SPORT NUTRITION CONFERENCE PROCEEDINGS

InternationalJournal of Sport Nutrition, 1996, 6, 80-92

0 1996 Human Kinetics Publishers, Inc.

Physical Activity, Fat Balance, and Energy Balance

James 0. Hill and Rene6 Commerford

In this paper, we review the impact of physical activity on energy and
macronutrient balances. Stability of body weight and body composition de-
pends on reaching a steady-state where the amount and composition of energy
ingested are equal to the amount and composition of energy expended. We
describe how aperson's level of physical activity can have a significant impact
on determining the level of body weight and body fatness at which that
steady-state is reached. First, physical activity can directly affect both total
energy intake and total energy expenditure. Physical activity can also affect fat
balance, and it is becoming clear that imbalances in total energy are largely
imbalances in fat. High levels of physical activity should help individuals reach
fat and energy balances at lower levels of body fatness than would have been
achieved at lower levels of physical activity.

Key Words: weight regulation, fat intake, exercise

There are many reasons for recommending that Americans increase their
level of physical activity. Increased physical activity has been associated with
improvement in risk factors for several chronic diseases (26,30,60) as well as with
maintenance of healthy levels of body weight and body fatness (31,41). Physical
activity appears to impact positively on health directly (e.g., via increased insulin
action) (19, 83) and indirectly via reductions in body weight and body fat content
(46,54). The purposes of this review are to define body weight regulation in terms
of energy and nutrient balance, emphasize the importance of fat balance in body
weight regulation, and describe the ways in which physical activity affects energy
and nutrient balance.

Regulation of Body Weight

While there appears to be a systematic increase in body weight across the adult
lifespan (81), the degree with which body weight is maintained constant suggests
that something is being regulated. This something couldbe body weight,body fatness,
temperature, energy intake, energy expenditure, or energy balance itself (11). Some
think of body weight as being regulated by a set-point,where physiologicalsystems act
to maintain a determined body weight (42). Others prefer the term settling-point,

The authors are with the Center for Human Nutrition, University of Colorado Health
Sciences Center, Denver, CO 80262.
 Fat and Energy Balance / 81

which suggests that there is not an optimal weight being maintained but rather that
the weight maintained over time is a result of inherent characteristics of the
individual combined with the environment in which that individual lives (33).

Regulation of Energy and Macronutrient Balances

An individual maintaining a constant body weight and body composition must be
in a steady-state where the amount of total energy ingested is equal to the amount
of total energy expended. In addition, over some period of time, the composition of
the energy ingested must equal the composition of substrate oxidized (33). Condi-
tions necessary for energy and nutrient balance are illustrated in Figure 1. Any
disruption of energy and macronutrient balances elicits compensatory responses to
restore these balances. The relative stability of body weight over time suggests an
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ability to compensate for small daily variations in intake or expenditure.

Any change in body weight must be a result of an imbalance in one or more
macronutrients and in total energy. However, since the organism strives to reachieve
homeostasis, changes in body weight and body composition will occur only until
homeostasis is reestablished. Steady-states of energy and macronutrient balances
can be reestablished at the same or at a different level of body weight and body fat
mass, depending on the nature of the challenge and characteristics of the subject.
It is becoming clear that changes in body weight and body composition are
more closely related to an imbalance in fat than to an imbalance in carbohydrate or
protein. Data in support of this conclusion come from several investigators working
on a model first proposed by Flatt (22,23). Flatt proposed that unlike carbohydrate
and protein balance, fat balance is not regulated acutely. In support of this

FAT + + FAT
CARBOHYDRATE + + CARBOHYDRATE
PROTEIN -+ + PROTEIN

Figure 1 - In order to achieve zero energy and macronutrientbalance, each factor to

the left of the figure must exactly equal the correspondingfactor on the right side of the
figure.
 82 / Hill and Commerford

hypothesis, he found that increasing the carbohydrate content of a single meal

immediately increased carbohydrate oxidation, while increasing the fat content of
a meal did not increase fat oxidation (24). In several studies, the carbohydrate and
fat contents of the diet have been varied over periods ranging from a single meal to
a 2-week period (2,7,38,64,70,7 1).A consistent finding has been that alterations
in carbohydrate intake produce rapid and substantial changes in carbohydrate
oxidation that serve to maintain carbohydrate balance. Alterations in fat intake
produce little if any immediate changes in fat oxidation, so that there is little effort
to maintain fat balance acutely. Thus, weight changes following challenges to body
weight are due primarily to disruptions in fat balance, which account for most of the
imbalance in total energy.
While the majority of evidence suggests that fat balance is not well regulated
acutely, it must be regulated over the long term, since the body weight regulation
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system must eventually reachieve energy and nutrient balance. Reachieving fat
balance in many situations may require substantial alterations in body fat mass,
which can alter fat oxidation to bring it into balance with fat intake and stabilize
body weight.

