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OCR for page 107
5
Energy
SUMMARY
Energy is required to sustain the body’s various functions, includ-
ing respiration, circulation, physical work, and maintenance of
core body temperature. The energy in foods is released in the
body by oxidation, yielding the chemical energy needed to sustain
metabolism, nerve transmission, respiration, circulation, and physical
work. The heat produced during these processes is used to maintain
body temperature. Energy balance in an individual depends on
his or her dietary energy intake and energy expenditure. Imbalances
between intake and expenditure result in gains or losses of body
components, mainly in the form of fat, and these determine changes
in body weight.
The Estimated Energy Requirement (EER) is defined as the average
dietary energy intake that is predicted to maintain energy balance
in a healthy, adult of a defined age, gender, weight, height, and
level of physical activity consistent with good health. To calculate
the EER, prediction equations for normal weight individuals were
developed from data on total daily energy expenditure measured
by the doubly labeled water technique. In children and pregnant
or lactating women, the EER includes the needs associated with
the deposition of tissues or the secretion of milk at rates consistent
with good health. While the expected between-individual variabil-
ity is calculated for the EER, there is no Recommended Dietary
Allowance (RDA) for energy because energy intakes above the
EER would be expected to result in weight gain. Similarly, the
Tolerable Upper Intake Level (UL) concept does not apply to
107
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108 DIETARY REFERENCE INTAKES
energy, because any intake above an individual’s energy require-
ment would lead to undesirable (and potentially hazardous)
weight gain.
BACKGROUND INFORMATION
Humans and other mammals constantly need to expend energy to
perform physical work; to maintain body temperature and concentration
gradients; and to transport, synthesize, degrade, and replace small and
large molecules that make up body tissue. This energy is generated by the
oxidation of various organic substances, primarily carbohydrates, fats, and
amino acids. In 1780, Lavoisier and LaPlace measured the heat produc-
tion of mammals by calorimetry (Kleiber, 1975). They demonstrated that
it was equal to the heat released when organic substances were burned,
and that the same quantities of oxygen were consumed by animal metabo-
lism as were used during the combustion of the same organic substrates
(Holmes, 1985). Indeed, it has been verified by numerous experiments on
animals and humans since then that the energy produced by oxidation of
carbohydrates and fats in the body is the same as the heat of combustion
of these substances (Kleiber, 1975). The crucial difference is that in organ-
isms oxidation proceeds through many steps, allowing capture of some of
the energy in an intermediate chemical form—the high energy pyrophos-
phate bond of adenosine triphosphate (ATP). Hydrolysis of these high-
energy bonds can then be coupled to various chemical reactions, thereby
driving them to completion, even if by themselves they would not proceed
(Lipmann, 1941). Typically, the rates of energy expenditure in adults at
rest are slightly less than 1 kcal/min in women (i.e., 0.8 to 1.0 kcal/min or
1,150 to 1,440 kcal/d), and slightly more than 1 kcal/min in men (i.e., 1.1 to
1.3 kcal/min or 1,580 to 1,870 kcal/d) (Owen et al., 1986, 1987). One
kcal/min corresponds approximately to the heat released by a burning
candle or by a 75-watt light bulb (i.e., 1 kcal/min corresponds to 70 J/sec
or 70 W).
Energy Yields from Substrates
Carbohydrate, fat, protein, and alcohol provide all of the energy sup-
plied by foods and are generally referred to as macronutrients (in contrast
to vitamins and elements, usually referred to as micronutrients). The amount
of energy released by the oxidation of carbohydrate, fat, protein, and
alcohol (also known as Heat of Combustion, or ∆H) is shown in Table 5-1.
