CHAPTER

anand_mani16 Dr. Anand Mani https://www.anandmani.com https://discord.io/anandmani t.me/anandmani001

 • Oxidation of food materials (breaking of C-C bonds of complex
molecules) within the cell to release energy for ATP synthesis
is called cellular respiration.
• This energy is used for absorption, transport, movement,
reproduction, breathing etc.
• Ultimate source of food that is respired is photosynthesis.
• The compounds that are oxidized during respiration are called
respiratory substrates. E.g. Carbohydrates (most common),
proteins, fats and organic acids.
• The energy released is not used directly but is used to
synthesize ATP. When energy is needed, ATP is broken down.
Hence, ATP acts as energy currency of the cell.

• For respiration, plants get O2 and give out CO2.

• In plants, gas exchange occurs via stomata & lenticels.
• Plants need no specialized respiratory organs because Each plant part takes care of its own gas-exchange needs.
So gas transport is very limited. Very low gas exchange as compared to that of animals. Leaves are adapted for
maximum gas exchange during photosynthesis. During this, O2 is released within the cell. Most living cells have
contact with air. They are located close to plant surface. In stems, living cells are organized in thin layers beneath
the bark. They also have lenticels. In leaves, stems & roots, parenchyma cells are loosely packed that provides
interconnected air spaces.
• Complete combustion of glucose yields energy most of which is given out as heat.

6H12O6 + 6O2 → 6CO2 + 6H2O + Energy

• This energy is utilized to synthesize other molecules.
• During the glucose catabolism, not all the liberated energy goes out as heat. Glucose is oxidized in several small
steps. It enables some steps to couple released energy to ATP synthesis.
• During respiration, oxygen is utilized, and CO2, water & energy are released.
• Certain organisms are adapted to anaerobic conditions. Some are facultative anaerobes. Others are obligate.

• It is the partial oxidation (breakdown) of glucose to 2 molecules of

pyruvic acid (C3H4O3) in the absence of O2.
• It occurs in cytoplasm of all living organisms.
• Its scheme was given by Gustav Embden, Otto Meyerhof & J. Parnas.
So it is also known as EMP pathway.
• In anaerobes, it is the only process in respiration.
• In plants, glucose is derived from sucrose (end product of
photosynthesis) or from storage carbohydrates. Sucrose is converted
into glucose & fructose by an enzyme, invertase. .
These 2 monosaccharides readily enter glycolytic pathway.
• Glucose & fructose are phosphorylated to form glucose-6-phosphate
by the enzyme hexokinase. It is then isomerized to produce
fructose-6-phosphate. Subsequent steps of metabolism of glucose
and fructose are same.

• It includes 10 steps under the control of different enzymes.

• ATP is utilized at 2 steps:
• In the conversion of glucose into glucose 6-phosphate. In the
conversion of fructose 6-phosphate to fructose 1, 6-diphosphate.
• Fructose 1, 6-diphosphate is split into dihydroxyacetone phosphate
and 3-phosphoglyceraldehyde (PGAL).

anand_mani16 Dr. Anand Mani https://www.anandmani.com https://discord.io/anandmani t.me/anandmani001

 - PGAL is oxidized and with inorganic phosphate get converted
to 1, 3-bisphosphoglycerate (DPGA). During this, 2 redox-
equivalents (2 H-atoms) are removed from PGAL and
transferred to NAD+ forming NADH + H+ .
- DPGA becomes 3-phosphoglyceric acid (PGA) yielding
energy. This energy is trapped by the formation of ATP.
- ATP is also formed when PEP converts to pyruvic acid.
- In glycolysis, 4 ATP molecules are directly synthesized
from one glucose molecule. In different cells, pyruvic acid
is handled in 3 ways:
Lactic acid fermentation.
Alcoholic fermentation.
Aerobic respiration (Krebs’ cycle).

- It is the incomplete oxidation of glucose under anaerobic condition.

