Photosynthesis in
Higher Plants
INTRODUCTION
01 Photosynthesis is a physico-chemical process by which green plants use
light energy (solar energy) to synthesise organic compounds.
6CO2 + 12H2O
Chlorophyll
Light C6H12O6 + 6H2O + 6O2 02
Photosynthesis is a redox reaction.
03
Oxidation of H2O and reduction of CO2 take place.
Photosynthesis is Anabolic and Endergonic process.
It is the primary source of all food on earth.
It releases oxygen into the atmosphere
04
EXPERIMENTS RELATED WITH PHOTOSYNTHESIS
Moll’s Half Leaf Experiment
Starch
Portion of leaf in test tube shows
negative test for starch.
Conclusion : CO2 is required for
starch synthesis (photosynthesis) KOH
No Starch
KOH in tube absorbs all the CO2
Variegated leaf experiment
Part of the leaf is covered Starch absent
by black strip to avoid light
exposure.
Covered portion of leaf shows
negative test for starch.
Conclusion: Light is essential Starch present
for starch synthesis
(photosynthesis)
EARLY EXPERIMENTS
1. J. Priestley (1770) [oxygen discovery-1774]
Animal dies due to suffocation as burning of candle and breathing of animal damages
the air.
Hint : Plant restores to air whatever breathing animal and burning candle removed.
2. Jan Ingenhousz
He concluded that light is essential to plant process that somehow purifies the fouled
air in Priestley’s experiment.
He also observed that, in the presence of sunlight aquatic plants release some gas,
later it was found, that gas is oxygen.
He showed that – Only green plants release oxygen.
3. Julius von Sachs (1854)
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.
4. T.W. Engelmann [1 st action spectrum] 400 500 600
Cladophora
700
Aerobic bacteria
Growth of aerobic bacteria was high in blue & red region. alga
Chloroplast
(Odisha NEET 2019)
This indicates release of more O2 in blue & red light.
Conclusion : Rate of photosynthesis was high when blue V I B G Y O R
or red light was supplied to plants.
Prism
Light
5. Van Neil
Studied photosynthesis in green & purple Bacteria. (NEET 2018)
Conclusions :
Hydrogen from some suitable oxidisable compound reduces CO2.
Light
2H2A + CO2 2A + CH2O + H2O
In bacteria “H2S” donates H for CO2 reduction.
In green plants “H2O” donates H for CO2 reduction.
O2 evolved by plants come from H2O.
Anoxygenic photosynthesis occurs in Bacteria.
PHOTOSYNTHESIS: SITE AND PIGMENTS
Photosynthesis occurs in green leaves & other green parts of the plant.
Chloroplasts are present in the mesophyll cells of leaves so that they get
optimum quantity of incident light.
Chloroplast contains a membranous system. It consists of grana, stroma
lamellae and fluid stroma.
The membrane system traps light energy and synthesizes ATP and NADPH. It is called
light reaction. (AIPMT 2015)
In stroma, enzymatic reactions incorporate CO2 into the plant for synthesizing sugar,
which in turn forms starch. It is called dark reaction. It does not mean that they occur in
darkness or that they are not light dependent. (AIPMT 2009)
Lamella
Ribosome
Thylakoid
Stroma
Lipids of Drop
Granum
Inner membrane
Intermembrane space
Outer membrane
Starch granule Chloroplast DNA
PIGMENTS INVOLVED IN PHOTOSYNTHESIS
Pigments are substances that have the ability to absorb light, at specific wavelengths.
Chromatography shows the following leaf pigments:
Chlorophyll a (bright or blue green in chromatogram) (NEET 2018)
Chlorophyll b (yellow green)
Xanthophylls (yellow)
Carotenoids (yellow to yellow-orange)
Absorption Spectrum
Action Spectrum
Absorbance of light by
(Rate of photosynthesis
chloroplast pigments
Chlorophyll b
O2 evolved)
Carotenoids
Chlorophyll a
400 500 600 700
Wavelength (nm) 400 500 600 700
Absorption of light by chlorophyll a Rate of photosynthesis is
is maximum at blue and then in red maximum at blue and red light.
wavelength.
Rate of photosynthesis
Absorption
Light absorbed
400 500 600 700
Wavelength of light in nanometres (nm)
Hence, we can conclude that chlorophyll a is the chief pigment and others are accessory
pigment.
Functions of Accessory Pigments
They absorb light at different wavelength and transfer the energy to chlorophyll .
They protect chlorophyll a from photo-oxidation.
LIGHT REACTION (PHOTOCHEMICAL PHASE)
Light reactions include light absorption, water splitting, oxygen release and formation of
ATP & NADPH (high-energy chemical intermediates).
Photosystems
(Reaction centre + LHC)
Chlorophyll-a is chief pigment for photosynthesis.
