate

of the lower-than-expected the fact that an RQ

Dreon Wood

DM, Frey-Hewitt B, Ellsworth PD. Dietary fat:carbohydrate

N, Williams PT, Terry RB, ratio and obesity in middle-

the

initial

4-wk

interval

for

the

standard

the last 4 wk for the low-fat diet, the mean energy with 19% and the mean body weight decreased

intake include under

and normal

changes living

in body conditions.

composition. of energy

Future expenditure

studies for wk 1

should 22.

direct

measurement

#{163}3

Acheson KJ. Flatt IP, l#{233}quier Glycogen E. synthesis versus after a 500 gram carbohydrate meal in man. Metabolism I 234-40.

lipogenesis 1982:31:

7645
23.

WESTERTERP diture in humans: effects of dietary fat and carbohydrate. Am I Physiol 1990:258:E347-5 I. Hill 10. Peters IC. Reed GW. Schlundt DG. Sharp T. Greene HL. Nutrient balance in humans: effects ofdiet composition. Am I Clin Nutr 1991:54:10-7. Dallosso HM. James WPT. Whole-body calorimetry studies in adult men: 1 The effect of fat over-feeding . on 24 h energy expenditure. Br I Nutr 1984:52:49-64. Schutz Y. Flatt IP. J#{233}quier Failure ofdietary E. fat intake to promote fat oxidation: a factor favoring the development of obesity. Am I Clin Nutr 1989:50:307-14. Acheson KI. Schutz Y. Bessard T, Anantharaman K. Flatt IP, and J#{233}quier Glycogen E. storage capacity and de novo lipogenesis during massive carbohydrate overfeeding in man. Am I Clin Nutr I 988:48:
240-7.

Acheson KI. Schutz Y. Bessard T. Ravussin E. l#{233}quier Flatt IP. E. Nutritional influences on lipogenesis and thermogenesis after a carbohydrate meal. Am I Physiol 1984:246:E62-70. 30. 24. Flatt JP. Ravussin E. Acheson 10. l#{233}quier Effects of dietary E. fat on postprandial substrate oxidation and on carbohydrate and fat 31. balance. I Clin Invest 1985:76:1019-24. 25. Verboeket-van de Venne PHG. Westerterp KR. Influence of the feeding frequency on nutrient utilization in man: consequences for energy metabolism. Eurl Clin Nutr 1991:45:161-9. 32. 26. Hunri M. Burnand B, Pittet Ph. lequier E. Metabolic effects ofa mixed and a high-carbohydrate low-fat diet in man. measured over 24 h in a respiration chamber. Br I Nutr 1982:47: 33. 33-43. 27. Livesey G. Elia M. Estimation of energy expenditure, net carbohydrate utilization. and net fat oxidation and synthesis by indirect calorimetry: evaluation of errors with special reference to the detailed 34. composition offuels. Am J Clin Nutr 1988:47:608-28. 28. Lean MEl, James WPT. Metabolic effects of isoenergetic nutrient exchange over 24 hours in relation to obesity in women. Int I Obes 35. 1988: 12:15-27. 29. Abbott WGH, Howard BV, Ruotolo G. Ravussin E. Energy expen-

Zurlo F. Lillioja S. Esposito-Del Puente carbohydrate oxidation as predictor of RQ. Am I Physiol l990:259:E650-7. Prewitt TE. Schmeisser D. Bowen PE. et both composition. and energy intake in fat diets. Am J Clin Nutr 199 1:54:304-10.

A. et al. Low ratio weight gain: study

of fat to of 24-h

al. Changes in body weight. women fed high- and low-

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Discussion Michael
interested test tube

animal studies

studies. because we have

but got

you

cant

really have far reason

compare a big intestine lower energy

those

with for food

human storage per energy of car-

we as humans a relatively

A Crawford:
in the and the analogy barrel.

This
the

may
that you test

be a naive
gave tube

question.

but
models

I was and

metabolism

us-your of carbohydrate out of showed

of the unit body and the expenditure

weight. Maybe the at a high-carbohydrate to fat.

