A-level Biology P530│ CELLULAR RESPIRATION

WHAT IS THE EXACT MEANING OF CELLULAR RESPIRATION?

A series of multi-enzyme catalysed reactions in cells during which the chemical-bond energy
of complex organic substances (e.g. carbohydrates, lipids, proteins) is released and
converted into the usable form called adenosine triphosphate (ATP).

WHERE IS CHEMICAL ENERGY STORED IN ORGANIC SUBSTANCES (FOOD),

AND HOW DO ORGANISMS HARVEST IT?
Organic substances especially carbohydrates and lipids have C-H covalent bonds which are
formed by sharing pairs of fast-moving energetic electrons, and therefore contain potential
energy.
When the C-H bonds are broken, some of the energy of hydrogen electrons powers the
formation of ATP – a compound which can readily hydrolyse to provide energy that powers
cellular activities. But the breakdown of C-H bonds occurs in a series of enzymatically-
catalysed reactions. Since lipids contain very many C-H bonds, they yield energy
about twice that of carbohydrates of same mass on hydrolysis.

WHAT IS THE FATE OF THE HIGH ENERGY ELECTRONS AND HYDROGEN

IONS RELEASED FROM BREAKDOWN OF C-H BONDS
A molecule cannot simply lose its electrons; they have to go someplace to prevent them from
combining with other molecules, which would be fatal! If oxygen is present, the electrons are
accepted by electron carrier molecules which pass them along the electron transport
chain until they get attached to oxygen, which becomes negatively charged, O2-. As the
electrons are transferred along the transport chain, energy is gradually extracted from them
to power ATP formation. To avoid PH becoming acidic, which would be fatal, hydrogen ions,
H+ combine with O2- to form neutral water.

WHAT IS ADENOSINE TRIPHOSPHATE (ATP) AND WHY IS IT CONSIDERED

TO BE AN ENERGY CARRIER?
ATP is a compound made up of a molecule of adenine – a nitrogenous base, a molecule of
ribose sugar, and three phosphate molecules.
ATP is an energy carrier because it stores chemical energy, which is released as free energy
on hydrolysis of the covalent phosphate to phosphate bonds. Hydrolysis of ATP to form
adenosine diphosphate (ADP) releases 30.6kJmol-1 of free energy, and further
hydrolysis of the terminal phosphate bond of ADP to form adenosine monophosphate
(AMP) yields another 30.6kJmol-1 of free energy, but hydrolysis of the phosphate-ribose
bond in AMP releases very little energy.
WHY ARE THE TWO TERMINAL PHOSPHATE BONDS OF ATP REFERRED TO
AS HIGH-ENERGY BONDS / ENERGY-RICH BONDS?
It is because their hydrolysis proceeds with the release of an unusually large amount of free
energy (about 7.3kcal/mol or 30.6kJ per mol from each phosphate bond)
(Suzan and Glenn Toole 1997: advanced human and social biology, student’s art notebook pg.66)

Triphosphate

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Note:
(1) Phosphorylation of AMP (addition of phosphate molecules to AMP) forms ADP,
while Phosphorylation of ADP yields ATP. (2) The addition of each phosphate molecule
requires 30.6kJ, and therefore energy released from any chemical reaction if less than
30.6kJ cannot be stored as ATP but is lost as heat. (3) High-energy bonds are symbolized
by the squiggle (~) i.e. solid curved line. (4) Potential energy increases whenever things
experiencing a repulsive force are pushed together such as adding the 3rd phosphate to an
ADP molecule. Potential energy also increases whenever things that attract each other are
pulled apart as in the separating of protons from the electrons.

HOW IS ATP FORMED IN CELLS?

1. Directly by substrate-level Phosphorylation i.e. direct transfer of a phosphate group
from high energy phosphorylated compounds to ADP. Examples of high energy phosphate
compounds include Phosphoenolpyruvate, 1, 3-Bisphosphoglycerate, acetyl
phosphate and phosphocreatine.
2. Indirectly by use of energy supplied by transmembrane proton concentration gradients
e.g. oxidative Phosphorylation I the mitochondria and photophosphorylation during
photosynthesis.

IS ATP THE ONLY HIGH-ENERGY COMPOUND IN CELLS?

No! There are other compounds with even higher energy than ATP, only that ATP provides
just the right amount of energy at the right time and can be moved to any place when need
arises. For example in muscles and nerve cells where ATP is continually hydrolysed at a rate
faster than respiration can provide due to high metabolic activity, phosphocreatine
provides the phosphate for regeneration of ATP from ADP. See the table below.

Standard Free Energies of Phosphate Hydrolysis

of Some Compounds of Biological Interest 1. ATP has an intermediate phosphate-group
Compound DG0’ (kJ/mol) transfer potential. Under standard conditions,
Phosphoenolpyruvate -61.9 the compounds above ATP in the table on the
1, 3-Bisphosphoglycerate -49.4 left can spontaneously transfer a phosphate
group to ADP to form ATP, which can in turn
Acetyl phosphate -43.1 spontaneously transfer a phosphate group to the
Phosphocreatine -43.1 appropriate groups to form the compounds
ATP (+ H2O Û ADP + Pi) -30.6 listed below it.
Glucose-1-phosphate -20.9 2. The negatives of the values listed in the table
Fructose-6-phosphate -13.8 are often referred to as phosphate group-
Glucose-6-phosphate -13.8 transfer potentials; compounds with large
-9.2 negative values readily transfer their phosphate
Glycerol-3-phosphate group to form compounds with small negative
values by first forming ATP

WHERE DOES THE ENERGY USED IN THE FORMATION OF ATP COME

FROM?
From reduction-oxidation reactions that occur in cells. Reduction is the addition of
electrons to compounds while Oxidation is the loss of electrons from compounds.
In addition to gaining electrons, reduced compounds also gain high potential energy and act
as electron donors, while oxidised compounds lose potential energy and act s electron
acceptors.

WHY IS ATP REFERRED TO AS “THE UNIVERSAL ENERGY CURRENCY”

It is because ATP’s structure as the energy supplier molecule is the same in all living
organisms.