(1) PS I-P700 (NEET 2023) (2) PS II – P680 (NEET 2023)
PS-II (Present in appressed
part of thylakoid membrane)
PS-I (Present in non-appressed part of
thylakoid and stroma lamella)
Stroma lamella lacks NADP-reductase enzyme & PS II
CHl-a is reaction centre
Accessory pigments form light harvesting complex (LHC).
ELECTRON TRANSPORT
01 When PS II absorbs red light of 680 nm wavelength, electrons are excited
and transferred to an electron acceptor (uphill).
02 The electron acceptor passes them through a chain of electron transport
system consisting of cytochromes.
03 This movement of electrons is downhill, in terms of redox potential scale.
04 Then electrons are transferred to the pigments of PS I.
Simultaneously, electrons in PS I are also excited when they receive red
05 light of 700 nm and are transferred to another acceptor molecule having
a greater redox potential (uphill).
06 These electrons are moved downhill to a molecule of NADP+. As a result,
NADP+ is reduced to NADPH + H+. (NEET 2022)
Transfer of electrons from PS II to PS I and finally downhill to NADP+ is
07 called the 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.
Photo-Phosphorylation
Photosystem II Photosystem I
The synthesis of ATP by cells (in
e– acceptor NADPH mitochondria & chloroplasts) is
Light e acceptor
–
ADP+ Pi ATP NADP+ called phosphorylation.
Photo-phosphorylation is the
Electron
transport synthesis of ATP from ADP in
system
chloroplasts in presence of light.
It occurs in 2 ways: Non-cyclic
LHC
and Cyclic.
(AIPMT 2010)
LHC
H 2O 2e– + 2H+ + [O]
Non-cyclic photophosphorylation and Chemiosmosis
H+
ht 2e– PQ
Lig Pheo PQH2
EC PQH2
PS-II O Cyt
1 b
– +
2H + O2 Cyt 6 &
2e 2 f
H+ 2e–
Lig
H2O
ht
PC
Thylakoid 2e–
H+H+H+
membrane H+H+H+H+
FRS
PS-I
Lumen H+H+
FD
2e– NADP+ + 2H+
FNR (Ferredoxin
NADP reductase)
Stroma NADPH + H+
H+
ADP + Pi ATP
Thus, chemiosmosis requires a membrane, a proton pump, a proton gradient and ATPase.
(NEET 2023)
The ATPase (ATP synthase)
Consists of two parts:
F0 : It is embedded in the membrane and forms a transmembrane channel that
carries out facilitated diffusion of protons across the membrane.
F1: It protrudes on the outer surface of the thylakoid membrane. The energy
due to breakdown of gradient causes a conformational change in the
F1 particle. It makes the enzyme to synthesise ATP molecules.
Cyclic
Photophosphorylation
Only PS I is used. (NEET 2021)(AIPMT 2010) Photosystem I
Only ATP is synthesised (AIPMT 2009)
No NADPH is formed
e– acceptor
No O2 is evolved
Occurs in stroma lamella (AIPMT 2010) Light
ATP
i
+P
Electron
ADP
transport
system
Chlorophyll
P700
DARK REACTION (BIOSYNTHETIC PHASE)
Products of light reaction are ATP, NADPH and O2.
In dark reaction ATP and NADPH is used to drive the processes for the synthesis of food
(sugars).
It is the biosynthetic phase of photosynthesis.
This phase does not directly depend on light but is dependent on the products of light
reaction. (i.e. ATP and NADPH besides CO2 and H2O)
It can be verified as follows: Immediately after light becomes unavailable, the biosynthetic
process continues for some time, 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 2 types :
1. C3 pathway: In this, first stable product of CO2 fixation is a C3 acid (PGA). Melvin
Calvin studied algal photosynthesis using 14C. He discovered that the first CO2 fixation
product was 3-phosphoglyceric acid (PGA), a 3-carbon organic acid.
2. C4 pathway: In this, first stable product is oxaloacetic acid (OAA), a 4-carbon (C4)
organic acid.
1. The CALVIN CYCLE (C 3 PATHWAY)
It occurs in all photosynthetic plants.
First discovered in Chlorella (a green alga) by M. Calvin.
It has 3 stages: carboxylation, reduction and regeneration.
Three Steps of C3 Cycle
(I) Carboxylation : (II) Reduction : (III) Regeneration :
Most crucial step Multistep reactions Regeneration of CO2
Enzyme involved – 2ATP and 2NADPH acceptor RuBP (5C)
RuBisCO are used for fixation 1 ATP is required to
First stable product – of 1CO2 molecule regenerate 1 RuBP.
