CHAPTER 4

B A S I C S O F M E TA B O L I S M

■ Energy Transductions in the Biosphere

M etabolism can be defined as the sum total of
processes occurring in a living organism. Be-
cause heat is produced by those processes, the meta- Our lives depend on conversions of chemical en-
bolic rate is indicated by the rate of heat production. ergy to other forms of energy. These conversions, or
All processes of metabolism ultimately depend on transductions, of energy are limited by the two laws
biological oxidation, so measuring the rate of O2 of thermodynamics, which apply to physical as well
consumption yields a good estimate of the rate of as biological energy transductions.
heat production, or metabolic rate. The maximum In the biological world (the biosphere), there are
capability of an individual to consume oxygen three major stages of energy transduction: photo-
1VO2 max 2 is highly related to that individual’s ability
#
synthesis, cell respiration, and cell work.
to perform hard work over prolonged periods. A The photosynthesis of sugars is illustrated by
high capacity to consume and utilize O2 indicates a Equation 4-1. In photosynthesis, the G is positive
high metabolic capacity. in sign. Energy is put in.

Figure 4-1 A photo from the classical study of

human metabolic and cardioventilatory responses
to exercise by H. M. Smith, 1922.

43
44 BASICS OF METABOLISM

Energy 1sunlight 2  6 CO2  6 H2O S C6H12O6  6 O2 (4-1)

C6H12O6  6 O2 S 6 CO2  6 H2O  Energy 1heat  work 2 (4-2)

ATP  Actin  Myosin ¡ Actomyosin  Pi  ADP  Energy 1Heat  Work2

Ca 2
(4-3)

Cell respiration can be illustrated by Equation is the sum of all transformations of energy and mat-
4-2. In cell respiration, the G is negative in sign. ter that occur within an organism. In other words,
Energy is given up and the process is associated by this definition, metabolism is everything going
with the production of the important high-energy on. It is not possible to measure that. Therefore, an-
intermediate compound, ATP. other operational definition has been developed,
There are many types of cellular work, includ- stating that metabolism is the rate of heat produc-
ing mechanical, synthetic, chemical, osmotic, and tion. This definition takes advantage of the fact that
electrical forms. Muscle contraction (a chemical- all the cellular events result in heat. By determin-
mechanical energy transduction) was described in ing the heat produced, one can obtain a measure of
Chapter 3 and is again illustrated by Equation 4-3. metabolism.
Here, actin and myosin are the contractile proteins The basic unit of heat measurement is the calo-
and the release of Ca2 within the muscle cell trig- rie. Simply defined, a calorie is the heat required to
gers the reaction. raise the temperature of 1 gram of water 1 degree
Although it may appear that our functioning Celsius. The calorie is a very small quantity, so the
depends on only two of these three major energy term kilocalories (kcal) is frequently used instead. A
transductions (respiration and cell work), in real- kilocalorie represents 1000 calories. Because heat
ity we are ultimately dependent on photosynthesis. must be measured to determine metabolic rate, this
The products of photosynthesis give us the oxygen procedure is termed calorimetry. Several types of
we breathe and the food we eat. Cell respiration is a calorimetry are currently used. They are dia-
reversal of photosynthesis. Have you thanked a grammed in Figure 4-3.
green plant today? Direct calorimetry, involving the direct measure-
ment of heat, is technically very difficult. However,

■ Metabolism and Heat Production

Metabolism Calorimetry
in Animals

One characteristic of living animals is that they give

off heat. As illustrated in Figure 4-2, for a body at Direct Indirect
rest, life processes result in heat production.
Scientists have developed two definitions of me-
tabolism. A functional definition is that metabolism O2 consumption Carbon
and
nitrogen
Respiration balance
Foodstuffs + O2 ATP + Heat
Open Closed
Cell work
circuit circuit

Figure 4-3 Relationship between metabolism and dif-

Heat ferent methods of calorimetry. Because the processes
of metabolism result in heat production, measuring
Figure 4-2 Metabolism and heat production. heat production gives an estimate of the metabolic
In a body at rest, all metabolic processes rate. Heat production can be measured directly (direct
eventually result in heat production. Measur- calorimetry), or it can be estimated from O2 consump-
ing heat production (calorimetry) gives the tion or from the carbon and nitrogen excreted (indirect
metabolic rate. calorimetry).
 Early Attempts at Calorimetry 45

Foodstuffs + O2 Heat + CO2 + H 2 O Insulation consisting

of ice water

Indirect Direct Ice water

calorimetry calorimetry Ice
Measure either Air Air
in out
Figure 4-4 The principle of indirect calorimetry
(measuring O2 consumption) as a basis of esti-
mating heat production. Instead of measuring
the heat produced as the result of biological
reactions, we measure the O2 used to support
biological oxidations.

it has been determined that indirect calorimetry, the

measurement of oxygen consumption, is also a valid
and technically reliable procedure for measuring
Ice
metabolic rate. The principle of indirect calorimetry
is illustrated in Figure 4-4. Another form of indirect
calorimetry involves determining the carbon and ni-
trogen content of excreted materials.

