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When the terminal phosphate group is removed from ATP by hydrolysis, two negatively
charged products are formed, ADP3− and the phosphate group HPO 2− (reaction [47]).
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These products are electrically more stable than the parent molecule and do not readily
recombine. The total free energy (G) of the products is much less than that of ATP; hence,
energy is liberated (i.e., the reaction is exergonic). The amount of energy liberated under
strictly defined conditions is called the standard free energy change (ΔG′). This value for
the hydrolysis of ATP is relatively high, at −8 kilocalories per mole. (One kilocalorie is the
amount of heat required to raise the temperature of 1,000 grams of water one degree
Celsius.) Conversely, the formation of ATP from ADP and inorganic phosphate (P ) is an
i
energy-requiring (i.e., endergonic) reaction with a standard free energy change of +8
kilocalories per mole.
mole); AMP hydrolysis liberates less energy (the standard free energy change is −2.2
kilocalories per mole).
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The free energy of hydrolysis of a compound thus is a measure of the difference in energy
content between the starting substances (reactants) and the final substances (products).
ATP does not have the highest standard free energy of hydrolysis of all the naturally
occurring phosphates but instead occupies a position at approximately the halfway point in
a series of phosphate compounds with a wide range of standard free energies of hydrolysis.
Compounds such as 1,3-diphosphoglycerate and phosphoenolpyruvate (PEP), which are
above ATP on the scale, have large negative ΔG′ values on hydrolysis and are often called
high-energy phosphates. They are said to exhibit a high phosphate group transfer potential
because they have a tendency to lose their phosphate groups. Compounds such as glucose
6-phosphate and fructose 6-phosphate, which are below ATP on the scale because they
have smaller negative ΔG′ values on hydrolysis, have a tendency to hold on to their
phosphate groups and thus act as low-energy phosphate acceptors.
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Both ATP and ADP act as intermediate carriers for the transfer of phosphate groups (which
are more precisely called phosphoryl groups), and hence of energy, from compounds lying
above ATP to those lying beneath it. Thus, in glycolysis, ADP acts as an acceptor of a
phosphate group during the synthesis of ATP from PEP (reaction [10]), and ATP functions
as a donor of a phosphate group during the formation of fructose 1,6-diphosphate from
fructose 6-phosphate (reaction [3]).
The first step in glycolysis, the formation of glucose 6-phosphate (G6P), illustrates how an
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coupling it to ATP.
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Reaction [48] has a positive ΔG′ value, indicating that the reaction tends to proceed in the
reverse direction. It is therefore necessary to use the standard free energy generated byAdtheclose
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breaking of the first phosphate bond in ATP (reaction [48a]), which is −7.3 kilocalories per
mole, to move reaction [48] in the forward direction. Combining these reactions and their
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standard free energies gives reaction [48b] and a standard free energy value of −4
kilocalories per mole, indicating that the reaction will proceed in the forward direction.
There are many intracellular reactions in which the formation of ADP or AMP from ATP
provides energy for otherwise unfavourable biosyntheses. Some cellular reactions use
equivalent phosphorylated analogues of ATP—for example, guanosine triphosphate (GTP)
for protein synthesis.
Energy conservation
The amount of ATP in a cell is limited, and it must be replaced continually to maintain
repair and growth. This is achieved by using the energy liberated during the oxidative
stages of catabolism to synthesize ATP from ADP and phosphate. The synthesis of ATP
linked to catabolism occurs by two distinct mechanisms: substrate-level phosphorylation
and oxidative, or respiratory-chain, phosphorylation. Oxidative phosphorylation is the
major method of energy conservation under aerobic conditions in all nonphotosynthetic
cells.
Substrate-level phosphorylation
donor (e.g., 1,3-diphosphoglycerate) to ADP to yield a molecule of ATP. This type of ATP
synthesis (reactions [7], [10], and [43]) does not require molecular oxygen (O ), although it
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Four types of hydrogen or electron carriers are known to participate in the respiratory
chain, in which they serve to transfer two reducing equivalents (2H) from reduced
substrate (AH ) to molecular oxygen (reaction [49]); the products are the oxidized
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substrate (A) and water (H O).
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The carriers are NAD+ and, less frequently, NADP+; the flavoproteins FAD and FMN
(flavin mononucleotide); ubiquinone (or coenzyme Q); and several types of cytochromes.
Each carrier has an oxidized and reduced form (e.g., FAD and FADH , respectively), the
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two forms constituting an oxidation-reduction, or redox, couple. Within the respiratory
chain, each redox couple undergoes cyclic oxidation-reduction; i.e., the oxidized
component of the couple accepts reducing equivalents from either a substrate or a reduced
carrier preceding it in the series and in turn donates these reducing equivalents to the next
oxidized carrier in the sequence. Reducing equivalents are thus transferred from substrates
to molecular oxygen by a number of sequential redox reactions.
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The fourth type of carrier, the cytochromes, consists of hemoproteins—i.e., proteins with a
nonprotein component, or prosthetic group, called heme (or a derivative of heme), which is
an iron-containing pigment molecule. The iron atom in the prosthetic group is able to carry
one electron and oscillates between the oxidized, or ferric (Fe 3+), and the reduced, or
ferrous (Fe 2+), forms. The five cytochromes present in the mammalian respiratory chain,
designated cytochromes b, c , c, a, and a , act in sequence between ubiquinone and
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molecular oxygen. The terminal cytochrome of this sequence (a , also known as
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cytochrome oxidase) is able to donate electrons to oxygen rather than to another electron
carrier; a is also the site of action of two substances that inhibit the respiratory chain,
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potassium cyanide and carbon monoxide. Special Fe-S complexes play a role in the activity
of NADH dehydrogenase and succinate dehydrogenase.
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respiratory chain
The respiratory chain.
Image: Encyclopædia Britannica, Inc.
In each redox couple, the reduced form has a tendency to lose reducing equivalents (i.e., toclose
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act as an electron or hydrogen donor); similarly, the oxidized form has a tendency to gain
reducing equivalents (i.e., to act as an electron or hydrogen acceptor). The oxidation-
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