3PGA (3C)
RuBP binds to CO2
(RuBP)(5C)
RuBP (ribulose bisphosphate - a 5-carbon Ribulose–1, 5- CO2 + H2O
bisphosphate
ketose sugar) is the primary CO2 acceptor. RUDISCO
1 Carboxylation
Since this enzyme also has an oxygenation
activity it is called RuBP carboxylase-oxygenase
(RuBisCO). Regeneration 3 Calvin cycle 2 × 3-phosphoglycerate
C3 cycle
To make 1 glucose molecule, 6 turns of cycle 2 Reduction
2ATP
are needed as fixation of 6CO2 molecule is ATP
2ADP+2Pi
2×
required. ADP
Triose 2NADPH
phosphate
2NADP+
Sucrose
C3-Cycle/dark reaction/biosynthetic phase
WHAT GOES IN AND COMES OUT OF THE CALVIN CYCLE?
In Out
6 CO2 1 glucose
18 ATP 18 ADP
12 NADPH 12 NADP
2. C 4 PATHWAY (HATCH & SLACK PATHWAY)
C4-Plants
Can tolerate high temperature
Show response to high light intensity
Do not show photorespiration
More efficient (AIPMT 2010)
Low CO2 saturation point.
They lack photorespiration.
They have greater productivity of biomass.
C4-plants show Kranz anatomy (Green bundle sheath cells in wreath like (circular)
arrangement around vascular bundles of leaf). (NEET-I 2016)
Phosphoenol pyruvate is Ist CO2 acceptor. (NEET 2022)
First product formed – OAA (4C) (NEET 2021)
C4-plants – Maize, Sorghum, Sugarcane. (NEET 2021)(AIPMT 2010)
Optimum temperature = 30 – 40°C
Atmospheric CO2
Plasma
Mesophyll membrane
cell
Cell wall
• Granal thylakoid Phosphoenol
• Lack RuBisCO –
pyruvate
HCO3
(NEET 2022)
Regeneration
Fixation
Plasmodesmata C4 acid
C3 acid
Transport
Bundle
sheath cell
C4 acid
• Large number of
Fixation
Chloroplast Transport
by
• Wall impermeable calvin
for gaseous exchange cycle
• No inter cellular space
• Agranal chloroplast CO2
• Site for C3 cycle
(Karnatka NEET 2013) Decarboxylation C3 acid
(AIPMT 2011)
(AIPMT 2010)
Diagrammatic representation of the Hatch and Slack Pathway
PHOTORESPIRATION : C2-CYCLE/GLYCOLATE - CYCLE
RubisCO binds O2 with RuBP
3C–PGA and 2C – phosphoglycolate are formed
Neither synthesis of sugar nor of ATP and NADPH takes place.
Loss of carbons is in the form of CO2 molecule
Takes place at high light intensities, at high temperature and in low CO2 or high O2
Most abundant enzyme in the world
RuBisCO
Has an affinity for both oxygen and carbon dioxide
Both O2 and CO2 show competitive binding for the active site of RuBisCO
Has a greater affinity for CO2
The relative concentration of O2 and CO2 decides which one will bind to
(O2/CO2) enzyme. (NEET 2020)
Differences between C 3 and C 4 plants
C3 Plants C4 Plants
Photosynthesis occurs in mesophyll In mesophyll and bundle sheath cells
cells
Kranz anatomy absent Present
RuBP is the primary CO2 acceptor PEP is the primary CO2 acceptor
3-PGA, a 3-C compound in the first OOA, a 4-C compound is the first stable
stable product product
Chloroplast are of only one type Dimorphic (granal in mesophyll and
(granal) agranal in bundle sheath)
Photo respiratory loss is high (NEET-II Photo respiration is absent or negligible
2016) (AIPMT 2012) (NEET-II 2016) (AIPMT 2012)
Optimum temperature for About 30°C–40°C
photosynthesis is about 25°C
Photosynthetically less efficient and Photosynthetically more efficient and
productivity low productivity high
E.g. Rice, wheat, bean, potato, etc. E.g. Maize, sugarcane, amaranth,
Sorghum, etc.
FACTORS AFFECTING PHOTOSYNTHESIS
Internal (plant) factors: The number, size, age and orientation of leaves, mesophyll
cells and chloroplasts, internal CO2 concentration and amount of chlorophyll.
External factors: Sunlight, temperature, CO2 concentration and water.
Blackman’s Law of Limiting Factors (1905): “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.”
E
.g. a plant with green leaf, optimal light & CO2 conditions may not photosynthesize
if the temperature is very low. If optimal temperature is given, it will start
photosynthesis.
External Factors
(1) Light
(i) Quality
Rate of photosynthesis
Maximum photosynthesis at blue and red
PAR→400–700 nm.
(ii) Quantity
At high intensity, Rate does not show further
increase as other factors become limiting.