- It occurs in many prokaryotes and unicellular eukaryotes.
- It is 2 types:
Alcoholic fermentation: Here, the pyruvic acid formed from glucose is
converted to CO2 and ethanol. The enzymes, pyruvic acid decarboxylase
and alcohol dehydrogenase catalyze these reactions. E.g. Yeast.
Yeasts poison themselves to death when the concentration of alcohol
reaches about 13%.
Lactic acid fermentation: Here, pyruvic acid is converted to lactic acid.
E.g. Some bacteria.
- The reducing agent (NADH+H+) is reoxidized to NAD+ in both the
processes.
- In animals, when oxygen is inadequate during exercise, pyruvic acid
in muscle cells is reduced to lactic acid by lactate dehydrogenase.
- Net ATP production from fermentation of one glucose
molecule = 2. (4 ATP from glycolysis – 2 ATP utilized).
- The steps involved in fermentation are shown below:

Energy production is limited. Less than 7% of the energy in glucose is released

and not all of it is trapped as high energy bonds of ATP.
Hazardous products (acid or alcohol) are formed

- It is a complete oxidation of organic substances in the presence of oxygen releasing CO2, water &energy.
- It occurs in mitochondria.
- For this, the pyruvate (final product of glycolysis) is transported from the cytoplasm into the mitochondria.
- The crucial events in aerobic respiration are:
Complete oxidation of pyruvate by stepwise removal of all the hydrogen atoms, leaving 3 CO2 molecules. It takes
place in the matrix of mitochondria.
Passing on of electrons removed as part of H-atoms to molecular O2 with simultaneous synthesis of ATP. It
occurs on the inner membrane of mitochondria.
- Pyruvate (pyruvic acid) enters mitochondrial matrix and undergoes oxidative decarboxylation in presence pyruvic
dehydrogenase. It needs coenzymes, NAD+ & Coenzyme A.
- During this process, 2 NADH molecules are produced from 2 pyruvic acid molecules.
- During this process, 2 NADH molecules are produced from 2 pyruvic acid molecules.

- Acetyl CoA then enters tricarboxylic acid (TCA) cycle.

anand_mani16 Dr. Anand Mani https://www.anandmani.com https://discord.io/anandmani t.me/anandmani001

 TCA cycle was first elucidated by Hans Krebs.
Steps:
1. Condensation of acetyl group with oxaloacetic acid (OAA)
& water to form citric acid in presence of citrate synthase
enzyme. A CoA molecule is released.
2. Citrate is isomerized to isocitrate.
3. Decarboxylation of isocitrate to a -ketoglutaric acid.
4. Decarboxylation of a-ketoglutaric acid to succinyl-CoA.
5. Succinyl-CoA is converted to succinic acid and a GTP
molecule is synthesized (substrate level phosphorylation).
In a coupled reaction, GTP is converted to GDP with
simultaneous synthesis of ATP from ADP.
6. Oxidation of succinate to Fumarate and then to Malate.
7. Oxidation of malate to OAA.
• At 3 points of TCA cycle, NAD+ is reduced to NADH + H+. At
one point, FAD+ is reduced to FADH2.
• Continued oxidation of acetic acid via TCA cycle requires
continued replenishment of OAA. It also requires
regeneration of NAD+ & FAD+ from NADH & FADH2.
6. Summary equation for this phase is given aside
Thus, a glucose is broken down to give 6 CO2, 8 NADH + H+, 2
FADH2 and 2ATP.

- Electron transport system (ETS) is the metabolic pathway present

in the inner mitochondrial membrane through which electron passes
from one carrier to another.
- This is to release and utilize energy stored in NADH + H+ 2 (formed
during TCA cycle) by oxidation.
- The electrons are passed on to O2 to form H2O.
- Electrons from NADH are oxidized by an NADH ehydrogenase (complex I).
- Electrons are then transferred to ubiquinone (Q) located within the
inner membrane. Ubiquinone also receives reducing equivalents via
FADH2 (complex II) that is generated during oxidation of succinate in
citric acid cycle.
- The reduced ubiquinone (ubiquinol or QH2) is then oxidized with the
transfer of electrons to cytochrome c via cytochrome bc1 complex
(complex III). Cytochrome c is a small protein attached to the outer
surface of the inner membrane. It acts as a mobile carrier of electrons
between complex III and IV.
- Complex IV refers to cytochrome c oxidase complex containing
cytochromes a & a3, and 2 copper centers.
- When the electrons pass from one carrier to another via complex I to
IV, they are coupled to ATP synthase (complex V) for the ATP
production.
- In aerobic respiration, the role of oxygen is limited to the terminal stage. Number of ATP
Yet, oxygen is vital since it drives the whole process by removing molecules produced
hydrogen from the system. Oxygen acts as the final hydrogen acceptor depends on nature of
electron donor.
- In respiration, energy of oxidation-reduction is utilized for the
phosphorylation. So this process is called oxidative phosphorylation. It is Oxidation of
not as photophosphorylation (Here, light energy is utilized for the 1 NADH→ 3 ATP
production of proton gradient for phosphorylation). 1 FADH2→ 2 ATP

anand_mani16 Dr. Anand Mani https://www.anandmani.com https://discord.io/anandmani t.me/anandmani001

