Chapter 4
RESPIRATION
In Chapter 3, you have studied that light energy
is converted into chemical energy, that is stored
in the bonds of complex organic molecules of
carbohydrates like glucose and starch. The
breaking of the C-C bonds of such compounds
through oxidation releases a considerable
amount of energy for utilisation by plants. This
phenomenon of release of energy by oxidation
of various organic molecules, for cellular use, is
known as respiration. The compounds that
are oxidised during this process, are known as
respiratory substrates. The whole of energy
contained in respiratory substrates is not
released all at once. It is released slowly in a
stepwise series of reactions controlled by
enzymes. During the process of respiration,
oxygen is utilised, and carbon dioxide, water
and energy are released as products. This energy
is utilised in various energy-requiring processes
of the organisms, and the carbon skeleton
produced during respiration is used as
precursors for biosynthesis of other molecules
in the cell. The reaction that occurs in common
respiration of glucose can be summed up in the
following equation :
C6H12O6 + 6O2 −→ 6CO2 + 6H2O + Energy
(2870 kJ)
In this chapter, you will study different
types of respiration, respiratory quotient and
the mechanism of respiration.
4.1 TYPES OF RESPIRATION
Depending upon the availability of oxygen,
respiration is classified into two major types:
(i) Aerobic respiration
(ii) Anaerobic respiration
Aerobic Respiration : This process of
respiration, which leads to a complete oxidation
of organic substances in the presence of
oxygen, and releases CO2, water and a large
amount of energy present in the substrate.
This type of respiration is the most common in
higher organisms.
Anaerobic Respiration : This type of
respiration takes place in the complete absence
of oxygen. It generally occurs in lower
organisms, such as bacteria and fungi. It also
occurs in higher plants and animals under
certain conditions, particularly when O2 is
limiting. In anaerobic respiration, the
carbohydrate is incompletely oxidised into
some carbonic compounds, such as ethyl
alcohol or acetic acid or lactic acid and CO2,
and the amount of energy released is also much
less as compared to aerobic respiration. This
process can be shown by the following
equation :
C6H12O6 −→ 2C2H5OH + 2CO2 + Energy (247 kJ)
This process of oxidation in microbes is
known as fermentation, which is very much
similar to that of anaerobic respiration in the
case of higher plants.
4.2 RESPIRATORY QUOTIENT
As you know, during aerobic respiration, O2 is
consumed and CO2 is released. The ratio of the
volume of CO2 evolved to the volume of O2
consumed in respiration is called respiratory
quotient (RQ) or respiratory ratio.
Volume of CO2 evolved
RQ =
Volume of O2 consumed
 RESPIRATION
39

The respiratory quotient depends upon the 4.3 MECHANISM OF RESPIRATION

type of respiratory substrate used during
respiration. This is different for different In the process of respiration, the carbohydrates
substrates. are converted into pyruvic acid through a series
Carbohydrates : When carbohydrates are
used as substrate and are completely oxidised,
Glucose
the RQ will be 1, because equal amounts of
CO 2 and O 2 are evolved and consumed,
Cytosol
respectively, as shown in the equation below :
C6H12O6 + 6O2 −→ 6CO2 + 6H2O + Energy Glycolysis

O2 absent O2 present
(anaerobic) (aerobic respiration)
6CO2
RQ = = 1.0 Pyruvic acid
6O2 (C 2H5OH+CO2)
Fermentation
Fats : When fats are used in respiration, the TCA
RQ is less than 1. For example, it is explained ATP Cycle CO 2
below with tripalmitin (2 molecules) as a NADH H2O
substrate.
2(C51H98O6) + 145O2 −→ 102CO2 + 98H2O + Energy Mitochondria
Tripalmitin

