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Food

Klaas R

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

respiratory

quotient,

and energy

balance12

ABSTRACT nutrient balance lose more Descriptive fat but

reviews diet and total and gain energy

evidence the studies weight intake

that show on

the that

macroof energy subjects diets. relatively

Dietary Eneigi Dietary

fat-carbohydrate balance fat is the main

ratio

maintenance

Intervention overweight

weight

high-fat eat

determinant energy 16.

ofthe

energy

density

of has

subjects

as nonoverweight

our diet. The metabolizable protein. and fat is. respectively. an important Fat can density is the Thus. plus they Duncan low density ofthe energy in most is mainly studies et al
(5)

for dietary carbohydrate. 16. and 37 kJ/g (3). Fat depot in the weight. eight times (glycogen) can and energy to satiety a diet high diets. body The

subjects. The body has a limited ability with its ability to oxidize carbohydrate elusion is that becoming overweight the fat content a ready increase

to oxidize fat compared and protein. The conby reducing

function with

as an energy minimal

as well. energy

be stored

additional

can be prevented

of the diet. Studies on nutrient utilization show in carbohydrate oxidation whereas fat oxidation meals enriched with, respectively, in the long term, the respiratory quotient diets, must weight the mobilize (FQ) eating RQ for subjects high-carbohydrate is lower body changing than from is the is supported a standard fat. This carboquotient highdiets. FQ.

fat stores is approximately density of the carbohydrate circumstances bulk stored show allowed Each diet to a diet density consuming as body that with energy fat. people a lower subjects (3 kJ/g) and change to eat from balance high-fat

higher than stores (4). with surwhen density. a diet in energy ate nearly or. conMJ/d). intake Suron an energy intake

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be reached

does not change after hydrate or fat. However, (RQ) For is closer than that on body fat diets indicating by data maintenance increased
it

the minimum Some

to the food

eating

is for subjects

change in energy

or a higher

energy from

high-carbohydrate subjects

loss in subjects

(6.5 kJ/g).

was provided

for 5 d in a randomized

diet to a low-fat diet, with nearly 20% . Direct is not

even while energy intake was evidence for a higher energy yet available. ..1,;i J (liii

cross-over design with a weekend twice as much on the high-energy versely. prisingly. twice there as little was on the no trend

in between. Subjects diet (12.5 MJ/d), diet a higher (6.5 energy

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

low-energy toward

WORDS diets,

Food carbohydrate,

quotient, protein.

respiratory fat

quotient,

energy

subsequent days on intake on subsequent (6) provided one with exchanging subjects a medium. carbohydrate of 8.7

the low-energy diet or to a lower energy days on the high-energy diet. Lissner et al with and sequence. three one and on the fat. different with Energy low-fat diets. one with density diet from en% fat) a low. by for a to a high intake diet energy increased ( 1 5-20

All subjects

got each

consume

food in the of our

to maintain form diet. The

energy distribution between can

balance. protein, of food cultures be and

Food and and

enfat,

14 d in a balanced mean value

is consumed macronutrients the There (c:p:f)

ofcarbohydrate, differs

MJ/d

energy counas I). The

between

macronutrients is a fairly at which intake wide energy studies ranges culture

9.8 MI/d on the medium-fat diet (30-35 en% fat) and MJ/d on the high-fat diet (45-50 en% fat). Again. there systematic any tation subjects weight 14-d trend dietary diet to the ofintake treatment. during the over time indicating observation in any that ofthe there period. subjects was On

to I 1.4 was no during no adapaverage,

range

of carbohydrate-protein-fat maintained. (Table the intake fat intake is at least take availStates from has will between maintenance the 32 often discuss the of intake protein dietary In the steadily I). The (Fig United increased shift incidence paper the of fats

balance of nutrient

by nationwide from with

carbohydrate Within

from

3 to 82 en% the There has

6 to 54 en#{176}% . whereas one changes of fats in food to the in supply. countries. diet 1985 for the energy evidence ofthe

were in energy balance on the low-fat diet. and like the laboratory

on the medium-fat gained weight on rat compensate

diet. lost the high-fat ifthe en-

or country.

changes

is an increasing

diet. Animals

in time

of fats 1910

in Western to 43 en% diet as the and reason

contribution position been weight. some energy

ergy density of the food is changed to achieve intake and reach the same hod weight as controls

the same (7). The

energy failure

in the comof over-

of the quoted

to a higher

contribution increasing This and balance. for the diet

ie, of a positive of reasons balance.