■ Early Attempts at Calorimetry

To understand the relationship between heat pro-

Figure 4-5 Lavoisier’s calorimeter of 1780. The animal’s
duction and O2 consumption as alternative meth- body heat melts the ice. Knowing that 80 kcal of heat
ods for determining metabolic rate, let us consider melts 1000 grams of ice, we can measure the amount
some of the work of the eighteenth-century genius, of water formed to estimate the heat produced. The ice
French chemist Antoine Lavoisier. water surrounding the calorimeter provides a perfect
(adiabatic) insulation because it is at the same tempera-
Because of his interest in studying living crea-
ture as the ice in the inner jacket around the animal’s
tures, Lavoisier came to recognize certain character- chamber. The insulation will neither add heat to nor take
istics of living animals: They give off heat and they heat from the calorimeter. Based on original sources and
breathe. Dead animals do not give off heat and do Kleiber, 1961. Used with permission.
not breathe. Lavoisier’s calorimeter, diagrammed in
Figure 4-5, is simple but beautiful in its design. By equal amounts. Lavoisier also determined that mat-
allowing the animal’s warmth to melt the ice, and ter gains weight when it burns. It had been thought
knowing the quantity of heat required to melt a previously that burning represented the loss of sub-
given quantity of ice, Lavoisier could calculate the stance, sometimes called phlogiston.
heat produced by the animal by measuring the vol- With information obtained from his experi-
ume of water produced. Such a device is called a di- ments, Lavoisier was able to interpret some earlier
rect calorimeter because it determines metabolism by findings. For instance, Boyle had shown that air
measuring heat produced. was necessary to have a flame, and Mayow had
Lavoisier’s respirometer (Figure 4-6) was an- observed that a burning candle and an animal to-
other device that was novel for its time. With it gether in an airtight container expired at the same
Lavoisier could establish that something in the air time. The fire of life and the fire of physical burning
(O2) was consumed by the animal and that some- depended on the same substance in the air, which
thing else (CO2) was produced in approximately Lavoisier called oxygène.
46 BASICS OF METABOLISM

5 5 5 0

0 0 0 5

5 5 5

NaOH

(a) (b) (c) (d)

Figure 4-6 Lavoisier’s respirometer of 1784. (a) A glass bell jar rests on a bed of mercury.
(b) An animal is placed in the jar from beneath the mercury seal and is left there for several
hours. The apparent respirometer volume increases when the animal enters, but then the
volume decreases very slowly if at all because O2 is being replaced by CO2. (c) The animal
is removed, and the volume is observed to have decreased slightly. (d) Addition of NaOH (a
CO2 absorber) into the jar results in a decrease in the measured volume. From these volume
# # #
changes, O2 consumption (VO2) and carbon dioxide production (VCO2) can be measured: VO2 
#
Va  Vd ; VCO2  Vc  Vd. Based on original sources and Kleiber, 1961. Used with permission.

The belief of Lavoisier and others that biological The Atwater and Rosa device (Figure 4-8) is
oxidation took place in the lungs has led to some an important apparatus for the study of metabo-
confusion. Although it is true that breathing, or ven- lism. Large enough to accommodate a person, it has
tilation, takes place in the lungs and associated the capability of determining heat production, O2
organs, respiration, or biological oxidation, takes consumption, and CO2 production simultaneously.
place in most of the body’s cells. Therefore, in this Through this device, the relationship between di-
text, we shall use the term respiration to denote cel- rect and indirect calorimetry was established. Thus,
lular oxidations and ventilation to denote pulmo- it is now possible to predict metabolic rate (heat
nary gas exchange. production) on the basis of determinations of
Devices such as Lavoisier’s respirometer are O2 consumption and CO2 production in resting
called indirect calorimeters because they estimate individuals.
heat production by determining O2 consumption or The calorimeter illustrated in Figure 4-9 is called
CO2 production. Lavoisier’s device is also referred a bomb calorimeter. In this device, foodstuffs are ig-
to as a closed-circuit indirect calorimeter because nited and burned in O2 under pressure. Through
the animal breathes gas within a sealed system. this device, the heats of combustion (H) of partic-
Haldane’s respirometer (Figure 4-7) is an ex- ular foods can be determined.
ample of an open-circuit indirect calorimeter. This Table 4-1 presents the relationships among
system is open to the atmosphere, and the animal caloric equivalents for combustion of various food-
breathes air. Today, the type of calorimeters most stuffs as determined by indirect and direct calo-
frequently used are open-circuit indirect designs. rimetry as well as by bomb calorimetry. Perhaps the
 Early Attempts at Calorimetry 47