When alcohol (ethanol or ethyl alcohol) is consumed, it promptly
appears in the circulation and is oxidized at a rate determined largely by
its concentration and by the activity of liver alcohol dehydrogenase. Oxi-
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109
E NERGY
TABLE 5-1 Heat of Combustion of Various Macronutrients
Heat of
Combustiona Atwater Factord
kcalb/L O 2 RQc (CO2/O2)
Macronutrient (kcal/g) (kcal/g)
Starch 4.18 5.05 1.0 4.0
Sucrose 3.94 5.01 1.0 4.0
Glucose 3.72 4.98 1.0 4.0
Fat 9.44 4.69 0.71 9.0
Protein by 5.6
combustiona
Protein through 4.70 4.66 0.835 4.0
metabolisma
Alcohole 7.09 4.86 0.67 —
a The energy derived by protein oxidation in living organisms is less than the heat of
combustion of protein, because the nitrogen-containing end product of metabolism in
mammals is urea (or uric acid in birds and reptiles), whereas nitrogen is converted into
nitrous oxide when protein is combusted. The heat liberated by biological oxidation of
proteins was long thought to be 4.3 kcal/g (Merrill and Watt, 1973), but a more recent
demonstration showed that the actual value is 4.7 kcal/g (Livesey and Elia, 1988).
b One calorie is the amount of energy needed to increase the temperature of 1 g of
water from 14.5˚ to 15.5˚C. In the context of foods and nutrition, “large calorie” (i.e.,
Calories, with a capital C), which is more properly referred to as “kilocalorie” (kcal),
has been traditionally used. In the International System of Units, the basic energy unit
is the Joule (J). One J = 0.239 calories, so that 1 kcal = to 4.186 kJ. A daily energy
expenditure of 2,400 kcal corresponds to the expenditure of 10,000 kJ, or 10 MJ (Mega
Joules)/d.
c RQ = respiratory quotient, which is defined as the ratio of CO produced divided by O
2 2
consumed (in terms of mols, or in terms of volumes of CO2 and O2).
d Atwater, a pioneer in the study and characterization of nutrients and metabolism,
proposed to use the values of 4, 9, and 4 kcal/g of carbohydrate, fat, and protein,
respectively (Merrill and Watt, 1973). This equivalent is now uniformly used in nutrient
labeling and diet formulation. Nutrition Labeling of Food. 21 C.F.R. §101.9 (1991).
e Alcohol (ethanol) content of beverages is usually described in terms of percent by
volume. The heat of combustion of alcohol is 5.6 kcal/mL. (One mL of alcohol weighs
0.789 g.)
dation of alcohol elicits a prompt reduction in the oxidation of other
substrates used for ATP regeneration, demonstrating that ethanol oxida-
tion proceeds in large part via conversion to acetate and oxidative phos-
phorylation. The phenomenon has been precisely measured by indirect
calorimetry in human subjects, in whom ethanol consumption was found
to primarily reduce fat oxidation (Suter et al., 1992). About 80 percent of
the energy liberated by ethanol oxidation is used to drive ATP regenera-
tion, so that the thermic effect of ethanol comes to about 20 percent (Siler
et al., 1999). The thermic effect of food is the increase in energy expendi-
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110 DIETARY REFERENCE INTAKES
ture as measured by heat produced upon ingestion of that food. The
thermic effect of alcohol is about twice the thermic effect of carbohydrate,
but less than the thermic effect of protein (see later section, “Thermic
Effect of Food”).
Reported food intake in individuals consuming alcohol is often similar
to that of individuals who do not consume alcohol (de Castro and Orozco,
1990). As a result, it has sometimes been questioned whether alcohol con-
tributes substantially to energy production. However, the biochemical and
physiological evidence about the contribution made by ethanol to oxidative
phosphorylation is so unambiguous that the apparent discrepancies
between energy intake data and body weights must be attributed to
inaccuracies in reported food intakes. In fact, in individuals consuming a
healthy diet, the additional energy provided by alcoholic beverages can be
a risk factor for weight gain (Suter et al., 1997), as opposed to alcoholics in
whom the pharmacological impact of excessive amounts of ethanol tends
to inhibit normal eating and may cause emaciation.