\zuir

correlation

macronutnient

composition

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

.Ini J (l/ii

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

Printed

in USA.

1993 American

Society

for Clinical

Nutrition

759S

760S TABLE 1 Ratio of macronutrients Country

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

exercise studies, were

(15,

16).

We using

do

not

know

whether to measure intake

the diwhile living The

of those intake. freely with it is nearly

mainly

self report phenomenon. habitual under

influenced impossible their

by this own

On the other ordinary technique.

to measure foods accurate as used feasible subjects Romieu the role intake.

choose a more

Nigeria Japan India CSSR United States Greenland


*

82 77 77 59 46 3 from reference I.

6 11 12
30

conditions

alternative

double-portion technique ofthe best but is not really with daily routines when they came eat something. Thus, that of energy to the conclusion

by Jiang and Hunt ( 1 1) is one in large groups and interferes are not at home every time et al ( 17) and offat intake Miller (18) may in obesity

12 43

42 54

be independent Adapted Substrate Knowing of humans the food Energy to compensate may dilution be a way with for a change for them carbohydrate to avoid in the energy becoming density overweight. like asparof do the subjects with

utilizatioui that obese subjects but to the generally do differ do not from ratio eat more of their than diet,

normal-weight respect

subjects

normal-weight

fat-carbohydrate

or fat analogues

tame (8, 9) and sucrose new market for light indications that humans not sions (10). all high-energy-density by increasing Subjects failed substituting the

polyesthers has its application in the products. On the other hand. there are compensate for energy dilution when food consumption part ofthe items are reduced-energy in energy with veritems intake high-energy of nonmanipulated for increases food items

next question is. why is an isoenergetic diet fattening when it contains relatively more fat and less carbohydrate? In terms ofenergy, carbohydrate nutrients. consuming we can and However and make carbohydrate from protein and fat from thus fat and carbohydrate the conversion processes a lot ofwaste products are not essential are very energy such as ammonia a mixture use carboearwith

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produce

to compensate

by covertly versions. Obesiti There

and urea. Ideally. the body covers its energy needs of fat and carbohydrate. Some tissues preferentially hydrate whereas fat is less bulky to consume, her. Additionally, humans metabolizers (ie. part ofthe

as mentioned

are periodic eaters and continuous energy intake is stored before usage for fat and carbohydrate for with fat store An the the are very difof carin the

are quite

a few

studies

on the

macronutrient

ratio

of

the diet and obesity or being overweight. Here we restrict ourselves to studies where subjects were allowed to freely choose their density. double own foods under overweight ordinary subjects living take conditions. diets with Under a higher those energy circumstances, Jiang portion

as well). The storage ferent body bohydrate of nutrient and weight. this

capacities may have store and Flatt

consequences a model with utilization. and

regulation as a function increase

( 1 9) proposed nutrient

a regulated

(glycogen) intake

and Hunt ( 1 1) asked 1 1 adult men of all they ate over 7 d for analysis.

to collect a The energy

density of the normal-weight prisingly, diet in overweight fat content, and

diet, including drinks, ranged from 1 .8 kJ/g in subjects to 3.9 kJ/g in overweight subjects. Sunmention that the energy density of the subjects was not higher no alternative explanation diet composition, composition. as middle-aged men. because of a higher is presented. Dreon

fat content of the diet needs an increase in fat oxidation and it is hypothesized that the latter needs an expansion of the body-

the researchers

et al ( 12) related records, to body weighing, in 1 55

as measured with 7-d food measured with hydrostatic Subjects with a higher permore fat and a comparable with a 3-

a) a)
a)