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DO CELLS STORE ATP FOR A LONG TIME?
No! ATP is continually hydrolysed and regenerated. The metabolic half-life of an ATP
molecule varies from seconds to a few minutes depending on the cell type and its metabolic
activity. For instance, brain cells have only a few seconds’ supply of ATP – which partly
explains why brain tissue deteriorates rapidly if deprived of oxygen. However, muscle cells
can store phosphocreatine (PCr) that acts as a reservoir of phosphate groups that can be
used to produce ATP. This ATP/PCr store although small, is important in
providing instant energy e.g. during sprinting.

WHY IS SO MUCH POTENTIAL ENERGY RELEASED WHEN ATP IS

HYDROLYSED TO ADP AND INORGANIC PHOSPHATE (Pi)?
(1) The three phosphate molecules in ATP bear four negative charges that repel each other,
raising the potential energy of the electrons. Hydrolysis of the terminal phosphate bond
forms ADP and Pi that have reduced electrical repulsion in comparison with the repulsion in
ATP. (2) The negative charges on ADP and Pi are stablised by much more efficiently by
interactions with the partial positive charges on surrounding water molecules. For these and
other reasons, ADP and Pi have lower potential energy than does ATP. The difference in the
potential energy of the reactant and the products is released as heat, or visible light e.g.
fireflies and certain bacteria are able to bioluminesce as some of this chemical-bond energy
is released as visible light, or some other form of energy.

USES OF ENERGY OF ATP IN CELLS:

(1) Enables transport of materials (2) Enables movement of cilia or flagella and muscle
contraction (3) Allows active transport to be carried out (movement of substances against
concentration gradient) e.g. ion pumps (4) Drives endergonic reactions e.g. assembly of
amino acids into proteins, synthesis of polysaccharides from monosaccharides, and DNA
replication (5) Activates chemicals to become more reactive e.g. Phosphorylation of sugar
during Glycolysis (6) Enables formation of vesicles during secretion of cell products.

AN ELEMENTARY UNDERSTANDING OF THE ROLE OF ATP

Each ATP molecule used in the cell is like a rechargeable AAA battery of 1.5v. Each contains
just the right amount of energy to power a small radio. When the power has been drained,
the AAA battery can be recharged many times using a small amount of energy from
hydroelectric power, but the small radio cannot be plugged directly into the high voltage
hydroelectric power. Cells of organisms operate in much the same manner. When the cell’s
“batteries”, ATP, are drained while powering a job like muscle contraction, the discharged
“batteries”, ADP can be recharged back to full ATP power using (1) chemical-bond energy
released from cellular respiration (2) sunlight during photosynthesis in green plants. In cells
therefore, ATP provides just the right amount of energy at the right time and place, and is
constantly used and synthesised.

WHERE DOES RESPIRATION OCCUR IN CELLS?

Cell type Location of pathway in cell
All prokaryotic cells Infoldings of cell membrane (mesosomes) and in cytoplasm
All eukaryotic cells Cytoplasm and inner membranes of mitochondria

STAGES OF CELLULAR RESPIRATION

(1) Glycolysis (2) Krebs cycle (3) Electron transfer system
Each step will be discussed individually, but remember that each is part of the whole
process.
GLYCOLYSIS (glyco = carbohydrate; lys = splitting; sis = the process of)
Definition:
A series of enzymatically controlled reactions in the cytoplasm of cells during which one
molecule of a six-carbon sugar glucose, is split into two molecules of the three-carbon
compound Pyruvate, with a net out put of two ATP molecules.
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DESCRIPTION OF THE PROCESS OF GLYCOLYSIS
Description through reactions
Glucose A friendlier description for exams
(6-Carbon) Glycolysis starts with phosphorylation of glucose by
ATP ATP to form glucose-6-phosphate. This process (1)
chemically reactivates glucose (2) traps glucose in
the cell because the phosphate group bears a
ADP negative charge yet the cell membrane is
Glucose-6-phosphate impermeable to ions.
(6-Carbon) Glucose-6-phosphate isomerizes to form fructose-
6-phosphate to ease another Phosphorylation.

Fructose-6-phosphate is phosphorylated by ATP to

Fructose-6-phosphate form fructose-1, 6-bisphosphate (NOT fructose-1,
(6-Carbon) 6-diphosphate - the old name!)

ATP
ADP
Fructose-1, 6-bisphosphate splits at once into two
glyceraldehyde-3-phosphates (3-
Fructose-1, 6-bisphosphate phosphoglyceraldehyde/3-PGAL), each with three-
(6-Carbon) carbons.

2 [Glyceraldehyde-3-phosphate]
(3-Carbon)
Pi Each 3-PGAL is dehydrogenated by nicotinamide
2NAD +

2 NADH adenine dinucleotide (NAD+) to form reduced

nicotinamide adenine dinucleotides (NADH)
2 [1, 3-Bisphosphoglycerate] Each 3-PGAL molecule is phosphorylated by
phosphates present in the cytoplasm to form 1, 3-
(3-Carbon) bisphosphoglycerate, which later donates the
2ADP
2ATP phosphate to ADP to form ATP and 3-
phosphoglycerate, which has 3-carbons.

2 [3-Phosphoglycerate]
(3-Carbon)
Each 3-phosphoglycerate isomerizes to form 2-
phosphoglycerate,
2 [2-Phosphoglycerate]
(3-Carbon) Each 2-phosphoglycerate loses a water
molecule to form 3-phosphoenolpyruvate
2H2O (PEP).

2 [Phosphoenolpyruvate]/PEP
Each 3-phosphoenolpyruvate (PEP) loses a
(3-Carbon) phosphate to ADP to form ATP and pyruvate
2ADP which has three-carbons
2ATP
Pyruvate
(3-Carbon)

SIGNIFICANCE OF GLYCOLYSIS
Glycolysis forms (1) ATP which is used to power cell activities (2) NADH and Pyruvate which
may be further oxidized to generate additional ATP. However in oxygen deficiency, both
NADH and pyruvate undergo fermentation to regenerate NAD+.