Energy Requirements Versus Nutrient Requirements
Recommendations for nutrient intakes are generally set to provide an
ample supply of the various nutrients needed (i.e., enough to meet or
exceed the requirements of almost all healthy individuals in a given life
stage and gender group). For most nutrients, recommended intakes are
thus set to correspond to the median amounts sufficient to meet a specific
criterion of adequacy plus two standard deviations to meet the needs of
nearly all healthy individuals (see Chapter 1). However, this is not the case
with energy because excess energy cannot be eliminated, and is eventually
deposited in the form of body fat. This reserve provides a means to main-
tain metabolism during periods of limited food intake, but it can also
result in obesity.
The first alternate criterion that may be considered as the basis for a
recommendation for energy is that energy intake should be commensu-
rate with energy expenditure, so as to achieve energy balance. Although
frequently applied in the past, this is not appropriate as a sole criterion, as
described by the FAO/WHO/UNU publication, Energy and Protein Require-
ments (1985):
The energy requirement of an individual is a level of energy intake
from food that will balance energy expenditure when the indi-
vidual has a body size and composition, and level of physical activity,
consistent with long-term good health; and that would allow for
the maintenance of economically necessary and socially desirable
physical activity. In children and pregnant or lactating women
the energy requirement includes the energy needs associated with
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111
E NERGY
the deposition of tissues or the secretion of milk at rates consis-
tent with good health (p. 12).
This definition indicates that desirable energy intakes for obese indi-
viduals are less than their current energy expenditure, as weight loss and
establishment of a steady state at a lower body weight is desirable for them.
In underweight individuals, on the other hand, desirable energy intakes
are greater than their current energy expenditure to permit weight gain
and maintenance of a higher body weight. Thus, it seems logical to base
estimated values for energy intake on the amounts of energy that need to
be consumed to maintain energy balance in adult men and women who
are maintaining desirable body weights, taking into account the incre-
ments in energy expenditure elicited by their habitual level of activity.
There is another fundamental difference between the requirements
for energy and those for other nutrients. Body weight provides each indi-
vidual with a readily monitored indicator of the adequacy or inadequacy of
habitual energy intake, whereas a comparably obvious and individualized
indicator of inadequate or excessive intake of other nutrients is not usually
evident.
Energy Balance
Because of the effectiveness in regulating the distribution and use of
metabolic fuels, man and animals can survive on foods providing widely
varying proportions of carbohydrates, fats, and proteins. The ability to
shift from carbohydrate to fat as the main source of energy, coupled with
the presence of substantial reserves of body fat, makes it possible to accom-
modate large variations in macronutrient intake, energy intake, and energy
expenditure. The amount of fat stored in an adult of normal weight com-
monly ranges from 6 to 20 kg. Since one gram of fat provides 9.4 kcal,
body fat energy reserves thus range typically from approximately 50,000 to
200,000 kcal, providing a large buffer capacity as well as the ability to
provide energy to survive for extended periods (i.e., several months) of
severe food deprivation. Large daily deviations from energy balance are
thus readily tolerated, and accommodated primarily by gains or losses of
body fat (Abbott et al., 1988; Stubbs et al., 1995). Coefficients of variation
for intra-individual variability in daily energy intake average ± 23 percent
(Bingham et al., 1994); variations in physical activity are not closely syn-
chronized with adjustments in food intake (Edholm et al., 1970). Thus,
substantial positive as well as negative energy balances of several hundred
kcal/d occur as a matter of course under free-living conditions among
normal and overweight subjects. Yet over the long term, energy balance is
maintained with remarkable accuracy. Indeed, during long periods in the
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112 DIETARY REFERENCE INTAKES
life of most individuals, gains or losses of adipose tissue are less than 1 to 2
kg over a year (McCargar et al., 1993), implying that the cumulative error
in adjusting energy intake to expenditure amounts to less than 2 percent
of energy expenditure.
Components of Energy Expenditure
Basal and Resting Metabolism
The basal metabolic rate (BMR) describes the rate of energy expendi-
ture that occurs in the postabsorptive state, defined as the particular con-
dition that prevails after an overnight fast, the subject having not consumed
food for 12 to 14 hours and resting comfortably, supine, awake, and motion-
less in a thermoneutral environment. This standardized metabolic state
corresponds to the situation in which food and physical activity have minimal
influence on metabolism. The BMR thus reflects the energy needed to
sustain the metabolic activities of cells and tissues, plus the energy needed
to maintain blood circulation, respiration, and gastrointestinal and renal
processing (i.e., the basal cost of living). BMR thus includes the energy
expenditure associated with remaining awake (the cost of arousal), reflect-
ing the fact that the sleeping metabolic rate (SMR) during the morning is
some 5 to 10 percent lower than BMR during the morning hours (Garby
et al., 1987).