50

centage body fat consumed less carbohydrate. Tremblay study in 244 male adults, also and d food record. They positively correlated correlated (14) made

a diet with relatively et al ( 1 3) performed measuring diet composition found that the en% the en% carbohydrate

40

fat of the diet was was negatively Miller 50% et al men conwith

30
0 C 0

with the fatness of the subjects. Finally, similar observations in 2 16 adult subjects,

20

and 50% women, and found that adiposity was positively related with dietary fat content and negatively correlated dietary above and carbohydrate ( 1 1 - I 4), there

10
C 0
0

.-

-4--

S.

consumption. In all four studies mentioned was no correlation between energy intake intake Reported tends intakes simulsubjects engaged to

0 1920
I 940

indexes for overweight or obesity. There have been indications that self-reported underestimate of true energy labeled energy habitual intake.

1 960

1 980

be an

year
FIG I . Distribution (H). protein (#{149}). and (from ref 2). of dietary energy intake between fat (5) in the United States from carbohydrate 1910 to 1985

tend to be lower than taneously with doubly and subjects with

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

a high

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

RQ,

AND
consumintake measured carbon

ENERGY
a test sumed hydrate diet. h after higher

BALANCE
comparable diet. the higher consumption group,
it was

76 1S
to the higher the one diet, the carbohydrate of 500 respectively. in the mentioned a mixed oxidation g in the low, carbohydrate above. diet, content rate medium, possibly Subjects ofthe after the and coninitial carhigh was because

in subjects in nutrient

ing high-fat diets. on nutrient with indirect utilization. calorimetry utilization can be oxygen consumption,

a low-carbohydrate The

or a high-carbo-

bohydrate carbohydrate than

meal: 177 5. 241 11, and 258 9 g over the 14 Carbohydrate (22), oxidation

dioxide production, and times protein oxidation 3C-labeled in respiratory (2 1). Studies studies. term with usually amino acid: gas provides of nutrient measuring

urinary nitrogen excretion is measured by primed measuring a more utilization the effects the rapid, can ofa elimination more single be split on

(20). Someinfusion of a ofthe label index and long-

first experiment

responsive meal,

in short-term a fixed diet The indirect-

of the use of a different the carbohydrate surplus was and the a significant 9.0 lipid However. high 1 .0 g in the

dietary carbohydrate. was stored as glycogen, synthesis mediumthe limited from and carbohydrate

Again, most of although there of 3.4 0.6 groups, with to be uncombined judged

studies. covering at least one or without an adaptation period system lying m2 offloor

24-h cycle beforehand.

high-carbohydrate lipogenesis was

respectively. extremely

calorimetry with subjects in a respiration on 10-20

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

is a ventilated hood, studies are performed to move around

carbohydrate

intake

chamber

subjects

important in daily life. Finally, Flatt of the addition of fat to a standard, bohydrate consumption oxidation (Table begins.

et al (24) studied the effects mixed meal. Fat and carby the state increased the main food fat fuel when con-

was not influenced 2). In the postabsorptive a rapid of the shift

Short-term
Short-term in the consumed

studies
studies

of nutrient
of nutrient condition after

utilization
utilization after which baseline are usually fast. started Food is

of the feeding sumed.

body

is fat (2 1 ), with independent

to carbohydrate of the

fat content

Downloaded from ajcn.nutrition.org by guest on March 9, 2014

postabsorptive

an overnight

as a breakfast,

measurements Acheson et al meal (9 Mi.