for having a higher diet is the conversion

barrel of fat. What seems the size of the input you One ofcarbohydrate and

Eric Ravussin: would I like to expand a little on this comment. and I think that in men there is less and less evidence that de novo different diets. It seems to me that if you had represented those lipogenesis plays an important role. and in the study that you inputs in proportion to the daily amount ofstuffthat was going showed from Kevin Acheson, in which he gave an enormous in, one would have seen that the amount of carbohydrate that amount ofcarbohvdrate (500 g). We tried that first as one meal. went in over a 24-hour period would be really very much larger but after the subject had problems, we offered it at 250 grams than the amount of carbohydrate that was stored in the test and at two times 125 grams. It was very, very difficult to induce tube. A 24-hour input of carbohydrate is quite a lot in a higha significant amount ofde novo lipogenesis. I agree that indirect carbohydrate diet. At the same time. the barrel is full of fat, so calorimetry doesnt differentiate between oxidation and going that carbohydrate. although coming through the carbohydrate into fat. but I think that the point is that there is little room for route, is actually filling up the fat barrel because it doesnt store de novo lipogenesis in men. Therefore. you are right when you that amount of carbohydrate. So. it has to be converted to fat say that maybe 50- 100% of the glycogen stores are replaced on and fill up the fat barrel. The energy process ofconverting that a daily basis vs< 1% for fat. I think that most ofthe carbohydrate carbohydrate to fat needs to be a sort of negative consideration ingested is either stored as glycogen or oxidized into CO2. I in terms ofthe overall energy balance ofthe individual. I wonder think that the enzymatic activities ofthe lipogenic enzymes are to what extent that contributes to disturbing these relationships very low in men, and recent studies using stable isotope showed and explaining why fat is going straight into the barrel and doesnt that de novo lipogenesis is very insignificant in man. really have any effect of energy. The carbohydrate going in demands energy to convert it into fat and destorts it as such. DoesWesterterp: I agree with that, but on the other hand. the studies
one offat ofdifferent proportions that confound vs a rich-fat Klaas I dont the diet? equation with regard to a rich-carbohydrate ofAcheson consume amount were short-term studies. diet. in your In the you cant glycogen long term. when store because this they you a high-carbohydrate of carbohydrate really stores

to be slightly showed. You

focus two

to me was bohydrate little pots.

R Westerterp: know whether

I think this

you model

are stressing was really

an interesting on the right proportion is not carbohydrate maybe

point.

and, as you say. we eat our carbohydrate stores. information on whether to or fat, two apart others from in the the one

a lot of carbohydrate in On the other hand, there we really have to convert study I showed in which in substrate part ofthe use. That you you, can

are just not big enough. So. you have to do something with it scale when you are in a positive energy balance. but apparently you to tend to be in a negative energy balance on a high-carbohydrate much diet. We do not yet have any reason for that. Maybe you have for why you tend diet. to be in a negative Why would your energy energy balance carbohydrate expenditure

and

literature

there is a day-night rhythm suggests that we might store during the day for overnight

utilization. carbohydrate has been

a suggestion one on a high see that at least go up? That rhythm we consume Ravussin: seen in small- study. you

In all the studies

need to push

ofoverfeeding.
a lot ofcarbohydrate

for example.
through the

Schutzs
system.

FQ. When viously a daily fore There might process. we basis. carbohydrate was be the higher I dont British talk about There weight energy is limited is oxidized: school gain balance. storage maybe in the l960s in obese but by there that it capacity

RQ. people.

AND they amount and lipogenesis. are

ENERGY obthan no on lication

BALANCE change from the for the pilot rest study of the for year. the There

765S
is a recent pub-

in positive

a minute in men, is a little said that

breast-cancer-dietary-fat that weight the year. trial. see There indeed the but nothing trials, also the that man are

there- trial, women like again

where on the something

you see a similar the very-low-fat diet you like would 1 or

phenomenon. lost some from the

lipogenesis

amounts

expect 2 kg over

short-term

in lean people because know what is the status

is an energy-costly of that now.

data from the I think is still only the more

Seattle dietary in press, where

alternative again you vs 40 en% mechanisms to resist

a 2-year study in middleaged

Nlartijn effectiveness cause the been

Katan: of final

Could

I sum

a little

note

of

caution

about

I kg weight loss on 20 going on than just the about. The carbohydrate-rich Maybe body seems diets. are right.

of fat. that weight

I think there is we have been loss even on the

high-carbohydrate diets in reducing tests are long-term control trials In were weight 1 or at 1970 where given loss most diets in the National large of the 2 kg groups 40% in the or 20-en%-fat first Diet of

a few

ofthose? in (irculation, men

obesity betalking and there have more Heart Study was middleaged energy group few but months as

published American There was was very

Westerterp:

you

The

only On costly.

problem the other Therefore,

is the control hand. longwe have

20%

some minor,

fat. offood intake in these types ofstudies. that term experimental feeding studies are and accept to the results as they are.

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