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NOTE:
Glycolytic degradation of glucose to two molecules of pyruvate yields only about 5.2% of the
total energy that can be released from glucose by complete oxidation. The two molecules of
Pyruvate formed by glycolysis still contain most of the chemical potential energy of the
glucose molecule, energy that can be extracted by oxidative reactions in the citric acid cycle

THE FATE OF PYRUVATE, NADH AND ATP PRODUCED FROM GLYCOLYSIS

1. ATP:
It is hydrolysed to release energy to power the cell’s needs.

2. NADH:
Under aerobic conditions (in the presence of oxygen), NADH is converted into FADH2 which
is then shuttled into the mitochondria where it donates electrons to a series of electron
carriers until they reach the final oxidizing agent oxygen in a process called electron
transport system. During this process, the free energy of electron transport drives the
synthesis of ATP from ADP and NAD+ is regenerated such that it can participate in further
catalysis.
Under anaerobic conditions, NADH must be re-oxidised by other means in order to keep the
glycolytic pathway supplied with NAD+

3. PYRUVATE:
Under aerobic conditions, it is completely oxidised via the citric acid cycle to
carbondioxide and water. Under anaerobic conditions in the cytoplasm, pyruvate under
goes fermentation.

Types of fermentation
There are many types of fermentation, but the two common types are given below:
(a) Alcoholic fermentation: pyruvate is decarboxylated to yield carbondioxide which is
converted to a 2-carbon compound acetaldehyde. Acetaldehyde is then reduced by NADH
to ethanol and NAD+ also forms. NAD+ enables the continuation of glycolysis. Alcoholic
fermentation occurs in some bacteria and yeasts.

(b) Lactic acid fermentation: pyruvate is reduced directly by NADH to form lactic acid
as the end product. No carbondioxide is released. Lactic acid fermentation (1) is carried out
by certain fungi and bacteria during the formation of yoghurt and cheese (2) occurs during
oxygen scarcity in human skeletal muscle cells during sprinting. The lactic acid is gradually
carried away by blood to the liver and converted back to pyruvate by liver cells.
If ATP is abundant, pyruvate and lactate can be used as a substrate in the synthesis of
glucose.

Comparison of cellular respiration and fermentation

Similarities: Both (1) form ATP (2) use glycolysis to oxidise glucose to pyruvate (3) use
NAD+ as the oxidizing agent that accepts electrons from food during glycolysis (4) may be
carried out by same cells (e.g. muscle cells) or same organisms (e.g. yeasts and bacteria).

Differences:
Cellular respiration Fermentation
• Final electron acceptor from is oxygen • Final electron acceptor is an organic molecule
such as pyruvate (lactic acid fermentation) or
acetaldehyde (alcohol fermentation)
• Harvests much more energy from each glucose • Harvests much less energy from each glucose
molecule i.e. up to 38 ATP per glucose molecule. molecule i.e. 2 ATP per glucose molecule.
• Occurs in mitochondria. • Occurs in cytoplasm (cytosol).

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Evolutionary significance of glycolysis
The role of glycolysis in both fermentation and respiration suggests that ancient prokaryotes
probably used glycolysis to make ATP long before oxygen was present in the atmosphere.
This conclusion is based on the following observations: (1) The oldest bacterial
fossils date back 3.5 billion years, yet oxygen accumulated about 2.7 billion years ago.
Therefore early prokaryotes may have generated ATP exclusively from glycolysis, which does
not require oxygen. (2) Glycolysis is the most widespread metabolic pathway, which
suggests that it evolved very early in the history of life. (3) Glycolysis is located in the
cytoplasm where no membrane-bounded organelles are required in eukaryotic cells, which
evolved approximately 1 billion years after the prokaryotic cell.

FATE OF PYRUVATE IN AEROBIC CONDITIONS (TRANSITION STATE OF

PYRUVATE)
Each pyruvate molecule produced by glycolysis in the cell cytoplasm is transported across
the inner mitochondrial membrane by active transport (since it is a charged molecule)
into the matrix, where it is first decarboxylated and then oxidised (dehydrogenated) to
form a 2-C compound called acetate, carbondioxide and NADH. Carbondioxide, a waste
product is eventually excreted while NAD+ serves as a hydrogen carrier.
Finally, Acetate is attached to Coenzyme A to form acetyl coenzyme A, making the acetyl
group very reactive. Acetyl coenzyme A is now ready to feed its acetyl group into the citric
acid cycle for further oxidation. (A – stands for acetylation)

Note: the transition from pyruvate to acetyl coenzyme A is not usually considered as a
separate phase and is included with the first step of Krebs cycle.

THE ROLE OF CoA IN RESPIRATION

(1) Within the active centre of the enzyme citrate synthetase, CoA transfers the 2-carbon
acetyl group to a 4-carbon molecule of oxalocetate to make a molecule of citrate which
enters the Krebs cycle. (2) it serves as a link between many different pathways of
metabolism to provide a wide range of carbon compounds needed in the cell (3) during
energy deficiency, amino acids from proteins and fatty acids from lipids can be broken down
to provide acetyl CoA for use in respiration.

Acetyl- Coenzyme A: a central metabolic intermediate

All proteins, lipids, and carbohydrates must be converted to Acetyl- Coenzyme A prior to
participation in cellular respiration.
The fate of acetyl-CoA is dependent upon ATP needs. When ATP is prevalent, acetyl-CoA
serves as the basis for fatty acid synthesis, which forms the basis of your body's long-term
energy storage: triglycerides (i.e., fat). Acetyl-CoA is the starting point for anabolic
pathways that result in the synthesis of fatty acids.
Alternatively, acetyl-CoA may enter the Kreb's citric acid cycle.

KREBS CYCLE/ TRICARBOXYLIC ACID CYCLE / CITRIC ACID CYCLE

It is named:
1. Krebs cycle after the formulator Hans Krebs
2. Citric acid because citric acid is the first compound formed.
3. Tricarboxylic acid because citric acid which is the first compound formed has 3 carboxyl
(-COOH) groups
It is a multi-step reaction in the mitochondrial matrix during which an acetyl group is
completely oxidized to CO2 with the generation of ATP and reducing hydrogens in the form
of NADH and FADH2.