102CO2 Fig. 4.1 The broad scheme of respiration

RQ = = 0.7
145O2

The RQ in this case is less than 1 because

fats contain less oxygen than the Glucose
carbohydrates. Therefore, they require
relatively greater amount of O2 for oxidation. Pyruvate
Organic acids : When organic acids, such as CO 2
oxalic acid and malic acid, serve as respiratory Acetyl coenzyme A
substrates, then RQ is more than one. It is (2C)
because organic acids contain more oxygen
than carbohydrates. Therefore, relatively less 6C
4C
amount of oxygen is required for their oxidation.
2(COOH)2 + O2 −→ 4CO2 + 2H2O + Energy Citric Acid Cycle
(Oxalic acid)
CO 2
5C
4CO2 CO 2
RQ = =4
1O2
Note that in anaerobic respiration, CO2 is
evolved but oxygen is not used. Therefore, RQ, Coenzyme A Coenzyme A
(reduced) (oxidised)
in such a case, will be infinite. For example, ATP
H 2O
Zymase O2
C6H12O6 → 2C2H5OH + 2CO2 + Energy
(Glucose)
2CO2
RQ = = Infinity (∝) Fig. 4.2 The steps of respiration at a glance
0O2
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40

of enzymatic reactions. These reactions, put

together, are known as glycolysis and take Glucose
place in the cytosol. The pyruvic acid thus ATP
(6C)
formed enters mitochondria, where O2 and the
ADP
necessary enzymes are available, and pyruvic Glucose-6-phosphate
acid is finally converted into CO2 and H2O. This (6C)
reaction series is known as Krebs cycle or
tricarboxylic acid (TCA) or citric acid cycle. Fructose-6-phosphate
(6C)
The general scheme of respiration is shown in ATP
Figures 4.1 and 4.2. ADP
Fructose1, 6-phosphate
Glycolysis : The term glycolysis has originated (6C)
from the Greek words, glycos for sugar, and
lysis for splitting. The scheme of glycolysis was Triose phosphate
Triose phosphate
given by Gustav Embden, Otto Meyerhof, and (Dihydroxy acetone
(3-phosphate glyceraldehyde)
phosphate)
J. Parnas, and is often referred to as the EMP (3C) NAD +
(3C)
pathway, after the abbreviation of their last
NADH
names. Glycolysis is the first stage in the
2 x Triose bisphosphate
breakdown of glucose and is common to all (1,3-bisphosphoglyceric acid)
organisms. In anaerobic organisms, it is the (3C)
only process in respiration. Glycolysis occurs ADP

in cytoplasm of the cell. In this process, glucose ATP

undergoes partial oxidation to form two 2 x Triose phosphate
molecules of pyruvic acid. In plants, this (3-bisphosphoglyceric acid)
(3C)
glucose is derived from sucrose, which is the
end product of photosynthetic carbon
reactions, or from storage carbohydrates. 3-phosphoglycerate
Sucrose is converted into glucose and fructose
by the enzyme invertase, and these two H2O
monosaccharides can readily enter the 2-phosphoglycerate
glycolytic pathway. The various steps of
glycolysis are depicted in Figure 4.3. HH
2O2O

Glucose and fructose are phosphorylated

PEP
to give rise to glucose-6-phosphate and ADP
ADP
fructose-6-phosphate, respectively, by the ATP
activity of the enzyme hexokinase. The
phosphorylated form of glucose then 2 x Pyruvic acid
isomerises to produce fructose-6-phosphate. (3C)
Subsequent steps of metabolism of glucose and
fructose are same. Fructose-6-phosphate
is phosphorylated and fructose-1,
Fig. 4.3 Major steps of glycolysis
6-bisphosphate produced by the action of the
enzyme phosphofructokinase, is split into two
molecules of triose phosphate, that and NAD+ is reduced to NADH. In the next
is, 3-phosphoglyceraldehyde and step of glycolysis, phosphoglycerate kinase
dihydroxyacetone phosphate, which are catalyses the formation of 3-phosphoglycerate
interconvertible.Once 3-phosphoglyceraldehyde from 1, 3-bisphosphoglycerate, generating
is formed, the glycolytic pathway enters the ATP in the process. This type of ATP generation,
energy conserving phase, and it is oxidised to whereby a phosphate group is directly
a carboxylic acid (1, 3-bisphosphoglycerate), transferred from substrate to ADP to form ATP,