Air
in

(a) (b) (c) (d) (e)

Soda lime H2SO4 H2SO4 Soda lime H2SO4 trap Flow meter
(CO2 trap) (H2O trap) (H2O trap) (CO2 trap) to absorb and pump
H2O from
•
trap d
VCO2 = ∆d /time + ∆e /time

Figure 4-7 Haldane’s respirometer. This device is an open-circuit indirect calorimeter, in which carbon
dioxide and water vapor in air entering the system are removed by traps (a) and (b), respectively. Trap
(c) removes the animal’s expired H2O vapor. Increase in weight of the soda lime CO2 trap (d) gives the
animal’s CO2 production. Based on original sources and Kleiber, 1961. Used with permission.

most interesting feature of this table is that, with a tion 4-4 shows why the RQ of glucose, a sugar car-
single exception, the caloric equivalents for the bohydrate, is unity:
combustion of foodstuffs inside and outside the
body are the same. Protein is the exception because C6H12O6  6 O2 S 6 CO2  6 H2O
nitrogen, an element unique to protein, is not oxi- 6 CO2 produced/6 O2 consumed (4-4)
dized within the body but is eliminated, chiefly in  1.0  RQ
urine but also in sweat. Therefore, the caloric equiv-
For the neutral fat trioleate, the RQ approxi-
alent of protein metabolism is approximately 26%
mates 0.7:
less than in a bomb calorimeter.
Table 4-1 also gives the caloric equivalents of C57H104O6  80 O2 S 57 CO2  52 H2O
foodstuffs in kilocalories per liter of O2 consumed. 57 (4-5)
Although fat, because of its relatively high carbon RQ   0.71
80
and hydrogen content (reduction), contains more
potential chemical energy on a per-unit-weight ba- During hard exercise, an individual’s respira-
sis, carbohydrates give more energy when com- tory gas exchange ratio [R, an estimate of RQ (see
busted in a given volume of O2. page 53)] approaches 1.0, whereas during pro-
In addition to providing an estimate of meta- longed exercise, the R may be somewhat lower, 0.9
bolic rate, indirect calorimetry provides a means of or less. Figure 4-10 shows data on male subjects run-
estimating the composition of the fuels oxidized. ning a marathon on a treadmill where respiratory
Similarly, determining the ratio of CO2 produced gas exchange could be measured. In one case, sub-
# #
(VCO2) to O2 consumed (VO2) gives an indication of jects ran at their race pace, whereas in the other case
the type of foodstuff being combusted. This ratio subjects ran more slowly. Note that subjects’ race
# #
(VCO2/VO2) is usually referred to as the respiratory pace corresponds to an R of 0.95 to 1.0. Note also
quotient (RQ) and reflects cellular processes. Equa- that in later stages, R declined, but then rose at the
48 BASICS OF METABOLISM

Voltmeter
Water
temperature
1 ( T1 )
Water Heat production =
(water T2 – T1) (flow)
WallT2
Water flow
Wall T1

Electric Heat exchanger

heater

Air
Weight

Manometer

Heater coils
Insulation

Air
pump
Flow meter
H2SO4 Soda lime H2SO4 O2 consumption =
O2 Flow of O2
H2SO4 Soda lime H2SO4 source necessary
(expired (expired (H2O vapor to keep
H2O vapor CO2 trap) trap from manometer
trap) soda lime) level

Figure 4-8 Atwater–Rosa calorimeter. A direct calorimeter suitable to accommodate a resting human
and simultaneously determine that individual’s O2 consumption and CO2 production. Through this device,
direct and indirect calorimetry were correlated. O2 consumption is equal to the volume of O2 added to
keep the internal (manometer) pressure constant. In the calorimeter, heat loss through the walls is pre-
vented by heating the middle wall (wall T2) to the temperature of the inner wall (wall T1). Metabolic heat
production is then picked up in the water heat exchanger. Based on original sources and Kleiber, 1961.