At low intensity, Rate of photosynthesis
∝ Intensity of light (linear relation)
Light saturation for CO2-fixation occurs at
10% of full sunlight. Light intensity
(2) CO2
Rate of photosynthesis ∝ concentration of CO2
⇒ But only upto 500 ppm (NEET 2017)
Current atmospheric CO2 level (0.03% – 0.04%) is limiting for C3-plants but not
for C4-plants.
Saturation point
For C3-plants = beyond 450 µlL–1 (NEET 2017)
For C4-plants = 360 µlL–1 (NEET 2017)
CO2–fertilizing effect : High CO2 concentration
⇒ Higher yield. e.g. tomato (NEET 2017), Bell pepper
(3) Temperature
Dark reaction is affected more than light reaction
Optimum temp.
For C3 → 20° – 25°C For C4 → 30° – 40°C
(4) Water
Water stress ⇒ stomata closed ⇒ low CO2 availability
⇒ low rate of photosynthesis.
Water stress ⇒wilting of leaves ⇒ reduced surface area of leaves ⇒ low
metabolism ⇒ low rate of photosynthesis.
Respiration
in Plants
INTRODUCTION
Oxidation of food materials (breaking of C-C bonds of complex molecules) within the
Photosynthesis
cell to release energy for ATP synthesis is called
Light
energy
cellular respiration.
The energy released is not used directly
but is used to synthesize ATP. When Chloroplast
CO2 + H2O C6H12O6 + O2
energy is needed, ATP is broken down.
Hence, ATP acts as energy currency of
the cell.
The compounds that are oxidized Cell respiration
Chemical
Mitochondria energy
during respiration are called respiratory (ATP)
substrates. E.g. Carbohydrates (most common), proteins, fats and organic acids..
BREATHING IN PLANTS
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
01 transport is very limited. Very low gas exchange occurs as compared to
that of animals.
Leaves are adapted for maximum gas exchange during photosynthesis.
During this, O2 is released within the cell. 02
Most living cells have contact with air. They are located close to plant
03 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.
During respiration, oxygen is utilized, and CO2, water & energy are released.
C6H12O6 + 6O2 → 6CO2 + 6H2O + Energy
Certain organisms are adapted to anaerobic conditions.
GLYCOLYSIS (EMP PATHWAY) Glucose
(6C)
ATP
It is the partial oxidation
Hexokinase
ADP
Glucose-6-phosphate
(breakdown) of glucose to form 2 (6C)
Phospho
hexoseisomerase
molecules of pyruvic acid (C3H4O3) Fructose-6-phosphate
(6C)
ATP
in the absence of O2. ADP
Phospho fructokinase
Fructose1,6-bisphosphate
Site - cytoplasm. (6C)
Aldolase Triose Phosphate
This scheme was given by Gustav Triose phosphate
Isomerase
Triose
phosphate
Embden, Otto Meyerhof & J. Parnas. (glyceraldehyde-3-phosphate)
(3C) + (Dihydroxy
Glyceraldehyde 2NAD acetone
So it is also known as EMP pathway. -3-phosphate
dehydrogenase 2NADH+H + phosphate)
(3C)
2 × Triose bisphosphate
It is the only process in respiration in (1,3 bisphosphoglyceric acid)
(3C)
anaerobes. (2) ADP
Phosphoglycerate kinase
(2) ATP
In plants, glucose is derived
2 × Triose phosphate
from sucrose (end product of (3-phosphoglyceric acid)
(3C)
photosynthesis) or from storage Phosphoglyceromutase
carbohydrates. Sucrose is 2 × 2-phosphoglycerate
converted into glucose & fructose
(2) H 2O Enolase
by an enzyme, [Link] 2
2 × phosphoenolpyruvate
monosaccharides readily enter (2) ADP
Pyruvate kinase
(2) ATP
glycolytic pathway.
2 × Pyruvic acid
(3C)
STEPS OF GLYCOLYSIS
It includes 10 steps under the control of different enzymes.
ATP is utilized in 2 steps (total 2 ATP).
(NEET 2023)
4 ATP molecules are directly synthesized from one glucose molecule. so, there is net
gain of 2 ATP. (NEET 2022)
Pyruvic acid is the key product of glycolysis.
Metabolic fate of pyruvate depends on the cellular need.
There are three major ways :
01 02 03
Lactic acid fermentation Aerobic respiration (Krebs’
Alcoholic fermentation
cycle)
FERMENTATION (ANAEROBIC RESPIRATION)
It is the incomplete oxidation of glucose under anaerobic condition.
It occurs in many prokaryotes and unicellular eukaryotes, and in germinating seeds.
It is of 2 types:
Alcoholic Fermentation
Alcoholic fermentation is a type of fermentation where pyruvic acid formed from
glucose is converted to CO2 and ethanol (AIPMT 1997) with the help of enzymes such
as pyruvic acid decarboxylase and alcohol dehydrogenase.
