STEP in BIOLOGY

Chapter # 11 Bioenergetics
Topic
Bioenergetics
▪ Bioenergetics is the quantitative study of energy relationships and energy conversions in biological systems.
▪ Biological energy transformations obey the laws of thermodynamics.

Role of Solar Energy:

▪ All organisms need free energy for keeping themselves alive and functioning.
▪ All life on this planet Earth is powered, directly or indirectly, by solar energy.
▪ No organism can make direct use of sunlight as source of energy for metabolism.
▪ All can use chemical energy in the food such as sugars etc.
▪ The chloroplasts of the plants capture light energy coming from the sun and convert it into chemical energy
that gets stored in sugar and then in other organic molecules.
▪ The process of photosynthesis helps to understand some of the principles of energy transformation
(Bioenergetics) in living systems.
▪ Photosynthetic organisms (higher land plants for instance) use solar energy to synthesize organic compounds
(such as carbohydrates) that cannot be formed without the input of energy.
▪ Energy stored in these molecules can be used later to power cellular processes and can serve as the energy
source for all forms of life.

Role of Photosynthesis:

▪ With the emergence of photosynthesis on earth, molecular oxygen began to accumulate slowly in the
atmosphere.
▪ The presence of free oxygen made possible the evolution of respiration.
▪ Photosynthesis provides the carbohydrate as substrate for glycolysis and respiration.

Role of Respiration:

▪ Respiration releases great deal of energy, and couples some of this energy to the formation of adenosine
triphosphate (ATP) molecules.
▪ ATP is a kind of chemical link between catabolism and anabolism.
▪ Glycolysis and kreb’s cycle are the processes whereby the energy stored in carbohydrate is released in a
controlled manner.
❖ So the photosynthesis acts as an energy-capturing while respiration as an energy releasing process.

PHOTOSYNTHESIS
(CONVERSION OF SOLAR ENERGY INTO CHEMICAL ENERGY)

Photosynthesis can be defined as,

▪ The process in which energy-poor inorganic oxidized compounds of carbon like C02 and hydrogen mainly water
are reduced to energy-rich carbohydrate like sugar-glucose using the light energy that is absorbed and
converted into chemical energy by chlorophyll and some other photosynthetic pigments.
▪ Oxygen is released as by product.

The process of photosynthesis in green plants can be summarized as:

6CO2 + 12H2O + Light+ Chlorophyll → C6H12O6 + 6H2O + 6O2

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Above equation can be simplified as follows:

6CO2 + 6H2O + Light+ Chlorophyll → C6H12O6 + 6O2

Photosynthetic Reactants and Products:

Reactants of Photosynthesis:

Reactants of photosynthesis are as follows:

1. Carbon dioxide.
2. Water.
3. Light.

Products of Photosynthesis:

Products of photosynthesis are as follows:

1. Glucose.
2. Oxygen.

Status of Water:

▪ Water appears on both sides of the equation because water is used as reactant in some reactions and released
as product in other.
▪ However, because there is no net yield of H20 so water is not written in product side.

We can simplify the summary equation of photosynthesis for purpose of discussion as follows:

6CO2 + 6H2O + light+ chlorophyll →C6H12O6 + 6O2

Compensation Point:

▪ At dawn and dusk, when light intensity is low, the rate of photosynthesis and respiration may, for a short time,
equal one another.
▪ Thus the oxygen released from photosynthesis is just the amount required for cellular respiration.
▪ Also, the carbon dioxide released by respiration just equals the quantity required by photosynthesizing cells.
▪ At this moment there is no net gas exchange between the leaves and the atmosphere.
▪ This is termed as compensation point.

At High Light Intensity:

▪ As the light intensity increases, so does the rate of photosynthesis and hence the requirement for more carbon
dioxide increases which respiration alone cannot supply.
▪ Similarly, the oxygen produced during photosynthesis is more than the need of the respiring cells, so the result
is the net release of oxygen coupled with the uptake of carbon dioxide.

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Water and Photosynthesis
▪ Oxygen released during photosynthesis comes from water, and is an important source of atmospheric oxygen
which most organisms need for aerobic respiration and thus for obtaining energy to live.

Van Niel hypothesis:

▪ In 1930s, Van Neil hypothesized that plants split water as a source of hydrogen, releasing oxygen as a by-
product.
▪ Niel’s hypothesis was based on his investigations on photosynthesis in bacteria that make carbohydrate from
carbon dioxide, but do not release oxygen.
▪ Niel’s hypothesis that source of oxygen released during photosynthesis is water and not carbon dioxide, was
later confirmed by scientists.