is distinctly different from the ATP produced
by ATP synthase during oxidative
phosphorylation in mitochondria (described
later in this chapter) or photophosphorylation
in chloroplasts (see Chapter 3). Subsequently,
3-phosphoglycerate is successively
converted into 2-phosphoglycerate and
phosphoenolpyruvate (PEP). PEP is a good
donor for the formation of ATP. Using PEP as
substrate, the enzyme pyruvate kinase
catalyses the formation of pyruvate and
liberates ATP.
In the above pathway, the molecules of ATP
are produced in two ways:
(i) direct transfer of phosphate to ADP, and
(ii) oxidation of NADH produced during
glycolysis to NAD+.
Each molecule of NADH gives rise to three
molecules of ATP. In the glycolysis scheme
described, it is obvious that two triose
phosphate molecules are formed from one
glucose molecule, and 4 ATP molecules are
produced. Out of these 4 ATP molecules, 2
ATP molecules are consumed initially in
converting glucose to fructose-1,
6-bisphosphate. In addition, three molecules
of ATP are produced from the oxidation of each
of the two molecules of NADH produced during
catabolism of glucose. Therefore, a net gain of
8 ATP molecules occurs during glycolysis.
Fermentation
No doubt, glycolysis can also function without
O2, but further oxidation of pyruvic acid and
NADH by mitochondria requires oxygen.
Therefore, when O2 is limiting, NADH and
pyruvic acid begin to accumulate. Under this
condition, plants carry out fermentation
(anaerobic respiration), leading to the formation
of CO2 and either ethanol or lactic acid (usually
ethanol). During fermentation, the pyruvic acid
releases one molecule of CO 2 to produce
acetaldehyde. The acetaldehyde, then reoxidises
NADH, and is itself reduced to ethanol. These
reactions are catalysed by the enzymes, pyruvic
acid decarboxylase and alcohol dehydrogenase.
The steps involved are shown in Figure 4.4.
Aerobic Oxidation of Pyruvic Acid
Pyruvic acid, generated in the cytosol, is
transported to mitochondria, and thus initiates
RESPIRATION
41
Glycolysis Fermentation
Glucose process
Ethanol
Glyceraldehyde-3-P
ADP NAD +
ATP NADH+H+
Acetaldehyde
1,3 bisphosphoglycerate Pyruvate
CO 2
Fig. 4.4 Pathway of anaerobic respiration in
yeast (Fermentation)
the second phase of respiration. You have
already studied the structure of mitochondria
in Class XI. Before pyruvic acid enters Krebs
cycle, operative in the mitochondria, one of the
three carbon atoms of pyruvic acid is oxidised
to carbon dioxide in a reaction called oxidative
decarboxylation, that is, pyruvate is first
decarboxylated, and then oxidised by the
enzyme pyruvate dehydrogenase. The
combination of the remaining 2-carbon
acetate unit is readily accepted by a
sulphur-containing compound, coenzyme A
(CoA), to form acetyl-CoA. This is the
connecting link between glycolysis and Krebs
cycle. During the process, NAD+ is reduced to
NADH. The summary of the reaction is given
below :
+
NAD NADH
Pyruvic acid + CoA Acetyl CoA + CO2
2+
Mg
During this process, two molecules of
NADH are produced (from the metabolism of
two molecules of pyruvic acid produced during
glycolysis), and thus, it results in a net gain of
6 ATP molecules (2NADH × 3 = 6 ATP).
Citric acid cycle : This was elucidated by
the British biochemist, Hans Krebs, in 1937.
This is also known as tricarboxylic acid cycle
(TCA cycle) or Krebs cycle. The actual citric
acid cycle begins when acetyl-CoA enters into
a reaction to form citric acid. The elucidation
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42

SIR HANS ADOLF KREBS

(1900-1981)