end. In the slower marathon runners, R was lower, in Equation 4-6, in oxidizing carbohydrate rather
indicating less carbohydrate and more lipid use than fat the individual derives
than in the faster runners. The fuel mix was, how-
5.0  4.7 kcal # liter 1 O2
ever, still mostly carbohydrate. (4-6)
4.7 kcal # liter 1 O2
Table 4-1 shows why it is an advantage for these
changes in RQ to occur. During hard exercise, O2 or 6.4% more energy per unit O2 consumed. During
consumption can be limiting. Therefore, as shown prolonged exercise, however, it makes sense that
 Early Attempts at Calorimetry 49

Thermometer RQ decreases, indicating that more fat is com-

busted. In prolonged work, glycogen supply rather
than O2 consumption can be limiting. Table 4-1 in-
Water
stirrer dicates that on a mass basis, fats provide about
9.5/4.2 kcal g1, or 2.3 times as much energy as car-
#
bohydrate. Given this large difference, we can also
see why endurance training improves the ability to
use fat as a fuel during prolonged mild to moderate
#
intensity exercise (i.e., 40 – 60% VO2 max).
Water
The preceding type of discussion is sometimes
referred to as a “teleological argument,” meaning
Chamber
O2
that the purpose of something is assumed to explain
its operation. In actuality, as will be shown, the rea-
son why relatively more carbohydrate is used in
hard exercise is related to the quantity of activity
and regulation of glycolytic enzymes. There are also
enzymatic explanations for the preponderance of
fat used in prolonged exercise.
In order to obtain a precise estimate of meta-
Adiabatic water jacket
bolic rate and fuel used by means of indirect cal-
orimetry, we must know a few other details besides
Figure 4-9 Bomb calorimeter. A food substance the quantities of O2 consumed and CO2 produced.
is attached to the ignition wires and placed in the These additional parameters include the food in-
chamber under several atmospheres of O2 pres- gested and the nitrogen excreted. To provide a rela-
sure. The sample is then ignited and burns explo-
tively simple example of the utility of indirect cal-
sively. The stirrer distributes the heat of combus-
tion uniformly throughout the water surrounding orimetry, let us consider a starving man, in whom
the chamber. The thermometer detects the heat there is no food input to account for and no large ex-
released. Based on Kleiber, 1961. cretion of urinary nitrogen (Table 4-2).

TABLE 4-1

Caloric Equivalents of Foodstuffs Combusted Inside and Outside the Body

RQ Inside Body Outside Body

# #
Food #
kcal liter O21 (VCO2/VO2) #
(kcal g1) (kcal g1)#
Carbohydrate 5.05 1.00 4.2 4.2
Fat 4.70 0.70 9.5 9.5
Protein 4.50 0.80 4.2 5.7 a
Mixed diet 4.82 0.82
Starving individual 4.70 0.70

a The amount of protein combusted outside the body is greater than that combusted inside the body (see text):
5.7  4.2
 26% difference
5.7
50 BASICS OF METABOLISM

Figure 4-10 Calculated per-

centage of energy expenditure * * * *
contributed by carbohydrates 100 * Slow 1.00
(CHO) before, during, and after Fast
a treadmill marathon in fast and

Total energy as CHO (%)

0.95
slow groups. Values are means 80
 SEM; N  6 per group.
Source: O’Brien et al., 1993. 0.90
Used with permission.
60

Recovery Recovery 0.85

40
0.80

20
0.75

0.70
–10 40 90 140 190 240
Time (min)

TABLE 4-2

Calculation of Nitrogen-Free RQ on a Resting Starving Man

#
Given: (a) Protein is about 17% N by weight, or there is 1 g N 5.9 g1 protein (1 ⁄ 5.9  0.17).
(b) For protein RQ  4.9 ⁄ 5.9  0.83, or 4.9 liters CO2 are derived from the catabolism of the protein associated
with 1 g N, and 5.9 liters of O2 are required to catabolize the protein.

The total O2 consumption was 634 liters. The total CO2 production was 461 liters, and urinary N losses were 14.7 g over
24 hours. We can use these data to calculate the nitrogen-free RQ.

