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Nutrition Department, Pennsylvania State University, University Park, PA 16802-6501
| ABSTRACT |
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KEY WORDS: energy density fat intake human food intake hunger satiety
| INTRODUCTION |
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During the 1990s, studies of energy intake have tested the hypothesis
that the macronutrients can have different effects. Of particular
importance for this review is the suggestion that dietary fat is
overeaten because it affects satiety and satiation less than
carbohydrate and protein (Rolls and Hammer 1995
). The
methods used to study satiety (the effect on subsequent intake) and
satiation (the amount eaten in a meal) differ; therefore, the results
from these studies will be considered separately.
| Do macronutrients differ in their effects on satiety? |
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We have published several reviews of the studies comparing the effects
of fat and carbohydrate on food intake (Rolls and Hammer 1995
, Rolls and Bell 1999
). In studies of
satiety, the fat and carbohydrate content of a preload had similar
effects on subsequent food intake when the palatability and energy
density were matched. This was seen when the foods were ingested orally
or infused intravenously or intragastrically. In one study, however,
fat ingested in yogurt preloads suppressed lunch intake slightly (but
significantly) less than carbohydrate in some of the participants who
were obese or concerned with their body weight (Rolls et al. 1994
). Although the difference was small, even small
differences could have an influence on energy balance over time.
| The role of the energy density of food in satiety |
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To test the effect of the energy density of a food on satiety, the
palatability and the macronutrient and energy content of a preload
should be similar, whereas the energy density changes. For example, the
addition of water will increase the weight or volume, thus decreasing
the energy density. This was done in a recent study in which lean young
men were served a milk-based preload that had varying amounts of
water added to change the volume (300, 450 and 600 mL) but not
macronutrient content or palatability (Rolls et al. 1998
). Because the total energy content was held constant (2088
kJ), the preloads varied in energy density. The results showed that
energy density of the preload affected energy intake at lunch 30 min
later such that intake was significantly lower by 18% after the high
volume, low energy density drink than after the low volume, high energy
density drink. Furthermore, the men in the study did not compensate at
dinner for the reduced intake at lunch. These results demonstrate that
the weight or volume of food consumed, and thus the energy density, can
affect satiety.
In general, the water content of foods is a critical determinant of
energy density; it has a larger effect than other ingredients such as
fat or fiber. We saw in the previous experiment that adding water to a
food increased the effect of the food on satiety. It is of both
practical and theoretical interest to determine whether the addition of
water affects intake simply by increasing the volume in the stomach, or
if an increased volume of food affects satiety through sensory and
cognitive influences. For example, a food with added water will look
bigger and will likely cause greater stimulation of oropharyngeal
mechanisms. One way to separate these influences is to test subjects
with the food and water consumed separately and to compare the effects
with those when the food and water are mixed together. In a recent
study, we developed a chicken, rice and vegetable casserole containing
1128 kJ and weighing 263 g (Rolls et al. 1999b
).
The base of this casserole was undiluted cream of chicken soup; thus
water could be added to the casserole to make a soup. The casserole and
soup had exactly the same ingredients except for the additional
356 g of water in the soup. In a third condition, the casserole
was served with a glass of water equivalent to the amount of water that
had been added to make the soup. When the women in the study ate lunch
5 min after finishing the preloads, the water affected how much was
eaten, but only when it was incorporated into the casserole to make a
soup. The women ate similar amounts when the casserole was served with
and without a glass of water. When they consumed soup, however, they
felt fuller and less hungry and consumed ~27% less energy compared
with intake at lunch after the casserole. The subjects did not
compensate at dinner for this reduction in lunch intake. This study
showed that water had to be incorporated into a food, thereby
increasing its weight or volume, to influence satiety.
The effects of the energy density of foods on satiety are robust and of a magnitude that implies that they are an important influence on the regulation of energy intake. When the macronutrient content of foods is varied, but the energy density is kept constant, the effects of fat and carbohydrate on satiety are similar. When the palatability of foods is similar, energy density is the major determinant of how foods affect satiety regardless of macronutrient content.
| Do macronutrients differ in their effects on satiation? |
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Most of the early studies on the effects of the fat content of foods on
satiation varied the amount of fat in the available foods without
concern for the confounding effects of differences in palatability and
energy density (Rolls and Hammer 1995
, Rolls and Bell 1999
). In these studies, which lasted from a single meal
to 11 wk, the higher the fat content and the energy density, the
greater the daily energy intake. When the participants in these studies
were allowed free access to the manipulated foods, they ate a similar
amount by weight of food regardless of the composition (Duncan et al. 1983
, Kendall et al. 1991
, Lissner et al. 1987
, Miller et al. 1998
, Stubbs et al. 1995
). The results demonstrated consistently that the
weight of food consumed daily was more constant than energy intake over
periods as long as 11 wk (Kendall et al. 1991
).