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1st Reaction: Prior to entering the Krebs Cycle, pyruvate must be converted into acetyl CoA
Acetyl CoA adds its 2-C acetyl group to a 4-C oxaloacetate to form a 6-C citrate molecule.
2nd reaction: citrate isomerizes to a more reactive isocitrate by both removal and addition
of one water molecule.
3rd reaction: isocitrate is decarboxylated (loses a carbondioxide) and then oxidized (loses
hydrogen to NAD+ to form NADH) to form a-ketoglutarate.
4th reaction: a-ketoglutarate loses a carbondioxide (is decarboxylated) and is oxidised
(loses hydrogen to NAD+ to form NADH) and attached to coenzyme A to form succinyl-CoA.
5th step: succinyl-CoA causes phosphorylation of ADP to ATP and the formation of
succinate.
6th reaction: a 4-C succinate loses two hydrogens to FAD (is dehydrogenated), forming
FADH2 and a 4-C fumarate.
7th reaction: fumarate is hydrated (a water molecule is added) and rearranged to form
malate.
8th reaction: finally, malate loses hydrogen to NAD+ to form NADH (is oxidised)
regenerating oxaloacetate.

Note:
(1) Carboxylic acids are represented in their ionized forms as –COO- because the ionized
forms prevail at the PH within the mitochondrion. E.g. citrate is the ionized form of citric
acid.
(2) The regeneration of oxalocetate makes the process a cycle
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(3) For each acetyl group that enters the cycle, 3 NAD+ are reduced to NADH (reactions 3,
4, and 8)
(4) Most of the ATP out put of respiration results from oxidative phosphorylation, when the
NADH and FADH2 produced by the citric acid cycle relay the electrons extracted from food
to the electron transport chain.

Comparison of Krebs cycle and glycolysis

Similarities: In both: (1) reducing hydrogens are accumulated in NADH (2) ATP is
generated (3) there is a reduction in number of carbon atoms of organic compounds(4)
pyruvate participates (5) both occur in living cells

Differences:
Glycolysis Krebs cycle
• The electron acceptor FAD is not involved • The electron acceptor FAD is involved
• Carbondioxide doesn’t form • Carbondioxide is liberated
• Occurs in cell cytoplasm • Occurs in mitochondrial matrix
• Doesn’t necessarily depend on oxygen • Depends on oxygen availability to occur

ELECTRON TRANSPORT SYSTEM AND CHEMIOSMOTIC THEORY

Electron transport system: A process whereby a series of electron carriers operate
together to transfer electrons from donors to any of several different terminal electron
acceptors to generate a transmembrane electrochemical gradient in the mitochondrion
What are the components of the electron transport chain?
Complex Name Prosthetic Groups
Complex I NADH Dehydrogenase FMN, 9 Iron-Sulphur (Fe-S) centres
Complex II Succinate-Coenzyme Q Reductase FAD, cyt b560, 3 Fe-S centers
Coenzyme Q (CoQ) (also called ubiquinone) cyt bH, cyt bL, cyt c1, Fe-S
Complex III Cytochrome bc1 complex Cytochrome b1 heme, b2 heme
Cytochrome c cyt c
Complex IV Cytochrome Oxidase cyt a, cyt a3, copper (CuA)and (CuB)

1. NADH dehydrogenase (complex I) 2. Succinate coenzyme Q reductase (complex II) 3.

Coenzyme Q (CoQ) (also called ubiquinone) 4. Cytochrome bc1 complex (complex III) 5.
Cytochrome c (Cyt c) 6. Cytochrome oxidase (complex IV)

Cytochromes: are proteins with heme prosthetic groups. Heme contains an iron atom
embedded in a porphyrin ring system. They absorb light at characteristic wavelengths.
Iron-sulfur centers (Fe-S): are prosthetic groups containing 2, 3, 4, or 8 iron atoms,
complexed to a combination of elemental and cysteine sulfur atoms.
NAD+ (Nicotinamide Adenine Dinucleotide): is a coenzyme containing the B-
vitamin, niacin. NAD+ accepts 2 e- and one H+ (a hydride) in going to the reduced state, as
NAD+ + 2 e- + H+ « NADH. It may also be written as: NAD+ + 2 e- + 2H+ « NADH + H+
NAD+ is a coenzyme, that reversibly binds to enzymes.

FAD (Flavin Adenine Dinucleotide): is derived from the vitamin riboflavin (B2). The
protein to which it is attached is termed a flavoprotein (FP). FAD normally accepts 2 e-
and 2 H+ in going to its reduced state: FAD + 2 e- + 2 H+ « FADH2
FAD is an electron-carrier coenzyme like NAD+. However, unlike NAD+, FAD always occurs
as a prosthetic group, tightly bound at the active site of an enzyme, never as a free carrier.

FMN (Flavin MonoNucleotide): is a prosthetic group of some flavoproteins. It is similar in

structure to FAD, but lacking the adenine nucleotide.

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LOCATION OF THE CONSTITUENTS OF THE ELECTRON TRANSPORT CHAIN
Most constituents of the respiratory chain are embedded in the inner mitochondrial
membrane. Coenzyme Q (Ubiquinone): is located within the lipid core of the inner
membrane. As its name suggests, is very widely distributed in nature. CoQ acts as a bridge
between enzyme complex 1 and 3 or between complex 2 and 3.Cytochrome c resides in the
intermembrane space (within the lumen of the cristae). It alternately binds to Complex III or
Complex IV during electron transfer.

H+

FAD FADH2

The production of ATP during electron transport involves two separate but connected
processes i.e. Chemiosmosis and oxidative phosphorylation

Description of the process of electron transport system

The electrons released during glycolysis and carried by NADH are converted to FADH2 in
order to shuttle them from the cytoplasm into the mitochondrial matrix.

In Complex I (also called NADH reductase), reduced nicotinamide adenine dinucleotide

(NADH) donates electrons to the coenzyme Flavin mononucleotide (FMN) which then
passes electrons to an iron-sulphur (Fe-S) protein and the electrons lose some energy.
NADH is oxidized to NAD+, while FMN and Fe3+ are reduced to FMNH2 and Fe2+
respectively. Each electron is transferred with a proton.
Electrons from the reduced Fe-S proteins are then passed to Coenzyme Q along with
protons. Coenzyme Q is thus reduced while the Fe-S proteins are oxidised back to Fe3+ state.

In complex II (succinate dehydrogenase), electrons from FADH2 are passed on to Fe-S

proteins then to Coenzyme Q which transfers them to complex III. FADH2 becomes
oxidised to FAD+. During this process, four protons (H+) are translocated across the inner
mitochondrial membrane, from the matrix to the intermembrane space. This creates a
proton gradient that will be later used to generate ATP through oxidative phosphorylation.
During oxidation of FADH2 complex I is bypassed because complex II has only enough
reducing potential to pass electrons to Coenzyme Q.