Calculations Total CO2 (liters) Total O2 (liters)

461 634
In the urine, there were 14.7 g N. The CO2 produced by
protein catabolism was (14.7) (4.9)  72.0 liters CO2. 72
The O2 consumed associated with protein catabolism
was (14.7) (5.9)  86.7 liters O2 86.7
389 547.3

389
Nonprotein RQ   0.71
547.3

Heat production
# #
From protein: (14.7 g N) (5.9 g protein g1 N) (4.2 kcal g1 protein)  364.3 kcal
From fat: The nonprotein RQ was 0.71, so fat comprised the remaining fuel. Therefore,
#
(547.3 liters O2) (4.7 kcal liter1 O2)  2572.3 kcal.

Total heat production  364.3  2572.3 kcal  2936.6 kcal

 The Utility of Indirect Calorimetry During Exercise 51

In exercise physiology, current estimates of fu-

els combusted are usually simplified by assuming
there is no increase in the basal amino acid and pro-
tein degradation during exercise. The ventilatory
exchange ratio R is then used to represent the non-
protein RQ. As we shall see later (Chapter 8), this
assumption is not quite valid.
Although both RQ and R are given by the same
# #
formula (VCO2/VO2), over any short period of mea-
surement of gas exchange at the lungs, changes in
CO2 storage may cause R not to equal RQ. Although
RQ does not exceed 1.0, R can reach 1.5 or higher.
For the present, let us consider RQ to be the ratio
# #
VCO2/VO2 in the cell, where O2 is consumed and CO2
produced. Further, let us consider R to be the ratio
# #
VCO2/VO2 measured at the mouth. Over time, R must
equal RQ, but during the onset and offset of exer-
cise, as well as during hard exercise, RRQ because
body CO2 storage changes (see Figure 4-14).

■ Indirect Calorimetry

For individuals at rest, indirect calorimetric deter- Figure 4-11 Bicycle ergometers are convenient, sta-
minations on the effects of body size, growth, dis- tionary laboratory devices to control the external work
rate (power output) while physiological responses to
ease, gender, drugs, nutrition, age, and environ-
standardized or experimental protocols are observed.
ment on metabolism are very useful. The resting Courtesy of Monark, Inc., Varberg, Sweden.
metabolic rate per unit body mass is greater in
males than in females, greater in children than in the
fore, devices to measure external work performed,
aged, greater in small individuals than in large ones,
such as bicycle ergometers (Figure 4-11) and tread-
and greater under extremes of heat and cold than
mills (Figure 4-12), are used.
under normal conditions.
During exercise, direct calorimeters such as the
Atwater–Rosa calorimeter (Figure 4-8) are of little
use for several reasons. First, such devices are very
■ The Utility of Indirect Calorimetry expensive. Second, the heat generated by an er-
During Exercise gometer, if it is electrically powered, may far exceed
that of the subject. Third, body temperature in-
Physical exercise represents a special metabolic sit- creases during exercise because not all the heat pro-
uation. As Figure 4-2 indicates, for a body at rest, all duced is liberated from the body. Therefore, the sen-
the energy liberated within appears as heat. If me- sors in the walls of the calorimeter do not pick up
tabolism is constant, the quantity of heat produced all the heat produced. Finally, the body sweats dur-
within the body over a period of time will be the ing exercise, which also affects the calorimeter and
same as that leaving the body. However, during ex- changes body mass. Changes in body mass and the
ercise, some of the energy liberated within the body unequal distribution of heat within the body make
appears as physical work outside the body. There- it very difficult to use direct calorimetry in exercise.
52 BASICS OF METABOLISM

Respiration
Foodstuffs ATP + Heat

(a)

Respiration
Foodstuffs ATP + Heat

(glycogen and Anaerobic

glucose)
Glycolysis

(b)

Figure 4-13 Respiration and ATP production. The validity

of indirect calorimetric measurements depends on the O2
consumption accurately representing the ATP formed.
This is not always the case in exercise. In (a), the mea-
surement is valid. In (b), it is invalid.

