All chemical reaction in a cell

Nutrient – Carbohydrate ( Glycosidic Bond )

Protein (Peptide)
Lipid (Ester)

Removal of CO2 from amino acid to form amine

Removal of amino group in a nucleotide base
Hydrolysis of glycosidic bond

The organic compounds taking part in metabolism is called metabolites. They are of two types:-

Primary Metabolites

They have identifiable functions in psychological processes and necessary for life.
Example amino acids, nucleic acids, sugars, lipids, vitamins, etc.

Secondary Metabolites

They are not directly involved in the normal growth, development or reproduction of organism.
Example essential oils, toxins, pigments, lectins, drugs, etc.

Metabolic pathways are similar to the automobile traffic in a city. Flow of metabolites through the
metabolic pathways has a definite rate and direction like automobile traffic. This metabolic flow is
called the dynamic state of body constituents.
There are two types of metabolic pathways:
Anabolic (biosynthetic) pathways and Catabolic pathways

Anabolic (Biosynthetic) pathways Catabolic pathways

Simpler molecules form complex structures Complex molecules become simple structures
(constructive process) (destructive process)
It consumes energy It releases energy
Example: formation of acetic acid from Example: formation of lactic acid from glycose
cholesterol, assembly of amino acids to protein, (glycolysis) respiration
photosynthesis

The energy released through catabolism is stored in the form of chemical bond. When needed, this
bond energy is utilised for biosynthetic, osmotic & mechanical works.

The most important energy currency in the living system is the bond energy in adenosine triphosphate
(ATP)
 Enzymes are biological catalysts which influence the speed of biochemical reactions OR They
are biological catalyst that speed up the rate of reaction without being involved
All enzymes are proteins but all proteins are not enzymes
The chemicals on which the enzyme acts are called substrate
Enzyme converts substrates into products
Enzymes are specific, i.e., each enzyme has its own substrate
Enzymes get damaged at high temperature
Enzymes lower the activation energy of the reaction
Enzymes isolated from thermophilic organism (live under high temperatures) are thermostable
Enzymes form tertiary structure (3D) with some crevices (pockets) called ‘active site’ into
which the substrate fits

Nucleic acids (RNA) that behave like enzymes are known as Ribozymes
Carbonic anhydrase is the fastest enzyme. It accelerates the following reaction 10 million times

In the absence of the enzyme, only 200 molecules of 𝐻2 𝐶𝑂3 are formed in an hour. In presence of
carbonic anhydrase, about 600,000 molecules are formed per second!

In a multistep chemical reaction (metabolic pathway), each step is catalysed by different enzymes

Example: In glycolysis {Glucose (𝐶6 𝐻12 𝑂6 ) → 2 pyruvic acid 𝐶3 𝐻4 𝑂3 )} 10 difference enzymes take
part

In a chemical reaction, there will be breaking of bonds.

𝐵𝑎(𝑂𝐻)2 + 𝐻2 𝑆𝑂4 → 𝐵𝑎𝑆𝑂4 + 2𝐻2 𝑂

(inorganic chemical reaction)

𝑆𝑡𝑟𝑎𝑐ℎ + 𝐻2 𝑂 → Glucose
(organic chemical reaction)

Rate of reaction = amount of product formed / unit time

Growth and reproduction of all organisms
depend on the division & enlargement of cells.

The mechanisms of division and multiplication

of cells together constitute cell reproduction.

Cell Division in Cancer Cell

It is the life period of a cell during which a cell

synthesizes DNA (replication), grows in size and
divides into two daughter cells.
Cell growth (cytoplasmic increase) is a continuous
process but DNA synthesis occurs only at a specific
stage.
Duration of cell cycle varies in each organism and
each cell type.
Duration of a typical eukaryotic cell cycle (e.g. human
cell) is about 24 hrs. In Yeasts, it is only about 90
minutes.

Cell cycle has 2 basic phases:

Interphase
M Phase
It is the phase between two successive M
phases.
It includes cell growth & DNA synthesis.
It lasts more than 95% of the duration of cell
cycle.
Interphase has 3 phases:
G1 phase (Gap 1 or Antephase)
S (Synthetic) phase
G2 (Gap 2) phase

First growth phase. It is the interval between mitosis

and DNA replication.
Main events:
Continuous growth of cell.
Cell becomes metabolically active.
Prepares machinery for DNA replication.
Synthesizes RNA and proteins.

It is the longest phase.

DNA replication takes place.
Amount of DNA per cell doubles.
But, chromosome
number is not increased.
In animal cells,
replication begins in the
nucleus, and the centriole
duplicates in the
cytoplasm.
Second growth phase.
Cell growth continues.
Synthesis of RNA and proteins continues.
Cell is prepared for mitosis.

It represents the actual cell division

(mitosis).
In human cell cycle, it lasts for only about
an hour.
M Phase includes
Karyokinesis: Nuclear division
Cytokinesis: Division of cytoplasm.
Some cells do not show division. E.g.
heart cells.
Many other cells divide only occasionally
to replace damaged or dead cells.
The cells that do not divide further exit G1
phase and enter an inactive stage called
quiescent stage (G0). Such cells remain
metabolically active but do not
proliferate.
It is the cell division occurring in somatic cells.
It is also called as equational division as the
number of chromosomes in the parent and
progeny cells is same.
Mitosis is generally seen in diploid cells. It also
occurs in haploid cells of some lower plants and
some social insects.
It involves major reorganization of all cell
components.

The karyokinesis of mitosis has 4 stages:

Prophase
Metaphase
Anaphase
Telophase

It is the longest phase in mitosis.

Early Prophase

The tangled chromatin fibres condense to chromosomes.

