International Textbook of Obesity. Edited by Per Bjorntorp.

Copyright © 2001 John Wiley & Sons Ltd

Print ISBNs: 0-471-988707 (Hardback); 0-470-846739 (Electronic)

11

Energy Expenditure at Rest and

During Exercise
Björn Ekblom
Karolinska Institute, Stockholm, Sweden

ENERGY METABOLISM phate (CrP). CrP is stored in the muscle for immedi-
ate use, but it can also be regarded as an energy
Alactacid Energy transport system between the mitochondria and the
Metabolism—Adenosine Triphosphate myofibrillar system as well as an ‘energy buffer’ for
phosphorylation of ATP, when the capacity and
The immediate source for muscle contraction is speed of the other energy regeneration systems can-
delivered from splitting of adenosine triphosphate not keep up with an acceptable ATP concentration
(ATP) to adenosine diphosphate (ADP) and a ‘free’ in the myofibrillar system. Phosphorylation of
phosphate ion (P). This reaction is very fast and ADP from CrP (CrP ; ADP to Cr ; ATP) is fast.
does not normally limit energy turnover and muscle The total CrP pool can be used up by several sec-
performance. However, the ATP stores in the onds of heavy exercise.
muscles are very limited. The whole ATP pool At rest and during light submaximal exercise the
would be emptied in only a few seconds of muscle CrP concentration is not different from resting con-
contraction. Therefore ATP has to be continuously centration due to continuous rephosphorylation of
regenerated through other energy systems. Cr in the mitochondria. During normal dynamic
These supporting energy systems are very effec- heavy exercise the CrP pool may be lowered to 50%
tive and can keep the ATP concentration un- or less of resting and completely emptied during
changed or only marginally lowered, even during extreme exercise conditions.
heavy exercise. In essence, there are four such sys- The amount of CrP in the muscle as well as ATP
tems, with different speeds of reaction and capaci- and ADP concentrations can only be measured by
ties, which release energy for active phosphoryla- sophisticated laboratory methods.
tion of ADP to restore the ATP pool. They are (1)
creative phosphate, (2) lactacid anaerobic metab-
olism, (3) aerobic metabolism and (4) adenylate Lactacid Anaerobic Metabolism
kinase reactions.
In this energy pathway the chemically bound en-
ergy in carbohydrates, mainly muscle glycogen, can
Creatine Phosphate be utilized for ATP regeneration during stepwise
degradation of glycogen or glucose to lactate (Hla)
The most immediate energy system to restore ATP and hydrogen ions. No oxygen is used. This reac-
from rephosphorylation of ADP is creatine phos- tion is fairly fast but normally limited by the

International Textbook of Obesity. Edited by Per Björntorp.

© 2001 John Wiley & Sons, Ltd.
146 INTERNATIONAL TEXTBOOK OF OBESITY
Fat Carbohydrates

RQ % %

1.00 0 100

0.90

50 50

0.80

0.70 100 0

20 40 60 80 100
% of maximum oxygen uptake
Figure 11.1 Metabolism of fat and carbohydrate at rest and during submaximal exercise, and during heavy exercise