Krebs, born in 1900, started his career as assistant to Warburg in Kaiser Wilhelm Institute
of Physiology, Berlin. He served the universities of Sheffield and Oxford as Professor of
biochemistry.
He established the central role of pyruvate in conversion of glucose hydrogens into
fumarate. He also discovered catalytic role of pyruvate. The citric acid cycle for production of
energy in the cell was described by him and is known with an alternative name, Tricarboxylic
Cycle (TCA) or Krebs Cycle. Krebs proposed this cycle based on the experimental findings in
1937, which showed that respiration was inhibited by malonate. He also demonstrated that
it is a cycle and not a collection of unrelated reactions. He shared in 1953 Nobel prize for
physiology and medicine with Fritz Lipmann mainly for discovery of TCA in living organisms.
This added to the basic understanding of cell metabolism. His publication, Energy
Transformation in Living Matter (1957), co-authored with Hans Kornberg, encouraged research
in this field.

of this cycle explained how pyruvate is broken

down to CO2 and H2O. It also highlighted the
Pyruvate
concept of cycles in metabolism. For this (3C)
pioneering work, Hans Krebs was awarded the
coveted Nobel Prize in 1953.
In the first reaction of citric acid cycle, one
molecule of acetyl CoA combines with Acetyl coenzyme A
4-carbon oxaloacetic acid (OAA), to form (2C)
6-carbon citric acid, and CoA is released. The
reaction is catalysed by the enzyme citrate
synthase. Citrate is then isomerised to Oxaloacetic acid
(4C) Citric acid
isocitrate. It is followed by two successive steps + (6C)
NADH+H CO 2
of decarboxylation, leading to the formation of NAD+
α-ketoglutaric acid and succinyl-CoA. In the NAD+
NADH+H+
remaining steps of citric acid cycle, succinyl-
α-ketoglutaric acid
CoA is oxidised to OAA (Fig. 4.5), allowing the Malic acid CITRIC ACID CYCLE (5C)
cycle to continue. (4C)
During this cycle, 3 molecules of NADH CO 2
and one molecule of FADH 2 are reduced NAD +
FADH 2
to produce NADH and FADH 2, respectively. NADH+H+
These reduced electron carriers pass on FAD
Succinic acid ADP
the hydrogen atoms to oxygen through (4C) ATP
electron transport system, yielding 11 more
ATP molecules for each molecule of
pyruvic acid. In addition, one more ATP
molecule is generated directly during the
cycle, to give a total of 12 ATP molecules per Fig. 4.5 The Citric acid cycle
 RESPIRATION
43

pyruvic acid molecule (3C). As two molecules

of pyruvic acid are produced from each
molecule of glucose (6C), a total of 24 molecules
of ATP are formed during the citric acid cycle.
NADH and FADH2 so produced during the
citric acid cycle are linked with the electron
transport system and produce ATP by
oxidative phosphorylation. The summary
equation for this phase of respiration may
therefore be written as follows :
Pyruvic acid + 4NAD+ + FAD + 2H2O + ADP + Pi
Mitochondrial
Matrix
3CO2 + 4NADH + 4H+ + FADH2 + ATP
Electron Transport System (ETS) and
Oxidative Phosphorylation
The glucose molecule is completely oxidised by
the end of the citric acid cycle. But the energy is
not released unless NADH and FADH 2 are
oxidised through the electron transport system.
At this stage, it is better to explain the meaning of
oxidation in terms of electrons. Here, oxidation
of a compound means removal of electrons from
it. This is usually accompanied by removal of
hydrogen. Reduction means addition of electrons
to a compound, usually accompanied by
addition of hydrogen. The metabolic pathway
through which the electron passes from one
carrier to another, is called the electron
transport system (ETS) (Fig. 4.6) and it is
operative in the inner mitochondrial membrane.
Electrons from NADH produced in the
mitochondrial matrix during citric acid cycle are
oxidised by an NADH dehydrogenase
(complex I), and electrons are then transferred Fig. 4.6 Electron transport system
to ubiquinone located within the inner
membrane. Ubiquinone also receives reducing complex containing cytochromes a and a3, and
equivalents via FADH2 that is generated during two copper centers.
oxidation of succinate, through the activity of the When the electrons pass from one carrier to
enzyme, succinate dehydrogenase (complex II), another via complex I to IV in the electron
in the citric acid cycle. The reduced ubiquinone transport chain, they are coupled to ATP
(ubiquinol) is then oxidised with the transfer of synthase (complex V) for the production of ATP
electrons to cytochrome c via cytochrome bc1 from ADP and inorganic phosphate. The
complex (complex III). Cytochrome c is a small number of ATP molecules synthesised depends
protein attached to outer surface of the inner on nature of the electron donor. Oxidation of one
membrane and acts as a mobile carrier for molecule of NADH gives rise to 3 molecules of
transfer of electrons between complex III and IV. ATP, while that of one molecule of FADH2
Complex IV refers to cytochrome c oxidase produces 2 molecules of ATP. During electron
 BIOLOGY
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transfer, the hydrogen atoms split into protons

