Chapter 11

Heat and Life

The degree of hotness, or temperature, is one of the most important environmen-

tal factors in the functioning of living organisms. The rates of the metabolic pro-
cesses necessary for life, such as cell divisions and enzyme reactions, depend on
temperature. Generally the rates increase with temperature. A 10 degree change
in temperature may alter the rate by a factor of 2.
Because liquid water is an essential component of living organisms as we
know them, the metabolic processes function only within a relatively narrow
range of temperatures, from about 2°C to 120°C. Only the simplest of living
organisms can function near the extremes of this range.1Large-scale living sys-
tems are restricted to a much narrower range of temperatures.
The functioning of most living systems, plants and animals, is severely lim-
ited by seasonal variations in temperature. The life processes in reptiles, for
example, slow down in cold weather to a point where they essentially cease
to function. On hot sunny days these animals must find shaded shelter to keep
their body temperatures down.
For a given animal, there is usually an optimum rate for the various
metabolic processes. Warm-blooded animals (mammals and birds) have
evolved methods for maintaining their internal body temperatures at near con-
stant levels. As a result, warm-blooded animals are able to function at an opti-
mum level over a wide range of external temperatures. Although this
temperature regulation requires additional expenditures of energy, the adapt-
ability achieved is well worth this expenditure.
In this chapter, we will examine energy consumption, heat flow, and tem-
perature control in animals. Although most of our examples will be specific
to people, the principles are generally applicable to all animals.

1. In deep oceans, the pressure is high and so is the boiling point of water. Here certain thermophilic
bacteria can survive near thermal vents at significantly higher temperatures.

Physics in Biology and Medicine. https://doi.org/10.1016/B978-0-12-813716-1.00011-2

© 2019 Elsevier Inc. All rights reserved. 153
154 Physics in Biology and Medicine

11.1 ENERGY REQUIREMENTS OF PEOPLE

All living systems need energy to function. In animals, this energy is used to
circulate blood, obtain oxygen, repair cells, and so on. As a result, even at com-
plete rest in a comfortable environment, the body requires energy to sustain its
life functions. For example, a man weighing 70 kg lying quietly awake con-
sumes about 70 Cal=hð1 cal ¼ 4:18J;1,000 cal ¼ 1 Cal;1Cal=h ¼ 1:16WÞ. Of
course, the energy expenditure increases with activity.
The amount of energy consumed by a person depends on the person’s weight
and build. It has been found, however, that the amount of energy consumed by a
person during a given activity divided by the surface area of the person’s body is
approximately the same for most people. Therefore, the energy consumed for
various activities is usually quoted in Cal=m2 -hr. This rate is known as the
metabolic rate. The metabolic rates for some human activities are shown in
Table 11.1. To obtain the total energy consumption per hour, we multiply
the metabolic rate by the surface area of the person. The following empirical
formula yields a good estimate for the surface area.
Areaðm2 Þ ¼ 0:202  M0:425  H0:725 (11.1)
Here M is the mass of the person in kilograms, and H is the height of the person
in meters.
The surface area of a 70-kg man of height 1.55m is about 1:70m2 . His met-
abolic rate at rest is therefore ð40Cal=m2 -hrÞ  1:70m2 ¼ 68Cal=hr, or about
70 Cal=hr as stated in our earlier example. This metabolic rate at rest is called
the basal metabolic rate.

TABLE 11.1 Metabolic Rates for Selected Activities

Activity Metabolic rate (Cal/m2-hr)
Sleeping 35
Lying awake 40
Sitting upright 50
Standing 60
Walking (3 mph) 140
Moderate physical work 150

Bicycling 250
Running 600
Shivering 250
 Heat and Life Chapter 11 155

11.1.1 Basal Metabolic Rate and Body Size

Larger animals have more cells requiring more energy to maintain them. There-
fore, we expect the metabolic rate to increase with the size of the animal.
Can this expectation be expressed mathematically? In 1883 the biologist
Max Rubner suggested that the basal metabolism, that is the energy consumed
by the animal at rest, ends up as heat and is therefore likely to be limited by the
amount of heat the animal can eliminate. Using a highly simplified model, he
proposed that the metabolic rate is proportional to M2=3 where M is the mass of
the animal. This expression is based on the assumption that the animal is spher-
ical in shape. Because the mass is proportional to the volume, the radius R of the
assumed sphere is proportional to M1=3 . The area that in this model sets the basal
metabolism, is proportional to R2 or M2=3 .
This simple model was a good starting point but it did not adequately match
subsequent experimental measurements. In 1932 Max Kleiber showed that for a
wide range of species the measured metabolic rate is proportional to M3=4 :This
relationship is obtained from a plot such as shown in Fig. 11.1. Here the met-
abolic rate is plotted versus the body mass for animals ranging in size from
mouse (0.05 kg) to elephant (5,000 kg); a factor of 105 in mass. On a log-log
scale the plot is best fit by a straight line with a slope of 0.75 yielding the