formation of hydrogen ions which decrease the body but also in the muscle, which is very import-
muscle pH and impair muscle performance in sev- ant. Glucose is stored as glycogen in the liver and
eral ways. muscles. Small but important amounts of glucose
If muscle glycogen stores are more or less empty are also found in the plasma.
this anaerobic energy system is impaired due to the During aerobic conditions—when oxygen is
reduced substrate availability. For each molecule of available at the site of the mitochondria—fatty
glucose or glycogen two and three ATP, respective- acids and glucose in combination are metabolized
ly, are formed. This energy system is activated only to carbon dioxide and water. For each molecule of
during heavy exercise. Measurements of lactate glucose 38 or 39 ATP can be formed depending on
concentration in blood are only able to indicate a whether glucose or glycogen is the substrate. For
qualitative involvement of this anaerobic energy each fatty acid about 130 ATP can be formed. The
system during physical stress. It is not possible to latter figure varies depending on the type of fatty
make quantitative calculations of the lactacid acid that is metabolized.
anaerobic energy yield during exercise from Since oxygen is a prerequisite for this reaction it
measurements of lactate concentrations in the is possible to calculate how much energy has been
blood due to dilution and transport of lactate in converted at rest and during exercise by measuring
different water compartments in the body, elimin- oxygen consumption. This is done at the level of
ation of lactate in the metabolism and other factors. pulmonary ventilation—see below. When fatty
acids are used as substrate 19.3 kJ (4.7 kcal) is trans-
formed for each litre of oxygen used. Corresponding
value for glucose is 21.0 kJ (5.0 kcal) and for protein
Aerobic Metabolism 18.8 kJ (4.5 kcal). However, for most calculations of
energy expenditure at rest and during exercise a
Quantitatively the most important energy system figure of 21 kJ (5 kcal) for each litre of oxygen can be
during exercise is the breakdown and splitting of used.
the energy-rich fat and carbohydrate molecules. Fat Respiratory quotient (RQ) is the volume of car-
is stored in large amounts in fat cells all over the bon dioxide formed divided by the volume of oxy-
 ENERGY EXPENDITURE AT REST AND DURING EXERCISE 147
gen used—for measurements see below. Since fat Summary
oxidation has a RQ of 0.7 and glucose 1.0 it is
possible using RQ determinations to evaluate the The energy metabolism for generating ATP to the
relative contribution of fat and carbohydrate, re- muscle contraction is complex and not fully under-
spectively, in energy metabolism. At rest and during stood. It includes reactions which can deliver en-
submaximal exercise RQ is normally about 0.80 to ergy very fast—such as the adenylate kinase reac-
0.85 and reaches 1.0 during heavy exercise. Thus, at tion, creatine phosphate and glucose/glycogen
rest and during submaximal exercise, fat and carbo- spliting. The negative aspects of these reactions are
hydrate are combusted to about an equal extent the limited stores and negative effects of the reac-
while during heavy exercise, when RQ is about 1.0, tions. Aerobic breakdown of fat and carbohydrates
only carbohydrates are metabolized (Figure 11.1). It provides a more patient and durable energy meta-
should be emphasized that carbohydrate- or fat- bolism. The aerobic energy systems are fairly slow
rich diets alter the RQ at rest and during submaxi- but stores are large. The metabolites of these reac-
mal exercise. tions have hardly any negative effects. In the dis-
The aerobic reaction of fat and carbohydrate cussion of energy balance and weight maintenance
metabolism is slower than other energy systems. On it is only the aerobic metabolism that is of interest.
the other hand, the stores are very large for fat and
intermediate for glycogen. The stores of glycogen
both in the liver and in the muscles can be increased
by carbohydrate-rich diets. Supplementation with MEASUREMENTS OF ENERGY
solutions containing carbohydrates but not fatty METABOLISM
acids increases physical and mental endurance dur-
ing prolonged exercise. Oxygen Consumption
The limitation of the aerobic energy system is the
maximal availability of oxygen at the site of the Oxygen in the ambient air is transported by the
mitochondria, delivered through the oxygen trans- pulmonary ventilation and in the main circulation
port system—see below. Endurance, defined as the to the muscle capillaries, from which through diffu-
capacity to carry out prolonged submaximal exer- sion it reaches the muscle mitochondria. To quan-