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Reduced coenzyme Q (CoQH2) transfers electrons to Complex III where they pass
through several cytochromes and Fe-S proteins and during the process Fe3+ is reduced to
Fe2+. The electrons lose additional energy and are passed on to cytochrome c which passes
electrons to Complex IV (cytochrome c oxidase), which finally transfers the electrons to
reduce molecular oxygen to form water. O2 + 4 H+ + 4 e 2 H2O. At the same time,
complex IV moves protons (H+) across the membrane into the intermembrane space,
producing a proton gradient.

As electrons lose energy in complex I, III and IV, additional protons are pumped into the
intermembrane space producing a proton gradient. Complex II (succinate
dehydrogenase) is not a proton pump. It only serves to funnel additional electrons into
coenzyme Q. Electron transfers involving Coenzyme Q and Cytochrome c do not release
enough free energy to pump any protons.
When the protons flow down the concentration gradient through the channels in the stalked
particles, ATP synthase enzymes are able to use the energy to generate ATP.

Note: If the oxygen supply is cut off, the electrons and hydrogen protons cease to flow
through the electron transport system. If this happens, the proton concentration gradient
will not be sufficient to power the synthesis of ATP. This is why we, and other species, are
not able to survive for long without oxygen!

Is the ETS a sequence?

No! The complexes move in the fluid membrane independently of one another, and
exchange electrons when they are in mutual proximity. Although textbooks show the ETS as
a physical sequence, the complexes and carriers are not locked in place.

CHEMIOSMOTIC COUPLING HYPOTHESIS AND OXIDATIVE

PHOSPHORYLATION
As proposed by Peter D. Mitchell, the chemiosmotic coupling hypothesis explains that the
electron transport chain and oxidative phosphorylation are coupled by a proton gradient
across the inner mitochondrial membrane.
The efflux of protons into the intermembrane space creates both a pH gradient and an
electrochemical gradient. This proton gradient is used by the ATP synthase complex to
make ATP via oxidative phosphorylation. The stalk component of ATP synthase
complex acts as an ion channel for return of protons back to mitochondrial matrix during
which the free energy produced during the generation of the oxidized forms of the electron
carriers (NAD+) is released and used to drive ATP synthesis, catalyzed by the head
component of the ATP synthase complex.

Definition of coupled reactions:

Reactions that occur in the same place, at the same time and in such a way that an energy
releasing-reaction can drive an energy requiring-reaction. Usually, the energy releasing
reaction is ATP breakdown. Below is a simplest representation of a coupled reaction.
ADP+P
ATP
C+D A+B
Coupling

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ACCOUNTING FOR THE ELECTRONS IN EUKARYOTIC ORGANISMS

Oxidation of NADH to NAD+ pumps 3 protons from the mitochondrial matrix into the
intermembrane space, which charges the electrochemical gradient with enough potential to
generate 3 ATP. Oxidation of FADH2 to FAD+ pumps 2 protons into the intermembrane
space, which charges the electrochemical gradient with enough potential to generate 2 ATP.
In some text books however, recent information suggests that 1NADH generates 2.5 ATP and
1FADH2 generates 1.5 ATP. The reasons for this are that not all of the energy stored in the
proton gradient is used to generate ATP. Some of the energy is used to power transport of
ions in and out of the mitochondria.

A total of 12 pairs of electrons and hydrogens are transported to the electron transport
system from glycolysis and Krebs cycle for each glucose molecule that enters the process:
• 4 pairs are carried by NADH and were generated during glycolysis in the cytoplasm, 8
pairs are carried as NADH and were generated within the mitochondrial matrix and 2
pairs are carried by FADH2 and were generated within the mitochondrial matrix.
• For each of the 8 NADHs generated within the mitochondrial matrix, enough energy is
released to produce 3 ATP molecules; therefore, 24 ATP molecules are released from these
electrons carried by NADH.
• The electrons carried by FADH2 are lower in energy, so during the oxidation-reduction
reactions, they release energy to produce only 8 ATP molecules.
• Therefore, a grand total of 32 ATP molecules are produced from hydrogen electrons that
enter the electron transport system.

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WHAT QUANTITY OF ATP IS GENERATED BY CHEMIOSMOSIS FROM ONE

MOLECULE OF GLUCOSE DURING THE ELECTRON TRANSPORT CHAIN?
The chemiosmotic model suggests that one ATP molecule is generated for each H+ pump
activated by the electron transport chain. Since the electrons from NADH activate three
pumps and those from FADH2 activate two, it would be expected that the numbers of ATP
molecules generated by each molecule of NADH and FADH2 are three and two
respectively. However, since the transportation of the two molecules of NADH produced
during Glycolysis into the mitochondrion requires two ATPs, the theoretical yield from
aerobic respiration = 36 molecules of ATP i.e. 4 (from substrate-level Phosphorylation) +
30 (3 from each of 10 molecules of NADH) + 4 (2 from each of 2 molecules of FADH2) – 2
(for transport of glycolytic NADH). The actual yield is less than 36 because (1) the inner
mitochondrial membrane allows some H+ to re-enter the matrix without passing though
ATP-generating channels (2) mitochondria often use the proton gradient generated by
chemiosmosis for purposes other than ATP synthesis e.g. transporting Pyruvate into the
matrix. As a result, the measured values of ATP generated are closer to 2.5 for each NADH
and 1.5 for each FADH2. The molecules of ATPs formed from one molecule of glucose = 30
i.e. 4 (from substrate-level Phosphorylation) + 25 (2.5 from each of 10 molecules of NADH)
+ 3 (2 from each of 2 molecules of FADH2) – 2 (for transport of glycolytic NADH)