cellular events are not always immediately repre-

sented in expired air. This is because the cells are
fluid systems, and they are surrounded by other
fluid systems on both the arterial and venous sides.
Figure 4-12 Treadmills are frequently used in the labo- When exercise starts, CO2 is frequently stored in
ratory to apply exercise stress and record physiological
responses on relatively stationary subjects during exer-
cells. When exercise is very difficult, the blood bi-
cise. Compared to the bicycle ergometer (Figure 4-11), carbonate buffer system buffers lactic acid, and ex-
it is difficult to quantify external work on the treadmill. tra nonrespiratory CO2 is produced (Chapter 11).
However, the treadmill does allow subjects to walk or As illustrated in Figure 4-14a, lactic acid (HLA)
run, which are perhaps more common modes of locomo- is a strong acid whose level in muscle and blood in-
tion than is bicycling. Photo: © David Madison.
creases during heavy work. It is known as a strong
acid in physiological systems because it can readily
dissociate a proton (H). To lessen the effect of pro-
As with direct calorimetry during exercise, tons generated from lactic acid during hard exer-
techniques of indirect calorimetry have certain lim- cise, the body has a system of chemicals that lessen,
itations. These are summarized in Figure 4-13a. In or buffer, the effects of the acid. In the blood, the bi-
#
order for determinations of VO2 to reflect metabo- carbonate (HCO3)– carbonic acid (H2CO3) system
lism accurately, the situation in Figure 4-13a must is the main system by which the effects of lactic acid
hold. If another mechanism is used to supply en- are buffered. As shown in Figure 4-14a and in Equa-
ergy, such as that shown in Figure 4-13b, then respi- tions 4-7 to 4-9, HCO3 neutralizes the H, but CO2
ratory determinations do not completely reflect all is produced. This is eliminated at the lungs and ap-
metabolic processes. As will be seen, the body has pears in the breath. Consequently, during hard ex-
the means to derive energy from the degradation of ercise, R  RQ (Figure 4-14b). After exercise, meta-
substances without the immediate use of O2. These bolic CO2 may be stored in cells, blood, and other
mechanisms include immediate sources and rapid body compartments to make up for that lost during
glycogen (muscle carbohydrate) breakdown. Use of exercise.
# #
the VCO2/VO2 ratio is also limited during exercise. Al-
though over time the O2 consumed by and CO2 lib- HLA S H  LA (4-7)
erated at the lungs (the respiratory, or ventilatory H  HCO3 S H2CO3

(4-8)
exchange ratio) equals the respiratory quotient, the H2CO3 S H2O  CO2 (4-9)
 The Utility of Indirect Calorimetry During Exercise 53

Lungs
CO2
Cell
CO2 and Metabolic
HCO3− pools and
Nonmetabolic
O2 CO2
Fuel +
H+ CO2
HCO3− +
H2O

Metabolic CO2 + Buffering = Total CO2 Excretion

(Nonmetabolic CO2)

(a)

Figure 4-14 Diagrammatic

representations of the formation
of metabolic CO2 (from fuel oxi-
dation) and nonmetabolic CO2
1.3 (from the buffering of lactic acid)
Cell RQ and their effects on CO2 excre-
1.2 Ventilatory R
tion in the lungs and the whole-
R > RQ
body respiratory exchange ratio
1.1 # #
(RER, or R  VCO2 /VO2 ). In panel
a, body cells produce metabolic
1.0 CO2 as well as protons (H) from
glycolysis. The effect of protons
0.9 is lessened (buffered) by the
bicarbonate (HCO3) buffering
0.8 system. Metabolic and nonmeta-
bolic CO2 from acid buffering
0.7 R < RQ are excreted in the lungs. In
panel b, the transient effects
0.6 of acid buffering at the onset
Time
of exercise and restoration of
Rest HCO3 reserves in recovery are
illustrated. The initiation and ces-
Exercise Recovery
sation of exercise are conditions
(b) when R  RQ.
54 BASICS OF METABOLISM