The tendency to consume a constant weight of food was confirmed by an
analysis of 4-d weighed food records from 45 nonobese adults
(Seagle et al. 1997
). The weight of food consumed
remained more constant over a day than either the intake of energy or
fat. Similarly, an analysis of reported intakes in large community
samples showed there was little adjustment in the amount of food
consumed to compensate for variations in the energy density of the
total diet (Poppitt and Prentice 1996
). All of these
studies showing that people eat a constant weight or volume of food
imply that energy density will be a critical determinant of energy
intake. Because high fat foods are often high in energy density, this
could be at least part of the explanation for the high intake of fat.
Of importance for the present discussion of the effects of dietary fat
on intake is whether fat affects satiation independently of the effects
on energy density.
Several studies have tested the effects of variations in the
fat content of foods while holding the energy density constant. In one
study, liquid diets varying in fat content (24 vs. 47%) but similar in
energy density were compared (van Stratum et al. 1978
).
Most of the energy consumed by the Trappist nuns in this study was from
the liquid diets that were offered ad libitum, with standardized snacks
contributing 25% of energy intake. Over the 2 wk on each diet, the
women consumed a constant weight of the manipulated liquid diet, and
thus energy intake remained constant. This was the first study to
demonstrate that the fat content of foods per se did not have an
independent effect on satiation. This finding was confirmed in another
study in which six men of normal weight were offered each of three 14-d
diets varying in fat content (20, 40 or 60% of energy) but similar in
energy density (Stubbs et al. 1996
). Again, subjects ate
a constant weight of food so that energy intake did not differ among
conditions despite the large difference in fat content. More recently,
Saltzman et al. (1997)
fed men diets containing foods
varying in fat content (20 vs. 40%) but matched for energy density.
The energy intakes when these diets were consumed were similar over the
9 d of each dietary condition, again suggesting that the level of
fat per se did not affect intake. These three studies indicate that the
amount of fat in the diet did not influence satiation when the energy
density was held constant.
| Does the energy density of foods affect satiation? |
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| Variations in the fat content or energy density of a portion of the diet |
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We manipulated the energy density and/or the fat content of part of the
diets of lean and obese women over 4 d to determine the effect on
energy intake (Rolls et al. 1999a
). In three of these
4-d test periods, subjects were served compulsory entrees which
represented 50% of their usual energy intake at breakfast, lunch and
dinner. Additional foods could be consumed ad libitum during and
between meals. The results showed that variations in the fat content
(16.4 vs. 36.5% energy) had no effect on energy intake. In contrast,
the energy density of the compulsory foods did affect energy intake
over the 4 d in both the lean and obese women. Intake of
self-selected foods at meals was reduced significantly by 16% in
the low (4.4 kJ/g) compared with the high energy density (6.7 kJ/g)
condition. Ratings of hunger did not differ between the diets. The
results showed that when a portion of the diet was manipulated, energy
density, but not the fat content, affected energy intake.
| How does energy density affect energy intake? |
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In addition to learned influences, the sensory cues associated with
changes in the amount of food consumed are likely to be important.
Seeing a large portion of food and then tasting, chewing and swallowing
it are likely to enhance satiety. Sensory-specific satiety has been
shown to be affected by the amount of food consumed rather than the
energy content (Bell et al. 1998b
). Gastric distension
and stomach emptying can also be affected by the energy density of
foods. The rate at which nutrients reach satiety receptors and release
satiety hormones could be affected by the energy density of food.
Future studies should determine how the behavioral and physiologic
effects of the energy density of food together lead to satiety and
satiation.
| Clinical implications |
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Because the critical importance of the energy density of foods
has been demonstrated only recently, there are few data available on
the typical energy densities of diets consumed by populations
(Poppitt and Prentice 1996
). In the future, community
studies should include data on energy density. This will require
weighed intakes and records of fluid consumption. Although laboratory
studies did not show a critical effect of water as a beverage on energy
intake, the effect of energy-containing beverages may be important
(Rolls et al. 1999b
). Assessment of diet records over
several days indicates that obesity is associated with the consumption
of foods high in energy density (McCrory et al. 1999
,
Westerterp-Plantenga et al. 1996
). Population studies
are required to assess the energy densities of the total diet, not only
in groups of different body weights, but also in other groups at
nutritional risk such as the elderly.
| FOOTNOTES |
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2 Supported by the National Institute of Diabetes and Digestive and Kidney Diseases (grants DK-39177 and DK-50156).
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