Elephant

1000 Bull
Horse
Boar Cow and steer

100 Man Sow

Woman
Metabolic rate (watt)

Chimpanzee
Sheep
Dog Goat
Goose
Wild birds Condor
10 Macaque
Hen Cat
Rabbit
Guinea pig Marmot

1 Small birds Rat

Pigeon and dove

Mouse
0.1

0.01 0.1 1 10 100 1000 10,000

Body mass (kg)
FIGURE 11.1 Metabolic rates for mammals and birds, plotted as a function of body mass on a
logarithmic scale. Adapted from Benedict (1938).
156 Physics in Biology and Medicine

Kleiber’s “law” that is, metabolic rate is proportional to M3=4 . (See problem
11.1). This relationship has been confirmed by many studies since the 1930’s.
Unlike the more obvious but less applicable M2=3 relationship derived by
Rubner, there is no easily derivable principle leading to Kleiber’s scaling
law. Several models of varying complexities have been proposed that yield
the 3/4 exponent but none of the models are sufficiently compelling to have
been universally accepted. The Kleiber’s scaling law remains subject of ongo-
ing research. The 2005 issue of the Journal of Experimental Biology, volume
208 is devoted to this field of inquiry.
It is universally observed that large animals live longer than small ones. For
example, the lifespan of a mouse is estimated to be 1 to 3 years while that of an
elephant is about 70 years. The power law sheds some light on this observation,
albeit only semi-quantitatively. The issue of lifespan is best examined in
terms of the specific metabolic rate that is the energy burned per unit mass.
This parameter is obtained by dividing the basal metabolic rate by the mass
of the animal. That is, the specific metabolic rate is proportional to M3=4 =M ¼
M1=4 : At this point the assumption is made that the total energy consumption
per unit mass of a creature during its lifetime is a constant. That is, (specific
metabolic rate)  (lifespan) ¼ constant. With this assumption, lifetime is
inversely proportional to the specific metabolic rate (i.e. ¼ M1=4 Þ. The mass
of an elephant being a factor of 105 greater than that of a mouse, its lifespan
is expected to be longer by (105 Þ1=4 ¼ 18. This estimate yields a lifespan for
an elephant of 18 to 54 years; not accurate, but in the right range.

11.2 ENERGY FROM FOOD

The chemical energy used by animals is obtained from the oxidation of food
molecules. The glucose sugar molecule, for example, is oxidized as follows:
C6 H12 O6 + 6O2 ! 6CO2 + 6H2 O + energy (11.2)
For every gram of glucose ingested by the body, 3.81 Cal of energy is released
for metabolic use.
The caloric value per unit weight is different for various foods. Measure-
ments show that, on the average, carbohydrates (sugars and starches) and pro-
teins provide about 4 Cal=g; lipids (fats) produce 9 Cal=g, and the oxidation of
alcohol produces 7 Cal=g.2

2. The high caloric content of alcohol presents a problem for people who drink heavily. The body
utilizes fully the energy released by the oxidation of alcohol. Therefore, people who obtain a sig-
nificant fraction of metabolic energy from this source reduce their intake of conventional foods.
Unlike other foods, however, alcohol does not contain vitamins, minerals, and other substances nec-
essary for proper functioning. As a result, chronic alcoholics often suffer from diseases brought
about by nutritional deficiencies.
 Heat and Life Chapter 11 157

The oxidation of food, which releases energy, does not occur spontaneously
at normal environmental temperatures. For oxidation to proceed at body
temperature, a catalyst must promote the reaction. In living systems, complex
molecules, called enzymes, provide this function.
In the process of obtaining energy from food, oxygen is always consumed. It
has been found that, independent of the type of food being utilized, 4.83 Cal of
energy are produced for every liter of oxygen consumed. Knowing this relation-
ship, one can measure with relatively simple techniques the metabolic rate for
various activities (see Exercise 11-3).
The daily food requirements of a person depend on his or her activities.
A sample schedule and the associated metabolic energy expenditure per square
meter are shown in Table 11.2. Assuming, as before, that the surface area of
the person whose activities are shown in Table 11.2 is 1:7m2 , his/her total
energy expenditure is 3940 Cal=day. If the person spent half the day sleeping
and half the day resting in bed, the daily energy expenditure would be only
1530 Cal.
For most people the energy expenditure is balanced by the food intake. For
example, the daily energy needs of the person whose activities are shown in
Table 11.2 (surface area 1:7m2 ) are met by the consumption of 400 g of carbo-
hydrates, 200 g of protein, and 171 g of fat.
The composition and energy content of some common foods are shown in
Table 11.3. Note that the sum of the weights of the protein, carbohydrates, and
fat is smaller than the total weight of the food. The difference is due mostly to
the water content of the food. The energy values quoted in the table reflect the
fact that the caloric content of different proteins, carbohydrates, and fats deviate
somewhat from the average values stated in the text.
If an excess of certain substances, such as water and salt, is ingested, the
body is able to eliminate it. The body has no mechanism, however, for elimi-
nating an excess in caloric intake. Over a period of time the excess energy is