and electrons. The electrons are carried by the
cytochromes. They recombine with their protons
before the final stage, when hydrogen atom is
accepted by oxygen to form water. Although the
aerobic process of respiration takes place only
in the presence of oxygen, the role of oxygen is
limited to the terminal stage of the process. Yet, H+
the presence of oxygen is vital, since it drives the
whole process by removing hydrogen from the
system. Oxygen acts as the final hydrogen
acceptor. The whole process by which oxygen
effectively allows the production of ATP by
phosphorylation of ADP, is called oxidative
phosphorylation.
The pathway of electron-transport system
is summarised in Figure 4.6. Fig. 4.7 ATP synthesis by inner membrane
As mentioned earlier, the energy released particles of mitochondrion
during the electron transport system is utilised
in synthesising ATP with the help of ATP Net gain of ATP : There is a net gain of 38
synthase (complex V). This complex consists ATP molecules during aerobic respiration of
of two major components, F1 and F0 (Fig. 4.7). one molecule of glucose. The details of ATP
The F1 headpiece is a peripheral membrane produced are given in the Table 4.1.
protein complex and contains the site for In most eukaryotic cells, 2 molecules of
synthesis of ATP from ADP and inorganic ATP are required for transporting the NADH
phosphate. F0 is an integral membrane protein produced in glycolysis into the mitochondrion
complex that forms the channel through which for further oxidation. Hence, the net gain of
protons cross the inner membrane. The ATP is 36 molecules.
passage of protons through the channel is
Significance of Citric Acid Cycle
coupled to the catalytic site of the F 1
component for the production of ATP. For each (i) During this pathway, carbon skeletons are
ATP produced, 2H+ pass through F0 from the obtained for use in growth and
intermembrane space to the matrix down the maintenance of the cell. Many intermediate
electrochemical proton gradient. compounds are formed, which are used in

Table 4.1 : ATP Molecules Produced during Respiration

Stage of Respiration Source Number of ATP

Molecules Produced
Glycolysis Direct 2
2-molecules of NADH 6
(one molecule of NADH
yields 3 molecules of ATP)
Pyruvic acid to acetyl-CoA 2 molecules of NADH 6
Citric acid cycle 6 NADH 18
2FADH2(FADH produces 4
only 2 molecules of ATP)
Direct 2
Total yield of ATP molecules 38
 RESPIRATION
45