The nucleolus is seen attached to the chromosome at the
nucleolar organizer.
Late prophase

Each chromosome splits into two chromatids attached together at the

centromere.
Condensation of chromosomes continues.
In animal cells, the centrioles move to opposite poles. They radiate
out astral rays (microtubular fibrils). Astral rays along with its
centriole pair is called aster. The 2 asters move to opposite poles and
start spindle formation.
The nuclear envelope, nucleolus, Golgi complexes and endoplasmic
reticulum disappear.
Spindle fibres originate from microtubular proteins (tubulin).
The nuclear envelope completely
disintegrates. Hence the chromosomes
spread through the cytoplasm of the cell.
Chromosome condensation is completed.
They can be studied easily under the
microscope. They will have two sister
chromatids.
Chromosomes come to lie at the equator.
The plane of alignment of the
chromosomes at metaphase is called the
metaphase plate.
The spindle fibres from both poles are
connected to chromatids by their
kinetochores in the centromere.

It is the shortest phase in the mitosis.

Centromere of each chromosome divides
longitudinally resulting in the formation of two
daughter chromatids (chromosomes of the future
daughter nuclei).
As the spindle fibres contract, the chromatids move
from the equator to the opposite poles.

Chromosomes cluster at opposite

poles and uncoil into chromatin
fibres.
Nuclear envelope assembles
around the chromatin fibres. Thus
2 daughter nuclei are formed.
Nucleolus, Golgi complex and ER
reappear.
The spindle fibres disappear.
It is the division of cytoplasm resulting in the
formation of 2 daughter cells.
It starts when telophase is in progress.

Here, a cleavage furrow is appeared in plasma

membrane.
It gradually deepens and joins in the centre
dividing the cytoplasm into two.

It is different from the cytokinesis in animal cells

due to the presence of cell wall.
In plant cells, the vesicles formed from Golgi
bodies accumulate at the equator. It grows
outward and meets the lateral walls. They fuse
together to form the cell-plate. It separates the 2
daughter cells. Later, the cell plate becomes the
middle lamella.
During cytokinesis, organelles like mitochondria
and plastids get distributed between the daughter
cells.
In some organisms, karyokinesis is not followed by
cytokinesis. As a result, multinucleate condition
(syncytium) arises. E.g. liquid endosperm in
coconut.
It produces diploid daughter cells with identical genome.

It helps to retain the same chromosome number in all somatic cells.

It helps in the body growth of multicellular organisms. Mitosis in the meristematic tissues
helps in a continuous growth of plants throughout the life.

It restores the nucleo-cytoplasmic ratio that disturbed due to cell growth.

It helps in cell repair & replacement. E.g. cells of the upper layer of the epidermis, lining of the
gut & blood cells.
It is the division of diploid germ cells that
reduces the chromosome number by half
forming haploid daughter cells (gametes).

It occurs during gametogenesis.

It leads to haploid phase in the life cycle of

sexually reproducing organisms. Fertilization
restores the diploid phase.
It involves two cycles (meiosis I & meiosis II)
but only a single cycle of DNA replication.
It involves pairing of homologous
chromosomes and recombination between
them.
Meiosis I begins after replication of parental
chromosomes to form identical sister
chromatids at the S phase.
4 haploid cells are formed at the end of
meiosis II.

Meiosis I Meiosis II

Prophase I Prophase II

Metaphase I Metaphase II

Anaphase I Anaphase II

Telophase I Telophase II
It is typically longer and more complex.
It includes 5 phases based on chromosomal
behaviour:
Leptotene
Zygotene
Pachytene
Diplotene
Diakinesis

Leptotene {Leptonema}

Chromatin fibres become long slender chromosomes.

Nucleus enlarges.
Zygotene {Zygonema)

Chromosomes become more condensed. Similar chromosomes start pairing together (synapsis) with the help
of a complex structure called synaptonemal complex.
The paired chromosomes are called homologous chromosomes. Each pair of homologous chromosomes is
called a bivalent.
Pachytene (Pachynema)

Comparatively longer phase.

Bivalent chromosomes split into similar chromatids. This stage is called tetrads.
Appearance of recombination nodules at which crossing over* occurs. It leads to recombination of genetic
material on the homologous chromosomes.
Recombination is completed by the end of pachytene.

Diplotene {Diplonema)

Dissolution of the synaptonemal complex occurs. The recombined homologous chromosomes of the bivalents
separate from each other except at the sites of crossovers. These X-shaped structures are called chiasmata.
In oocytes of some vertebrates, diplotene lasts for months or years.

Diakinesis

Terminalisation of chiasmata.
Chromosomes are fully condensed.
The meiotic spindle fibres originate from the poles to prepare the homologous chromosomes for separation.
Nucleolus & nuclear envelope disappear.
Spindle formation is completed.

The chromosomes align on the equatorial

plate.

The microtubules from the spindle attach to

the pair of homologous chromosomes.

The homologous chromosomes separate, while

sister chromatids remain associated at their
centromeres.

The nuclear membrane and nucleolus reappear

and 2 haploid daughter nuclei are formed. This
is called diad.

After this, cytokinesis may or may not occur.

After a short interphase, it is followed by

meiosis II. This short stage between the two
meiotic divisions is called interkinesis.

DNA replication does not occur in this phase.

It resembles the mitosis.
It has 4 phases:
Prophase II
Metaphase II
Anaphase II
Telophase II

It is initiated immediately after cytokinesis. The

chromosomes again become compact.

Nucleolus and nuclear membrane disappear in both

nuclei.

The chromosomes align at the equator and the microtubules from opposite poles of the
spindle get attached to the kinetochores of sister chromatids.
It begins with the simultaneous splitting
of the centromere of each chromosome
(which was holding sister chromatids
together). Thus they move toward
opposite poles of the cell.

The two groups of chromosomes once

again get enclosed by a nuclear
envelope; cytokinesis follows resulting
in the formation of tetrad of cells i.e., 4
haploid daughter cells.
It conserves the chromosome number of each species.

It causes genetic variation (due to crossing over) in the population of

organisms. It is important for evolution.
Photosynthesis is a physio-chemical process by
which green plants use light energy {solar energy}
to synthesis organic compounds

It is the basis of all life on earth

Ultimately, all living forms depend on sunlight for

energy.

