10.

4 Thermodynamics of Living Systems

It is obvious that animals need food to live, but the reason for this is less
obvious. The idea that animals need energy because they consume energy is,
Section 10.4 Thermodynamics of Living Systems 141
strictly speaking, incorrect. We know from the First Law of Thermodynamics
that energy is conserved. The body does not consume energy, it changes it
from one form to another. In fact, the first law could lead us to the erroneous
conclusion that animals should be able to function without a source of external
energy. The body takes in energy that is in the chemical bonds of the food
molecules and converts it to heat. If the weight and the temperature of the
body remain constant and if the body performs no external work, the energy
input to the body equals exactly the heat energy leaving the body. We may
suppose that if the heat outflow could be stoppedby good insulation, for
examplethe body could survive without food. As we know, this supposition
is wrong. The need for energy is made apparent by examining the functioning
of the body in the light of the Second Law of Thermodynamics.
The body is a highly ordered system. A single protein molecule in the
body may consist of a million atoms bound together in an ordered sequence.
Cells are more complex still. Their specialized functions within the body
depend on a specific structure and location. We know from the Second Law
of Thermodynamics that such a highly ordered system, left to itself, tends
to become disordered, and once it is disordered, it ceases to function. Work
must be done on the system continuously to prevent it from falling apart. For
example, the blood circulating in veins and arteries is subject to friction, which
changes kinetic energy to heat and slows the flow of blood. If a force were not
applied to the blood, it would stop flowing in a few seconds. The concentration
of minerals inside a cell differs from that in the surrounding environment.
This represents an ordered arrangement. The natural tendency is toward an
equalization with the environment. Work must be done to prevent the contents
of the cell from leaking out. Finally, cells that die must be replaced, and if the
animal is growing, new tissue must be manufactured. For such replacement
and growth, new proteins and other cell constituents must be put together
from smaller, relatively more random subcomponents. Thus, the process of
life consists of building and maintaining ordered structures. In the face of the
natural tendency toward disorder, this activity requires work. The situation is
somewhat analogous to a pillar made of small, slippery, uneven blocks that
tend to slide out of the structure. The pillar remains standing only if blocks
are continuously pushed back.
The work necessary to maintain the ordered structures in the body is
obtained from the chemical energy in food. Except for the energy utilized
in external work done by the muscles, all the energy provided by food is ultimately
converted into heat by friction and other dissipative processes in the
body. Once the temperature of the body is at the desired level, all the heat
generated by the body must leave through the various cooling mechanisms
of the body (see Chapter 11). The heat must be dissipated because, unlike
heat engines (such as the turbine or the steam engine), the body does not have
the ability to obtain work from heat energy. The body can obtain work only
142 Chapter 10 Thermodynamics
from chemical energy. Even if the body did have mechanisms for using heat
to perform work, the amount of work it could obtain in this way would be
small. Once again, the second law sets the limit. The temperature differences
in the body are smallnot more than about 7 C between the interior and the
exterior. With the interior temperature T1 at 310 K (37C) and the exterior
temperature T1 at 303 K, the efficiency of heat conversion to work would be
(from Eq. 10.1) at most only about 2%.
Of all the various forms of energy, the body can utilize only the chemical
binding energy of the molecules which constitute food. The body does not
have a mechanism to convert the other forms of energy into work. A person
could bask in the sun indefinitely, receiving large quantities of radiant energy,
and yet die of starvation. Plants, on the other hand, are able to utilize radiant
energy. As animals use chemical energy, so plants utilize solar radiation to
provide the energy for the ordering processes necessary for life.
The organic materials produced in the life cycle of plants provide food
energy for herbivorous animals, which in turn are food for the carnivorous
animals that eat them. The sun is, thus, the ultimate source of energy for life
on Earth.
Since living systems create order out of relative disorder (for example, by
synthesizing large complex molecules out of randomly arranged subunits), it
may appear at first glance that they violate the Second Law of Thermodynamics,
but this is not the case. To ascertain that the second law is valid, we must
examine the whole process of life, which includes not only the living unit but
also the energy that it consumes and the by-products that it rejects. To begin
with, the food that is consumed by an animal contains a considerable degree
of order. The atoms in the food molecules are not randomly arranged but
are ordered in specific patterns. When the chemical energy in the molecular
bindings of the food is released, the ordered structures are broken down. The
eliminated waste products are considerably more disordered than the food
taken in. The ordered chemical energy is converted by the body into disordered
heat energy.