the synthesis of other biomolecules like
amino acids, nucleotides, chlorophyll,
cytochromes and fats. For example,
succinyl CoA is the starting molecule for
synthesis of chlorophyll; amino acids are
synthesised from α-ketoglutaric acid,
pyruvic acid, and oxaloacetic acid.
(ii) This is the major pathway for generation
of ATP molecules, the energy currency of
the cell.
How Efficient is the System?
The total energy yield from 38 ATP molecules
comes to 1292 kJ (one ATP molecule yields
34 kJ of energy). Energy released by one
molecule of glucose on complete oxidation
corresponds to 2870 kJ. Thus, the efficiency
is 45 per cent. It shows that only a part of this
energy is used to make ATP, and much of the
energy generated during respiration is released
in the form of heat.
4.4 PENTOSE PATHWAY
You have seen that glucose is broken down into
CO2 and water during aerobic respiration. This
is the principal pathway of respiration. But
sometimes, oxidation of glucose takes place by
another pathway, which is called pentose
phosphate pathway (PPP).
In pentose pathway, glucose-6-phosphate
(6C) produced during the early stages of
glycolysis, or the photosynthates produced
during photosynthesis, are oxidised to give rise
to 6-phosphogluconate. This reaction takes place
in the presence of the enzyme, glucose-
6-phosphate dehydrogenase, and generates
NADPH. The 6-phosphogluconate molecule is
further oxidised by the enzyme,
6-phosphogluconate dehydrogenase. As a result
of this, one molecule each of ribulose-5-
phosphate, carbon dioxide and NADPH are
produced. Ribulose-5-phosphate undergoes
many changes to produce glycolytic intermediate,
such as glyceraldehyde-3-phosphate and
fructose-6-phosphate. The various reactions of
this cycle take place in the cell cytoplasm.
4.5 COMPENSATION POINT
At given low concentration of CO2 and non-
limiting light intensity, the photosynthetic rate
of a given plant will be equal to the total amount
of respiration (true respiration plus
photorespiration). The atmospheric
concentration of CO2 at which photosynthesis
just compensates for respiration, is referred to
as the CO 2 compensation point. The CO 2
compensation point is reached when the
amount of CO2 uptake is equal to that generated
through respiration at a non-limiting light
intensity. Net photosynthesis under these
conditions is zero. In C 3 plants, the CO 2
compensation point is usually much higher (25
to 100 µl.L–1) than in C4 plants (less than 5 µl.L–1).

SUMMARY

Respiration is a metabolic process in which carbon compounds generated

during photosynthesis are oxidised, in the presence of various enzymes, into
carbon dioxide and water with the release of energy. The ratio of CO2 evolved
to the volume of O2 consumed during the oxidation of carbon compounds is
called respiratory quotient. The respiratory quotient depends upon the
respiratory substrate being used in respiration. Respiration is basically of
two types : (i) aerobic respiration that occurs in the presence of oxygen, and
(ii) anaerobic respiration that does not require O2.
Anaerobic respiration takes place in the absence of oxygen. It occurs in the
cytoplasm of the cell. The end products are ethanol or lactic acid, and very little
energy is released. The initial steps, constituting glycolysis, are common to both
 BIOLOGY
46

anaerobic and aerobic respiration. During glycolysis, glucose is converted into

pyruvic acid. There is a net gain of 8 ATP molecules during this process.
Aerobic respiration occurs in the mitochondria. In the mitochondria, the
pyruvic acid is oxidised to acetyl-CoA, which then enters the TCA or citric acid
cycle where it is oxidised further. During this process, CO2, and NADH and
FADH2 are produced and take part in the electron transport system (ETS) that
carries the electrons in a chain to molecular oxygen, ultimately producing
water. The energy released during ETS is utilised to synthesise ATP. This
process is termed as oxidative phosphorylation. The energy trapped in ATP is
utilised in various metabolic processes by the living organisms. Most organisms
contain an alternate route also for glucose metabolism, called the oxidative
pentose phosphate pathway. This pathway also operates in cytosol of the cell
and produces pentose sugars and NADPH.

EXERCISES

1. Define the following :

(a) Respiration
(b) Respiratory substrate
(c) Respiratory quotient
(d) Anaerobic respiration
(e) Aerobic respiration
(f) Fermentation
2. Fill in the blanks with suitable words :
(a) Pyruvic acid is oxidised into _______ before entering citric acid cycle.
(b) The RQ is _________ if respiratory substrate is oxalic acid.
(c) Glycolysis takes place in __________.
(d) F0-F1 complex participate in the synthesis of _________.
(e) The Acetyl CoA is accepted by ________ in the citric acid cycle.
3. Distinguish between the following :
(a) Aerobic respiration and anaerobic respiration
(b) Glycolysis and fermentation
(c) Glycolysis and citric acid cycle
4. Why does anaerobic respiration produce less energy than aerobic
respiration?
5. What is oxidative phosphorylation?
6. Describe the process and role of citric acid cycle in living organism.
7. Describe the pentose phosphate pathway.
8. Calculate the efficiency of respiration in the living system.
9. Write the significance of citric acid cycle.
10. Illustrate the mechanism of electron transport system.