It is the primary source of all food on earth.

It releases oxygen into the atmosphere

Take a variegated leaf {or leaf partially

covered with black paper} that was exposed
to light
Test the leaves for starch
it shows that photosynthesis occurs only in
green parts of the leaves in presence of light
A part of a leaf is enclosed in a
test tube containing KOH soaked
cotton {which absorbs CO2}
The other half of the leaf is
exposed to air
Place this setup in light for
sometime .
Test the leaf for presence of
starch
Exposed part shows positive for
starch and portion in the tube
shows negative
This proves that CO2 is required
Hugo von Mohl
for photosynthesis

Priestley performed experiments

to prove the role of air in the
growth of green plants
He discovered oxygen in 1774
He observed that a candle
burning in a closed bell jar gets
extinguished. Similarly, a mouse
is suffocated in a closed jar

He continued that a burning candle or a breathing animal

damage the air
He placed a mint plant in the bell jar
He found that the mouse stayed alive and the candle
continued to burn
He hypothesised that plants restore to the air whatever
breathing animals and burning candles remove
He conducted the same experiment by placing it
once in the dark and once in the sunlight
He showed that sunlight is essential to the plant for
purifying the air
He repeated this experiment with an aquatic plant.
It showed that in bright sunlight, small bubbles were
formed around green parts while in the dark, they
did not

Later, he identified
these bubbles to be of
oxygen. Thus, he
showed that only the
green part of plants
release O2

He proved that
Glucose is produced when
plants grow and it is usually
stored as starch
Chlorophyll is located in
special bodies {chloroplasts}
Glucose is made in the green
parts of plants
He split the light using a prism into its spectral components
and illuminated a green alga {Cladophora} placed in a
suspension of aerobic bacteria. The bacteria were used to
detect the sites of O2 evolution
He observed that the bacteria accumulated mainly in the
region of blue & red light of the split spectrum
It was the first described action spectrum of photosynthesis.
It resembles the absorption spectra of chlorophyll a and b.

By the middle of the 19th century, it is discovered that plants use the light energy to make
carbohydrates from CO2 & H2O
The empirical equation of the process of photosynthesis is

Where CH2O represents a carbohydrate {example: glucose}

Van Niel {microbiologist} conducted some studies on purple
and green bacteria
He demonstrated that photosynthesis is a light-dependent
reaction in which hydrogen from a suitable oxidisable
compound reduces CO2 to carbohydrates
This can be expressed by

In plants, H2O is the hydrogen donor and is oxidised to O2

Purple and green sulphur bacterial use H2S as H- donor. So the
‘oxidation’ product is sulphur or sulphate and no O2 is
produced
Thus, he inferred that the O2 evolved by green plants comes
from H2O, not from CO2. this was later proved by using radio-
isotopic techniques
Therefore overall correct equation for photosynthesis is

Photosynthesis takes place in green leaves and other

green parts of the plants
Chloroplast is present in the walls of the mesophyll cells
of leaves. It helps to get optimum quality of incident
light
Chloroplast contains a membranous system. It consists
of grana, stroma lamellae and fluid stroma
Each granum Is a group of membrane-bound sacs called
thylakoids {lamellae}
They contain leaf pigments
The membrane system traps light energy and synthesis
ATP and NADPH. It is called light reactions
In stroma, enzymatic reactions incorporate CO2 into the
plant for synthesizing sugar, which in turn forms starch. It
is called dark reactions or that they are not light
dependent

Pigments are substances that have the ability to absorb

light at specific wavelengths
Chromatography shows the following leaf pigments in
chromatogram:
Chlorophyll a : bright or blue green
Chlorophyll b : yellow green
Xanthophylls : yellow
Carotenoids : yellow to yellow – orange

Chlorophyll b, Xanthophylls & carotenoids are accessory

pigments

Functions of accessory pigments

They absorb light at different wavelength and transfer

the energy to chlorophyll a
They protect chlorophyll a from photo-oxidation
The absorption spectrum and action spectrum coincide closely showing that photosynthesis
us maximum at blue and red regions of the spectrum
The graphs below also show that chlorophyll a is the chief pigment associated with
photosynthesis

Graph showing Graph showing action Graph showing action

absorption spectrum of spectrum of spectrum
chlorophyll a, b & photosynthesis superimposed on
carotenoids absorption spectrum of
chlorophyll a

Pigments are organised into two

photosystems called Photosystem I {PSI} and
Photosystem II {PSII}. These are named in the
sequence of their discovery
Each photosystem has a chlorophyll a and
accessory pigments bound by proteins
All pigments {except one molecule of
chlorophyll a} form a light harvesting complex
{LHC or antennae}
Single chlorophyll a acts as reaction centre
In PS I, the reaction centre absorbs light at
700 nm, and so called P700
In PS II, the reaction centre absorbs light at
680nm, and so called P680
When PSII absorbs red light of 680 nm wavelength,
electrons are excited and transferred to an electron
acceptor
The electron acceptor passes them to a chain of
electrons transport system consisting of cytochromes
This movement of electrons is downhill, in term of
redox potential scale
The electrons are transferred to the pigments of PSI
Simultaneously, electrons in PSI are also excited when
they receive red light of 700 nm and are transferred
to another acceptor molecule having a greater redox
potential
These electrons are moved downhill to a molecule of
NADP+. As a result, NADP+ is reduced to NADPH + H
Transfer of electrons from PSII to PSI and finally
downhill to NADP+ is called Z scheme, due to its
zigzag shape. This shape is formed when all the
carriers are placed in a sequence on a redox potential
scale
 The water splitting complex in PSII is
located on the inner side of the thylakoid
membrane
Water is split into H, [O] & electrons

So PSII can supply electrons continuously

by replacing electrons from water
splitting
Thus, PSII provides electrons needed to
replace those removed from PSI
The protons {H+} are used to reduce
NADP to NADPH+
O2 is liberated as a by-product of
photosynthesis