Regulation of Fat Balance

While carbohydrate balance can be achieved with little alteration of endogenous
carbohydrate stores, changes in body fat mass can contribute substantially to
achieving fat balance. Individual differences in achieving fat balance may play a
primary role in determining body weight and level of body fatness. Fat intake, fat
oxidation, and the size of the body fat mass all may play a role in determining how
fat balance is achieved. A prevailing view is that body fat mass changes in size to
allow an individual to achieve fat balance. If, for example, fat intake exceeds fat
oxidation, the fat mass will increase. Increases in the fat mass will be associated with
increased availability of lipid substrate and increased fat oxidation. When the fat
mass has increased sufficiently, fat balance will be reestablished and the fat mass
will stabilize. Similarly, when fat intake is less than fat oxidation, the body fat mass
will decline to bring about a lower fat oxidation.
Body fat mass is maintained at widely different levels in different people. This
suggests that some individuals are able to achieve fat balance with a small body fat
mass while others require a larger fat mass. Theoretically, any factor that reduces
fat intake or increases fat oxidation should contribute to differences in the level of
body fat mass at which fat balance can be achieved.

Influences on Fat Intake

There is controversy concerning regulation of specific macronutrients in human
subjects. Both the composition and energy density of the diet may influence
total energy intake and thus intake of specific macronutrients. Specifically,
high-fat diets have been shown to increase total energy intake when compared with
low-fat diets (43,48,69,71). Since high-fat diets are also high in energy density,
a question is whether it is the high fat content or the high energy density of the diet
that leads to increased total energy intake. It has been suggested that dietary fat
intake is higher in obese subjects than in lean subjects, but the data are conflicting
(49,73).
 Fat and Energy Balance / 83

There is a great deal of interest in assessing the capacity of human subjects to

regulate intake of specific macronutrients. Relevant data come from studies
assessing the ability of subjects to compensate for a caloric preload (58). Results of
these studies are mixed and often difficult to interpret. Rolls et al. (58) found that
lean, unrestrained male subjects can compensate equally well for carbohydrate and
fat preloads, while obese, restrained female subjects show poor compensation for
carbohydrate and fat preloads. Other groups show a mixed response. Studies with
the fat substituteOlestra suggest that subjects have the ability to compensate for the
calories substituted by Olestra but that the compensation is not specific for the fat
calories (9). Thus, after short-term consumption of Olestra, subjects eat a similar
amount of calories as with the control diet but consume a diet lower in fat.

Influences on Fat Oxidation

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Very little is known about reasons for individual differences in fat oxidation. There
is a modest positive relationship between body fat mass and fat oxidation (6), but
many other factors likely influence fat oxidation. Availability of substrate, insulin
action, and characteristics of fuel-consuming tissues are factors that have been
proposed to act either individually or together to account for individual differences
in fat oxidation. The proportion of total oxidation coming from fat will generally
increase with increased circulating lipid levels (17). However, the proportion of
total oxidation from fat versus carbohydrate will also depend on insulin action (65).
Finally, it has been suggested that characteristics of skeletal muscle may influence
the compositionof the fuel mixture oxidized (1). It is likely that substrate availabil-
ity, insulin action, and skeletal muscle characteristics all interact to influence fat
oxidation.

Physical Activity and Fat Balance

Physical activity can influence energy intake and substrate oxidation, and thus it
likely contributes substantially to achieving fat balance. Negative relationships
between measures of physical activity (usually self-reports) and indices of obesity
(usually body mass index, or BMI) are seen in most data sets obtained from the
general U.S. population (20,52,67,79). At least two studies using doubly labeled
water to measure energy expended in physical activity have shown a significant
negative relationship between physical activity and indices of fatness (16, 63).
Figure 2 shows the relationship between physical activity level (PAL; determined
from doubly labeled water) and percent body fat in children between 1.5 and 4.5
years of age (from Reference 16).
While there are strong associations between physical activity and obesity,
there is insufficient data to conclude that a low level of physical activity is a cause
of obesity. It is equally possible that a low level of physical activity is a consequence
of obesity. There are a few data sets in which changes in physical activity and
changes in BMI have been assessed longitudinally. Williamson et al. (80) per-
formed follow-up studies on 3,5 15men and 5,8 10 women in the NHANES I study.
BMI and level of physical activity were obtained 10 years later in subjects who
participated in NHANES I (1971-75). While a significant relationship was
obtained between BMI and level of physical activity at baseline and follow-up,
physical activity at baseline was not significantly correlated with change in BMI.
 84 / Hill and Cornrnerford
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5- 0 Boys 0
- 0 Girls
0 1 . 1 . 1 ' 1 ' 1 ' 1 . 1 ' 1

0.8 1.0 1.2 1.4 1.6 1.8 2.0 2.2 2.4

PAL

Figure 2 - Physical activity level (PAL), determined using doubly labeled water, is
negatively related to percentage body fat in children ages 1.5 to 4.5 years. Reprinted
with permission from P.S.W. Davies, J. Gregory, and A. White. Physical activity and
body fatness in pre-school children. Znt. J. Obes. 19:6-10,1995.

