Energy Production: Hapter 3
Energy Production: Hapter 3
Energy Production
The reason that we eat, besides the fact that food can be so delicious, is
for energy and building blocks. Although energy cannot be created or
destroyed, its form can change. During metabolism, our bodies break down
fuel molecules and trap the energy released within the molecule adenosine
triphosphate (ATP). In this chapter, we examine the energy transformations
that occur during metabolism.
ATP
OH H
ADP Inorganic phosphate
The energy requirements of an individual vary widely with the activity
being performed (Table 1). During periods of heavy exertion, energy
demands increase dramatically, particularly by muscles. However, the
Exercise Physiology
J. T. Millard
Phosphocreatine
Phosphocreatine is a secondary reserve of energy that can quickly
generate more ATP from ADP as follows:
NH2 NH2
N N
N HN PO3- N NH2
+
N
C NH2 C NH2+
O O N O O O N N
+ N CH3 + N CH3
- -
O P O P O O P O P O P O
O O
CH2 CH2
O -
O - H H O -
O -
O - H H
H H COO- H H COO-
OH H OH H
ADP Phosphocreatine ATP Creatine
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Exercise Physiology
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Carbohydrates
Animals store carbohydrates as glycogen, a polysaccharide consisting of
thousands of glucose units covalently linked together. Glycogen reserves,
which represent the body’s “quick” form of energy, are found in the muscles
and the liver. They are first broken down to glucose before undergoing
further metabolism. Glucose can then be further processed to release
energy under both anaerobic and aerobic conditions.
Fats
Fats are stored as triglycerides, molecules that contain a single glycerol
unit linked to three fatty acid units. Most of our fat stores are found in
adipose tissue, but other organs, including muscles, also have small reserves
to fuel their ongoing needs. Triglycerides, which account for about 84% of
the total energy stores in the average individual, are degraded to fatty acids
(and glycerol) before being further metabolized. Fatty acids can undergo
only aerobic metabolism.
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Exercise Physiology
J. T. Millard
Proteins
While dietary protein
can also be degraded for
energy, we have very little
expendable protein
reserves. In dire
circumstances, such as a
precipitous drop in blood Further
sugar levels, skeletal metabolism
muscle will actually be Figure 1. The three classes of fuel molecules and
broken down despite how they are broken down during metabolism.
potentially disastrous
consequences. This occurs primarily to fuel the brain, which needs a constant
source of energy in the form of glucose. Animals lack a metabolic pathway
that can make glucose from fats, but most amino acids can undergo
conversion to glucose. Thus, skeletal muscle may be sacrificed when glycogen
stores are depleted.
Energy Content
Glucose and triglycerides are normally broken down to release energy
through oxidation to carbon dioxide, with oxygen undergoing reduction to
water in the process. Recall that redox reactions involve the transfer of
electrons from one substance to another. In reactions involving electron
transfer there cannot be an oxidation without a reduction.
During aerobic metabolism, animals obtain energy for essential life
processes through the oxidation of carbon-containing nutrients such as
glucose (C6H12O6):
C6H12O6 + 6 O2 à 6 CO2 + 6 H2 O
This reaction illustrates that there are several ways to think about redox
reactions. We can think about oxidation as loss of electrons, as gain of
oxygen (hence the name), or as loss of hydrogen. Reduction is just the
opposite: gain of electrons, loss of oxygen, or gain of hydrogen. C6H12O6
(glucose) is oxidized in the last reaction because carbon gains bonds to
oxygen and loses bonds to hydrogen in its conversion to CO2. If glucose is
oxidized, then O2 (oxygen) must be reduced as it is the only other reactant,
and oxidation is always paired with reduction. Indeed, oxygen gains hydrogen
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atoms during the conversion to H2O, meeting the third criterion for
reduction.