BMR is commonly extrapolated to 24 hours to be more meaningful,
and it is then referred to as basal energy expenditure (BEE), expressed as
kcal/24 h. Resting metabolic rate (RMR), energy expenditure under rest-
ing conditions, tends to be somewhat higher (10 to 20 percent) than under
basal conditions due to increases in energy expenditure caused by recent
food intake (i.e., by the “thermic effect of food”) or by the delayed effect
of recently completed physical activity (see Chapter 12). Thus, it is impor-
tant to distinguish between BMR and RMR and between BEE and resting
energy expenditure (REE) (RMR extrapolated to 24 hours).
Basal, resting, and sleeping energy expenditures are related to body
size, being most closely correlated with the size of the fat-free mass (FFM),
which is the weight of the body less the weight of its fat mass. The size of
the FFM generally explains about 70 to 80 percent of the variance in RMR
(Nelson et al., 1992; Ravussin et al., 1986). However, RMR is also affected
by age, gender, nutritional state, inherited variations, and by differences
in the endocrine state, notably (but rarely) by hypo- or hyperthyroidism.
The relationships among RMR, body weight, and FFM are illustrated in
Figures 5-1 and 5-2 (Owen, 1988), which show that differences in RMR
relative to body weight among diverse individuals such as men, women,
and athletes mostly disappear when RMR is considered relative to FFM.
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E NERGY
3,000
RMR (k cal/24 h)
2,000
1,000
0
Weight (kg)
FIGURE 5-1 Resting metabolic rates (RMR) are contrasted against the weights of
44 lean ( ) and obese (●) healthy women, 8 of whom were athletes (⊕), and 60
lean (∆) and obese ( ) healthy men. Reprinted, with permission, from Owen
(1988). Copyright 1988 by W.B. Saunders.
3,000
2,000
RMR (k cal/24 h)
1,000
0
FFM (kg)
FIGURE 5-2 Resting metabolic rates (RMR) are contrasted against the fat-free
masses (FFM) of 44 lean ( ) and obese (●) healthy women, 8 of whom were
athletes (⊕), and 60 lean (∆) and obese ( ) healthy men. Reprinted, with permis-
sion, from Owen (1988). Copyright 1988 by W.B. Saunders.
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114 DIETARY REFERENCE INTAKES
BEE has been predicted from age, gender, and body size. Prediction
equations were developed for each gender (WN Schofield, 1985) by pool-
ing and analyzing reported measurements made in 7,393 individuals. A
recent re-evaluation of all available data performed by Henry (2000) has
led to a new set of predicting equations.
Thermic Effect of Food
It has long been known that food consumption elicits an increase in
energy expenditure (Kleiber, 1975). Originally referred to as the Specific
Dynamic Action (SDA) of food, this phenomenon is now more commonly
referred to as the thermic effect of food (TEF). The intensity and duration
of meal-induced TEF is determined primarily by the amount and composi-
tion of the foods consumed, mainly due to the metabolic costs incurred in
handling and storing ingested nutrients (Flatt, 1978). Activation of the
sympathetic nervous system elicited by dietary carbohydrate and by sensory
stimulation causes an additional, but modest, increase in energy expendi-
ture (Acheson et al., 1983). The increments in energy expenditure during
digestion above baseline rates, divided by the energy content of the food
consumed, vary from 5 to 10 percent for carbohydrate, 0 to 5 percent for
fat, and 20 to 30 percent for protein. The high TEF for protein reflects the
relatively high metabolic cost involved in processing the amino acids
yielded by absorption of dietary protein, for protein synthesis, or for the
synthesis of urea and glucose (Flatt, 1978; Nair et al., 1983). Consumption
of the usual mixture of nutrients is generally considered to elicit increases
in energy expenditure equivalent to 10 percent of the food’s energy con-
tent (Kleiber, 1975). Since TEF occurs during a limited part of the day
only, it can result in noticeable increases in REE if energy expenditure is
measured during the hours following meals.