The process of alcoholic fermentation is carried out by the brewing yeast Saccharomyces
cerevisiae (unicellular eukaryote).
Yeast poison themselves to death when concentration of alcohol reaches about 13%.
Pyruvate Alcohol
decarboxylase dehydrogenase
Pyruvate Acetaldehyde Ethanol
(C3) (C2) (C2)
NADH + H+ NAD+
CO2
Lactic Acid Fermentation
Lactic acid fermentation is another type of fermentation in which pyruvic acid is converted to lactic acid without releasing CO2 (AIPMT 2014), as seen in some bacteria, e.g., Lactobacillus.
Prokaryote, as well as animal cells like muscles during exercise, also produce lactic acid from pyruvic acid because O2 is inadequate.
Lactate dehydrogenase reduces pyruvic acid to lactic acid using NADH + H+ as the reducing agent, which is subsequently re-oxidized to NAD+.
Lactate
dehydrogenase
Pyruvate Lactate
(C3) (C3)
NADH + H+ NAD+
The reducing agent (NADH + H+) is reoxidized to NAD+ in both the processes.
Net ATP production from fermentation of one glucose molecule = 2. (Out of 4 ATP from
glycolysis 2 ATP are utilized in fermentation).
The steps involved in fermentation are shown below:
Glucose Glyceraldehyde
Lactic
3-Phosphate
acid
NAD+ NAD+
NADH + H+
NADH + H+
3-Phosphoglyceric
acid Pyruvic
acid
NADH + H+
NAD+
Phosphoenol Ethanol + CO2
Pyruvic acid
DRAWBACK OF FERMENTATION
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. (NEET 2022)
AEROBIC RESPIRATION
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 the 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 of pyruvic dehydrogenase (NEET 2023). It needs several
enzymes, NAD+ & Coenzyme A. (NEET 2018)
During this process, 2 NADH molecules are produced from 2 pyruvic acid molecules.
Mg2+
Pyruvic acid + CoA + NAD +
Acetyl CoA + CO2 + NADH + H+
Pyruvate dehydrogenase
(NEET 2023)
Acetyl CoA then enters tricarboxylic acid (TCA) cycle.
Tricarboxylic acid cycle (Kreb’s Cycle or Citric Acid Cycle)
TCA cycle was first elucidated by Hans Krebs.
01 At 3 points of TCA cycle, NAD+ is reduced to NADH + H+. At one point, FAD+
is reduced to FADH2. (NEET 2017)
Succinyl-CoA is converted to succinic acid and a GTP molecule is
synthesised (substrate level phosphorylation) (NEET 2020). In a coupled
reaction, GTP is converted to GDP with simultaneous synthesis of ATP from 02
ADP. (NEET 2020)
Continued oxidation of Pyruvic acid via TCA cycle requires continued
03 replenishment of OAA. It also requires regeneration of NAD+ & FAD+ from
NADH & FADH2
Mitochondrial
04
Matrix
Pyruvic acid 4NAD FAD 2H2O ADP Pi 3CO2
4NADH 4H FADH2 ATP
Pyruvate
(3C)
NADH + H+
CoA
CO2
Acetyl coenzyme A
(2C)
Citrate
synthase
NADH + H+
Malate
dehydrogenase
Isocitrate
dehydrogenase
NADH + H+
Fumarase
a-ketoglutarate
dehydrogenase
NADH + H+
Succinate
dehydrogenase S
ucc
syn inyl-C
the
taseoA
GDP
GTP
Electron Transport System (ETS) & Oxidative Phosphorylation
Inter-membrane Inner Mitochondrial Matrix
Electron transport system (ETS) is the metabolic space membrane
pathway present in the inner mitochondrial NADH + H+
4H+
2e-
membrane through which electron passes from (Fe-S) FMN
NAD+
Complex I
one carrier to another. (Odisha NEET 2019)(NEET e- (NADH dehydrogenase)
2018) e-
UQH2
e-
This is to release and utilize energy stored in
Complex III
4H+
NADH + H+ and FADH2 (formed during TCA cycle) Cyt C1 Fe-S Cyt b (Cytochrome bc1)
e-
by oxidation. Cyt c UQH2
Complex II
The electrons are passed on to O2 to form H2O. (Succinate dehydrogenase)
e- Succinate
Electrons from NADH are oxidised by an NADH (Fe-s) FAD
Fumarate
dehydrogenase (complex I). Cyt c
e-
CuA
Electrons are then transferred to ubiquinone (UQ) 2H+
IV
2H+
Cyta Cyta3 CuB
located within the inner membrane. Ubiquinone
H O
2
Complex IV
(Cytochrome c oxidase)
also receives reducing equivalents via FADH2 F0 ADP + Pi
F1
H+
(complex II) that is generated during oxidation ATP
synthase ATP
of succinate in citric acid cycle. Electrochemical
gradient
The reduced ubiquinone (ubiquinol or UQH2) 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. (AIPMT 2015 cancelled)
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 (NEET 2022). 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 utilized for the phosphorylation. So, this
process is called oxidative phosphorylation (NEET 2023). While in photophosphorylation,
where light energy is utilized for the production of proton gradient for phosphorylation.