Long-term
Observations a 24-h pattern tories these sequences

studies

of nutrient
balance

balance
should ideally cover of the several at least diurnal laboracon-

and observations (22) measured

are continued for several hours. the effect of a large carbohydrate

of nutrient

Table
lowered

2) on nutrient
the fat oxidation

utilization.
rate:

The

high-carbohydrate
there were

load

however.

no indications

interval or a multiple of nutrient utilization have studies. facilities There of nutrient such have been

of 24 h because (25). Nowadays. chambers four studies at least

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

of carbohydrate to fat. Assuming complete meal in the subsequent 10-h observation incapacity for carbohydrate the diet intake the In their over as glycogen surplus next 3-6 experiment. d preceding was because to accommodate was oxidized. controlled the

as respiration for nutrient

to perform on the as meacon-

exchange

utilization in subjects

1 33 g carbohydrate

sured over 24 h (Table 3). Hunni Ct al (26) measured nutrient suming a mixed diet and. subsequently.

utilization

a high-carbohydrate,

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. flavored with fruit juice. test. 14:1 1:75 en%. test. 60: 12:28 en. triglycerides.

t Dextrin-maltose

solution :t c:p:fdiet 3-6 d preceding c:p:f diet 3-6 d preceding

II c:p:fdiet 3-6 d preceding test. 80:1 1:9 en%. #{182} Three-quarters of the fat in the form of medium-chain

762S

WESTERTERP
was higher than fat intake, assuming nutrient diets: balance: there utilization a mixed they did they diet, not presented was no net protein after diet, data whether only

TABLE 3 Mean 24-h food quotient (FQ) and respiratory quotient (RQ) for diets with different carbohydrate:protein:fat ratios (c:p:f) from four studies Study (ref) and c:p:f Duration ofdiet d 26 44: 16:40 en% 78:16:6 en%
28

synthesis or oxidation. Hill et al (30) measured 3 and 7 d on three a high-fat subjects diet. were different and the

in subjects a low-fat report

Energy balance .41J/d

FQ

RQ

Unfortunately. in energy

on diet composition was a tendency for 0.4


0.5

and nutrient utilization. RQ to be lower than FQ.

For all diets there Hill et als explacal-

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

1.4

0.85
0.95

0.80
0.88

0.0 1
0.01* 0.01

-1.9

nation orimetry

for the discrepancy combined with

is the difficulty in using indirect food-table nutrient analysis (27).

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

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

-0.1 -0.1 -0.3 -0.1


-

0.85 0.96 0.83 0.91


0.77 0.77 0.83

0.82 0.87 0.84 0.88


0.75 0.75 0.79

0.01 0.01

Nutrient
Dallosso 50% with

addition
and fat, lames mainly

and

nutrient
(3 1 ) increased

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

0.3

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

0.2

0.011
0.01 0.01 0.01

as double diet days with when

cream.

observation

on a maintenance were observed on on days when they and protein balances the

a c:p:f of 57: 1 3:30 en%. Subjects they had a low activity level and activity affected level. The carbohydrate by the addition of fat to was mainly 33 g/d The these from stored on days mean figures days. they 2-d net as fat,

had a high were not fat -14

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 terms did

5-MI from and high

supplement in fat balance activity. and 19 to +151

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judging low storage Mi et al (32)

the change

from

+ 14 25 to +163 with fat is 5.5 Schutz intervals 50: On

*t Significantly

FQ: *P

0.05,

tP

0.001.

activity

respectively. 5. 1 Ml

of energy days and

calculated experiment substrate

on low-active expenditure

on high-active in which utilization over

a comparable

measured

low-fat washout lization tunately.

diet. period took

Both place

diets on were

were the than protein diet. of the than last

consumed and measurements day of each energy

for

7 d, with of nutrient 7-d period. intake the lower results (27). The 24-h

a 2-wk utiUnforwas respibaltheir still carbothe isothe meaa highexexwere

energy

in between,

in subjects getting a maintenance diet on the 1 5:35 en%) and the diet with the same amounts and protein but twice the amount the first day, the mean difference was 0.1 0.2 MI. indicating The 24-h FQ and mean RQ respectively. indicating well. On the second utilization were not

first day (c:p:f of carbohydrate day.

