S Y S T E M A N D S U R R O U N D IN G S 1 1

F i g . 1 .9 D i f f e r e n t t y p e s o f s y s t e m .
( A ) A c lo s e d s y s t e m . T h e s t o p p e r
in h ib it s e v a p o r a t io n o f t h e s o lv e n t ,
s o e s s e n t ia lly n o m a t t e r is
e x c h a n g e d w it h t h e s u r r o u n d in g s
( t h e a ir s u r r o u n d in g t h e t e s t t u b e ) ,
b u t h e a t e n e rg y c a n b e e x c h a n g e d
w it h t h e s u r r o u n d in g s t h r o u g h t h e
g la s s . ( B ) A n o p e n s y s t e m . A ll liv in g
o r g a n is m s a r e o p e n s y s t e m s . A c a t
is a r a t h e r c o m p le x o p e n s y s t e m .A
s i m p l i fi e d v i e w o f a c a t i s s h o w n i n
F i g . 1 .1 0 . ( C) A s c h e m a t i c d i a g r a m
o f a sy ste m .

C. S y s t e m a n d s u r r o u n d in g s

We need to define some important terms. This is perhaps most

easil y done b y w ay of ex ampl e. C onsider a b ioc hemic al reac tion that
is c arried ou t in aq u eou s sol u tion in a test tu b e ( F ig . 1 .9 A ) . The system
c onsists of the sol v ent, w ater, and al l c hemic al s dissol v ed in it,
inc l u ding b u f f er sal ts, enz y me mol ec u l es, the su b strate rec og niz ed
b y the enz y me, and the produ c t of the enz y matic reac tion. The
sy stem is defined as that part of the u niv erse c hosen f or stu dy . The
su r r o u n d i n g s are simpl y the entire u niv erse ex c l u ding the sy stem.
The sy stem and su rrou nding s are separated f rom eac h other b y a
b o u n d a r y, in this c ase the test tu b e.
A sy stem is at any time in a c ertain thermody namic sta te or
c ondition of ex istenc e ( w hic h ty pes of mol ec u l e are present and in
w hat amou nt, the temperatu re, the pressu re, etc .) . A sy stem is said
to b e c l o sed if it c an ex c hang e heat w ith the su rrou nding s b u t not
matter. That is, the b ou ndary of a c l osed sy stem is impermeab l e to
matter. A l eak y tire and a dial y sis b ag in a b u c k et of sol v ent – ob j ec ts
permeab l e to smal l mol ec u l es b u t not to l arg e ones – are not c l osed
sy stems! I n ou r test tu b e il l u stration, as l ong as no matter is added
du ring the period of ob serv ation, and as l ong as ev aporation of the
sol v ent does not c ontrib u te sig nific antl y to any ef f ec ts w e mig ht
ob serv e, the sy stem c an b e c onsidered c l osed. M oreov er, the sy stem
w il l b e c l osed ev en if the b ioc hemic al reac tion w e are stu dy ing
resu l ts in the rel ease or ab sorption of heat energ y ; energ y transf er
b etw een sy stem and su rrou nding s c an oc c u r in a c l osed sy stem.
A nother ex ampl e of a c l osed sy stem is E arth itsel f . O u r pl anet
c ontinu al l y rec eiv es radiant energ y f rom the S u n and g iv es of f heat,
b u t b ec au se E arth is neither v ery heav y nor v ery l ig ht, the pl anet
ex c hang es prac tic al l y no matter w ith its su rrou nding s. B y c ontrast,
b l ac k hol es hav e su c h a l arg e g rav itational attrac tion that l ittl e or
nothing c an esc ape, b u t asteroids hav e no atmosphere.
12 ENERGY TRANSFORMATION

