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Am J Clin Nutr 1987 Leveille 158 63
Am J Clin Nutr 1987 Leveille 158 63
Am J Clin Nutr 1987 Leveille 158 63
Gilbert
A Leveille,
Phi)3
and Paul
F Cloutier,
Discussion
158
Am
J C/in
Nuir
l987;45:
158-63.
Printed
planning
diets. (For research
purposes
it must
be clear that the Atwater
factors are rounded
values.)
The discussion
thus far would suggest that
energy intake can be conveniently
partitioned
and that the energy value of foods is a constant.
Unfortunately, while it is a fact that the absolute energy content of a food is essentially
constant,
the efficiency
with which it is used
varies and is influenced by numerous factors.
In the experimental
design of animal
studies, it is important
to recognize
the complexities associated
with comparing
diets. Because
of the greater caloric density
of fat compared
to carbohydrate
or protein
(9 kcal/g
vs 4
kcal/g), alteringthe percentage of fat in a diet
changes
the ratio of nutrients
to energy.
An
increase in percentage of One component, eg,
fat, necessitates
a decrease
in one or more of
the
other
components,
eg, carbohydrate
and/or
protein.
A change
in the relative
proportion
of protein,
carbohydrate,
or fat may
influence
eating patterns.
For example,
rats
fed diets with low or inadequate
protein
tend
to eat more in an apparent
effort to obtain
sufficient
protein.
Thus, studies involving
dietary manipulations
often require
the use of
pair-feeding.
If control of caloric intake and/or
meal pattern
is considered
necessary,
yokefeeding
may be used. In this procedure,
when
the animal
fed ad libitum
presses a lever to
obtain
food, the pair-fed
partner
simultaneously receives the same amount
of food (7).
Forbes and colleagues
(8, 9) published
several papers in 1946 on the relationship
between
the level of fat in the diet of adult rats and the
From Nutrition
Corporation,
White
2Adcfr
reprint
Foods
PhD,
General
Foods Corporation,
250 North
Street, White
Plains, NY 10625.
3Present
address:
Nabisco
Brands
Inc., Technology
Center, P.O. Box 1943, East Hanover,
NJ 07936-1943.
in USA.
1987 American
Society
for Clinical
Nutrition
PhD
ISOCALORIC
DIETS
159
GROSS ENERGY
100
Fecal energy
I:
Urinary
energy
METABOLfl A8LE
ENERGY
25
NET ENERGY
Maintenance
Work
Productive
Functions
10
30
Percent
of
Fat
in Diet
efficiency
of use of dietary
energy.
In those
studies
dietary
fat varied
from 2% to 30%,
while intake of gross energy, protein,
and essential
nutrients
remained
constant.
Researchers
observed
that as the dietary fat content increased,
relative
heat production
decreased.
Thus, with increasing
fat intake the
apparent
efficiency
of food use increased,
allowing for more energy to be stored in the animal carcass (see Fig 2). In essence,
more energy is stored as fat in the body as the proportion
of fat to carbohydrate
increases
in the
diet.
Increasing
the protein level of the diet at the
expense of carbohydrate
also results in changes
in apparent
energy efficiency.
Donald et al (10)
observed
that adult rats fed a high-protein
diet
gained more weight than a similar
group fed
TABLE
1
Basis for Atwater
factors*
Gross
energy
Digestible
energy
5.65
5.20
4.00
4.15
4.00
8.90
4.00
8.90
Metabolizable
energy
Atwater
factors
a low-protein
diet when the content
of dietary
fat was maintained
at a constant
level and levels of protein
varied from 5% to 25%. The animals had free access to food and water. Absolute food consumption,
ie, g/rat, was not
significantly
different
between
the 5% and 25%
groups.
Yet total body weight gain and body
fat were higher in the 25% group (Table 2).
Humans
appear
to respond
in much
the
same way as rats to changes
in dietary
composition.