cise, is to a large extent limited by the glycogen tify the amount of oxygen involved in metabolism
stores in the muscles and liver. in the mitochondria, the oxygen uptake is measured
at the site of the pulmonary ventilation using the
Douglas bag method or automatic analysis systems.
Adenylate Kinase Reaction The volume of expired air and the oxygen and
carbon dioxide concentration in the expired air are
This special energy pathway is not very well inves- measured. Since the oxygen and carbon dioxide in
tigated but is believed to be used only during ex- the inspired air normally is 20.94 and 0.03%, re-
treme physical stress conditions. In this reaction spectively, it is easy to calculate the amount of
two ADP react to form one mole of ATP and one oxygen that has been taken up and carbon dioxide
mole of AMP (aminomonophosphate) in an at- that has been produced in litres per minute, both at
tempt to produce ATP very quickly and to reduce rest and during exercise (1). The error in measuring
the amount of ADP in the muscle. To keep this oxygen uptake during submaximal and maximal
reaction going forward AMP has to be degraded by exercise with these methods is now less than 2%.
deamination. In several subsequent degrading reac- At rest oxygen uptake in a normal trained or
tion steps uric acid will be formed and can be used untrained young man with body mass 70 to 75 kg is
as a marker for the net loss of ATP. But more about 0.25 litres per minute. Corresponding value
importantly, during this last reaction step oxygen for a young woman of the same age is somewhat less
free radicals can be formed, which may negatively due to smaller body mass. With increasing age rest-
influence cell membranes, several biochemical and ing oxygen uptake decreases mainly due to decreas-
other functions and structures in the muscle. Under ing muscle mass. During exercise oxygen uptake
normal conditions this reaction is of little value in normally increases linearly with increasing rate of
the total energy output. work up to maximal exercise.
148 INTERNATIONAL TEXTBOOK OF OBESITY
Table 11.1 Maximal aerobic power and energy expenditure this is that there is roughly a linear relationship
during 1 hour of exercise between oxygen uptake and heart rate for most
V  types of physical work under normal conditions (1).
  However, it must be emphsized that the heart
(litres per minute) kJ per hour
rate for a given absolute and even relative (per cent
Untrained women of maximum) oxygen uptake can vary extensively,
25 years 2.3 1400
50 years 1.9 1200 for example with age, different peak heart rates,
75 years 1.4 900 training status, diseases, psychological status and
stress, medication (beta-blockers) and many other
Untrained men
25 years 3.3 2100 factors. Therefore, each estimation of energy expen-
50 years 2.7 1700 diture from heart rate recordings should be done
75 years 2.0 1300 individually, taking all these variations into con-
Endurance athletes
sideration.
women 4.0—4.5 4200—4800 The estimation of energy expenditure from heart
men 5.0—7.4 5400—7800 rate recordings is done first by establishing the rela-
tionship between heart rate and oxygen uptake dur-
ing increasing rates of work on a cycle ergometer,
The maximal aerobic power is defined as the treadmill or the type of exercise that the subject is
peak oxygen uptake during dynamic exercise with performing. The energy expenditure can thereafter
large muscle groups under normal atmospheric be estimated by interpolation from heart rate re-
conditions. In order to ensure that the maximal cordings during the actual activity.
oxygen uptake has been reached the linear relation If all these measurement are done properly, the
between oxygen uptake and submaximal work load error of the method for estimation of energy expen-
should ‘level off’ at maximal oxygen uptake. High diture from heart rate recordings during the actual
values for blood lactate concentration and heart work is in the range of < 15%. However, this
rate can only be used as indications that maximal method is less accurate than the direct measure-
oxygen uptake have been reached. Maximal aerobic ment of oxygen uptake due to the temporary vari-
power in most healthy mean and women is limited ation in heart rate caused by static work, psycho-
by the capacity of the heart to pump blood during logical stress etc. Therefore, estimations of energy
maximal exercise (maximal cardiac output). How- expenditure from heart rate recordings must be
ever, maximal oxygen uptake can be modified by done with great caution.
many factors such as lowered oxygen carrying ca-