SUMMARY OF THE RESPIRATORY STAGES OF ONE GLUCOSE MOLECULE

Molecules of ATP
Number of Number of
gained
Process Where it occurs Molecules of Molecules of
Net/
Reactants Products Theoretical Actual
Cytoplasm 1 Glucose 2 Pyruvate
(most organisms) 2 ATP 2 ADP + 2 Pi 04** 02
Glycolysis Glycosomes* 4 ADP + 4 Pi 4 ATP
(trypanosomes) 2 NAD+ + 2 H+ 2 NADH
2 H2O
Krebs cycle/ 2 Pyruvate 6 CO2
Tricarboxylic acid Mitochondrial matrix 2 ADP + 2 Pi 2 ATP 02 02
cycle / citric acid 8 NAD+ + 8 H+ 8 NADH
cycle 2FAD + 4 H 2 FADH2
Electron transfer 8NADH+24ADP+24Pi 8NAD++24ATP+16H
system & Inner mitochondrial 2FADH2+8ADP+8Pi 4FAD+8ATP+8H 34 30
chemiosmosis membrane 6 O2 + 24 H+ 12 H2O
(oxidative
Phosphorylation)

Glycosomes*: Are organized cytoplasmic organelles in trypanosomes, the parasitic

protozoans that cause African sleeping sickness

04**: Because two molecules of ATP are used to start the process and a total of four ATPs
are generated, each glucose molecule that undergoes Glycolysis produces a net yield of two
ATPs.

The purpose of the several steps in the electron transport chain:

1) to pass along 2H+ ions and 2e- to eventually react with oxygen;
2) to conserve energy by forming three ATP's; and
3) to regenerate the coenzymes back to their original form as oxidizing agents.

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Inhibitors of electron transport
Inhibitors Action
Cyanide and Carbon monoxide Block cytochrome oxidase enzyme in complex IV
Rotenone Blocks complex I. It’s a common rat poison
Antimycin Blocks electron transfer in complex III
Oligomycin Blocks the proton channel in ATP synthase

Inhibitors bind to the components of the electron transport chain and block electron
transfer. All components before the block are stuck in a reduced state and all components
after in an oxidised state. No electron transfer is possible and proton pumping stops. The
proton gradient is quickly run down and ATP synthesis stops. Inhibitors may also block the
proton channel of ATP synthase.

EFFICIENCY OF RESPIRATION
Not all the energy present in the high-energy hydrogen atoms is conserved as ATP. Part of
the energy is released as heat used for the maintenance of body temperature, but if it is in
excess then it can be dissipated to the external environment.
The efficiency of energy conserved in aerobic respiration, alcoholic fermentation and lactic
acid fermentation are thus as follows:

Aerobic respiration Alcoholic fermentation Lactic acid fermentation

A total of 38 molecules of Alcohol fermentation Lactic acid fermentation
ATP are formed while the releases 210KJ with the releases 150KJ with the
amount of energy released is formation of 2ATP. To form 1 formation of 2ATP. To form 1
2880KJ. To form 1 molecule molecule of ATP requires molecule of ATP requires
of ATP requires 30.6kj. Thus 30.6kj. Thus the amount of 30.6kj. Thus the amount of
the amount of energy used to energy used to form 2 energy used to form 2
form 38 molecules of ATP is molecules of ATP is equal to molecules of ATP is equal to
equal to 38 x 30.6 = 2 x 30.6 = 61.2KJ. 2 x 30.6 = 61.2KJ.
1162.8KJ.
Efficiency of energy Efficiency of energy Efficiency of energy
conserved= conserved = conserved =

(38ATP x 30.6KJ) x 100 (2ATP x 30.6KJ) x 100 (2ATP x 30.6KJ) x 100

2880 210 150
= 40.375 ≈ 40.4% = 29.1% = 40.8%
The remaining The remaining The remaining
1717.2KJ(59.6%) is released 148.8KJ(70.9%) is released 88.8KJ(59.2%) is released as
as heat as heat heat
However, considering that glucose on complete oxidation releases 2880KJ of energy, the
yield from anaerobic respiration is given by: (2ATP x 30.6KJ) x 100 = 2.1%
2880
Therefore, on a whole anaerobic respiration is 2% efficient.

ENERGY FROM LIPIDS, PROTEINS AND HEXOSES OTHER THAN GLUCOSE

(e.g. FRUCTOSE ,GALACTOSE AND MANNOSE)
Energy from lipids (fat and oil):
In the gut, the enzyme lipase catalyses the hydrolysis of lipids into fatty acids and glycerol
which enter the lacteal and finally gain entry into liver cells.
Glycerol is phosphorylated with ATP, dehydrogenated with NAD and converted to triose
phosphate (glyceldehyde-3-phosphate) which is fed into the glycolysis pathway. There is a
net yield of 19 molecules of ATP from the oxidation of triose phosphate and of the NADH
formed.

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The fatty acid component is progressively broken in the matrix of the mitochondria into
fragments of 2 carbons each which are converted to acetyl coenzyme A. This then enters the
Krebs cycle with subsequent release of energy.

Carbohydrates versus Fats in energy release

Aspect Explanation
Gram for gram, fats provide more energy than carbohydrates. The reason for this is
the amount of oxidation that takes place as these compounds are converted to
carbon dioxide and water. Carbon for carbon, fats require more oxidation to
become CO2 and H2O than do carbohydrates. Roughly, carbohydrates already have
one oxygen for every carbon atom, thus each carbon atom needs only one more
oxygen and each pair of hydrogen atoms needs one more oxygen. However, almost
every carbon atom in a fat molecule needs two oxygens instead of just one
Amount of
additional one, and each pair of hydrogen atoms still needs one more oxygen. So,
energy
just from counting the number of oxygens needed to be added, fats require about
released
half again as much oxygen for the same number of carbon atoms. Because of this,
the oxidation of fats takes longer, but it also gives off more energy.
When comparing gram to gram, instead of carbon to carbon, the effect is
exaggerated. When you weigh a carbohydrate, more oxygen is included in that
weight. When you weigh a fat, you get more carbon atoms per gram and therefore,
gram for gram, the fats will give even more energy (over twice as much) than will
the carbohydrates.
Carbohydrates enter into the oxidation process much more quickly and provide energy
Time spent more rapidly than fats. This is because fats go through several more steps than do
carbohydrates to become acetyl CoA and enter the citric acid cycle.