Danforth
(11) noted that lean subjects
gained
weight
relatively
easily
when
overfed
fat but not when fed a mixed diet of
carbohydrate
and fat. In an earlier study by
Miller
and Mumford
(12), researchers
observed that students
overfed a high-protein
diet
gained more weight than a group of students
fed a low-protein
diet with a similar
number
of calories.
The weight gain for both diets was
less than the predicted
value.
TABLE 2
Final body weight and body fat of rats fed ad libitum
low- and high-protein diets of equal caloric contents
Protein
Protein (kcal/g)
Carbohydrate
(kcal/g)
Fats (kcal/g)
*
9.40
Atwater
and
Bryant
9
(6).
Based
in diet
Body wt
Body fat
5
25
397
487
16
24
on ref(l0).
Heat including
Specific dynamic action
or
Heat
increment
of foods
LEVEILLE
160
AND
PROTEIN
CARBOHYDRATE
Ji
Ketogenic
CO2
ACIDS
+
+
ENERGY
FIG 4. Metabolic
bohydrate.
pathways
with intake
of excess
car-
PROTEIN
CAOHYDRATE
FAT(Triglycerides
GLrOSE
GLU ,SE
Glucogenic
AMINO
ACIDS
FAT Trig1ycerjj
FATTY
>P9LJVATE
<
Glucogenic
PYRIATE
> GLYCEROL
AMINO
ACIDS
1<
ii IKI__
5 GLYCEROL
>ACETYL
C0A<
>FATTY
ACIDS
Ketogenic
ACE
CoA
> FATTY
ACIDS
KREBS
CYCLE
#{231}LE
V
El
CO2
CO2
H20
H2O
ENERGY
ENERGY
FIG 3. Metabolic
tein.
pathways
with intake
of excess
pro-
FIGS. Metabolic
triglycerides.
pathways
with intake
of excess
fat as
Level of dietary
energy
intake
has been
shown to influence
the use of dietary protein
in humans
(13). Inadequate
energy intake reduces the apparent
efficiency
of nitrogen
use.
On the other hand, excess energy intake may
increase
retention
of nitrogen.
Most studies in
humans
indicate
that the effects of energy and
protein
intakes
on fat metabolism
are interrelated
(14-16).
For example,
in studies
of
isocaloric
exchange
of fat for carbohydrate,
the
efficiency
of the use of dietary protein
was improved on diets in which more energy was derived
from
carbohydrates
than
from
fats.
This protein-sparing
effect was more noticeable under
situations
where energy
intakes
were marginal
and individuals
were losing
weight (17).
At least part of the observed
differences
in
efficiency
between
fat, carbohydrate,
and protein can be explained
on theoretical
grounds.
Figures 3,4, and 5 illustrate
schematically
the
metabolic
fate of protein,
carbohydrate,
and
fat, respectively.
For example,
as depicted
in
Figure 3, protein
that is not needed to replenish body protein
is degraded
to amino
acids,
which are deaminated
and oxidized
to provide
energy or are converted
to fat. The energy cost
CLOUTIER
ISOCALORIC
DIETS
161
L.
C
VA
.
.
162
LEVEILLE
AND
CLOUTIER
physical
formance
the per-
Conclusions
4,
S.
Protein
C
C
I.
4,
Fat
C
Lu
ii
Time
Meal
in Hours
effect of food
duction
and maintain
normal
body temperatures without
shivering.
The source
of this
heat is located in the specialized
mitochondria
of brown adipose
tissue, which is distributed
predominantly
around
critical
organs
(24).
Another
example
of AT found in humans
and
animals
is the response
of RMR to changes
in
intake of nutrients. During prolonged
periods
of food restriction,
RMR is gradually
reduced.
When
intake
of nutrients
is excessive,
the
RMR increases.
In both cases, the changes are
more than can be predicted
from calculations
when energy condition
is in balance
(19).
The extra heat produced
from the ingestion
of food, as exemplified
by TEF and AT, is a
waste product
as far as the economy
of the
animal is concerned. One possible function
for this waste energy,
however,
is regulation
of body weight as a supplement
to appetite
control.