The synthesis of ATP by cells {in mitochondria and chloroplasts} is called phosphorylation
Photophosphorylation is the synthesis of ATP from ADP in chloroplasts in presence of light
It occurs in 2 ways

Cyclic
Non cyclic

It occurs in stroma lamellae when only PSI is functional

The electron is circulated within photosystem and ATP
synthesis occurs due to cyclic flow of electrons
The lamellae of grana have PSI and PSII. The stroma
lamellae membranes lack PSII and NADP reductase
The electron does not pass on to NADP but is cycled
back to PSI through electron transport chain
Here, only ATP is synthesised {no NADPH +H+}
Cyclic photophosphorylation also occurs when only light
of wavelengths beyond 680nm are available
It occurs when the two photosystems work in a series, {first PSII and then PSI} through an
electron transport chain as seen in the Z scheme
Here, ATP & NADPH + H+ are synthesised
It is a non-cyclic process because the electrons lost by PSII do not come back to it but pass on
to NADP

It explains the mechanism of ATP synthesis in chloroplast

Chemiosmosis : movement of ions across a semipermeable membrane. It occurs in chloroplast
and mitochondria
Chemiosmosis needs a membrane, a proton pump, a proton gradient {across thylakoid
membranes} and ATP synthase
Splitting of water takes place on the inner side of the membrane. So the protons accumulate in
the lumen of thylakoids

As electrons move through the photosystems, protons are transported across the membrane.
It is due to the removal of protons from stroma for the following reasons:

The primary electron acceptor is located towards the outer side of the membrane and
transfer its electron to an H carrier. So this molecule removes a proton from the stroma
while transporting an electron. When this molecule passes on its electron to the electron
carrier on the inside of the membrane, proton is released into the lumen of the
membrane.
The NADP reductase enzyme is located on the stroma side of the membrane. Along with
electrons coming from PSI, protons are necessary to reduce NADP. These protons are
also removed from stroma

Hence, protons in stroma are

decreased but in lumen, protons
are accumulated. It creates a
proton gradient across the
thylakoid membrane and
decrease in pH in lumen
Breakdown of proton gradient
leads to synthesis of ATP by ATP
synthase enzyme
ATP Synthase consists of 2 parts

CF0 : It is embedded in the membrane and forms a trans membrane channel. It carries
out facilitated diffusion of protons across the membrane to th stroma. It results in
breakdown of proton gradient
CF1 : It protrudes on the outer surface of the thylakoid membrane. The energy due to
breakdown of gradient causes a conformational change in CF1 particle. It makes the
enzyme to synthesise ATP.

Energy is used to pump protons across a membrane, to create a gradient or a high

concentration of protons within the thylakoid lumen
ATP Synthase has a channel for the diffusion of protons back across the membrane. This
releases energy to activate ATP synthase that catalyses formation of ATP

Products of light reaction are ATP, NADPH & O2

Dark reaction is the use of ATP & NADPH to drive the processes for the synthesis of food
{sugars}
This phase does not directly depend on the light but is dependent on the products of the light
reaction
It can be verified as follows
Immediately after light becomes unavailable, the biosynthetic process continues for sometime,
and then stops. If light is available, the synthesis starts again

CO2 combines with H2O to form (CH2O)n or sugars

CO2 assimilation during photosynthesis is of two types
C3 Pathway : in this, first product of CO2 fixation is a C3 organic acid {phosphoglyceric acid –
PGA}. Melvin Calvin discovered this using 14C in algal photosynthesis
C4 pathway : In this, first stable product is Oxaloacetic acid {OAA}, a 4-carbon {C4} acid
It Occurs in all photosynthetic plants {C3 or C4 pathways }
It has three stages
Carboxylation
Reduction
Regeneration

RuBP {ribulose bisphosphate – a 5 Carbon ketose

sugar} is the primary CO2 acceptor
It is the most crucial step. CO2 is fixed by RuBP to two
3 PGA in presence of the enzyme RuBP carboxylase

Since this enzyme also has an oxygenation activity, it is also called RuBP carboxylase –
oxygenase {RuBisCO}
RuBisCO is the most abundant enzyme in the world

It is a series of reactions leading to glucose

formation
Here, 2 ATP molecules for phosphorylation and 2
NADPH for reduction per CO2 molecule are used
Fixation of 6 CO2 molecules and 6 turns of the cycle
are needed to remove one glucose molecule from
the pathway

It is crucial for continuation of the cycle

It requires one ATP for phosphorylation to form RuBP
Hence for every CO2 molecule, 3 ATP molecules and 2 NADPH are required
It is probably to meet this difference in number of ATP and NADPH used in the dark
reaction that the cyclic phosphorylation takes place
To make one glucose molecule, 6 turns of the cycle are needed
It is present in plants adapted to dry tropical regions
They also use C3 pathways as main biosynthetic pathway
The large cells around the vascular bundles of the C4 plants are called bundle sheath cells.
Such anatomy is called Kranz anatomy
The bundle sheath cells may form several layers around the vascular bundles
They have large number of chloroplasts, thick walls impervious to gas exchange and no
intercellular spaces
 Primary CO2 acceptor is phosphoenol
pyruvate {PEP}
It is a 3-carbon molecule seen in mesophyll
cells. The enzyme for this fixation is PEP
carboxylase (PEPcase}
The mesophyll cells lack RuBisCO enzyme
The C4 acid OAA is formed in mesophyll cells
It then forms other 4-carbon acids like malic
acid or aspartic acid. They are transported to
bundle sheath cells

In the bundle sheath cells, C4 acids are broken down to release CO2 & a C3 molecule
The C3 molecule is transported back to mesophyll where it is converted to PEP again
The released CO2 enters the C3 pathway
Bundle sheath cells are rich in RuBisCO, but lack PEPcase. Thus, C3 pathway is common to
C3 & C4 plants

C4 plants are special because :

They have a special type of leaf anatomy {Kranz}

They tolerate higher temperatures
They show a response to high light intensities
They lack photorespiration
They have greater productivity of biomass
In calvin pathway, RuBP combines with CO2

