section 5.5 
Energy Metabolism
77
the kinetic theory of reaction rates. From the Arrhenius 
equation, it is apparent that the larger the value of 
Es
(or 
Ea),
the smaller the value of 
k.
Hence, the basis for 
kinetic irreversibility is the same as that for thermody-
namic irreversibility; the more energy a process takes (the 
more positive the value of AG or 
Ea),
the less likely the 
reaction is to occur.
Although values for A are not well understood, the 
Arrhenius equation can, on rearrangement, be used to 
calculate 
Es:
Es
—R
In 
k =
------
R
In A.
T
The slope of a plot o f
—R
In 
k
versus (1/7/) equals 
Es,
and 
A 
Ha = Es —
RT.
5.5 Energy Metabolism
In order to carry out the body’s essential functions (e.g., 
growth and repair, pregnancy, lactation, physical activ-
ity, maintenance of body temperature), food must be con-
sumed and utilized and body constituents synthesized. 
The term “metabolism” encompasses the numerous chem-
ical transformations that occur within the human body. 
Metabolism comprises 
anabolism
and 
catabolism.
An-
abolism is concerned with the synthesis of new molecules, 
usually larger than the reactants, and is an energy-requiring 
process. Catabolism is concerned with degradation pro-
cesses, usually the breaking down of large molecules into 
smaller ones, and is an energy-yielding process. 
Inter-
mediary metabolism
refers to all changes that occur in a 
food substance beginning with absorption and ending with 
excretion.
Two main sources of energy for metabolism are carbo-
hydrates and fats (lipids). Proteins have less importance as 
an energy supplier. Knowledge of the total energy (A
H)
content of the major body fuels is necessary to understand 
how energy requirements are met by different fuels.
The reactions for representative food fuels, glucose and 
palmitic acid, will be used to indicate differences in A 
H
values. The subscripts s, 
£,
and g (solid, liquid, and gas, re-
spectively) indicate the phase of the material under 
1
atm, 
the pressure at which the reaction is carried out, and at 
the given temperature. The A 
H°
values given are 
stan-
dard heats of formation,
since all reactants are in their 
standard (natural) state for the temperature given.
Oxidation of 1 mol of glucose
(a carbohydrate) to
carbon dioxide and water:
C6H120 6(S) + 602(g) —►
6C02(g) + 6H20(£)
At 20°C, 
AH
= —673 kcal/mol or —2816 kJ/mol.
Oxidation of 1 mol of palmitic acid (a fatty acid) to
carbon dioxide and water:
Ci6H3
2
0 2(S) + 2302(g)—>16C02(g) + 16H20(i)
At 20°C, 
A
H°
= 2380 kcal/mol 
or 
-9.96 MJ/mol 
(1 MJ = 1 megajoule = 1000 kJ).
The average 
A H
values in kilocalories per gram are 
as follows: carbohydrate, 4.1; lipid, 9.4; and protein, 5.6. 
These values vary within each class of food depending 
on chemical structure. For example, the value for starch 
is 4.1 kcal/g, whereas for glucose it is 3.8. Furthermore, 
in humans the end product of protein metabolism is urea 
instead of C 0
2
and nitrogen, which are the end products 
obtained after complete oxidation in the bomb calorime-
ter. Therefore, a realistic estimate of energy derived from 
protein in the body, taking into account incomplete oxida-
tion and specific dynamic action, is 4.1 kcal/g. In human 
nutrition, the commonly used values for energy yield in 
kilocalories (or kilojoules) per gram are as follows: car-
bohydrate, 4 (16.7); lipid, 9 (37.7); and protein, 4 (16.7). 
During the metabolism of food substances in the body, 
oxygen is consumed and C 0
2
is produced. The molar ra-
tio of C 0
2
produced to oxygen consumed is known as 
the 
respiratory quotient
(RQ) and is characteristic of a 
given substrate. For example, in the complete oxidation of 
glucose presented above,
volume of C 0
2
produced 
6C0
2
RQ = ------------------— ----------= -------
-
= 1.0.
volume of 
0
2
consumed 
602
For the corresponding oxidation of palmitic acid,
RQ =
16C02
2302
= 0.7.
The RQ for protein is difficult to measure because it is not 
oxidized completely in the body and part of its carbon, hy-
drogen, and oxygen is lost in the urine and feces. However, 
the RQ is usually taken to be 0.8. The measurement of RQ 
thus provides a means for assessing the type of food that 
is being metabolized. An overall RQ of 0.85 is obtained in 
a normal adult who consumes a mixed diet. This value is 
enhanced by carbohydrate feeding and decreased by lipid 
feeding. Heat production can be calculated from oxygen 
consumption and RQ as follows. For glucose oxidation:
C6Hi20 6 + 60 2 = 6C02 + 6H20
180 g 
192 g
(134L)
Since the energy yield of 1 g of glucose is 3.8 kcal, 180 g 
of glucose yields 180 x 3.8 or 684 kcal. The energy yield 
per liter of oxygen consumed equals (180 x 3.8)/134 = 
5.1 kcal (21.3 kJ). Again, for palmitate oxidation,
Ci6H3
2
0
2
+ 2302
256 g 
236 g
16C02 + 16H20.
 
    