subjects

in negative expenditure

balance: during were these

of fat on the second

on average ration-chamber ance, intake, within hydrate diet flexibility Lean energetic consumption sured fat diet. penditure penditure, fasting

14% lower and range more with body James the (28)

measurements. on the error mixed

Despite oxidations However, applied doubled

the negative

energy than are

between intake and expenditure subjects were in energy balance. values were 0.87 and 0.85 0.01, were in nutrient balance as expenditure and substrate fat supplement. The energy with the extra intake was stored as observed subjects for I wk with diet to deplete diet was 6.7 Ml/d. expenditure on the got a diet with a over 7 d from I 5.5 the energy surplus lipogenesis started the fifth day of over-

carbohydrate even the compared ofthe and exchange and, plus with

methods diet.

that subjects day the energy changed by the

oxidation

on the high-carbohydrate illustrating metabolism. ofan after were and energy energy to nutrient comparing 2). in subjects ward. between were intake range of effect interval diet fasting mean for energy metabolic one meal. on than all
<

low-carbohydrate measured evening after based the over

to use carbohydrate of nutrients

balance became +4. 1 0.3 Mi, compared of4. I 0. 1 Ml, that is, the fat supplement before (3 1 ). Acheson et al (33) overfed carbohydrate the glycogen with c:p:fof third day was after stores. 10:15:75 9.6 Ml. a 3-d interval with

24-h

a restricted

of a standard subsequently, intake, Mean energy 5%, was individual

Subjects

a low-fat 24-h was

The energy-restricted en%. whereas energy Subsequently. subjects

1 .7% higher differences

10% (ie, subjects

in energy balance). Nutrient utilization intake for the high-fat diet than for the the after calculated Abbott 5-43 d on FQ with et al (29) high-fat started the with the and presented energy low-fat measured

was closer low-fat diet. RQ value (Table metabolism

c:p:fof86: 1 1:3 en%, increasing energy intake to 2 1 .0 MI/d. On the first day ofoverfeeding. was fully stored as glycogen: then, de novo making up all of the energy surplus from feeding onwards. was a glycogen representing nance
75%

At the end of the overfeeding period. there gain of 0.7 kg and a fat gain of 1 . 1 kg, together of the energy consumed in excess of mainte-

diets diet. than within

in a metabolic The between the interval Subjects energy error

Part
and first and

of the subjects
the others and energy second balance: expenditure

ate the high-fat


the low-fat was mean (< 5%) more was difference

diet on the first admission


4 wk.

requirements.

admission

Discussion
Based change bining on

and
the

conclusion
literature referred to. there is evidence that a

in energy

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

FQ and RQ were that nutrient intake the high-carbohydrate FQ (P < 0.00

not different and nutrient diet, RQ that

on the highutilization was systemfat oxidation

to a fattier diet leads to an increase this with the fact that obese people

in body weight. Comtend to eat more fat be prethere

1 ), indicating

leads to the conclusion vented by reducing the

that becoming overweight can fat content of the diet. Secondly.

FQ,
is evidence compared that with the the body ability that this individuals obese hand. has a limited ability

RQ.

AND fat in are are support up to is inof fat

ENERGY

BALANCE

763S

to oxidize and protein. pronounced individuals diet do not than

References
I . Fabry

to oxidize

carbohydrate

It is often suggested obesity-susceptible more others. likely On to become the other

limitation is more (34). Thus some consuming intervention a high-fat studies

2.

unequivocally Short-term
14 h after

the bodys limitations to burn fat. studies, measuring substrate utilization a meal, does of the not show influence substrate opposite. how the carbohydrate meal fat oxidation utilization At diets between FQ over higher and RQ whereas (Table
24

over addition

3.

oxidation

creased to a meal term. dications

after measuring

a high-carbohydrate

2). In the long


h, there

4. 5. is 6.

