ATP AND COUPLED REACTION PROCESSES

Adenosine triphosphate or ATP is the energy currency of the cell. ATP hydrolysis
releases the energy present in the high-energy terminal phosphate bonds, which is
utilised to carry out various cellular reactions, such as muscle contraction, carbon
fixation, etc. Various reactions are coupled with ATP hydrolysis.
ATP hydrolysis is an exergonic process. It produces ADP (Adenosine diphosphate), Pi
(inorganic phosphate) and energy. ADP can undergo further hydrolysis in some cases to
produce AMP (Adenosine monophosphate) and Pi.

ATP Structure and Hydrolysis Mechanism

ATP is ribonucleic acid. It is made up of a nitrogenous base, pentose sugar and

phosphate. In ATP, the nitrogenous base is adenosine and the sugar is ribose, which is
linked to phosphate.

ATP Hydrolysis

The terminal phosphoanhydride bonds are known as high-energy bonds. They release a
large amount of energy on hydrolysis to power the energy-requiring cellular processes.
The reaction of ATP hydrolysis is as follows:
ATP + H2O ⇋ ADP + Pi + Energy
This reaction is reversible. The reversible reaction requires energy to produce ATP. ADP
and Pi regenerate ATP. Thus, energy gets stored in the form of ATP and when energy is
required, ATP is hydrolysed. The reverse reaction of ATP hydrolysis or the reaction for
ATP synthesis is as follows:
ADP + Pi + Energy ⇋ ATP + H2O
The enzyme ATP synthase catalyses the synthesis of ATP.

Energy Output in ATP Hydrolysis

The hydrolysis of 1M of ATP into ADP and inorganic phosphate, releases −7.3 kcal/mol
of energy. The energy released in the living cell is almost double the value in standard
conditions, it is equal to −14 kcal/mol.
ATP + H2O → ADP + Pi ΔG° = −7.3 kcal/mol (−30.5 kJ/mol)
The hydrolysis of ATP into AMP and pyrophosphate (PPi) releases −10.9 kcal/mol of
energy.
ATP + H2O → AMP + PPi ΔG° = −10.9 kcal/mol (−45.6 kJ/mol)

Significance of ATP Hydrolysis

Energy is stored in the form of ATP in living organisms. Most ATP is produced during
respiration. The energy released by the oxidation of carbohydrates and respiratory
substrates is trapped and stored in the form of ATP, which can later be utilised as and
when the requirement arises.
The energy released during ATP hydrolysis is utilised to power the cellular processes.
There are many processes that require energy, such as muscle contraction, active
transport, cell signalling, DNA RNA synthesis, etc.
There are many reactions that are coupled to ATP hydrolysis. These reactions are
endergonic, i.e. they require energy. An example of reaction coupling is phosphorylation,
e.g. glucose to glucose 6 phosphate catalysed by the enzyme hexokinase in the
glycolysis process. The phosphate group released in ATP hydrolysis is used to
phosphorylate glucose. Sodium-potassium pump is another example where ATP
hydrolysis is coupled to the change of shape of transport proteins, leading to transport
of ions across the membrane.

Reaction coupling

How is the energy released by ATP hydrolysis used to power other reactions in a cell? In most
cases, cells use a strategy called reaction coupling, in which an energetically favorable reaction
(like ATP hydrolysis) is directly linked with an energetically unfavorable (endergonic) reaction.
The linking often happens through a shared intermediate, meaning that a product of one
reaction is “picked up” and used as a reactant in the second reaction.

When two reactions are coupled, they can be added together to give an overall reaction, and the
ΔG of this reaction will be the sum of the ΔG values of the individual reactions. As long as the
overall ΔG is negative, both reactions can take place.

ATP in reaction coupling

When reaction coupling involves ATP, the shared intermediate is often a phosphorylated
molecule (a molecule to which one of the phosphate groups of ATP has been attached). As an
example of how this works, let’s look at the formation of sucrose, or table sugar, from glucose
and fructose^{3,4}3,4start superscript, 3, comma, 4, end superscript.

In this process, ATP transfers one of its phosphate groups to the pump protein, forming ADP and
a phosphorylated “intermediate” form of the pump. The phosphorylated pump is unstable in its
original conformation (facing the inside of the cell), so it becomes more stable by changing
shape, opening towards the outside of the cell and releasing sodium ions outside. When
extracellular potassium ions bind to the phosphorylated pump, they trigger the removal of the
phosphate group, making the protein unstable in its outward-facing form. The protein will then
become more stable by returning to its original shape, releasing the potassium ions inside the
cell.

Although this example involves chemical gradients and protein transporters, the basic principle
is similar to the sucrose example above. ATP hydrolysis is coupled to a work-requiring
(energetically unfavorable) process through formation of an unstable, phosphorylated
intermediate, allowing the process to take place in a series of steps that are each energetically
favorable.