pacity in the blood (anaemia), medication and other
factors. Values of maximal oxygen uptake in trained
and untrained men and women are given in Table Core Temperature
11.1.
Direct measurements of oxygen uptake can only Determination of core temperature during or im-
be done with specialized laboratory or field test mediately after exercise can also be used for estima-
equipment. Furthermore, in some activities such as tion of energy expenditure during dynamic exercise.
prolonged work or in many work situtions direct The background is that there is a close relationship
measurements of oxygen uptake are more or less between core temperature and the relative oxygen
impossible. Therefore, energy expenditure must uptake (1). Thus, if the exercise has persisted for
usually be evaluated by other methods. longer than 15 to 20 minutes and has been per-
formed under normal conditions (e.g. within the air
temperature range of approximately from 5 to
35°C), a core temperature of 38.0°C indicates a
Heart Rate relative energy expenditure of about 50% of maxi-
mal aerobic power. At an average energy expendi-
Measurement of heart rate during physical activity ture of 75% of maximal aerobic power the core
is one possible way to estimate oxygen consump- temperature is approximately 38.8°C. These figures
tion and energy expenditure. The background for are consistent for men and women, irrespective of
 ENERGY EXPENDITURE AT REST AND DURING EXERCISE 149
whether the individual is untrained or well trained one possibility to estimate total energy turnover
or has a maximal aerobic power of 3 or 6 litres per over a prolonged period (weeks). However, under-
minute. and overreporting is very common, especially in
This method has its limitations, such as the iner- obese subjects (3). Furthermore, there are normal
tia of the core temperature with time and changes in variations in body weight of < 1—2 kg even over
energy expenditure. Furthermore, core temperature fairly short periods of time. One kilo body mass
rises for a given energy expenditure during hy- change due to for instance body water shifts can
pohydrated conditions, with extreme adiposity and indicate a fat mass change of about 27 MJ
some other conditions. Nevertheless, this method (6500 kcal), which equals more than 2 days’ normal
may be very useful in some situations, such as dur- free-living energy expenditure in most individuals.
ing intermittent exercise, in which the rate of work Therefore, one must be cautious about making as-
changes rapidly and also during physical exercise sumptions based on estimations of energy expendi-
with high levels of psychological stress. In this latter ture from individually reported dietary intakes.
situation the heart rate is increased due to the effect
of catecholamines and, thus, the normal relation
between heart rate and oxygen uptake is changed
Summary
and not valid. In this and some other situations
measurements of core temperature may be the best
Energy expenditure is best estimated by measuring
method to estimate the relative energy expenditure
oxygen consumption, since direct calorimetry is not
during exercise. In addition, maximal aerobic
a practical method. For calculation of energy ex-
power must be measured or estimated.
penditure for a fairly short period of time aerobic
power times duration can simply be used. However,
this procedure is not useful and possible in many
Double Labelled Water
practical situations outside laboratory settings.
Therefore, calculations of energy expenditure from
The doubly labelled water method is one of the best
indirect estimations of oxygen consumption by
and in many situations the only possible method for
heart rate recordings and core temperature
estimation of energy expenditure over long periods
measurements during and after physical activity,
(2). The method makes it possible to measure the
respectively, are well-accepted methods. For calcu-
total energy expenditure during periods up to 2
lations of energy expenditure for longer periods of
weeks under free-living conditions with a minimum
time (days and weeks) only the diary intake and
of inconvenience for the individual. At the start the
doubly labelled water methods are valid and poss-
individual drinks water containing two isotopes
ible. In all these methods there are many different
(H and O). The two isotopes will then be diluted
 possibilities for erroneous recordings and calcula-
in the total body water pool. Both leave the body as
tions. Therefore these methods must be used with
water but in addition the oxygen also disappears as
caution.
carbon dioxide as a result of the energy metabolism.
By measuring the concentration of H and O in