The mechanisms
involved
are not
understood
completely.
Some of the mechanisms for which evidence
exists include
sympathetic
nervous
system activity,
futile metabolic cycles, uncoupling of oxidative phosphorylation, and Na-K
ATPase
activity
(sodium
pump)
(20, 24).
The metabolizable
energy
from
food is
transformed
into heat and net energy. The degree of heat production
from all mechanisms
influences total net energy. Following the laws
of conservation
of energy, as less heat is produced,
more energy
becomes
available
for
References
1. Baldwin
RL, Bywater AC. Nutritional
energetics
of
animals.
Ann Rev Nutr 1984;4:10l-l4.
2. Buskirk ER, Mendez J. Energy: caloric requirements.
In:Alfin-Slater
RB, Kiitchevsky
D, eds. Nutrition
and
the adult: macronutrients.
New York, NY: Plenum
Press, 1980:49.
3. Dale HE. Energy metabolism.
In: Swenson MJ, ed.
Dukes physiology
of domestic
animals.
Ithaca, NY:
Comstock
Publishing
Associates,
1970:6 19.
4. Kleiber M. The fireof life. Huntington,
NY: Robert
E Krieger Publishing,
1975.
5. Mitchell HH. Comparative nutritionof man and domestic animals.
Vol 2. New York, NY: Academic
Press, 1964:471-565.
6. Maynard
LA. The Atwater system of calculating
the
caloric value of diets. J Nutr l944;28:443-52.
7. Cox JE, Powley TL. Development of obesity in diabeticmice pair-fedwith lean siblings.J Comp Physiol
Psychol l977;9l:347-58.
8. Forbes EB, Swift RW, Elliott RF, James WH. Relation
of fatto economy of food utilization.
IIBy the mature
albino rat. J Nutr l946;3l:2l3-27.
9. Forbes EB, Swift RW, Thacker El, Smith VF, French
CE. Further experiments on the relation of fat to
economy of food utilization.IIBy the mature albino
rat.J Nutr 1946;32:397-403.
10. Donald P, Pius GC, Pohi SL. Body weight and composition in laboratory rats:effectsof diets with high
or low protein concentrations.
Science l98l21l:185-6.
11. Danforth E. Diet and obesity.
Am J Clin Nutr
l985;4l(suppl):1132-45.
12. Miller DS, Mumford P. Gluttony. 1 An experimental
study of overeating low- or high-protein diets.Am J
Clin Nutr l967;20:12l2-22.
FIG 8. Schematic
after a meal (adapted
After
1) Research
involving
dietary
manipulations must consider
and control the nutrientto-energy
ratio of the diets and total caloric
intake.
2) The composition
of the diet with respect
to the proportion
of the major
nutrientsprotein,
fat, and carbohydrate-has
a profound and variable
effect on the efficiency
of
food energy use and ultimately
on the results
of an experiment.
3) The level of energy intake relative to
maintenance
can significantly
influence
energy
expenditure
and hence the net energy value of
diets and foods.
ISOCALORIC
13. Van Es AJH, Vogt JE, Niessen CH, et al. Human
energy metabolism below, near, and above energy
equilibrium.
Br J Nutr l984;52:429-42.
14. Dallosso HM, James WPT. Whole-body
calorimetry
studies in adult men. I The effect of fat overfeeding
on 24-h energy expenditure.
Br J Nutr 1984;52:4964.
15. Garz.a C, Scrimshaw
NS, Young VR. Human protein
requirements:
the effect of variationsin energy intake
within
the maintenance
range. Am J Clin Nutr
1976;29:280-7.
16. MacLean
WC, Graham GG. The effect of level of
protein
intake in isoenergetic
diets on energy utilization. Am J Clin Nutr 1979;32:l38l-7.
17. Richardson
DP, Wayler AH, Scrimshaw
NS, Young
VR. Quantitative
effect of an isoenergetic
exchange
of fat for carbohydrate on dietary protein utilization
in healthy young men. Am J Clin Nutr l979;32:22l726.
DIETS
163