Active site of RuBisCO can bind to CO2 and O2

RuBisCO has greater affinity for CO2 than for O2
This binding is competitive
Relative concentration of O2 and CO2 determines which one will bind to the enzyme

In C3 plants, some O2 bind to RuBisCO. Hence, CO2 fixation is decreased. Here, RuBP binds
with O2 to form one molecule of phosphoglycerate and phosphoglycolate. This pathway is
called photorespiration

In this pathway, there is no synthesis of sugars, ATP and NADPH. Hence photorespiration is
a wasteful process. Rather it causes the release of CO2 by using ATP

In C4 plants, photorespiration does not occur because they can increase CO2 concentration
at the enzyme site. This takes place when C4 acid from the mesophyll is broken down in the
bundle cells to release CO2. this minimises the oxygenase activity of RuBisCO

Due to lack of photorespiration, productivity and yields are bettere in C4 plants. In

addition, these plants show tolerance to higher temperatures.
 C3 plants C4 plants
Photosynthesis occurs in In mesophyll cells and bundle
mesophyll cells sheath cells
Kranz anatomy is absent Present
RuBP is the primary CO2 acceptor PEP is the primary CO2 acceptor
3-PGA, a 3-C compound is the OAA, a 4-C compound is the first
first stable product stable product
Chloroplasts are of only one type Chloroplasts are dimorphic
{granal} {granal in mesophyll and agranal
in bundle sheath}
Photorespiratory loss is high Photorespiration is absent or
negligible
High CO2 compensation point Low CO2 compensation point {0-
{25-100 ml} 10ml}
Optimum temperature for About 35 C – 45 C
photosynthesis is about 25 C
Photosynthetically less efficient Photosynthetically more efficient
and productivity is low and productivity is high

Example : Rice, wheat, bean, Example : maize, sugarcane,

potato amaranth, sorghum

Marshall Hatch Roger Slack Melvin Calvin

 Internal {plant} factors
Number, size, age and External factors
orientation of leaves, Sunlight

mesophyll cells and Temperature

chloroplasts CO2 concentration

Internal CO2 concentration Water

Amount of chlorophyll

Plant factors are dependent on the genetic predisposition an the growth of the plant

“ If a biochemical process is affected by more than one

factor, its rate is determined by the factor nearest to its
minimal value : it is the factor which directly affects the
process if its quantity is changed”

Example :- Frederick F. Blackman

A plant with green leaf, optimal light and CO2 may not photosynthesise in very low
temperature
If optimal temperature is given, it will start photosynthesis
 Light quality, light intensity and duration of exposure
to light influence photosynthesis
There is a linear relationship between incident light
and CO2 fixation rates at low intensities
At higher intensities, the rate does not further increase
because other factors become limiting
Light saturation occurs at 10% of the full sunlight
Hence, except for plats in shade or in dense forests,
light is rarely a limiting factor in nature
High increase in incident light breaks down
chlorophyll. It decreases photosynthesis

Major limiting factor for photosynthesis

CO2 concentration is very low in the atmosphere {0.03% -
0.04%}
Increase up to 0.05% cause increase in CO2 fixation rates.
Beyond this level, it becomes damaging over longer periods
At low light conditions, C3 and C4 plants do not respond to
high CO2 conditions. At high light intensities, they show
increased rate of photosynthesis

C4 plants show saturation at about 360 μ1L -1

C3 plants respond to increased CO2 concentration and
saturation is seen only beyond 450 μ1L -1
Thus, currently availability of CO2 levels is limiting to the C3
plants
Due to response to higher CO2 concentration, C3 plants
show increased photosynthesis and higher productivity. This
fact is used for some greenhouse crops {tomatoes, bell
pepper etc.}
they hare grown in CO2 enriched atmosphere that leads to a
higher yield
Dark reactions, being enzymatic, are
temperature controlled. Influence of
temperature on light reactions is very less
The C4 plants respond to higher temperatures
and show higher rate of photosynthesis
C3 plants have a much lower temperature
optimum
The temperature optimum for plants also
depends on their habitat. Tropical plants have
a higher temperature optimum than the plants
adapted to temperate climates

Water stress closes the stomata hence reduce

the CO2 availability
Water stress also wilts leaves, thus reduce the
surface area of the leaves and their metabolic
activity
All plant cells are descendants of the zygote (fertilized egg).
The zygote develops into a mature plant through growth and differentiation forming roots,
leaves, branches, flowers, fruits and seeds. Then they eventually die.
Growth is an irreversible permanent increase in size of an
organ or its parts or an individual cell.
It involves metabolic processes that consume energy.

Plant growth continues throughout the life due to the presence of

meristems.
Meristematic cells have the capacity to divide and self-perpetuate.
The growth where new cells are always added to the plant body by the
activity of the meristem is called the open form of growth.

Primary growth : It occurs due to root apical meristem & shoot

apical meristem. It causes the elongation of plants along their axis.
Secondary growth (In gymnosperms & dicots): It occurs due to
lateral meristems, vascular cambium & cork-cambium. These cause
the increase in girth of the organs

At cellular level, growth occurs due to

increase in the amount of protoplasm.
Increase in protoplasm is difficult to measure
directly. So growth is measured by parameters
like increase in fresh weight, dry weight,
length, area, volume & cell number.
Cell number: A maize root apical meristem
can produce more than 17,500 new cells per
hour.
Cell size: Cells in a watermelon may increase
in size by up to 3,50,000 times.
Length: Growth of a pollen tube.
Surface area: Growth in a dorsi-ventral leaf.
Meristematic phase: It occurs in the
meristems at the root apex & the shoot
apex. Cells in this region have rich
protoplasm and large nuclei. Cell walls are
primary, thin & cellulosic with abundant
plasmodesmata.