Mechanism of membrane linked ATP synthesis is explained by chemiosmotic hypothesis.
The energy released during the ETS is utilized to synthesize ATP by ATP synthase (complex
V). (NEET 2022)
ATP synthase has two major components: F1 & F0.
Outer ATP F1 F1 headpiece (peripheral
side membrane protein
complex): Site for ATP
2H+ production from ADP and
F0 F1
inorganic phosphate.
Inner
mitochondrial ADP Pi
membrane F0
Matrix F0 (integral membrane
protein complex): It
creates a pathway for
protons to pass through
the inner membrane.
The transport of protons
is coupled to the F1
component’s catalytic
site for ATP synthesis.
For each ATP produced, 4H+ passes through F0 from the inter-membrane space to the
matrix down the electrochemical proton gradient. (AIPMT 2011)
THE RESPIRATORY BALANCE SHEET
Net gain of ATP from each glucose molecule is calculated based on the following
assumptions. (NEET 2013)
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 synthesize
other compounds.
Only glucose is being respired. Other alternative substrates do not enter in the pathway
at any stages.
Such assumptions are not valid because,
01 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. 02
03 ATP is utilised as and
when needed.
Enzymatic rates are controlled by
multiple means. 04
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 4 ATP/6 ATP
Oxidative decarboxylation 2 NADH 6 ATP
6 NADH 18 ATP
2 FADH2 4 ATP
TCA cycle
2 GTP 2 ATP
TOTAL 36/38 ATP
Comparison between fermentation & aerobic respiration.
Fermentation Aerobic respiration
Partial breakdown of glucose. Complete breakdown of glucose to CO2 &
H2O.
Net gain of only 2 ATP. Net gain of 36 ATP / 38ATP.
NADH is oxidised to NAD+ rather slowly NADH is oxidised to NAD+ very vigorously.
AMPHIBOLIC PATHWAYS
Glucose is the favored substrate for respiration. So, all carbohydrates are
01 first converted to glucose for respiration. Other substrates are also respired,
however they do not enter at the first step in the respiratory pathway.
Fats break down into glycerol & fatty acids. Fatty acids are degraded to
acetyl CoA and enter the pathway. Glycerol is converted to PGAL and
enters the pathway. (NEET-II 2016)
02
Proteins are degraded by proteases into amino acids. Each amino acid
03 (after deamination) enters the pathway at some stage in the Krebs’
cycle or as pyruvate or acetyl CoA. (NEET-II 2016)
The respiratory pathway is generally considered as a catabolic pathway.
However it also involves both anabolism (synthesis) and catabolism
(breakdown). So it is better called as an amphibolic pathway. (AIPMT 04
2009)
Fats Carbohydrates Proteins
Simple sugars Amino acids
Fatty acids and glycerol
e.g., glucose
Glucose 6-phosphate
Fructose 1, 6 biphosphate
Dihydroxy Acetone Phosphate Glyceraldehyde 3-phosphate
Pyruvic acid
Acetyl CoA
H2O Krebs'
cycle
CO2
RESPIRATORY QUOTIENT (RQ) OR RESPIRATORY RATIO
It is the ratio of the volume of CO2 evolved to the volume of O2 consumed in respiration.
Volume of CO2 evolved
RQ =
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.
6 CO2
C6H12O6 6O2 6 CO2 6H2O energy RQ = =1
; 6 O2
RQ for fats = < 1. Calculations for a fatty acid, (e.g. tripalmitin) are shown:
2 C51H98O6 145O2 102 CO2 98 H2O energy
102 CO2
=RQ = 0.7
145 O2 (NEET 2019)
RQ for proteins = 0.9.
In living organisms, respiratory substrates are often more than one. Pure proteins or fats
are never used as respiratory substrates.
Plant growth and
development
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
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 is generally indeterminate
Plant growth continues throughout the life due to the presence of meristems (open
form of growth).
PLANT GROWTH
Primary Growth Secondary Growth
e.g. Growth due to root apical e.g. In dicots and gymnosperms
meristem and shot apical the lateral meristem is responsible
meristem for secondary growth
Contribute to the elogation of Vascular cambium
plants along their axis. Cork cambium
Growth is Measurable
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
Number : A maize root apical meristem can produce more than 17,500 new cells per
hour.