Furthermore, during and immediately after ex-

TABLE 4-3
ercise, urine production by the kidney is inhibited.
Also, during exercise considerable nitrogen can be Estimates of Caloric Expenditures
lost as urea in sweat. Therefore, it is difficult to de- of Sports Activities for a 70-kg Person
termine the nitrogen excreted during exercise.
Caloric Expenditure
Determinations of indirect calorimetry are Activity #
(kcal min1)
somewhat limited in their use by the fact that the
respiratory gases give no specific information on the Archery 4.6
fuels used. If, for example, RQ is 1.0, then although Badminton 6.4
we know that carbohydrate was the fuel catabo- Basketball 9.8
Canoeing 7.3
lized, we do not know specifically which carbohy-
Cycling 12.0
drate was involved. The possibilities could include, Field hockey 9.5
among others, glycogen, glucose, lactic acid, and Fishing 4.4
pyruvic acid. However, radioactive and nonra- Football 9.4
dioactive tracers to study metabolism at rest and Golf 6.0
Gymnastics 4.7
during exercise have come into use in conjunction
Judo 13.8
with indirect calorimetry to provide more detailed Resting 1.2
information on specific fuels. Running
Exercise is also a special situation in that the #
8 min mi1 14.8
metabolic responses persist long after the exercise #
6 min mi1 17.9
Squash 15.1
itself may have been completed. Consequently,
Swimming
physical activity results in an excess postexercise O2 Backstroke 12.0
consumption (EPOC). This EPOC has sometimes in Crawl 11.1
the past been called the “O2 debt” and has been Tennis 7.7
used as a measure of anaerobic metabolism during Volleyball 3.6
Walking, easy 5.7
exercise. A more detailed explanation of the O2 debt
is given later (Chapter 10); suffice it to say here that
the mechanisms of the O2 debt are complex and
cannot be used to estimate anaerobic metabolism work. This fraction is frequently reported as a per-
during exercise. centage and is called efficiency.
Whereas the body does present certain prob- An example of how the efficiency of the human
lems in determining metabolic rate during exercise, body is calculated during bicycle ergometer exer-
careful consideration of those various factors allows cise is given in Equation 4-10. In Figure 4-15, we see
us to obtain important information about metabolic that the O2 consumption of an individual increases
#
responses to exercise. Estimations of VO2 , for ex- in direct response to increments in work load while
ample, provide information on the cardioventila- pedaling at constant speed. In this case, efficiency
tory response to exercise. The caloric cost of various can be calculated as in Equation 4-10.
exercises can be estimated (Table 4-3), and informa- The plateau steps in Figure 4-15a are referred to
tion about the fuels used to support the exercise can as steady-rate exercise. During the steady rate, the
#
be obtained. oxygen consumption (VO2) is relatively constant and
Knowing that part of the energy liberated dur- is directly proportional to the constant submaximal
ing exercise appears as external work is useful. By work load.
measuring the respiratory response to graded, sub- The calculation of body efficiency during bicy-
maximal exercise at specific external work rates, we cle exercise is given in Table 4-4. Here the calculated
can determine the fraction of the energy liberated value of efficiency is 29.2%, which is close to a max-
within the human machine that appears as external imum value for bicycle ergometer work. Cycling at
 The Utility of Indirect Calorimetry During Exercise 55

10

800
8 kg . m . min–1
Caloric output (kcal . min–1)

600
6 kg . m . min–1

400
kg . m . min–1
4
200
kg . m . min–1
2

Rest

0 3 6 9 12 0 200 400 600 800

Exercise time (min) Work rate (kg . m . min–1)
(a) (b)

Figure 4-15 Respiratory response to graded submaximal bicycle ergometer work.

Every 3 minutes
# the work rate is increased 200 kg m min1. The observed O2 con- # #
#
sumption (VO2) is converted to kcal min1. These values are then plotted as (a) a function
of time and (b) a function of the steady-rate work load. Note that a plot of the caloric cost
of exercise against work rate (b) yields a straight line, or one that bends upward slightly.

greater speeds and working at greater loads results

TABLE 4-4 in decrements in calculated efficiency. The efficiency
Calculation of Body Efficiency of walking is slightly higher than that of cycling, but
During Cycling Exercise responds similarly to increments in speed and resis-
tance. The reason it is usually easier to cycle from
Given: one place to another than to walk is that the rolling
#
VO2 at 200 kg m min1  0.76 liter min1 # # # and wind resistance to cycling at a particular speed
#
VO2 at 400 kg m min1  1.08 liters min1 # # # are far less than the work done in accelerating and
R  RQ  1.0
decelerating the limbs during walking—that is, less
work is done in cycling. Attempting to bicycle in
When RQ  1.0, 1 liter O2  5 kcal
soft sand will reveal that the work done in covering
1 kg m  0.00234 kcal # a given distance is far greater; yet measurements of
Change in work output the efficiency of movement would reveal no change
Efficiency  # or only a relatively small decrement.
Change in VO2
In contrast to the bicycle ergometer, where the
Efficiency
work done is the product of the pedaling speed and
400  200 kg # m # min1
 the resistance, the calculation of work done in walk-
1.08  0.76 liter # min1
ing is more involved. This is because the body walk-
200 kg # m # min1 0.00234 kcal # kg1 # m1
 ing on a level treadmill does no external work. Es-
0.32 liter # min1 5 kcal # liter1 O2
timates of the work done in walking, therefore,
 0.292 or 29.2%
depend on applying an external work load that can
56 BASICS OF METABOLISM