Elongation phase: It occurs in cells

proximal (just next, away from the tip) to
meristematic zone. Cells have increased
vacuolation, size and new cell wall Detection of zones
deposition. of elongation by the
parallel line
Maturation phase: It occurs in the cells technique.
further away from the apex (more Zones A, B, C, D
proximal to the phase of elongation). The immediately behind
the apex have
cells attain maximal size in terms of wall
elongated most.
thickening and protoplasmic modifications.

It is the increased growth per unit time.

The growth rate may be arithmetic or geometrical.

In this, following
mitotic cell division, Mathematically, it is
only one daughter cell expressed as
continues to divide 𝐿𝑡 = 𝐿0 + 𝑟𝑡
while the other 𝐿𝑡 = length at time ‘t’
differentiates and 𝐿0 = length at time
matures. ‘zero’
On plotting the length 𝑟𝑡 = growth rate /
of the organ against elongation per unit
time, a linear curve is time.
obtained.
In most systems, the initial growth is slow (lag phase), then it increases rapidly (log or
exponential phase).
Here, both the daughter cells continue and retain the ability of mitotic cell division.
If nutrient supply is limited, the growth slows down leading to a stationary phase.

On plotting the parameter of growth against

time, we get a typical sigmoid (S) curve.
A sigmoid curve is a characteristic of living
organism growing in a natural environment.
It is typical for all cells, tissues and organs of a
plant.
The exponential growth can be expressed as

𝑊1 = 𝑊0 𝑒𝑟𝑡

𝑊1 = final size (weight, height, number etc.)

𝑊0 = initial size at the beginning of the
period
𝑟 = relative growth rate
𝑡 = time of growth
𝑒 = base of natural logarithms
Relative growth rate (r) is the measure of the ability of plant to produce new plant material,
referred to as efficiency index. Hence, the final size of 𝑊1 depends on the initial size, 𝑊0 .
Quantitative comparisons between the growth can also be made in 2 ways:
Absolute growth rate: Measurement and comparison of total growth per unit time.
Relative growth rate: Measurement of growth of the given system per unit time expressed on a
common basis, e.g., per unit initial parameter.

Diagrammatic comparison of absolute

and relative growth rates. Both leaves A
and B have increased their area by 5
cm2 in a given time to produce A1, B1
leaves.

Water: For cell enlargement. Turgidity of cells helps in extension growth. Water provides
medium for enzymatic activities needed for growth.
Oxygen: It helps to release metabolic energy for growth.
Nutrients: Macro & micro elements are needed for the synthesis of protoplasm and act as
source of energy.
Temperature: Plants have an optimum temperature at which growth is maximum. Deviation
from this range could be detrimental to its survival.
Light & gravity: Affect certain phases/stages of growth.
Differentiation is a process in which the meristem cells (root apical & shoot-apical) and
cambium differentiate and mature to perform specific functions.
In this, cell walls & protoplasm undergo structural changes. Capacity of cell division is lost.
E.g. Loss of protoplasm to form a tracheary element. They also develop very strong, elastic,
lignocellulosic secondary cell walls, to carry water to long distances even under extreme
tension.

Under certain conditions, living differentiated cells

regain the capacity of division. This is called
dedifferentiation.
E.g. formation of meristems (interfascicular
cambium & cork cambium) from differentiated
parenchyma cells.

Dedifferentiated cells can produce cells that again

lose the capacity to divide but mature to perform
specific functions. It is called redifferentiation.
Plant growth is open, i.e., it can be indeterminate
or determinate. Differentiation is also open,
because cells/tissues arising out of the same
meristem have different structures at maturity.
Final structure at maturity of cell/tissue is also
determined by the location of the cell.
E.g. cells positioned away from root apical
meristems differentiate as root-cap cells, while
those pushed to periphery mature as epidermis.
It is a process that includes
all changes in the life cycle
of an organism from seed
germination to senescence.
It is the sum of growth and
differentiation.
It is represented as follows:

Plants follow different pathways in response to

environment or phases of life to form different kinds of
structures. This ability is called plasticity. E.g.
Heterophylly due to phases of life:
E.g. In cotton, coriander and larkspur, the leaves of the
juvenile plants and mature plants are different in shape.
Heterophylly due to environment:
E.g. Difference in shapes of leaves produced in air and
water (e.g. buttercup).
 Plant growth regulators (PGRs) are small, simple molecules that regulate plant growth.
Based on the functions, PGRs are 2 groups:
Plant growth promoters: For growth promoting activities like cell division & enlargement, tropic
growth, pattern formation, flowering, fruiting & seed formation. E.g. auxins, gibberellins & cytokinins.
Plant growth inhibiters: For growth inhibiting activities like dormancy & abscission. Respond to
wounds & stresses of biotic & abiotic origin. E.g. abscisic acid & ethylene.
Ethylene fits either of the groups, but it is largely a growth inhibitor.

Charles Darwin & his son Francis Darwin

observed that the coleoptiles of canary grass
responded to unilateral illumination by
growing towards the light source
(phototropism). It was concluded that the tip of
coleoptile caused the bending of the entire
coleoptile.
F.W. Went isolated Auxin from tips of
coleoptiles of oat seedlings.
Auxin was first isolated from human urine.

Charles Darwin Francis Darwin

Auxins are produced by the growing apices of stems & roots, from where they migrate to regions
of their action.
 Types of Auxins
Natural: E.g. Indole-3-acetic acid (IAA) & indole butyric acid (IBA). They are isolated
from plants.
Synthetic: E.g. NAA (naphthalene acetic acid) & 2, 4-D (2, 4-dichlorophenoxyacetic).

Functions
To initiate rooting in stem cuttings
for plant propagation.
Promote flowering. E.g. in
pineapples.
To prevent fruit and leaf drop at
early stages.
Promote the abscission of older
leaves & fruits.
Induce parthenocarpy, e.g., in
tomatoes.
They are used as herbicides. 2, 4-D
is used to kill dicot weeds. It does
not affect monocot plants. It is used
to prepare weed-free lawns.
Controls xylem differentiation and
helps in cell division.