are inthere RQ

in carbohydrate such that

is a bigger

discrepancy

lower than FQ (Table 3). This should mean lization is closer to substrate intake for diets for diets readily Under must ance, near1 not and use energy 8 Ml. protein, equal higher explained. conditions RQ. Under in the and The ofperfect conditions adult form stores energy and ofenergy or mobilizes, fat. in carbohydrate. a phenomenon

that substrate utihigher in fat than that cannot balance. be FQ imbalterm. does or of liver

P. Feeding pattern and nutritional adaptations. Prague: Academia. and London: Butterworths. 1969. National Research Council (US). Committee on Diet and Health. Diet and health: implications for reducing chronic disease risk. Washington. DC: National Academy Press. 1989. Atwater WO. Bryant AP. The availability and food values of food materials. In: The I 2th annual report ofthe Storrs. CT. Agricultural Experimental Station. Storrs, CT: Storrs. CT. Agricultural Experiment Station, 1900:73-I 10. Bray GA. Bethune JE. eds. Treatment and management of obesity. Hagerstown. MD: Harper & Row, 1974. Duncan KH. Bacon IA. Weinsier RL. The effects of high and low energy density diets on satiety. energy intake. and eating time of obese and nonobese subjects. Am I Clin Nutr 1983:37:763-7. Lissner L, Levitsky DA. Strupp BJ. Kalkwarf HI. Roe DA. Dietary fat and the regulation of energy intake in human subjects. Am J Clin Nutr 1987:46:886-92. to eat and drink in rats. Am I Physiol 1947:151:

nutrient

or nutrient in the long the body Then.

7. Adolph EF. Urges 1 10-25.


8.

a normal-weight all energy protein mobilization. muscle glycogen

of body

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.

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carbohydrate carbohydrate fluctuates

reserves store 250 between

for energy and

storage

in the form

500 g, or 4 and

Reference man has a muscle mass of 30 kg with 7.5 kg or an energy equivalent of 120 Mi. but changes in musin terms of energy compared with body fat is nearly depleted. Thus, in conversion lower than RQ measured by

cle mass are insignificant changes in fat mass, unless the long term, of carbohydrate the FQ,

9. Porikos KP. Control offood intake in man: response to covert caloric dilution of a conventional and palatable diet. In: Cioffi LA, James WPT, Van Itallie TB, eds. The body weight regulatory system: normal and disturbed mechanisms. New York: Raven Press, 198 1:83-7. 10. Foltin RW. Fischman MW, Emurian CS, Rachlinski. Compensation for caloric dilution in humans given unrestricted access to food in a residential laboratory. Appetite I 988:10:13-24.
I 1. Jiang

an RQ higher than the FQ implicates or protein to body fat. and an RQ ofenergy observation diet interval fat leads with from body fat. of the lower-than-expected the fact that an RQ

body
12.

C-L. Hunt IN. The relation between freely habitus. Am I Clin Nutr 1983:38:32-40. DM, Frey-Hewitt B, Ellsworth PD. Dietary fat:carbohydrate

chosen

meals

and

a mobilization

Combining the for a high-carbohydrate over a long-term mobilizing body drate diets
(5. 6), a recent

Dreon Wood

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

can only be lower than the FQ to the conclusion that high-carbohy-

induce body fat loss. Apart from studies study suggests that the macronutrient plays a role in the energy Prewitt et al (35) reported

referred earlier composition

of the diet maintenance.

requirement for weight data on 18 women conen%), Energy weight


>

suming a standard by 20 wk of low-fat the body subjects weight was period. the the study Comparing

diet for 4 wk (c:p:f44:19:37 diet (c:p:f 60: 19:2 1 en%). to maintain was interval or 4-wk increased body increased by for the is. intake

followed intake of throughout when diet with

adjusted that initial

or decreased 1 kg, standard

decreased

respectively.