The amount of disorder in a system can be expressed quantitatively by
means of a concept called entropy. Calculations show that, in all cases, the
increase in the entropy (disorder) in the surroundings produced by the living
system is always greater than the decrease in entropy (i.e., ordering) obtained
in the living system itself. The total process of life, therefore, obeys the second
law. Thus, living systems are perturbations in the flow toward disorder. They
keep themselves ordered for a while at the expense of the environment. This
is a difficult task requiring the use of the most complex mechanisms found in
nature. When these mechanisms fail, as they eventually must, the order falls
apart, and the organism dies.
Section 10.5 Information and the Second Law 143
10.5 Information and the Second Law
We have stressed earlier that work must be done to create and maintain the
highly ordered local state of life. We now turn to the question, what else is
needed for such local ordering to occur? Perhaps we can get an insight into
this issue from a simple everyday experience. In the course of time, our apartment
becomes disordered. Books, which had been placed neatly, in alphabetical
order, on a shelf in the living room, are now strewn on the table and some
are even under the bed. Dishes that were clean and neatly stacked in the cupboard,
are now dirty with half-eaten food and are on the living room table. We
decide to clean up, and in 15 minutes or so the apartment is back in order. The
books are neatly shelved, and the dishes are clean and stacked in the kitchen.
The apartment is clean.
Two factors were necessary for this process to occur. First, as was already
stated, energy was required to do the work of gathering and stacking the books
and cleaning and ordering the dishes. Second, and just as important, information
was required to direct the work in the appropriate direction. We had to
know where to place the books and how to clean the dishes and stack them
just so. The concept of information is of central importance here.
In the 1940s, Claude Shannon developed a quantitative formulation for
the amount of information available in a given system. Shannons formula for
information content is shown to be equivalent to the formula for entropythe
measure of disorderexcept, with a negative sign. This mathematical insight
formally shows that if energy and information are available, the entropy in
a given locality can be decreased by the amount of information available to
engage in the process of ordering. In other words, as in our example of the
messy living room, order can be created in a disordered system by work that
is directed by appropriate information. The second law, of course, remains
valid: the overall entropy of the universe increases. The work required to
perform the ordering, one way or another, causes a greater disorder in the
surroundings than the order that was created in the system itself. It is the
availability of information and energy that allows living systems to replicate,
grow, and maintain their structures.
The chain of life begins with plants that possess information in their genetic
material on how to utilize the energy from the sun to make highly ordered complex
structures from the simple molecules available to them: principally water,
carbon dioxide, and an assortment of minerals. The process is, in essence,
similar in human beings and other animals. All the information required for
the function of the organism is contained in the intricate structure of DNA. The
human DNA consists of about a billion molecular units in a well-determined
sequence. Utilizing the energy obtained from the food that is consumed by
144 Chapter 10 Thermodynamics
the organism, the information in the DNA guides the assembly of the various
proteins and enzymes required for the functioning of the organism.
_ EXERCISES _
10-1. Explain how the second law of thermodynamics limits conversion of heat
to work.
10-2. From your own experience, give an example of the second law of thermodynamics.
10-3. Describe the connections between information, the second law of
thermodynamics, and living systems.
_ Chapter 11
Heat and Life
The degree of hotness, or temperature, is one of the most important environmental
factors in the functioning of living organisms. The rates of the
metabolic processes 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 2C to 120C. Only the simplest of living
organisms can function near the extremes of this range.1 Large-scale living
systems are restricted to a much narrower range of temperatures.
The functioning of most living systems, plants and animals, is severely
limited 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 constant levels.
As a result, warm-blooded animals are able to function at an optimum
level over a wide range of external temperatures. Although this temperature
regulation requires additional expenditures of energy, the adaptability
achieved is well worth this expenditure.
1Indeep 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.
145
146 Chapter 11 Heat and Life
TABLE 11.1 _ Metabolic Rates for Selected Activities
Metabolic rate
Activity (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
In this chapter, we will examine energy consumption, heat flow, and temperature
control in animals. Although most of our examples will be specific
to people, the principles are generally applicable to all animals.
11.1