In higher plants, the growing apical bud inhibits the

growth of lateral (axillary) buds. It is known as apical
dominance.
Removal of shoot tips (decapitation) results in the
growth of lateral buds. It is applied in tea plantations,
hedge-making.
These are acidic PGR.
E. Kurosawa treated the sterile
filtrates of Gibberalla fujikuroi (a
fungus that causes ‘bakanae’ disease
or foolish seedling in rice) to healthy
rice seedlings. As a result, it showed
the symptoms of ‘bakanae’ disease.
Later, the active substances were
identified as gibberellic acid.
There are more than 100 gibberellins
(GA1, GA2, GA3 etc.) in fungi and
higher plants.
Gibberellic acid (GA3 or Terpenes) is
one of the first discovered and most
intensively studied gibberellins.

Functions
They cause increase in length of axis. So they are used
to increase the length of grapes stalks.
To elongate and improve the shape of fruits such as
apple.
They delay senescence. So the fruits can be left on the
tree to extend the market period.
GA3 is used to speed up malting process in brewing
industry.

Sugarcane stores sugar in stems. Spraying

sugarcane crop with gibberellins increases the
length of the stem. It increases the yield by as
much as 20 tonnes per acre.

Spraying juvenile conifers with GAs hastens the

maturity period. It leads to early seed
production.
For bolting (internode elongation just prior to
flowering) in beet, cabbages and many plants
with rosette habit.
F. Skoog and co-workers observed that from
intermodal segments of tobacco stems, the
callus (a mass of undifferentiated cells)
proliferated only if the nutrients medium
was supplemented with extracts of vascular
tissues, yeast extract, coconut milk or DNA.
Skoog & Miller later identified and
crystallized the active substance and termed
as kinetin.
Cytokinins were discovered as kinetin (N6-
furfurylamino purine - an Adenine
derivative) from the autoclaved herring
sperm DNA.
Kinetin does not occur naturally in plants.

Functions
Zeatin (from corn-kernels & coconut milk) is the natural
substances with cytokinin-like activities.
There are some synthetic compounds with cell division
promoting activity.
Natural cytokinins are synthesized in regions of rapid cell
division (root apices, shoot buds, young fruits etc).
Functions
They play a role in cytokinesis.
They help to produce new leaves, chloroplasts in leaves,
lateral shoot growth and adventitious shoot formation.
They help overcome the apical dominance.
They promote nutrient mobilization which helps in the
delay of leaf senescence.
Folke Skoog
Cousins confirmed the release of a volatile
substance from ripened oranges that hastened the
ripening of stored bananas. Later this substance
was identified as ethylene.
Ethylene is a simple gaseous PGR.
It is synthesized in large amounts by tissues
undergoing senescence and ripening fruits.

Functions
It influences horizontal growth of seedlings, swelling of axis and apical hook formation in dicot
seedlings.
It promotes senescence and abscission of plant organs especially of leaves and flowers.
It promotes fruit ripening. It enhances respiration rate during fruit ripening. This is called
respiratory climactic.
It breaks seed and bud dormancy, initiates germination in peanut seeds, sprouting of potato
tubers.
Functions
It influences horizontal growth of seedlings, swelling of axis and apical hook formation in dicot
seedlings.
It promotes senescence and abscission of plant organs especially of leaves and flowers.
It promotes fruit ripening. It enhances respiration rate during fruit ripening. This is called
respiratory climactic.
It breaks seed and bud dormancy, initiates germination in peanut seeds, sprouting of potato
tubers.
The most widely used source of ethylene is ethephon.
Ethephon in an aqueous solution is readily absorbed and transported within the plant and
releases ethylene slowly.
Ethephon hastens fruit ripening in tomatoes & apples and accelerates abscission in flowers and
fruits (thinning of cotton, cherry, walnut). It promotes female flowers in cucumbers thereby
increasing the yield.
During mid-1960s, it was reported 3 kinds of
inhibitors: inhibitor-B, abscisin II & dormin.
They were chemically identical and now
known as abscisic acid.
ABA is the derivatives of carotenoids.
It regulates abscission and dormancy

Functions
It acts as an inhibitor of plant growth &
metabolism.
It inhibits seed germination.
It stimulates closure of stomata in the
epidermis.
It increases the tolerance of plants to various
kinds of stresses. Therefore, it is also called
the stress hormone.
It has an important role in seed development,
maturation and dormancy. Seed dormancy by
ABA helps to withstand desiccation and other
factors unfavourable for growth.

PGRs play individualistic or synergistic role. Such

roles may be complimentary or antagonistic.
PGRs interact to affect dormancy in seeds/ buds,
abscission, flowering, senescence, vernalisation,
apical dominance, seed germination, plant
movements etc.
In most cases, ABA acts as an antagonist to GAs.
It is the response of plants to periods of
day/night.
Some plants require light to induce
flowering.
Based on light duration, plants are 3
groups:
Long day plants: They require the
exposure to light for a period exceeding
a well-defined critical duration.
Short day plants: They require the
exposure to light for a period less than
the critical duration before the
flowering is initiated in them.
Day-neutral plants: They have no
correlation between exposure to light
duration and induction of flowering.

Long day plant Short day plant Day neutral plant

It is the phenomenon in which some plants depends quantitatively or qualitatively on
exposure to low temperature for flowering.
Advantages:
It prevents precocious reproductive development late in the growing season.
It enables the plant to have sufficient time to reach maturity.

Examples for Vernalisation

Some food plants, wheat, barley, rye have 2 varieties:

Spring varieties: These are normally planted in the spring and come to flower and produce
grain before the end of the growing season.
Winter varieties: If they planted in spring would normally fail to flower or produce mature
grain within a span of a flowering season. Hence, they are planted in autumn. They germinate,
and over winter come out as small seedlings, resume growth in the spring, and are harvested
usually around mid-summer.