Thermoregulation
Birds and mammals, including humans, regulate their body tempera-
ture within narrow limits. This process, termed thermoregulation, can elicit
increases in energy expenditure that are greater when ambient tempera-
tures are below the zone of thermoneutrality. The environmental tem-
perature at which oxygen consumption and metabolic rate are lowest is
described as the critical temperature or thermoneutral zone (Hill, 1964).
Because most people adjust their clothing and environment to maintain
comfort, and thus thermoneutrality, the additional energy cost of thermo-
regulation rarely affects total energy expenditure to an appreciable extent.
However, there does appear to be a small influence of ambient tempera-
ture on energy expenditure as described in more detail below.
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Physical Activity
The energy expended for physical activity varies greatly among indi-
viduals as well as from day to day. In sedentary individuals, about two-
thirds of total energy expenditure goes to sustain basal metabolism over
24 hours (the BEE), while one-third is used for physical activity. In very
active individuals, 24-hour total energy expenditure can rise to twice as
much as basal energy expenditure (Grund et al., 2001), while even higher
total expenditures occur among heavy laborers and some athletes.
The efficiency with which energy from food is converted into physical
work is remarkably constant when measured under conditions where body
weight and athletic skill are not a factor, such as on bicycle ergometers
(Kleiber, 1975; Nickleberry and Brooks, 1996; Pahud et al., 1980). For
weight-bearing physical activities, the cost is roughly proportional to body
weight. In the life of most persons, walking represents the most significant
form of physical activity, and many studies have been performed to deter-
mine the energy expenditures induced by walking or running at various
speeds (Margaria et al., 1963; Pandolf et al., 1977; Passmore and Durnin,
1955). Walking at a speed of 2 mph is considered to correspond to a mild
degree of exertion, walking speeds of 3 to 4 mph correspond to moderate
degrees of exertion, and a walking speed of 5 mph to vigorous exertion
(Table 12-1, Fletcher et al., 2001). Over this range of speeds, the increment
in energy expenditure amounts to some 60 kcal/mi walked for a 70-kg
individual, or 50 kcal/mi walked for a 57-kg individual (see Chapter 12,
Figure 12-4). The exertion caused by walking/jogging increases progres-
sively at speeds of 4.5 mph and beyond, reaching 130 kcal/mi at 5 mph for
a 70-kg individual.
The increase in daily energy expenditure is somewhat greater, how-
ever, because exercise induces an additional small increase in expenditure
for some time after the exertion itself has been completed. This excess
post-exercise oxygen consumption (EPOC) depends on exercise intensity
and duration and has been estimated at some 15 percent of the increment
in expenditure that occurs during exertions of the type described above
(Bahr et al., 1987). This raises the cost of walking at 3 mph to 69 kcal/mi
(60 kcal/mi × 1.15) for a 70-kg individual and to 58 kcal/mi (50 kcal/mi ×
1.15) for a 57-kg individual. Taking into account the dissipation of 10 percent
of the energy consumed on account of the thermic effect of food to cover the
expenditure associated with walking, then walking 1 mile raises daily energy
expenditure to 76 kcal/mi (69 kcal/mi × 1.1) in individuals weighing 70 kg,
or 64 kcal/mi (58 kcal/mi × 1.1) for individuals weighing 57 kg. Since the cost
of walking is proportional to body weight, it is convenient to consider that the
overall cost of walking at moderate speeds is approximately 1.1 kcal/mi/kg
body weight (75 kcal/mi/70 kg or 64 kcal/mi/57 kg). The effects of varia-
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116 DIETARY REFERENCE INTAKES
tions in body weights and the impact of various physical activities on energy
expenditure are considered in more detail in Chapter 12.