However, level of physical activity at follow-up was significantly negatively

correlated with major weight gain (>I3 kg). The authors hypothesize that a low
level of physical activity may be both a cause and a consequence of weight gain.
A few other studies of adults and children have found that a low level of physical
activity is associated with weight gain over time (45, 53).

Physical Activity and Fat Intake

Total energy expenditure increases during exercise, requiring additional fuel to
support the physical activity. The additional fuel comes largely from endogenous
carbohydrate and fat stores, and the relative contribution of each depends on
characteristics of the physical activity and characteristics of the subject (14,28).
Depletion of endogenous energy stores may, in turn, stimulate ingestive behavior
that replenishes these energy stores.
Although the effects of physical activity on substrate oxidation have been
studied extensively, much less is known about the effects of physical activity on
food intake. Since more active people generally have higher levels of energy
expenditure, their energy intake should be higher. This is generally found in
cross-sectional studies (50,68). Far less is known about the time course over which
 Fat and Energy Balance / 85

increases in physical activity lead to increases in energy intake. Verger et al. (75,
76) found increases in voluntary energy intake 2 hr after exercise. However,
several short-term training studies did not find increases in total energy intake
(determined from self-reports) over several weeks of exercise training (5, 12, 18,
35,47).
The way in which physical activity influences energy intake might vary with
characteristicsof the subjects. For example, there is some indication that nonobese
subjects show a greater increase in energy intake with increasing physical activity
than obese subjects (81, 82). Additionally, the individual's dieting history could
influence the effects of physical activity on energy intake. We previously proposed
that restrained eaters may respond differently to physical activity than unrestrained
eaters (32). For example, unrestrained eaters may be better attuned to any physi-
ological signal produced by increased energy expenditure and may increase energy
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intake in response to increased physical activity. Conversely,restrained eaters may

not be as attuned to physiological signals and may not alter energy intake in response
to increased physical activity.
Tremblay and colleagues demonstrated that exercise can promote negative
energy and fat balance and that this depends both on substrate oxidized during
exercise and the composition of postexercise food intake (4,72).
There is little knowledge of whether and how changes in physical activity
affect the composition of food eaten. Theoretically, the composition of fuel
oxidized during physical activity may influence the composition of food eaten
following physical activity, if the goal is to replace the stored fuel. This could
include replenishing intramuscular triglyceride stores as well as glycogen stores.
However, as previously mentioned, while carbohydrate intake and oxidation are
linked acutely, fat intake and fat oxidation do not appear to be linked. One could
speculate that the increased carbohydrate oxidation that occurs with increased
physical activity may drive carbohydrate intake while the increased fat oxidation
has no effect on fat intake. Thus, an increase in physical activity may promote intake
of a diet high in carbohydrate. There is some indication that this does occur in some
populations (62). There is a clear need for additional studies to identify the precise
relationship between increasing physical activity and food intake and to identify
factors influencing that relationship.

Physical Activity and Fat Oxidation

An increase in fat oxidation is seen in virtually all forms of physical activity.
However, the amount of fat oxidized during physical activity depends on character-
istics of the activity and of the subject. Fat oxidation during physical activity is
usually assessed by indirect calorimetry (40). This involves knowing both the total
amount of energy expended and the relative proportion that comes from fat. The
former is estimated from measurement of oxygen consumption and the latter from
the whole body nonprotein respiratory quotient (NPRQ) (40). The NPRQ generally
ranges from 0.70 (100% fat oxidation) to 1.0 (100% carbohydrate oxidation).
There is some indication that high levels of fitness or training may affect the
way that dietary fat is partitioned between oxidation and storage. Simsolo et al. (66)
studied regulation of lipoprotein lipase (LPL), a key enzyme in the uptake of
triglyceride-derived free fatty acids, in adipose tissue and skeletal muscle before
and after detraining in runners. After detraining, LPL was increased in adipose
 86 / Hill and Commerford