urine at the start and after some time the total
energy expenditure during the period can be es- ENERGY EXPENDITURE AT REST
timated. The only drawback of this method is the
high cost of the isotopes and analysis, so that it is In general medicine and medical practice the inter-
only feasible for studies with a small number of est in energy metabolism is often focused on basal
subjects. metabolism. This is easy to understand because
variations in basal metabolic rate (BMR) can be in
the range of 30—40%. This variation can account for
Dietary Intake large increases and decreases in body weight, es-
pecially if they persist for a long period of time. The
If total body mass and its composition is unchanged reason for the inter- but also intra-individual vari-
over time, then energy expenditure must equal en- ations in BMR can only partly be explained by
ergy intake. Thus, measuring dietary intake may be variations in active body mass—mainly muscle
150 INTERNATIONAL TEXTBOOK OF OBESITY
mass. Therefore, a mitochondria protein—the un- Summary
coupling protein (UCP), found in the mitochondria
in the brown adipose tissue—is of great interest in About two-thirds of the energy expenditure over 24
this respect. hours amounts to the resting energy metabolism.
Brown adipose tissues have many mitochondria. New findings regarding the uncoupling protein can
The energy released in the brown fat cells is to a shed new light on BMR and might to some extent
lesser degree than in other cells used for active explain the variations in BMR between individuals
phosphorylation of ADP to ATP and more for and perhaps also changes in BMR with time and
thermogenesis. Recently, proteins which have struc- ageing.
tures very like the UCP ones in brown adipose
tissue have also been found in muscle tissue. Al-
though there are many questions to be answered ENERGY EXPENDITURE DURING
regarding the presence of the UCP-like protein in EXERCISE
the muscle (exact function, regulation etc.), it can be
speculated that this protein might explain why only Intensity and Duration
about half of the oxygen used in metabolism in the
muscles is used for active phosphorylation of ADP One cannot apply strict mathematical principles to
at rest (4). The consequence could be that some part biological systems, but when analysing energy bal-
of the energy taken in is not stored in the body, if the ance for longer periods of time, energy metabolism
energy released in the metabolism is not used for during and after exercise must be taken into ac-
mechanical events in the muscle but only increases count. It is obvious that both the intensity and the
the thermogenesis. Of interest in this discussion is duration are the main determinants of energy ex-
that it has been shown that there are differences penditure during exercise. However, many factors
between overweight and normal-weight individuals may modify the energy expenditure for a given rate
in how this UCP-like protein is expressed in mRNA of work and the total cost for certain activities. For
(5). this reason it is difficult to give exact figures for the
Studies in rats have shown that regular endur- energy cost of exercise. Therefore the discussion of
ance training decreases the mRNA linked to the energy expenditure should be based on individual
UCP in muscles (6). On the other hand, after an conditions and values given for certain activities or
endurance exercise session the activity of UCP is for groups of subjects are subject to large uncertain-
increased (7), which might explain part of the in- ties.
creased post-exercise oxygen consumption. Regular During short-term (a few minutes) hard dynamic
physical training increases muscle and mitochon- muscular exercise carried out with large muscle
drial mass and as a consequence presumably also groups, the energy metabolism may increase to
the amount of UCP. Thus, both acute and chronic 10—15 times the BMR in untrained subjects and
exercise is of importance for the BMR and conse- 25—30 times the BMR in well-trained athletes from
quently the energy balance in both normal-weight endurance events. However, due to muscle fatigue
and overweight individuals. during heavy exercise the duration of exercise is
If UCP is downregulated by physical activity often fairly short. In such cases the total energy
then its activity should increase with physical inac- expenditure is relatively low. On the other hand,
tivity, leading to an increased BMR per kilo lean low-intensity exercise, which may require half or
body mass. On the other hand, muscle mass is two-thirds of the individual’s maximal aerobic
reduced as a result of physical inactivity. In any power, can be performed for a very long time even
case, when studying changes in body weight, diet by an untrained individual. In this case total energy
and eating habits and level of physical exercise in turnover can be fairly high.
individuals, in groups and also in population inves-
tigations, it is obvious that the energy turnover both
during and after exercise as well as the influence of Variations in Energy Expenditure
exercise on BMR must be considered. Thus, level of During Submaximal Exercise
physical exercise is therefore of vital importance in
the discussion of energy balance in humans. Variations in energy expenditure for a given sub-
 ENERGY EXPENDITURE AT REST AND DURING EXERCISE 151
18 than walking in both these aspects. The upper panel
of the figure also shows that the net energy cost for
running per kilometre is more or less independent
of speed. For a normal man with a body mass of 70
14
Net VO2 (L/km)