ENERGY FROM PROTEIN

The body resorts to protein as an energy source only during starvation.
Catalysed by the enzymes, protein is first hydrolysed to amino acids which are then
individually deaminated i.e. amino groups (-NH2) are removed and converted to ammonia,
urea or uric acid for excretion. The residual carbon compound (a keto acid) then enters the
respiratory pathway at a number of points depending on their number of carbon atoms. E.g.
5-carbon amino acids like glutamate are converted to α-ketoglutarate, 4-carbon amino acid
like aspartate are converted to oxaloacetate. Both α-ketoglutarate and oxaloacetate are Krebs
cycle intermediates. 3-carbon amino acids like alanine are first converted to pyruvate and
then acetyl coenzyme A. other amino acids with larger number of carbon atoms are
converted by transamination reactions into 3, 4 0r 5-carbon amino acids.

OTHER MONOSACCHARIDES ENTER THE GLYCOLYTIC PATHWAY AT

SEVERAL POINTS
In most organisms, hexoses other than glucose can undergo glycolysis after conversion to a
phosphorylated derivative.

1. FRUCTOSE: is present in free form in many fruits and is also formed by hydrolysis of
sucrose in the ileum of vertebrates. In the muscles and kidney fructose is phosphorylated to
fructose-6-phosphate by hexokinase enzyme while in the liver fructokinase enzyme catalyses
the phosphorylation of fructose to fructose-1-phosphate which then splits into
glyceraldehyde and dihydroxyacetone phosphate.
Dihydroxyacetone phosphate converts to glyceraldehyde 3-phosphate while glyceraldehyde
is phosphorylated by ATP to glyceraldehyde 3-phosphate. Thus both products of fructose 1-
phosphate hydrolysis enter the glycolytic pathway as glyceraldehyde 3-phosphate.

2. GALACTOSE: is a product of hydrolysis of the disaccharide lactose (milk sugar).

Galactose is first phosphorylated by ATP to galactose-1-phosphate and then converted to
glucose-1-phosphate through a series of reactions.
Galactosemia is a human genetic disease that results from disordered galactose
metabolism in which the overall conversion of galactose to glucose prevented.
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MANNOSE, which is released in the digestion of various polysaccharides and glycoproteins
of foods, can be phosphorylated at C-6 by hexokinase:
Mannose + ATP → mannose 6-phosphate + ADP
Mannose 6-phosphate then isomerizes to fructose 6-phosphate, an intermediate of
glycolysis.
Tissue respiration and its connections with the rest of metabolism

CONTROL OF RESPIRATION
Because the principle function of respiration is to produce ATP, it must be regulated so that
ATP is generated only when needed. This occurs in a number of ways:
1. At cellular level, the rate at which respiration occurs is regulated mainly by the energy
state of the cell (i.e. the ratio of ATP to ADP), acting via regulatory enzymes. High levels of
ATP (high energy level of the cell) inhibit the enzyme hexokinase that catalyses
phosphorylation of glucose at the start of glycolysis while low energy levels (high ADP levels)
stimulate hexokinase enzyme. Highly active cells utilize ATP very fast breaking it to ADP.
This has the effect of enhancing the rate of respiration. Such cells include liver cells,
striated muscle cells, spermatozoa and nerve cells. They are characterized by
presence of numerous mitochondria. Less active cells utilize ATP slowly and hence
respiration in them is slow e.g. fat cells.
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2. At the level of the whole organism, the respiratory rate is influenced by environmental
factors e.g. temperature, structural factors e.g. body size and physiological factors
such as level of activity, growth and dormancy.

Temperature: generally, very low temperature slows down respiration in both

homoiotherms and poikilotherms, although it can be observed that homoiotherms need
increased respiration rate to generate much heat for maintaining body temperature. In
poikilotherms temperature near to that of the body increases the respiration rate. This
partly explains why mosquitoes and tsetse flies are only found in the tropics where
environmental temperature is close to their optimal temperature. High temperature slows
down the respiration rate in homoiotherms. This explains why such animals tend to be
sluggish during hot weather. However, excessively high temperatures trigger increased
respiration rate and finally stop as a response by enzymes to temperature.

Body size: small organisms with a large surface area to volume ratio lose heat faster and
therefore respire faster than large organisms.
Level of activity: animals engaging in vigorous physical exercise require much energy and
so experience faster respiration rate e.g. sprinting, flying, etc
Growth: actively growing organisms e.g. young animals and germinating seeds respire
faster to generate much energy required to drive metabolic processes
Dormancy during extreme cold and hot seasons: respiration rate is always slow to
avoid depleting food reserves before the unfavourable season ends.

RESPIRATORY QUOTIENT (RQ)

It is the ratio of the volume of Carbondioxide produced to the volume of oxygen used in
respiration during the same period of time

RQ = volume of Carbondioxide given out

Volume of oxygen taken in

Importance of RQ:
(1) it can indicate the kind of substrate being respired (2) it can indicate whether the
respiration is aerobic or anaerobic.
RQ can be measured using a spirometer or respirometer.

RQ FOR HEXOSE SUGAR: like glucose, the equation for its complete oxidation is:

C6H12O6 + 6O2 6CO2 + 6H2O

Hence RQ is: 6CO2 = 1.0 (one)

6O2

R.Q FOR FATS: For a lipid like tripalmitin, the equation for its complete oxidation is:

RQ is: 102CO2 = 0.7 (less than one)

145O2
NB: the R.Qs for different fats will of course show slight variations because of differences in
molecular composition

R.Q FOR PROTEINS: no concrete value can be calculated since (1) they vary so much in
composition and (2) are difficult to separate in the pure state. Estimates for protein vary
between 0.5 and 0.8 for the complete oxidation of proteins.
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SUMMARY OF THE POSSIBLE INTERPRETATIONS OF R.Q VALUES:

Subject R.Q. Possible interpretations
Germinating starchy seeds 1.0 Complete oxidation of a carbohydrate
aves rich in carbohydrate 1.0 substrate
Wheat seedlings in nitrogen ∞ Anaerobic respiration
Germinating linseeds 0.64 Oxidation of a fatty substance
Slow entry oxygen causing some anaerobic
Germinating peas 3.0 to 4.0
respiration
Germinating peas (testa More rapid entry of oxygen, but some
1.5 to 2.5
removed) anaerobic respiration
Man (average) 0.8 to 0.85 Mixed fat and carbohydrates substrate
Lumbricus terrestris 0.75 Mainly fat substrate
Conversion some carbohydrate to fat :
Drosophila (at rest) 1.23
excess CO2 produce by decarboxylation
Drosophila (flying) 1.0 Complete oxidation carbohydrate
Nerve tissue (resting) 0.77 Possibly mainly fat substrate
Nerve tissue (active) 0.97 Almost entirely carbohydrate substrate

Measurement of R.Q. for small animals e.g. frog or a few earthworms

A, B, C, D and E are weighed. The animal(s) is/are placed in C and its weight determined.
Then the apparatus is connected, C is placed in a vessel of cold water to keep its temperature
constant, and the pump is run for 10 to 20 minutes. After the experiment, the weight of
Carbondioxide exhaled is the gain in the weight of E. the weight of water given off by the
animal is the gain in weight of D. the weight of substrate used is the loss in weight of the
animal. Hence, the weight of oxygen absorbed can be obtained thus:

Substrate + oxygen = Carbondioxide + water.