tissue and decreased in skeletal muscle. This suggests that detraining may have
influenced fat partitioning by shunting it away from muscle (presumably for
oxidation) toward adipose tissue (presumably for storage). Thus, a high level of
physical activity may route dietary fat to muscle for oxidation rather than to adipose
tissue for storage.
Characteristics of Physical Activity. The type of fuel used during physical
activity depends on the intensity and duration of the activity (14,28). Breakdown
of glycogenprovides fuel forphysical activity performed at maximal or supramaximal
work intensities(2100%maximal oxygen uptake, orV0,max) (14). With submaximal
exercise there is a balance between aerobic and anaerobic metabolism (61).
Carbohydrateis the main fuel source during submaximal exercise performed at high
work intensities (>70-75% V0,max) (13). As exercise becomes predominantly
more aerobic, lipid provides an increasing proportion of fuel (78). Protein oxidation
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during physical activity is small (36). Trained subjects can perform aerobic exercise
for long periods of time with lipid oxidation increasing as time progresses (3,21).
Characteristicsof the Exercising Individual. Both training status and gender
can influencefuel utilization during physical activity.Age and body fat composition
have been suggested as additional factors that can modify fat oxidation during
physical activity.
With increasing levels of fitness or training, a higher proportion of fat is used
during physical activity performed at the same absolute workload (39,44). This is
partly due to the lower relative intensity of the physical activity. Even so, fat
oxidation may be increased at the same relative work intensity in trained versus
untrained individuals (8,74). Increased fat oxidation in trained subjects is greatly
facilitated by morphological and enzymatic adaptations in skeletal muscle follow-
ing training (34, 37) combined with increased availability of lipid substrate (59).
Some studies have found that females oxidize proportionally more lipid
during physical activity of a similar intensity compared to males (10, 25), while
other studies have found no such gender differences (15, 56). Protein oxidation
during moderate-intensity physical activity appears to be lower in females versus
males, although for both genders its overall contribution to total fuel oxidation
remains small (57). The stage of menstrual cycle in females may also affect the
pattern of fuel oxidation. Although few adequately controlled studies have been
performed in this area, there is a suggestion of increased lipid oxidation during
physical activity performed in the luteal versus follicular phase of the menstrual
cycle (29,51).
It has been suggested that the ability to oxidize fat during physical activity
declines with age (55). If true, this could be due to changes in the morphology and
metabolism of skeletal muscles and/or to availability of lipid substrate during
physical activity.
Wade et al. (77) proposed that obese subjects used substrates differently from
lean subjects during exercise. Obese subjects oxidized less fat during exercise than
lean subjects. The authors related the lower fat oxidation to a lower proportion of
type I (oxidative) muscle fibers in the obese as compared to lean subjects. Geerling
et al. (27) replicated this study and failed to find a difference in respiratory quotient
(RQ) during exercise between lean and obese subjects. It remains to be determined
whether and how body composition influences substrate oxidation during exercise
and whether characteristics of skeletal muscle influence this process.
 Fat and Energy Balance / 87

Physical Activity and Body Weight Regulation

An increase in physical activity will increase total energy expenditure and fat
oxidation. This will lead to an overall energy imbalance if there is not complete
compensation in terms of increased energy intake and increased fat intake. What-
ever compensation does occur does not appear to be complete, so that negative
energy balance, negative fat balance, and weight loss should occur following an
increase in physical activity.
Body weight and body fat mass should decrease until the system reestablishes
fat balance and energy balance. Fat balance can be reachieved in several ways. First,
if energy intake remains unchanged, fat balance can only be reachieved by a
reduction in fat oxidation. This may occur subsequent to a substantial decline in
body fat mass that occurs as a result of negative fat and energy balance. Alterna-
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tively, when fat oxidation exceeds fat intake this may eventually stimulate food
intake, thus reestablishing fat and energy balance. There is little indication that
increases in physical activity stimulate fat intake, so it is unlikely that fat balance
is reestablished due to increased fat intake. Finally, there could be simultaneous
declines in fat oxidation (due to declining fat mass) and increases in energy intake
that reestablish fat and energy balance.
The way in which fat and energy balances are reestablished will determine the
period of time over which increases in physical activity lead to declines in body
weight and body fatness. If the major way that fat balance is reestablished is by a
decline in body fat mass, this may take months or years, depending on the magnitude
of increased activity. Alternatively, changes in energy intake can establish energy
and fat balance much more quickly.

Summary
Achieving fat balance is essential to achieving a stable body weight and body
composition. In this regard it is useful to examine reasons why fat balance is
achieved at such widely differing levels of body fatness in the U.S. population. A
prevailing view is that changes in the body fat mass itself help achieve fat balance.
Efforts to promote healthy body weights shouldfocuson reasons why some individuals
can achieve fat balance at low levels of body fat mass while others cannot.
Physical activity can have major effects on both intake and oxidation of total
energy and in particular on intake and oxidation of fat. Epidemiological data reveal
a consistent negative relationship between level of physical activity and indices of
obesity but do not provide clear evidence of a cause-and-effect relationship.
Increases in physical activity increase the proportion of total oxidation from fat and
may decrease the proportion of ingested energy from fat. For these reasons,
maintaining an active lifestyle should allow achievement of fat balance at a lower
level of body fat mass than would occur with a sedentary lifestyle.

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Acknowledgment
Supported in part by NIH grants DK 42549 and DK 38088.