Box 1.1 Hot viviparous lizard sex

Viviparous reptiles bear their offspring live. Skinks are any of the more than 1000
liz ard spec ies w hic h c onstitute the family Sc inc id ae. P resent in tropic al regions
ac ross the globe, these liz ard s are partic ularly d iverse in Southeast A sia. Some
spec ies lay eggs; others give birth to fully d eveloped progeny. Eulamprus
t y mpan um is a med ium- siz ed viviparous sc inc id liz ard w hic h inhabits alpine regions
in southeastern A ustralia. M others ac tively thermoregulate to stabiliz e the
temperature of gestation. T he litter siz e is 1 to 5 young. R ec ently, researc hers in
A ustralia found that the d eveloping embryos of E. t y mpan um are subj ec t to
temperature- d epend ent sex d etermination. I n other w ord s, the mother c an
infl uenc e the sex of her offspring and sex ratios in w ild populations. W armer
temperatures give rise to a higher perc entage of male progeny, the frac tion of
females falling from nearly 3 / 5 in the fi eld to 9 / 2 0 at 2 5  C , 1/ 4 at 3 0  C , and 0 at
3 2  C . I n the laboratory, females provid ed w ith unlimited c ond itions for
thermoregulation maintain a bod y temperature of 3 2  C and prod uc e male
offspring only, w hereas in the fi eld , eq ual sex ratios result from natural gestation.
T he w armer temperatures of low er altitud es c ould yield a prepond eranc e of male
young and the eventual inability of those populations to proc reate. G lobal w arming
c ould d rive E. t y mpan um into ex tinc tion. I n early 2 007 c limatologists announc ed
that the rec ent d rought in A ustralia w as likely to lead to an inc reased average
temperature of several d egrees ac ross the c ontinent for the nex t several years.

What if matter can be exchanged between system and

surroundings? Then the system is said to be open. An example of an
open system is a cat (Fig. 1.9B). It breathes in and exhales matter (air)
continually, and it eats, drinks, defecates and urinates periodically.
In barely sufferable technospeak, a cat is an open, self- regulating and
self- reproducing heterogeneous system. The system takes in food
from the environment and uses it to maintain body temperature,
“ power” all the biochemical pathways of its body, including those
of its reproductive organs, and to run, jump and play. The system
requires nothing more for reproduction than a suitable feline of the
opposite sex. And the molecular composition of the eye is certainly
very different from that of the gut; hence, heterogeneous. In the
course of all the material changes of this open system, heat energy
is exchanged between it and the surroundings, the amount
depending on the system’ s size and the difference in temperature
between its body and the environment. A schematic diagram of the
internal structure of this open system is shown in Fig. 1.10 . Whether
the living system is a cat, crocodile, baboon or bacterium, it is an
open system. It seems that it can only be the case that all living
systems that have ever existed have been open systems.
To wrap up this section, an isolated system is one in which the
boundary permits neither matter nor energy to pass through. The
system is constant with regard to material composition and energy.
A schematic diagram of a system, surroundings and boundary are
shown in Fig. 1.9C.
 ANIMAL ENERGY CONSUMP TION 13

Fig. 1.10 The plumbing of a higher animal. Food energy, once inside the body, gets
moved around a lot. Food is digested in the gut and then absorbed into the circulatory
system, which delivers it to all cells of the body. The respiratory system plays a role in
enabling an organism to acq uire the oxygen it needs to burn the fuel of food. Again, the
circulatory system is involved, providing the means of transport of respiratory gases.
W hen energy input to the body exceeds output (excretion þ heat), there is a net increase
in weight. In humans and other animals, the ideal time rate of change of body weight, and
therefore food intak e and physical activity, varies with age and physical condition. Based on
Fig. 1– 5 of P eusner (197 4 ).