aged men. Am I Clin Nutr 1988:47:995-1000. I 3. Trembla A. Plourde G. Despres I-P. Bouchard C. Impact of dietary fat content and fat oxidation on energy intake in humans. Am I Clin Nutr 1989:49:799-805. 14. Miller WC. Lindeman AK. Wallace I. Niederpruem M. Diet composition, energy intake. and exercise in relation to body fat in men and women. Am I Clin Nutr 1990:52:426-30. I 5. Schoeller DA. How accurate is self-reported dietary energy intake? Nutr Rev 1990:10:373-9. 16. Westerterp KR. Verboeket-van de Venne WPHG. Meijer GAL. Hoor F ten. Self-reported intake as a measure for energy intake. a validation against doubly labelled water. In: Ailhaud G. Guy-Grand B. Lafontan M. Riequier D. eds. Obesity in Europe 91 . London: John Libbey.
1992:17-22.

the last 4 wk for the low-fat diet, the mean energy with 19% and the mean body weight decreased a high-carbohydrate diet resulted in a significant

intake increased with 2 kg. Thus. reduction in

body weight

despite a substantial increase in energy intake aimed at weight maintenance. This review leads to the suggestion that energy expenditure is higher for low-fat, high-carbohydrate diets than for high-fat, lowcarbohydrate not yet available. expenditure chamber, intake include under and normal diets. Convincing Current studies evidence include for this measurement at this point of energy is

Romieu I. Willett WC. Stampfer MI, et al. Energy intake and other determinants of relative weight. Am I Clin Nutr I 988:47:406-12. 18. Miller WC. Diet composition. energy intake, and nutritional status in relation to obesity in men and women. Med Sci Sports Exerc I991:23:280-4. 19. Flatt IP. The difference in the storage capacities for carbohydrate and for fat. and its implications in the regulation of body weight. Ann NY Acad Sci 1987:499:104-23.
I 7. 20.

Frayn KN. Calculation of substrate oxidation gaseous exchange. I AppI Physiol 1983:55:628-34.

rates

in vivo

from

over short time intervals, and calculation of energy changes living in body conditions. composition. of energy direct measurement

up to 24 h in a respiration expenditure from energy Future expenditure studies for

2 1 . Garlick P1, McNurlan MA. McHardy KC. Factors controlling the disposition of primary nutrients. Proc Nutr Soc 1988:47:169-76. should 22. Acheson KJ. Flatt IP, l#{233}quier E. Glycogen synthesis versus lipogenesis 1 wk after a 500 gram carbohydrate meal in man. Metabolism 1982:31: #{163}3 I 234-40.

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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 E. Failure ofdietary 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 E. Glycogen storage capacity and de novo lipogenesis during massive carbohydrate overfeeding in man. Am I Clin Nutr I 988:48:
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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 reason

compare a big intestine energy

those

with

human storage per energy of car-

we as humans a relatively

for food metabolism

A Crawford:
in the and the analogy barrel.

This
that the

may
you test

be a naive
gave tube

question.

but I was
of the and the

and

far lower

us-your of carbohydrate

models

unit body expenditure bohydrate

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

to be slightly showed. You one offat

out of focus showed two

to me was little pots. and those going that larger

ofdifferent

proportions

Eric Ravussin:
I think lipogenesis showed from that plays

I would
in men an Kevin there

like important Acheson, had

to expand is less and role.

a little and

on this that study

comment. de novo that you

different diets. It seems to me that if you inputs in proportion to the daily amount in, one would have seen that went in over a 24-hour period

had represented ofstuffthat was

less evidence in the he gave

the amount of carbohydrate would be really very much

in which

an enormous

than the amount of carbohydrate that was stored in the test tube. A 24-hour input of carbohydrate is quite a lot in a highcarbohydrate diet. At the same time. the barrel is full of fat, so that carbohydrate. although coming through the carbohydrate route, that and is actually filling up the fat barrel because it doesnt store to fat that amount of carbohydrate. fill up the fat barrel. The to fat needs ofthe extent overall energy that contributes So. it has to be converted energy process ofconverting of negative balance ofthe to disturbing

amount ofcarbohvdrate but after the subject and at two a significant calorimetry into fat. but de novo say that a daily ingested think very that

