ENERGY BALANCE

Energy Input to the Body is the Sum of Energy Output and Storage
The first law of thermodynamics states that energy can neither be created nor destroyed; in a closed
system, total energy is constant. This concept is illustrated in Figure 57-1. Humans acquire almost all their
energy from ingested food, store it in different forms, and expend it in different ways. In the steady state,
energy uptake must equal energy output.
The gastrointestinal (GI) tract breaks down ingested carbohydrates, fats, and proteins into smaller
components and then absorbs them into the bloodstream for transport to sites of metabolism. For example,
the GI tract reduces ingested carbohydrates to simple sugars (e.g., glucose), which are then transported to
muscle cells and either oxidized to release energy or converted to glycogen for storage. Oxidation of fuels
not only generates free energy but also waste products and heat (thermal energy).
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Figure 57-1 Energy balance.
The body's energy inputs must balance the sum of its energy outputs and the energy stored. When the
body takes in more energy than it expends, the person is in positive energy balance and will gain weight.
Healthy children are in positive energy balance during growth periods. Conversely, when energy intake is
less than expenditure, this negative energy balance leads to weight loss. A person in negative energy
balance preferentially uses stored fat as fuel.
A person can gain or lose weight by manipulating energy intake or output. An optimal strategy to encourage
weight loss involves both increasing energy output as well as reducing energy intake. In most people, a
substantial decrease in energy intake alone leads to inadequate nutrient intake, which can compromise
bodily function.
Because of the Inefficiency of Chemical Reactions, the Body's Total Free Energy Falls
The second law of thermodynamics states that chemical transformations always result in a loss of free
energy. The total internal energy (E) of the human body is the sum of the disposable or "good" energy (i.e.,
the Gibbs free energy, G) plus the unavailable or "wasted" energy (i.e., the product of absolute
temperature, T, and entropy, S):
Imagine that you eat some glucose, which increases your total internal energy by a small amount (E).
Some of this energy will be stored as glycogen (G), and some will waste as heat (T S). According to
Equation 1, as long as the temperature is constant, the change in total internal energy will have two
components:

Thus, some of the increased total energy (E) will be stored as glycogen (G). However, because of the
inefficiencies of the chemical reactions that convert glucose to glycogen, some of the E is wasted as heat
(T S). Another way of stating the second law is that T S can never be zero or negative, and chemical
reactions can never be 100% efficient.
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If we add no energy to the body (i.e., E is zero), the body's total free energy must decline (i.e., G is
negative). This decline in G matches precisely the rise in T S, reflecting inefficiencies inherent in chemical
transformations. Consider, for example, what would happen if you took 1 mole of glucose (180 g), put it into
a bomb calorimeter with O
2
, and completely burned the glucose to CO
2
and H
2
O. This combustion would
yield 686 kcal in the form of heat but conserve no usable energy. Now consider what happens if your body
burns this same 1 mole of glucose. In contrast to the bomb calorimeter, your mitochondria not only oxidize
glucose to CO
2
and H
2
O but also conserve part of the free energy in the form of adenosine triphosphate
(ATP). Each of the many chemical-conversion steps from glucose to CO
2
and H
2
O makes available a small
amount of the total energy contained in glucose. Converting 1 mole of adenosine di-phosphate (ADP) and
inorganic phosphate (P
i
) to 1 mole of ATP under the conditions prevailing in a cell consumes approximately
11.5 kcal/mole. Therefore, if a particular step in glucose oxidation releases at least 11.5 kcal/mole, it can be
coupled to ATP synthesis. The conversion of the lower-energy ADP to the higher-energy ATP traps energy
in the system, conserving it for later use. The cellular oxidation of 1 mole of glucose conserves
approximately 400 kcal of the potential 686 kcal/mole; the remaining 286 kcal/mole are liberated as heat.
Because the body cannot reconvert this heat to energy, the body's total free energy falls.
Free Energy, Conserved as High-Energy Bonds in ATP, is Released in a Controlled Manner to Provide the
Energy for Cellular Functions
ATP consists of a nitrogenous ring (adenine), a five-carbon sugar (ribose), and three phosphate groups
(Fig. 57-2). The last two phosphates are connected to the rest of the molecule by high-energy bonds. The
same is true for a related nucleotide, guanosine triphosphate (GTP). If we compare the free energies of
phosphate bonds of various molecules, we see that the "high-energy" phosphate bonds of ATP lie toward
the middle of the free-energy scale. Thus, in the presence of P
i
, ADP can accept energy from compounds
that are higher on the free-energy scale (e.g., phosphocreatine), whereas ATP can release energy in the
formation of compounds that are lower on the free-energy scale (e.g., glucose-6-phosphate [G6P]). Thus,
ATP can store energy derived from energy-releasing reactions and release energy needed to drive other
chemical reactions.
Figure 57-2 Hydrolysis of ATP to ADP, inorganic phosphate, and H
+
. ADP, adenosine diphosphate; ATP, adenosine triphosphate.
Examples of the chemical reactions fueled by converting ATP to ADP and P
i
include the formation of
peptide bonds in protein synthesis, bridge formation between actin and myosin during muscle contraction,
and pumping Ca
2+
against its concentration gradient during muscle relaxation. Although the conversion of
ATP to ADP and P
i
is often referred to as a "hydrolysis reaction," because the traditional representation is
ATP + H
2
O ADP + P
i
+ H
+
, the reaction actually occurs in two steps. The first typically involves transfer
of a part of the phosphate group to an intermediate molecule, thus increasing the intermediate's free-energy
content. The second step involves displacing the phosphate moiety, releasing P
i
and energy. In contrast,
true hydrolysis reactions merely release heat, which cannot be trapped to drive chemical processes.
Printed from STUDENT CONSULT: Medical Physiology (on 28 August 2006)
2006 Elsevier