to 75 kg the energy expenditure during running is

about 280 to 300 kJ per kilometre independent of
speed, while walking for the same man may cost
.

10
between 150 and 350 kJ per kilometre depending on
speed. It must be emphasized that well-trained male
and female racewalkers and long-distance runners
have much lower values for energy expenditure
6 both per minute and net per kilometre than normal,
untrained individuals.
Women and children have lower energy cost for a
given speed in walking and running due to their
4.0 lower body mass. However, energy expenditure cal-
culated per kilo body mass is the same for men and
women whereas children have higher values. The
3.0
energy expenditure also increases with body weight.
Overweight individuals can have 50% and higher
VO2 (L/min)

energy expenditure for a given walking speed. For

example, during uphill treadmill walking (4—5 km
2.0 per hour, 4° elevation) the oxygen uptake in an
.

untrained overweight woman with a BMI of 35—40

may be maximal. Thus, for a given low walking
speed the variation in energy expenditure can be up
1.0 to 100% in a normal population.
The energy expenditure at a given speed varies
also with different conditions such as surface, uphill
and downhill walking and running, wind resistance
2 6 10 14 etc. People with joint disease, an amputation or
other physical handicaps have decreased locomo-
Speed km/h tion economy, that is the oxygen uptake for a given
Figure 11.2 Energy expenditure (as measured by oxygen up- submaximal rate of work is increased.
take) during walking and running In some types of exercise in which technique is
very important, such as swimming, the energy ex-
penditure at a given speed may vary by more than
maximal rate of work are due both to individual 100% for poor and good swimmers for the same
variations in economy of locomotion, such as differ- swimming stroke but also for different swimming
ent technique and body mass, and to temporary strokes in the same individual. On the other hand,
interindividual factors, such as changes in core tem- the energy expenditure for submaximal cycling is
perature and choice of substrate. about the same for well-trained cyclists and as it is
Energy expenditure (as evaluated from oxygen for runners for instance.
consumption) during walking and running is illus- In high speed activities in which wind resistance
trated in Figure 11.2. At low speeds—2—5 km per increases, the energy expenditure increases cur-
hour—walking costs less than running; that is oxy- vilinearly. In addition, the style, position and/or
gen uptake during walking is less than in running at equipment can influence the energy expenditure for
the same speed. This is true for both energy expen- a given speed. This is particularly true in cycling but
diture per minute of exercise and net cost of energy also for running. For example, running behind an-
per kilometre covered. However, at speeds greater other runner may save up to 6% in energy cost
than 6 to 8 km per hour running is more effective because of the wind protection.
152 INTERNATIONAL TEXTBOOK OF OBESITY
There are situations in which the energy expendi- Table 11.2 Average energy cost for different activities for a
ture for a given submaximal rate of work is in- 20- to 30-year-old man
creased such as in hypothermia due to shivering, in kJ per minute
very cold climates due to resistance of cold, stiff
clothes and when for instance running technique is Complete rest 4—7
impaired for various reasons. However, in most Sitting 6—8
Standing 7—9
such situations the magnitude of the increased en- Standing, light activity 9—13
ergy expenditure for a given rate of work is of little Light housework 13—30
quantitative importance. On the other hand, in Gardening activities 15—45
many situations the energy expenditure for a given Walking
rate of work does not change. There are no major 3 km per hour 15—30
changes in energy expenditure for a given rate of 5 km per hour 20—40
work with variations in hot or moderately cold 7 km per hour 30—60
climate (except for shivering), in moderate altitude Running
compared to sea-level, in anaemia and most dis- 7 km per hour 30—50
eases including most types of medication, although 9 km per hour 40—70
11 km per hour 50—90
in these conditions the physical performance can be
severely impaired. It should also be emphasized
that although the energy expenditure at submaxi-
mal work is not changed, the total energy expendi- mal aerobic power of about 3.3 litres per minute,
ture may be reduced due to the individual becoming 0.50 g of fat is used per minute at this intensity.
fatigued earlier. Suppose that this individual through physical train-
The average energy expenditures for different ac- ing increases his/her maximal aerobic power by 0.5
tivities performed for more than 10—15 minutes by a litres per minute, which is possible in 4 to 5 months
man aged 20—30 years are given in Table 11.2. It of endurance training. Compared to the situation
must be emphasized that these values are subject to before the training period, two observations can be
large interindividual variations, as discussed above. mentioned regarding the fat and carbohydrate con-
tribution to the energy expenditure. Firstly, for a
given submaximal relative but also absolute rate of
work the RQ is lowered (lower part of the shadowed
Substrate Use During Exercise and area in Figure 11.1). Thus, more fatty acids are used
Physical Training and the stores of carbohydrate are utilized less.
Secondly, the intensity for peak fatty acid contribu-
As stated above, fatty acids and carbohydrates in tion to the energy expenditure has increased from
combination are used during submaximal exercise. 60% to about 70% of maximal aerobic power. This
A common question in this discussion of substrate means that the peak contribution of fatty acids in
utilization is: Which is the best way to burn fat this individual has increased due to the training
during exercise? effects from 0.50 to 0.75 g per minute. In addition,
From Figure 11.1 it can be seen that the RQ for the individual can probably be active for longer
an untrained person (upper part of the shadowed periods of time after the training period and, thus,
area) is about 0.85 to 0.88 at exercise intensities increase the fatty acid turnover still more. For in-
from about 25 to 60% of maximal aerobic power. stance, if she/he increases the exercise time from 30
This means that the fat and carbohydrate contribu- minutes before to 45 minutes after the training per-