Caloric equivalent of change in external work

Efficiency  (4-10)
Caloric equivalent of change in O2 consumption
External work rate  3Body weight 1kg 2 4 3Speed 1m # min1 2 4 3 % grade/1004 (4-11)
External work rate  3 Body weight 1kg2 4 3Speed 1m # min 1
2 4 3sin ™ 4 (4-12)

be measured, or by estimating the work done inter- by Ralston, Zarrugh, and other mechanical engi-
nally in the body as a result of accelerating and de- neers at the University of California. They attached
celerating the limbs. sensitive transducers to the joints so that their
The most common way to apply external work movements during walking could be recorded;
during walking is to have a subject go up an incline. these recordings, coupled with estimates of the
In Figure 4-16, the vertical external work performed masses of different body parts, made it possible to
is in lifting the body mass the distance B–D. The calculate on a computer the work done in moving
work done is calculated according to either of two the body parts and the entire body. Because the var-
formulas as seen in Equations 4-11 and 4-12. ious techniques of estimating work done in walking
Where sin ∫ is the angle ACB  BD/CB. Re- give similar results, it appears that the efficiencies
cently, external work has been applied in studies of with which the body does internal, horizontal, and
energetics by having subjects walk against a hori- lifting work during walking are similar.
zontal impeding force (Figure 4-17). The work done Although the efficiency of the body during easy
against the horizontal impeding force is calculated cycling and walking may be as high as 30%, it
in Equation 4-13. can only be surmised that the efficiency of running
is somewhat lower. Evidence concerning the effi-
External work  3Speed 1m # min1 2 4 3Weight pulled 1kg2 4 ciency of running is lacking because running is not
(4-13) a true steady-rate situation. During running, the
metabolic rate is so high that both situations a and b
#
An example of how to calculate the efficiency of in Figure 4-13 occur. Because VO2 does not account
performing external work during incline walking is for all the ATP supplied during running, a proper
given in Table 4-5. estimation of efficiency during running awaits de-
Another innovation for estimating the work in- velopment of the technical ability to estimate non-
volved in horizontal walking has been established oxidative ATP supply during exercise.

TABLE 4-5

Estimation of the Whole-Body Efficiency of Doing Vertical Work

During Steady-Rate Treadmill Walking at 3.0 km h1 #
#
Given: (a) Steady-rate caloric equivalent of VO2 during horizontal, ungraded walking (i.e., zero vertical work) at
#
3.0 km h1  5 kcal min1 # #
# #
(b) Steady-rate caloric equivalent of VO2 while performing 375 kg m min1 of vertical work at
#
3.0 km h1  7.9 kcal min1 #
#
(c) 1.0 kg m  0.00234 kcal

Caloric equivalent of change in vertical work

Efficiency 
Caloric equivalent of change in respiration
1375 kg # m # min1  0 kg # m # min1 2 10.00234 kcal # kg 1 # m 1 2

7.9  5 kcal # min 1
 0.30 or 30%
 Summary 57

Figure 4-16 During horizontal treadmill walking, no

external work is done; therefore, it is impossible to cal-
culate a value for body efficiency. However, a way to
determine external work is to measure the work done
in lifting the body up a hill. Refer to Equations 4-11 and
4-12 in the text for details of work rate calculation.

φ
A
D

Figure 4-17 External work can be

determined during horizontal treadmill
walking by having subjects pull a training
weight. Refer to Equation 4-13 in the
Pulley text for work rate calculation.

Wide belt

Weight

Horizontal treadmill

SUMMARY

Metabolism can be estimated in two ways: by direct direct calorimetry must be used. However, during
determinations of heat production and by deter- hard and prolonged exercise, indirect calorimetry
minations of O2 consumption. Determinations of may not provide a precise estimate of metabolic
metabolic rate provide valuable information about rate. Under these conditions, determinations of O2
the status of an individual. In resting individuals, consumption still provide important information
both methods provide similar results. During exer- about the cardioventilatory systems.
cise, direct calorimetry is not feasible; therefore, in-
58 BASICS OF METABOLISM

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Atwater, W. O., and F. G. Benedict. Experiments on the Interscience, 1950.
metabolism of matter and energy in the human body. Kleiber, M. The Fire of Life: An Introduction to Ani-
U.S. Dept. Agr. Off. Exp. Sta. Bull. 136: 1–357, 1903. mal Energetics. New York: Wiley, 1961, pp. 116 –128,
Atwater, W. O., and E. B. Rosa. Description of new 291–311.
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