Oxygen used = (Carbondioxide + water) – substrate
= (gain in weight of E + gain in weight of D) – loss in weight of animal

The weights of Carbondioxide and oxygen thus obtained can be converted into
volumes, and thus the R.Q. obtained.

Precautions to be considered: (1) the soda-lime and sulphuric acid containers should
be doubled in each case (2) the animal container C must be carefully dried after the
experiment and before weighing (3) all vessels must not be touched by the hand but by
strong wooden forceps.

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COMPARISON OF RESPIRATION WITH PHOTOSYNTHESIS

Differences: Photosynthesis Respiration

Where they occur In chlorophyll-bearing cells In all cells
When they occur In the presence of light All the time
Reduced carbon compounds and
Input Carbon dioxide and water
oxygen
Reduced carbon compounds, oxygen, and
Output Carbon dioxide and water
water
Energy sources Light Chemical bonds
Energy result Energy stored Energy released
Chemical reaction Reduction of carbon compounds Oxidation of carbon compounds
Energy carrier(s) NADP NAD and FAD

Similarities
Both (1) involve converting energy from one form to another (2) occur in living cells (3)
involve the formation of ATP (4) require energy to occur (5) involve a series of multi-
enzyme catalysed reactions (6) involve flow of electrons along carriers.

ECONOMIC IMPORTANCE OF ANAEROBIC RESPIRATION

(1) Fermentation is applied in the manufacture of alcoholic drinks like wine making, beer
making and manufacture of spirits. (2) Fermentation of yeast is used in leavening of bread
i.e. production of raised bread (3) it is applied in the manufacture of milk products like sour
milk, yoghurt and cheese (4) is applied in the manufacture of organic acids e.g. citric acid,
oxalic acid and butyric acid all of which have several industrial applications especially in
food processing.

ATP PRODUCTION DURING EXERCISE

On average, a muscle contains only enough ATP to sustain about 15 seconds of intense
exercise. For muscle contractions to continue, massive amounts of ATP are required.
Depending on the level of and duration of activity, the muscles being exercised may produce
the ATP they need by cellular respiration or by fermentation.
Sustained periods of sub maximal activity like jogging are powered by aerobic respiration,
but in contrast short periods of intense activity like sprinting are powered by a combination
of aerobic and anaerobic respiration. “Anaerobic” here means a combination of
glycolysis and stored ATP/Phosphocreatine release.

The table below shows the relative contributions of anaerobic and aerobic respiration to
exercise during a work out.
Duration of maximal exercise
Seconds Minutes
10 30 60 2 4 10 30 60 120
Percent anaerobic 90 80 70 Percent anaerobic 50 35 15 5 2 1
Percent aerobic 10 20 30 Percent aerobic 50 65 85 95 98 99
[Quoted by Krogh David (2002): Biology; a guide to the natural world (2nd ed.), Prentice Hall; New Jersey,
adapted from Astrand, P. O., and Rodahl, K. (1977): textbook of work physiology; McGraw Hill, New York]

Observations:
(1) In the first minute, the energy supply from aerobic respiration is low but rapidly
increases while that of anaerobic respiration is high but rapidly decreases.
(2) In the second minute there is equal contribution to energy needs from both aerobic and
anaerobic respiration, followed by a rapid increase from aerobic respiration up to the 30th
minute and a gradual increase thereafter, while anaerobic respiration decreases rapidly up to
the 30th minute and gradually thereafter.

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Explanation:
(1) During the first minute, the small amounts of ATP and phosphocreatine stored in cells
provide instant energy. When glycolysis starts, it provides a proportionally smaller
contribution, and a smaller contribution yet comes from aerobic metabolism. These
differences reflect the time it takes for each of these systems to get going.
(2) As the duration of the exercise increases to the ATP/PCr reservoir reduce greatly while
the aerobic respiration now predominates.

NOTE:
Phosphocreatine (also called creatine phosphate), stores ~P bonds in nerve and
muscle cells.
Creatine Kinase catalyzes: phosphocreatine + ADP Û ATP + creatine
This is a reversible reaction, though the equilibrium constant slightly favors
phosphocreatine formation. Phosphocreatine is produced when ATP levels are high.
When ATP is depleted during exercise in muscle, phosphate is transferred from
phosphocreatine to ADP, to replenish ATP.
The figure below shows how the different energy sources are used at different stages in a
bicycle race.
ATP/phosphocreatine
100 Anaerobic respiration

Aerobic respiration
% of body’ s energy resources

80

60

40

20
used

0
Fast steady riding Hill climbing Final sprint
[Quoted by Krogh David (2002): Biology; a guide to the natural world (2nd ed.), Prentice Hall; New Jersey,
adapted from Kearney, J. T. (June 1996, p.54): Training the Olympic Athletes, scientific American]

Observations:
(1) During the fast steady riding, there is an overriding contribution of aerobic respiration,
but very little from ATP/phosphocreatine and anaerobic respiration.
(2) In the stretches of hill climbing, glycolysis predominates over ATP/phosphocreatine and
aerobic respiration in providing energy. The latter two make equal contribution, which
follows closely.
(3) In the final sprint, ATP/phosphocreatine takes the leading role in providing energy,
followed by aerobic respiration and leastly by anaerobic respiration. This is because during
the steady riding, ATP/phosphocreatine reservoirs replenished and therefore provides
energy fast to the cells.

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