Cell size: E.g. Cells in a watermelon can increase in size by up to 3,50,000 times.
Length: E.g. Growth of a pollen tube.
Surface area: E.g. Growth in a dorsi-ventral leaf.
Fresh weight Dry weight Volume
Phases of Growth
MERISTEMATIC PHASE
Constantly dividing cells at root and Cell wall is primary, thin and cellulosic.
shoot apex. Cells are rich in protoplasm.
Cells are small in size Large conspicuous nucleus.
Abundant plasmodesmatal
connections.
ELONGATION PHASE
Cells proximal to meristematic zone → Increased vacuolation
represent the phase of elongation → Cell enlargement
Cells in the region show → New cell wall deposition.
MATURATION PHASE
Cells more proximal to phase of → Maximal size
elongation represents the phase of
→ Wall Thickening
maturation
Cells in this zone show → Protoplasm modification
GROWTH Rates
It is the increased growth per unit time.
Arithmetic Growth Geometric Growth
Both the daughter cells follow
mitotic division (growth in all
Following mitotic Zygote cells). Initial growth is slow (lag
divided
cell division, only phase); then rapid growth
one daughter cell (exponential phase or log
continues division, phase) (NEET 2020), eventually
while the other due to limited nutrients growth
differentiates and Geometric becomes slow and stops
phase: (stationary phase).
matures.
All cells
divide
Mathematical Mathematical
expression of expression of
Arithmetic
arithmetic growth phase exponential growth
(linear curve) (Sigmoid or S-curve)
Lt = L0 + rt Cells capable W1 = W0 ert
of division
Cells that lose
capacity to divide
Fig. : Diagrammatic Where, W1 is final size whether it
Where, Lt is length representation of is in terms of weight, height, or
at time ‘t’; L0 is number
stages during embryo
W0 = Initial size at the beginning
length at time ‘0’; development showing of period; r = Relative growth rate
r is growth rate/ geometric and (measure of plant capacity to
elongation per arithmetic phases form new material) It is also called
efficiency index; t = Growth time;
unit time
e = Base of natural logarithms
Arithmatic growth Geometric growth
Quantitative comparisons between growth of living systems
Absolute Growth Rate →
Total growth per unit time
Relative Growth Rate →
Growth per unit time
Relative growth rate = ´100
Initial parameterrs
Growth rates Leaf A Leaf B
Absolute growth rate 5 cm2 5 cm2
Relative growth rate 100% 10%
CONDITIONS (ESSENTIAL ELEMENTS) FOR GROWTH
Water: Essential 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: At optimum temperature, growth is maximum. Deviation
from this may harm the plants.
Light & gravity: Affect certain phases/stages of growth.
DIFFERENTIATION, DEDIFFERENTIATION AND
REDIFFERENTIATION
DIFFERENTIATION
Cell loses its ability to divide.
Cells derived from root apical, shoot apical meristem and cambium diferentiate
and mature to perform specific functions.
Structural changes take place in cell wall and protoplasm. eg. Tracheary elements
DEDIFFERENTIATION
Differentiated cells regain the capacity to divide under certain conditions.
e.g. Formation of secondary meristem (inter fascicular and cork cambium) from
fully differentiated parenchyma cells.
REDIFFERENTIATION
Dedifferentiated cells once again lose the capacity to divide and mature to perform
specific function.
e.g. List of tissues in woody dicotyledonous plants that are the products of
redifferentiation.
Secondary xylem Cork or phellem
Secondary phloem Secondary cortex
Differentiation in plants is open because cells/tissues arising out of the same meristem
have different structures at maturity. The final structure is also determined by the
location of the cell within.
e
.g. Cells positioned away from root apical meristem differentiate as root cap cells,
while those pushed to the periphery mature as epidermis.
DEVELOPMENT
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.
Cell Division Death
Senescence
Plasmatic growth Differentiation
Meristematic cell
Mature cell
Expansion Maturation
(Elongation)
Sequence of the development process in a plant cell
Development = Growth + Differentiation
PLASTICITY
The ability of plants to follow different pathways in response to environment or phases
of life to form different structure (NEET 2021).
Example: Heterophylly in cotton, coriander and larkspur, in different phases of life.
Different shapes of leaves produced in air and those produced in water in Buttercup
(Ranunculus).
Juvenile Adult Terrestrial habitat Water habitat
(a) (b)
Heterophylly in (a) larkspur and (b) in buttercup
FACTORS CONTROLLING THE DEVELOPMENT
INTRINSIC FACTORS: Include intracellular (genetic) or intercellular factors (such as plant
growth regulators).
EXTRINSIC FACTORS: Include light, temperature, water, oxygen, nutrition, etc.