(500 g). We tried that problems, we offered

first as one meal. it at 250 grams

times 125 grams. It was very, very difficult to induce amount ofde novo lipogenesis. I agree that indirect doesnt I think 50<

differentiate that the point in men. 100% stored

between is that

oxidation and going there is little room for you most are ofthe right when you on I stores are replaced carbohydrate into CO2.

lipogenesis maybe basis vs is either

Therefore. that

of the glycogen as glycogen or ofthe using

carbohydrate in terms to what

to be a sort

consideration

1% for fat. I think

individual. I wonder these relationships

oxidized

and explaining why fat is going straight into the barrel and doesnt really have any effect of energy. The carbohydrate going in demands energy to convert it into fat and destorts it as such. Does that confound vs a rich-fat Klaas I dont the diet? equation with regard to a rich-carbohydrate

that the enzymatic activities low in men, and recent studies de novo lipogenesis I agree were with is very that, but studies.

lipogenic enzymes are stable isotope showed in man. hand. term. the studies when you this they it you have

insignificant on the other

Westerterp: ofAcheson consume amount are just when tend diet. you We

short-term

In the long

R Westerterp: know whether

I think this

you model

are stressing was really

an interesting on the right

point. scale

a high-carbohydrate of carbohydrate not are big enough. in a positive yet have you diet.

diet. you cant really store in your glycogen stores because So. energy any tend you energy reason would have balance to do something but apparently Maybe energy energy you on a high-carbohydrate for that. your balance.

with

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

a lot of carbohydrate in proportion to On the other hand, there is not much we really have to convert carbohydrate study I showed in which in substrate part ofthe use. That you you, can and maybe one see that at least

to be in a negative do not for why carbohydrate

a suggestion on a high go up?

to be in a negative Why

balance

literature

expenditure

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

utilization. carbohydrate has been

That rhythm we consume seen in small-

Ravussin:
study. you

In all the studies


need to push

ofoverfeeding.

for example.
through

Schutzs
the system.

a lot ofcarbohydrate

FQ. When viously a daily fore There we talk basis. about There weight energy is limited is oxidized: school gain balance. storage maybe in the in obese but capacity there that l960s people. in men, is a little said that

RQ. they and

AND are obon there-

ENERGY than lication

BALANCE no change from the for the pilot rest study of the for year. the There is a recent

765S
pub-

in positive

by a minute

amount lipogenesis. lipogenesis

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

carbohydrate was the British

trial, where women on like again the

you see a similar phenomenon. the very-low-fat diet lost some you like would expect from the 1 or 2 kg over

amounts

might be higher process. I dont

in lean people because know what is the status

it is an energy-costly of that now.

something

data from the Seattle dietary I think is still in press, where the only more

alternative trial. a 2-year study again you see in middleaged

Nlartijn effectiveness cause the been published American There was

Katan: final in men was some minor,

Could

I sum

a little

note

of caution

about

I kg weight loss on 20 vs 40 en% going on than just the mechanisms body seems diets. to resist

of fat. I think there is that we have been weight loss even on the

of high-carbohydrate tests are long-term In 1970 given loss (irculation, were weight where diets in the

diets in reducing obesity becontrol trials and there have Diet groups or 20% first few Heart of Study middleaged as fat. but that and was large of 40%

a few ofthose?

the National

talking about. The more carbohydrate-rich Westerterp: offood term to accept intake experimental the results Maybe

you are right. types as they feeding studies are.

The only are costly.

problem

is the control hand. longwe have

energy group months

in these

ofstudies.

On the other Therefore,

20-en%-fat

very

1 or at most

2 kg in the

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