D. Animal energy consumption

Now let’s take a more in-depth look at the relationship between

food, energy, and life. We wish to form a clear idea of how the
energy requirements of carrying out various activities, for instance
walking or sitting, relate to the energy available from the food we
eat. The discussion is largely qualitative, but a formal definition of
heat will be given.
Energy measurements can be made using a calorimeter.
Calorimetry has made a big contribution to our understanding of
the energetics of chemical reactions, and there is a long tradition
of using calorimeters in biological research. In the mid seven-
teenth century, pioneering experiments by R obert Boyle (16 2 7 –
16 91) in Oxford demonstrated the necessary role of air in com-
bustion and in respiration. Taking a breath is more like burning a
piece of wood than many people suspect. About 12 0 years later, in
17 8 0, Antoine L aurent L avoisier (17 4 3 –17 94 ) and P ierre Simon de
L aplace (17 4 9–18 2 7 ) used a calorimeter to measure the heat given
off by a live guinea pig. On comparing this heat with the amount
of oxygen consumed, the Frenchmen correctly concluded that
respiration is a form of combustion. Nowadays, a so-called bomb
14 ENERGY TRANSFORMATION

Fig. 1.11 Schematic diagram of a bomb calorimeter. A sample is placed in the reaction
chamber. The chamber is then filled with oxygen at high pressure (>20 atm) to ensure
that the reaction is fast and complete. Electrical heating of a wire initiates the reaction.
The increase in water temperature resulting from the combustion reaction is recorded,
and the temperature change is converted into an energy increase. The energy change is

1
1
divided by the total amount of substance oxidized, giving units of J g or J mol .
Insulation helps to prevent the escape of the heat of combustion, increasing the accuracy
of the determination of heat released from the oxidized material. Based on diagram on
p. 36 of Lawrence et al. (1996 ).

calorimeter9 (Fig. 1.11) is used to measure the heat given off in the
oxidation of a combustible substance like food, and nutritionists
refer to tables of combustion heats in planning a diet.
The study of energy transformations is called thermodynamics.
It is a hierarchical science – the more advanced concepts assume
knowledge of the more basics ones. To be ready to tackle the more
difficult but more interesting topics in later chapters, let’s use this
moment to develop an understanding of what is being measured in
the bomb calorimeter. We know from experience that the oxidation
(burning) of wood gives off heat. Some types of wood are useful for
building fires because they ignite easily (e.g. splinters of dry pine);
others are useful because they burn slowly and give off a lot of heat
(e.g. oak). The amount of heat transferred to the air per unit volume
of burning wood depends on the density of the wood and its struc-
ture. The same is true of food. Fine, but this has not told us what
heat is.
It is the nature of science to define terms as precisely as possible
and to formalize usage. Accepted definitions are important
for minimizing ambiguity of meaning. What we need now is a

9
But one of many different kinds of calorimeter. The instrument used to measure the
energy given off in an atom smasher is called a calorimeter. In this book we discuss a
bomb calorimeter, isothermal titration calorimeter, and differential scanning
calorimeter.
 ANIMAL ENERGY CONSUMPTION 15