TABLE 11.2 One Day’s Metabolic Energy Expenditure

Activity Energy expenditure (Cal/m2)
8 hr sleeping (35 Cal/m2-hr) 280
2
8 hr moderate physical labor (150 Cal/m -hr) 1200
2
4 hr reading, writing, TV watching (60 Cal/m -hr) 240
2
1 hr heavy exercise (300 Cal/m -hr) 300
2
3 hr dressing, eating (100 Cal/m -hr) 300
Total expenditure 2320
158 Physics in Biology and Medicine

TABLE 11.3 Composition and Energy Content of Some Common Foods

Total
Total Protein Carbohydrate Fat energy
Food weight (g) weight (g) weight (g) weight (g) (Cal)
Whole milk, 976 32 48 40 660
1 quart
Egg, 1 50 6 0 12 75
Hamburger, 1 85 21 0 17 245
Carrots, 150 1 10 0 45
1 cup
Potato 100 2 22 0 100
(1 med.,
baked)

Apple 130 0 18 0 70
Bread, rye,1 23 2 12 0 55
slice

Doughnut 33 2 17 7 135

used by the body to manufacture additional tissue. If the consumption of excess

food occurs simultaneously with heavy exercise, the energy may be utilized to
increase the weight of the muscles. Most often, however, the excess energy is
stored in fatty tissue that is manufactured by the body. Conversely, if the energy
intake is lower than the demand, the body consumes its own tissue to make up
the deficit. While the supply lasts, the body first utilizes its stored fat. For every
9 Cal of energy deficit, about 1 g of fat is used. Under severe starvation, once the
fat is used up, he body begins to consume its own protein. Each gram of con-
sumed protein yields about 4 Cal. Consumption of body protein results in the
deterioration of body functions, of course. A relatively simple calculation
(see Exercise 11-6) shows that an average healthy person can survive without
food but with adequate water up to about 50 days. Overweight people can do
better, of course. The “Guinness Book of World Records” states that Angus
Barbieri of Scotland fasted from June, 1965, to July, 1966, consuming only
tea, coffee, and water. During this period, his weight declined from 472 lb to
178 lb.
For a woman, the energy requirements increase somewhat during pregnancy
due to the growth and metabolism of the fetus. As the following calculation
indicates, the energy needed for the growth of the fetus is actually rather small.
Let us assume that the weight gain of the fetus during the 270 days of gestation
 Heat and Life Chapter 11 159

is uniform.3 If at birth the fetus weighs 3 kg, each day it gains 11 g. Because 75%
of tissue consists of water and inorganic minerals, only 2.75 g of the daily mass
increase is due to organic materials, mainly protein. Therefore, the extra calo-
ries per day required for the growth of the fetus is
2:75g protein 4 Cal
Calories required ¼  ¼ 11 Cal=day
day g protein
To this number, we must add the basal metabolic consumption of the fetus. At
birth, the surface area of the fetus is about 0:13m2 (from Eq. 11.1); therefore, at
most, the basal metabolic consumption of the fetus per day is about
0:13  40  24 ¼ 125Cal. Thus, the total increase in the energy requirement
of a pregnant woman is only about ð125 + 11Þ Cal=day ¼ 136 Cal=day. Actu-
ally, it may not even be necessary for a pregnant woman to increase her food
intake, as the energy requirements of the fetus may be balanced by decreased
physical activity during pregnancy. Various other aspects of metabolic energy
balance are examined in Exercises 11-4 to 11-7.

11.3 REGULATION OF BODY TEMPERATURE

People and other warm-blooded animals must maintain their body temperatures
at a nearly constant level. For example, the normal internal body temperature of
a person is about 37°C. A deviation of one or two degrees in either direction may
signal some abnormality. If the temperature-regulating mechanisms fail and the
body temperature rises to 44° or 45°C, the protein structures are irreversibly
damaged. A fall in body temperature below about 28°C results in heart
stoppage.
The body temperature is sensed by specialized nerve centers in the brain and
by receptors on the surface of the body. The various cooling or heating mech-
anisms of the body are then activated in accord with the temperature. The effi-
ciency of muscles in performing external work is at best 20%. Therefore, at least
80% of the energy consumed in the performance of a physical activity is con-
verted into heat inside the body. In addition, the energy consumed to maintain
the basic metabolic processes is ultimately all converted to heat. If this heat
were not eliminated, the body temperature would quickly rise to a dangerous
level. For example, during moderate physical activity, a 70-kg man may con-
sume 260 Cal=hr. Of this amount, at least 208 Cal is converted to heat. If this
heat remained within the body, the body temperature would rise by 3 C°/hr. Two
hours of such an activity would cause complete collapse. Fortunately, the body
possesses a number of highly efficient methods for controlling the heat flow out
of the body, thereby maintaining a stable internal temperature.

3. This is a simplification because the weight gain is not uniform. It is greatest toward the end of
gestation.