tion to the energy expenditure is 45 and 55%, re- iod at the exercise intensity at which she/he can
spectively. From these data the substrate use during exercise fairly easily, then the fatty acid breakdown
exercise can be calculated. increases from 15 g to 30 g for the exercise period.
The total fatty acid contribution to the exercise The increased use of fatty acids at a given rate of
expenditure is highest at around 60% of maximal work and the higher speed of exercise may be of
aerobic power, which is a pace that even an un- interest not only in conditioning exercise such as
trained person can exercise at for some time. This jogging and cycling but also in the everyday ‘behav-
means that for an untrained individual with a maxi- iour’ type of exercise (climbing stairs, walking short
 ENERGY EXPENDITURE AT REST AND DURING EXERCISE 153
distances etc.) as part of the energy expenditure in uncommon that body fat mass in many individuals
the discussion of energy balance. increases 2 kg in one year. This corresponds to a
daily energy imbalance of about 150 kJ. Unless net
energy intake is increased this corresponds to an
extra 10 minutes of walking per day. Furthermore,
Maximal Exercise
in order to maximize the beneficial effects of physi-
cal activity on health, and in prevention of diseases
Variations in maximal power are due to age, genetic
that are related to physical inactivity, the Surgeon
endowment, body size, physical activity and some
General in the USA has recommended accumulated
other factors and can partly explain differences in
low-intensity physical activity of at least 30 minutes
total energy expenditure for different reasons. Indi-
per day (8). Thus, regular low-intensity physical
viduals with high maximal aerobic power will more
activity such as walking and cycling to work two
likely walk distances or climb stairs than use cars
times 15—20 minutes a day may be a good base for
and elevators. They can more easily carry loads and
energy balance, body weight maintenance and good
they may in general be more physically active in
health.
normal life. In addition, due to increased energy
Sporting activities can generate quite a large total
intake when physically active they also have in-
energy expenditure. In male elite soccer matchplay
creased intake of essential nutrients. But the total
the heart rate is on average some 25 to 30 beats per
daily need and turnover for essential nutrients in-
minute lower than peak heart rate obtained during
creases less than the increased total daily energy
maximal exercise. Core temperature after the game
need and turnover when a person becomes more
is above 39°C as an average for the players in the
physically active. Therefore the difference between
team. Blood lactate concentration measured several
intake and turnover of essential nutrients widens
times during the match varies between 4 and
with increasing levels of physical activity under the
10 mM. Thus, from these figures it can be calculated
assumption that the individual is in energy balance
that the average energy expenditure during the
while trained and untrained.
game amounts to 75 to 80% of maximal aerobic
power. For an average male elite player with a
maximal oxygen uptake of 4.5 litres per minute the
Total Energy Expenditure total energy expenditure for a whole game including
some warm-up can be calculated to be about 7.5 MJ
As stated above, duration of exercise may be more (1800 kcal) which is about the same as the BMR for
important than intensity for total energy expendi- 24 hours. Corresponding values for total energy
ture. In Table 11.1 the total energy expenditure is expenditure for a female elite player are some 20%
given for one hour of exercise such as walking in less (9).
uneven terrain, cycling or playing a game of tennis, The energy cost of a marathon race (42 km) for a
volleyball or table tennis in a moderate fashion. The 30- to 40-year-old man who performs the race in 4
intensity of these types of physical activities is on hours is about 12—15 MJ (3000—3500 kcal). How-
average about 50 to 60% of maximal aerobic power ever, in order to be able to carry out the race in 4
when carried out as free-chosen physical activity. hours the training during the preceding 6 months
The rate of work of 50 to 60% is easily performed can be calculated to be about 400 MJ. It is obvious
even by an untrained person for one hour. The that regular physical training for sport is of import-
individual maximal oxygen uptake values for un- ance for energy balance and body weight control.
trained men and women at different ages and en-
durance athletes are also given in Table 11.1.
The table shows that one hour of leisure time
exercise yields an energy expenditure in an un- Summary
trained person which corresponds to about one-
quarter of 24 hour BMR, which is 7 MJ for men and Energy for physical activity is generated though
5—6 MJ for women. The importance of these types several complicated systems of which the aerobic
of regular physical exercise is illustrated when dis- splitting of fat and glucose is the most important
cussing body mass changes over time. It is not one. For most people physical activity amounts to
154 INTERNATIONAL TEXTBOOK OF OBESITY
about 30—40% of the total energy expenditure lar basis, create a daily energy expenditure well
during 24 hours. The amount of exercise energy above 3000 kJ and, thus, well above the level for
expenditure during 24 hours is dependent on inten- good health and body mass maintenance. Elite ath-
sity and duration but many other factors can influ- letes often have a daily energy expenditure of
ence energy expenditure. 14 000—16 000 kJ (3500—4000 kcal); in some sports it
In the population physical activity can be divided may be even higher. In addition to energy expendi-
into four main parts. The difference between them is ture during exercise, the effect of regular physical
often not very clear. The lowest one is spontaneous activity on resting metabolic rate is of interest.
activity, which is trivial activities such as moving Thus physical activity is very important for body
arms and legs, take small steps etc. The energy mass maintenance. All its different parts must be
needed for this type of activity is fairly small but for considered when discussing energy balance.
people who seldom sit still or move regularly the
whole day the total amount can reach some volume.
The physical stress in most jobs is nowadays
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