definition of heat. H eat, or thermal energy, q, is a form of kinetic

energy; that is, energy arising from motion. H eat is the change in
energy of a system that results from a temperature difference
between it and the surroundings. For instance, when a warm can of
Coke is placed in a refrigerator, it gives off heat continuously until
reaching the same average temperature as all other objects in the
fridge, including the air. The heat transf erred from the Coke can to
the air is absorbed by the other things in the fridge. H eat is said to
fl ow from a region of higher temperature, where the average speed
of molecular motion is greater, to one of lower temperature.
The fl ow of heat does indeed remind us of a liquid, but it does
not necessarily follow, and indeed we should not conclude, that
heat is a material particle. H eat is rather a type of energy transfer.
H eat makes use of random molecular motion. Particles that exhibit
such motion (all particles!) are subject to the usual mechanical laws
of physics. A familiar example of heat transfer is the boiling of water
in a saucepan. The more heat applied, the faster the motion of
water. The bubbles that form on the bottom of the pan give some
indication of how fast the water molecules are moving. This is about
as close as we get under ordinary circumstances to “seeing” heat
being transferred. But if you’ve ever been in the middle of a shower
when the hot water has run out, you will know what it’s like to f eel
heat being transferred! By convention, q > 0 if energy is transferred
to a system as heat, if the total energy of the system increases by way
of heat transfer. In the case of a cold shower, and considering the
body to be the system, q is negative.
Now we are armed for another look at the oxidation of mate-
rials in a bomb calorimeter and the relationship to nutrition. The
heat released or absorbed in a reaction is measured as a change in
temperature; calibration of an instrument using known quantities
of heat can be used to relate heats of reaction to changes in
temperature. One can plot a standard curve of temperature v ersus
heat, and the heat of oxidation of an unknown material can then
be determined experimentally. Table 1.2 shows the heats of oxi-
dation of different foodstuffs. Evidently, and important for phy-
siology, some types of biological molecule give off more heat per
unit mass than others. Some idea of the extent to which the
energy obtained from food is utilized in various human activities is
given in Table 1.3.
Animals, particularly humans, “consume” energy in a variety of
ways, not just by eating, digesting and metabolizing food. For
instance, most automobiles of the present day run on octane, and
electrical appliances depend on the generation of electricity. The
point is that energy transformation and consumption can be
viewed on many different levels. As our telescopic lens becomes
more powerful, the considerations range from one person to a
family, a neighborhood, city, county, state, country, continent, sur-
face of the earth, biosphere, solar system, galaxy . . . As the length
scale decreases, the microscope zooms in on an organ, a tissue,
16 ENERGY TRANSFORMATION

Table 1.2. Heat released upon oxidation to CO2 and H2O

E nergy yield

kcal
Substance kJ ( mol
1) kJ ( g
1) kcal ( g
1) ( g
1 wet wt)
Glucose 2 8 17 15.6 3.7 —
L actate 1 36 4 15.2 3.6 —
Palmitic acid 10 040 39.2 9.4 —
Glycine 979 13.1 3.1 —
Carbohydrate — 16 3.8 1.5
F at — 37 8 .8 8 .8
Protein — 23 5.5 1.5
Protein to urea — 19 4.6 —
E thyl alcohol — 29 6 .9 —
L ignin — 26 6 .2 —
Coal — 28 6 .7 —
O il — 48 11 —
D-glucose is the principal source of energy for most cells in higher organisms. It is converted to lactate in anaerobic homolactic
fermentation (e.g. in muscle), to ethyl alcohol in anaerobic alcoholic fermentation (e.g. in yeast), and to carbon diox ide and w ater in
aerobic ox idation. P almitic acid is a fatty acid. G lycine, a constituent of protein, is the smallest amino acid. C arbohydrate, fat and
protein are three different types of biological macromolecule and three different sources of energy in food. M etabolism in animals
leaves a residue of nitrogenous ex cretory products, including urea in urine and methane produced in the gastrointestinal tract. E thyl
alcohol is a maj or component of alcoholic beverages. L ignin is a plasticlik e phenolic polymer that is found in the cell w alls of plants; it
is not metaboliz ed directly by higher euk aryotes. C oal and oil are fossil fuels that are produced from decaying organic matter,
primarily plants, on a time scale of millions of years. T he data are from T able 2 .1 of W rigglesw orth (19 9 7 ) or T able 3 .1 of B urton (19 9 8 ).
S ee also T able A in A ppendix C .

Table 1.3. E nergy expenditure in a 7 0 k g human

Total energy expenditure

F orm of activity ( kcal h– 1)
L ying still, awake 77
Sitting at rest 100
Typewriting rapidly 140
D ressing or undressing 150
Walking on level, 2.6 mi/h 200
Sexual intercourse 28 0
B icycling on level, 5.5 mi/h 304
Walking on 3 percent 357
grade, 2.6 mi/h
Sawing wood or shoveling 48 0
snow
J ogging, 5.3 mi/h 570
Rowing, 20 strokes/min 8 28
Maximal activity ( untrained) 1440
The measurements were made by indirect calorimetry. Digestion increases the rate of
metabolism by as much as 30% over the basal rate. During sleep the metabolic rate is about
10% lower than the basal rate. The data are from Table 15 –2 of V ander, Sherman and Luciano
(1985 ).