PLANT GROWTH REGULATORS (PLANT HORMONES OR
PHYTOHORMONES)
Auxins
Growth Promoters Gibberellins
Cytokinins
Plant Hormones
Ethylene (also acts as
Growth Inhibitors growth promoter)
ABA
Hormone Name Chemical Composition Other Names
1. Auxin Indole compounds IAA, IBA (Natural); NAA,
2,4-D (Synthetic)
2. Cytokinin Adenine derivatives (N6- Zeatin (Natural), Kinetin
furfurylamino purine) (Not found naturally in
plants)
3. Gibberellin Terpenes GA1, GA2, GA3, (one of the
first to be discovored)
GA4 ... (more than 100)
4. Ethylene Gas (C2H4) Ethephon (aqueous
solution)
5. Abscisic Acid Carotenoids derivatives ABA
(AIPMT 2009)
Hormone Name Isolation by and From Discovery (Accidental
for all hormones)
1. Auxin F.W. Went Frances Darwin
Coleoptiles tips of oat and Charles Darwin
seedlings (AIPMT 2014). (Phototropism of
First isolated from human coleoptile tip) of canary
urine. grass (AIPMT 2012)
2. Cytokinin Kinetin : From the autoclaved F. Skoog and Co-workers
herring sperm DNA (not (Internodal segments of
naturally in plants) tobacco stems; the callus
Zeatin : Extracts of ‘vascular proliferated only when
tissues’, ‘yeast extract’, auxin was supplemented
‘coconut milk’, ‘corn-kernels’ with cytokinin)
3. Gibberellin Sterile filtrates of the fungus E. Kurosawa
‘Gibberela fujikuroi’ (Fungal pathogen
Gibberela fujikuroi
causing ‘bakanae’
(foolish seedling)
disease of rice seedling.
4. Ethylene Volatile substance released H.H. Cousins
from ‘ripened fruits’. (Volatile substance from
ripened orange hasten
the ripening of stored
unripened banana.
5. Abscisic Acid Stress Hormone (AIPMT 3 independent
2014) researchers
(3 different kinds of
inhibitors : Inhibitor-B,
Abcission-II, Dormin.
All were chemically
identical.
(Decapitation is done for Hedge marking and in tea plantation (AIPMT 2014)
(i) Apical Dominance
(vii) Weedicide for dicot
(ii) Cell division and
(e.g. 2, 4-D)
enlargement
(NEET 2021)
(vi) Parthenocarpy in AUXINS
(iii) Root initiation
tomato
(v) Flower initiation in (iv) Prevent leaf and fruit drop
pineapple and litchi at early stages but promote
(Karnatka NEET 2023) abscission of older mature
leaves and fruits (NEET-I 2017)
(i) Stem/Internode elongation (NEET 2023)
(ii) Elongation of
(vii) Bolting in beet
genetically Dwarf
and cabbage
plant
GIBBERELLINS
(vi) Hastens maturity (iii) Promotes seed
period of conifers germination
(NEET 2020, 2013)
(v) Delays senescence of fruits (iv) Increase height of
sugarcane stem (NEET 2020)
and hence increase in yield
• GA3 is used to speed up the malting process in brewing industry.
• GA3 increase the length of grapes stalks and cause fruits like apple to elongate
and improve its shape.
(i) Promotes Cell-Division
(v) Helps to produce new
leaves and chloroplast
(ii) Morphogenesis
(NEET-II 2016)
Low CK
⇒ Root
High Auxin
CYTOKININS
High CK
⇒ Shoot
Lo w Aux in
(NEET-II 2016)
(iii) Overcomes apical dominance
(iv) Delay leaf senescence
and promotes lateral growth
by promoting nutrient
and adventitious shoot
mobilisation
formation
(ii) Promotes senescence and abscission of fruits
and flower (Thinning of cotton, walnut and cherry)
(i) Ripening of fruits
(iii) Flowering and (NEET 2019)
(Respiratory
synchronising fruiting in
climactic)
pineapple and also induces
flowering in mango.
(Karnatka NEET 2023)
(viii) Internode/petiole
ETHYLENE (iv) Promotes root growth
elongation in deep
and root hair formation
water rice plants
(NEET 2022)
(NEET 2023)
(v) Triple response on stem
(vii) Breaks seed dormancy in (a) apical hook formation in
peanuts and bud dormancy dicot seedlings
in potato (b) Swelling of axis
(c) Horizontal growth of
seedlings
(i) Induces senescence and abscision
ABSCISIC (ii) Plays an important
(iv) Helps seed to withstand role in seed development,
dessication and other ACID
maturation and dormancy
factors unfavourable (ABA)
for growth
(iii) Closes stomata
• ABA acts as a general plant growth inhibitor.
• ABA acts as an antagonist to GAs (Gibberellins) (AIPMT 2012)