Vernalisation in biennial plants: Biennials are

monocarpic plants that normally flower and
die in the second season. E.g. Sugar beet,
cabbages, carrots etc. Subjecting the growing
of a biennial plant to a cold treatment
stimulates a subsequent photoperiodic
flowering response.
There are certain seeds which fail to germinate even when external conditions are favorable.
Such seeds are understood to be undergoing a period of dormancy which is controlled not by
external environment but are under endogenous control or conditions within the seed itself.
Impermeable and hard seed coat; presence of chemical inhibitors such as abscissic acids,
phenolic acids, para-ascorbic acid; and immature embryos are some of the reasons which
causes seed dormancy.
This dormancy however can be overcome through natural means and various other man-
made measures. For example, the seed coat barrier in some seeds can be broken by
mechanical abrasions using knives, sandpaper, etc. or vigorous shaking.
In nature, these abrasions are caused by microbial action, and passage through digestive tract
of animals.
Effect of inhibitory substances can be removed by subjecting the seeds to chilling conditions
or by application of certain chemicals like gibberellic acid and nitrates.
Changing the environmental conditions, such as light and temperature are other methods to
overcome seed dormancy.
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.
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 and lenticels
Why plants need no specialized respiratory organs?
Each plant part takes care of its own gas-exchange needs. So gas transport is very limited.
Gas exchange is very low as compared to animals.
Leaves are adapted for maximum gas exchange during photosynthesis. During this, O2 is released
within 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.
C6H12O6 + 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 oxidised 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 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 isomerised 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).

PGAL is oxidised 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 synthesised from one glucose molecule.
Pyruvic acid (pyruvate) is the key product of glycolysis. Its metabolic fate depends on the cellular
need.
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.

Here, the pyruvic acid formed from glucose is

converted to CO2 and ethanol. The enzymes,
pyruvic acid decarboxylase and alcohol
dehydrogenase catalyse these reactions. E.g.
Yeast.
Yeasts poison themselves to death when the
concentration of alcohol reaches about 13%.

Net ATP production from fermentation of

one glucose molecule = 2.
(4 ATP from glycolysis – 2 ATP utilized).
Here, pyruvic acid is converted to lactic
acid. E.g. Some bacteria.
In animals, when oxygen is inadequate
during exercise, pyruvic acid in muscle
cells is reduced to lactic acid by lactate
dehydrogenase.
The reducing agent (NADH+H+) is
reoxidised to NAD+ in both type of
fermentation

Net ATP production from fermentation of one

glucose molecule = 2.
(4 ATP from glycolysis – 2 ATP utilized).

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 mitochondria.
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
requires coenzymes such as
NAD+ and Coenzyme A.
During this process, 2 molecules
of NADH are produced from 2
molecules of pyruvic acid.
The acetyl CoA then enters
tricarboxylic acid cycle.
 TCA cycle was first elucidated by Hans Krebs.

Steps:
Condensation of acetyl group with oxaloacetic acid (OAA) & water to form citric acid in
presence of citrate synthase enzyme. A CoA molecule is released.
Citrate is isomerised to isocitrate.
Decarboxylation of isocitrate to -ketoglutaric acid.
Decarboxylation of  -ketoglutaric acid to succinyl-CoA.
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.
Oxidation of succinate to Fumarate and then to Malate.
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.

Summary equation for this phase is given below:

Pyruvic acid + 4 NAD+ + 2H2O + ADP + Pi

↓ Mitochondrial matrix
3CO2 + 4 NADH + 4H+ + FADH2 + ATP

Thus, a glucose is broken down to give 6 CO2, 8NADH + H+, 2 FADH2 and 2 ATP.
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+ and
FADH2 (formed during TCA cycle)
by oxidation.
The electrons are passed on to O2
to form H2O.

Electrons from NADH are oxidised

by an NADH dehydrogenase
(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 oxidised 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
centres.

Complex I NADH dehydrogenase

When the electrons pass from one carrier to
another via complex I to IV, they are coupled to Complex II FADH2
ATP synthase (complex V) for the ATP
production.
Complex III Cyt bc1
Number of ATP molecules produced depends on Complex IV Cytchrome c oxidase
the nature of the electron donor.
Oxidation of 1 NADH → 3 ATP. Complex V ATP synthase
Oxidation of 1 FADH2 → 2 ATP.
In aerobic respiration, the role of oxygen is limited to the terminal stage. Yet, oxygen is vital
since it drives the whole process by removing hydrogen from the system. Oxygen acts as the
final hydrogen acceptor.

In respiration, energy of oxidation-reduction is utilised for the phosphorylation. So this

process is called oxidative phosphorylation.

Oxidative phosphorylation is different from photophosphorylation (Here, light energy is

utilised for the production of proton gradient for phosphorylation).
The energy released during the ETS is utilized to synthesize ATP by ATP synthase (complex
V).

ATP synthase has 2 major

components: F1 & F0.

F1 headpiece (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-
membrane space to the matrix
down the electrochemical proton
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 synthesised in glycolysis is transferred into mitochondria and undergoes oxidative
phosphorylation.
Intermediates in the pathway are not used to synthesise 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 :

All pathways work simultaneously and do not take place one after another.
Substrates enter the pathways and are withdrawn from it as and when necessary.
ATP is utilized as and when needed.
Enzymatic rates are controlled by multiple means.
Such calculations are used to appreciate the efficiency of the living system in extraction
and storing energy.

2 ATP directly 2 ATP

Glycolysis 2 molecules of
6 ATP
NADH
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
 Fermentation Aerobic respiration
Complete breakdown of glucose to
Partial breakdown of glucose.
CO2 & H2O.
Net gain of only 2 ATP. Net gain of 36 ATP.
NADH is oxidised to NAD+ rather NADH is oxidised to NAD+ very
slowly. vigorously.

Glucose is the favoured substrate for

respiration. So all carbohydrates are
first converted into 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.
Glycerol is converted to PGAL and
enters the pathway.
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 synthesise fatty acids, acetyl CoA withdraw from the respiratory pathway.
Similarly, during breakdown and synthesis of protein, respiratory intermediates are involved.