Physical Activity Level
The level of physical activity is commonly described as the ratio of
total to basal daily energy expenditure (TEE/BEE). This ratio is known as
the Physical Activity Level (PAL), or the Physical Activity Index. Describ-
ing physical activity habits in terms of PAL is not entirely satisfactory
because the increments above basal needs in energy expenditure, brought
about by most physical activities where body weight is supported against
gravity (e.g., walking, but not cycling on a stationary cycle ergometer), are
directly proportional to body weight, whereas BEE is more nearly propor-
tional to body weight0.75. However, PAL is a convenient comparison and is
used in this report to describe and account for physical activity habits. The
effect of variations in activities on PAL is described in Chapter 12.
Total Energy Expenditure
Total Energy Expenditure (TEE) is the sum of BEE (which includes a
small component associated with arousal, as compared to sleeping), TEF,
physical activity, thermoregulation, and the energy expended in deposit-
ing new tissues and in producing milk. With the emergence of informa-
tion on TEE by the doubly labeled water (DLW) method (Schoeller, 1995),
it has become possible to determine energy expenditure of infants, chil-
dren, and adults under free-living conditions. TEE from doubly labeled
water does not include the energy content of the tissue constituents laid
down during normal growth and pregnancy or the milk produced during
lactation, as it refers to energy expended during oxidation of energy-
yielding nutrients to water and carbon dioxide.
It should be noted that direct measurements of TEE represent a dis-
tinct advantage over previous TEE evaluations, which had to rely on the
factorial approach and on food intake data, which have limited accuracy
due to the inability to reliably determine average physical activity cost and
nutrient intakes.
Estimated Energy Requirement
Information on energy expenditure obtained by DLW studies con-
ducted by a number of research units (see Appendix I) are used in this
report to estimate energy requirements, taking into account estimates of
the energy content of new body constituents during growth and preg-
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117
E NERGY
nancy and of the milk produced during lactation. Energy expenditure
depends on age and varies primarily as a function of body size and physical
activity, both of which vary greatly among individuals. Recommendations
about energy intake vary accordingly, and are also subject to the criterion
that an individual adult’s body weight should remain stable and within the
healthy range.
SELECTION OF INDICATORS FOR ESTIMATING
THE REQUIREMENT FOR ENERGY
Reported Energy Intake
The reported energy intakes of weight-stable subjects (i.e., those in
energy balance) could, in principle, be used to predict energy require-
ments for weight maintenance. However, it is now widely recognized that
reported energy intakes in dietary surveys underestimate usual energy
intake (Black et al., 1993).
The most compelling evidence about underreporting has come from
measurements of total energy expenditure (TEE) by the doubly labeled
water (DLW) method (Schoeller, 1995). The use of a measure or estimate
of TEE to validate instruments that measure food intake is dependent on
the principle of energy balance. That is, in weight-stable adults, energy
intake must equal TEE. By comparing reported energy intake to TEE, the
accuracy of food intake reporting can be assessed (Goldberg et al., 1991a).
A large body of literature documents the underreporting of food
intake, which can range from 10 to 45 percent depending on the age,
gender, and body composition of individuals in the sample population
(Johnson, 2000). Underreporting tends to increase as children grow older
(Livingstone et al., 1992b), is worse among women than in men (Johnson
et al., 1994), and is more pronounced among overweight and obese than
among lean individuals (Bandini et al., 1990a; Lichtman et al., 1992;
Prentice et al., 1986). Low socioeconomic status, characterized by low
income, low educational attainment, and low literacy levels increase the
tendency to underreport energy intakes (Briefel et al., 1997; Johnson et
al., 1998; Price et al., 1997; Pryer et al., 1997). Ethnic differences affecting
sensitivities and psychological perceptions relating to eating and body
weight can also affect the accuracy of reported food intakes (Tomoyasu et
al., 2000). Finally, individuals with infrequent symptoms of hunger under-
report to a greater degree than those who experience frequent hunger
(Bathalon et al., 2000).
There is some evidence suggesting that underreporters often fail to
report foods perceived to be bad or sinful, such as cakes/pies, savory
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Representative terms from entire chapter:
physical activity