 - The energy released during the ETS is utilized to synthesize ATP by
ATP synthase (complex V).
- ATP synthase has two major components: F1 & F0.
- F 1headpiece (peripheral membrane protein complex): Site for ATP
synthesis from ADP & inorganic phosphate.
- F0 (integral membrane protein complex): It forms a channel through
which protons cross the inner membrane. The passage of protons is
coupled to the catalytic site of the F1 component for ATP production.
- For each ATP produced, 2H+ passes through F0 from the inter-
Diagrammatic presentation of ATP
membrane space to the matrix down the electrochemical proton synthesis in mitochondria
gradient.

- Net gain of ATP from each glucose molecule is calculated based on the following assumptions.
All steps in Glycolysis, TCA cycle & ETS occur sequentially and orderly.
The NADH synthesized in glycolysis is transferred into mitochondria and undergoes oxidative phosphorylation.
Intermediates in the pathway are not used to synthesize other compounds.
Only glucose is being respired. Other alternative substrates are not entered in the pathway at any stages.
- Such assumptions are not valid because,
o All pathways work simultaneously and do not take place one after another.
o Substrates enter the pathways and are withdrawn from it as and when necessary.
o ATP is utilized as and when needed.
o Enzymatic rates are controlled by multiple means.
- Such calculations are useful to appreciate the efficiency of the living system in extraction and storing energy.

Net gain of ATP molecules from one glucose molecule

2 ATP directly 2 ATP

Glycolysis 2 molecules of NADH 6 ATP

Oxidative 2 NADH 6 ATP

decarboxylation
6 NADH 18 ATP
2 FADH 4 ATP
TCA cycle
2 GTP 2 ATP
Total 38 ATP

2 ATP molecules are spent for transporting 2 NADH molecules formed

during glycolysis to the mitochondria. Hence the net gain = 36 ATP molecules.
Comparison b/w fermentation & aerobic respiration

Fermentation Aerobic respiration

Partial breakdown of Complete breakdown of
glucose. glucose to CO2 & H2O.

Net gain of only 2 ATP. Net gain of 36 ATP.

NADH is oxidised to NADH is oxidised to NAD+ very

NAD+ rather slowly. vigorously.

anand_mani16 Dr. Anand Mani https://www.anandmani.com https://discord.io/anandmani t.me/anandmani001

 Glucose is the favored substrate for respiration. So, all
carbohydrates are first converted to glucose for
respiration.
- Other substrates are also respired.
- Fats breakdown into glycerol & fatty acids. Fatty acids
are degraded to acetyl CoA and enter the pathway.
Diagrammatic presentation of ATP
Glycerol is converted to PGAL and enters the pathway. synthesis in mitochondria
- Proteins are degraded by proteases into amino acids.
Each amino acid (after deamination) enters the
pathway at some stage in the Krebs’ cycle or as
pyruvate or acetyl CoA.
- The respiratory pathway is generally considered as a
catabolic pathway. But it involves both anabolism
(synthesis) and catabolism (breakdown). So it is better
called as an amphibolic pathway.
E.g. Fatty acids breakdown to acetyl CoA before entering
the respiratory pathway. But when the organism needs
to synthesize fatty acids, acetyl CoA withdraw from the
respiratory pathway.
Similarly, during breakdown and synthesis of protein,
respiratory intermediates are involved.

- It is the ratio of the volume of CO2 evolved to the volume of O2 consumed in respiration.
RQ = Volume of CO2 consumed
Volume of O2 consumed

- RQ depends upon the type of respiratory substrate.

- RQ for carbohydrates= 1, because equal amounts of CO2 and O2 are evolved and consumed, respectively.
C6H12 O6 + 6O2 → 6CO2 + 6 H2O + energy

RQ = 6 CO2 = 1.0
6 O2
- RQ for fats = < 1. Calculations for a fatty acid, (e.g. tripalmitin) are shown:
2 (C51H98 O6) + 145O2 → 102 CO2 + 98 H2O + energy

RQ = 102 CO2 = 0.7

145 O2
- RQ for proteins = 0.9.
- In living organisms, respiratory substances are often more than one. Pure proteins or fats are never used as
respiratory substrates.

anand_mani16 Dr. Anand Mani https://www.anandmani.com https://discord.io/anandmani t.me/anandmani001