If we examine the structure of a representative carbohydrate, like
glucose, in comparison to a representative fat, like lauric acid (C12H24O2), we
can account for the different energy contents of these fuels. Many of the
carbon atoms in glucose are bonded to oxygen, meaning that they are initially
more oxidized than the carbons in fat (Figure 2). Therefore, only 4 kcal of
energy per gram are released upon oxidizing carbohydrates versus 9 kcal per
gram for fats. That is, fats represent a more efficient means of storing
energy because they are more highly reduced, thereby releasing more
energy upon oxidation.
Mobilization of Fuels
When muscles exhaust their stores of readily available ATP and
phosphocreatine, the next fuel of choice is glucose, either from blood sugar
or the muscle’s own glycogen. Glucose is quickly mobilized to produce ATP to
power the muscles and allow activity to continue. The ultimate product of
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Figure 7. Electrons
from NADH and
FADH2 enter the
ETC at complex I
and II, respectively.
Electrons take the
path indicated by the
orange arrows,
causing pumping of
+
H where shown,
making the inside of
the mitochondria
less acidic. The
resulting gradient
can be used to make
+
ATP when H comes
back inside through
the ATP synthase.
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cycle (1) with the number produced in the ETC (27) yields a grand total of 30
molecules of ATP per molecule of glucose.
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Exercise Physiology
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Fat Metabolism
The average individual has enough stored fat to last for about two months
without eating. Recall that fats are stored primarily in fat cells (adipocytes).
These cells send out fatty acids when they receive a hormonal signal that
either the body’s blood sugar has dropped (the hormone glucagon) or that
the body needs energy for fight or flight (the hormone adrenaline).
During exercise, it takes about 20 minutes for fats to be sent out from
adipocytes and reach the skeletal muscle and heart. Once the fatty acids
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liver, which converts the NH3 to the molecule urea (CH4N2O) for excretion
because ammonia itself is toxic.
Interestingly, the liver lacks the appropriate enzyme to break down one
family of amino acids, the branched-chain amino acids (leucine, isoleucine,
and valine). On the other hand, not only can muscles use these amino acids
for energy, but there is also some evidence that suggests that the
branched-chain amino acids, particularly leucine, reduce muscle breakdown
during exercise.
Figure 12. Muscles have small amounts of ATP that are used up in a couple of seconds.
Phosphocreatine (PC) rapidly forms more ATP but is used up in about 10 seconds.
Anaerobic metabolism of glucose (the lactic acid system) operates quickly but is shut
down in a minute or two because of acidification of the muscle. Aerobic metabolism of
glucose, fats, and amino acids is slower but can be sustained for hours.
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Table 1.
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Lactate Threshold
Lactic acid is produced by working muscles and released into the
bloodstream for transport to the liver. As exercise intensity increases,
there is usually little increase in the lactic acid concentration of blood
initially because other tissues can burn it for energy, like the heart, or
reprocess it, like the liver. Once the intensity reaches a certain level,
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Fat Metabolism
Many aspects of lipid metabolism are also carried out in the liver. The
liver actively oxidizes triglycerides to meet its own energy demands. In fact,
the liver breaks down many more fats than it needs, producing and exporting
water-soluble derivatives of fats called ketone bodies that can be used by
other tissues for energy. The liver is also the major site for converting
excess carbohydrates and proteins into triglycerides, which are then
exported and stored in adipose tissue.
Protein Metabolism
Several critical aspects of protein metabolism also occur in the liver,
including processing of most ingested amino acids, followed by conversion of
the non-nitrogenous part of those molecules to glucose or lipids. The liver
also processes nitrogenous waste from proteins, helping to remove ammonia
from the body through the synthesis of urea. Ammonia is very toxic and can
affect the brain with a range of symptoms from mild confusion to possible
brain damage, coma, and death. The liver also makes many proteins found in
the bloodstream, including albumin, which escorts fats from adipose tissue
to muscle cells during exercise.
With all the beneficial chemistry that the liver is performing, it’s no
wonder that many athletes choose to go dry during their seasons. Your liver
is busy enough without making it metabolize a bunch of alcohol!
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