Experiment to Test Niel’s Hypothesis:

▪ During 1940s, scientist first used an isotopic tracer (O18) in biological research to test Niel’s hypothesis.
▪ Water and carbon dioxide containing heavy-oxygen isotope O18 were prepared in the laboratory.
▪ Experimental green plants in one group were supplied with H20 containing O18 and with C02containing only
common oxygen O16.
▪ Plants in the second group were supplied with H20 containing common oxygen O16 but with C02 containing O18.

Observations:

▪ It was found that plants of first group produced O18 but the plants of second group did not.

Group-1 Plants: CO2 + 2H2O18 + light+ chlorophyll →CH2O +H2O+ O182

Group-2 Plants: CO182 + 2H2O + light+ chlorophyll →CH2O18+H2O18 + O2

Conclusion:

▪ Hence, it is concluded from above experiment that oxygen comes from water during photosynthesis.

Important Information:

▪ Water is one of the raw materials of photosynthesis, other being carbon dioxide.
▪ Hydrogen produced by splitting of water reduces NADP to NADPH2 (NADPH + H+).
▪ NADPH is the “reducing power” which, along with ATP also formed during light reactions, is used to reduce
C02to form sugar during dark reactions.

CHLOROPLASTS - THE SITES OF PHOTOSYNTHESIS IN PLANTS

Occurrence:

▪ All green parts of a plant have chloroplasts, but the leaves are the major sites of photosynthesis in most plants.
▪ Chloroplasts are present in very large number, about half a million per square millimeter of leaf surface.
▪ Chloroplasts are present mainly in the cells of mesophyll tissue inside the leaf.
▪ Each mesophyll cell has about 20-100 chloroplasts.

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Structure of Chloroplast:

Chloroplast has following structure:

Envelope:

▪ Chloroplast has a double membrane envelope.

Stroma:

▪ Envelope encloses dense fluid-filled region, the stroma which contains most of the enzymes required to
produce carbohydrate molecules.

Thylakoids:

▪ Another system of membranes is suspended in the stroma.

▪ These membranes form an elaborate interconnected set of flat, disc like sacs called thylakoids.
▪ The thylakoid membrane encloses a fluid-filled thylakoid interior space or lumen, which is separated from the
stroma by thylakoid membrane.
▪ Chlorophyll (and other photosynthetic pigments) are found embedded in the thylakoid membranes and impart
green color to the plant.
▪ Electron acceptors of photosynthetic ‘Electron Transport Chain’ are also parts of these membranes.
▪ Thylakoid membranes are thus involved in ATP synthesis by chemiosmosis.

Granum:

▪ In some places, thylakoid sacs are stacked in columns called grana (sing granum).
▪ Chlorophyll (and other pigments) absorbs light energy, which is converted into chemical energy of ATP and
NADPH, the products which are used to synthesize sugar in the stroma of chloroplast.

Fig: A plant possesses thick layer of mesophyll cells rich in chloroplasts. Thylakoids in chloroplasts are stacked into
grana. Light reactions take place on the grana, and dark reactions in the stroma.

Photosynthetic Prokaryotes:
▪ Photosynthetic prokaryotes lack chloroplasts.
▪ They have unstacked photosynthetic membranes which work like thylakoids.

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PHOTOSYNTHETIC PIGMENTS
▪ Photosynthetic pigments are organic compounds that absorb visible light of 380-750 nm in wave length.
▪ Different pigments absorb light of different wave lengths (colors).
▪ The wave lengths that are absorbed disappear.

Types of Pigments:

Thylakoid membranes contain several kinds of pigments as follows:

1. Major Pigments:
▪ Chlorophyll-a is the main photosynthetic pigment.
2. Accessory Pigments:
▪ These broaden the absorption and utilization of light energy.

Accessory photosynthetic pigments present in the chloroplasts include following pigments:

1. Carotenoids:
▪ These are yellow and red to orange.

Carotenoids are of two types:

i. Carotenes:
▪ Carotenes are mostly red to orange.
ii. Xanthophylls:
▪ Xanthophylls are yellow to orange.
2. Chlorophyll-b:
▪ Chlorophyll-b is another accessory photosynthetic pigment.
▪ Chlorophyll b is found along with chlorophyll a in all green plants (embryophytes) and green algae.
▪ Chlorophyll b is yellow-green.

Spectrophotometer:

▪ An instrument called Spectrophotometer is used to measure relative abilities of different pigments to absorb
different wavelengths of light.

Chlorophylls
▪ There are known many different kinds of chlorophylls in prokaryotes and eukaryotes.
▪ Chlorophylls are insoluble in water but soluble in organic solvents, such as carbon tetrachloride, alcohol etc.

Chlorophyll-a:

▪ Of all the chlorophylls, chlorophyll a is the-most abundant and the most important photosynthetic pigment.
▪ It takes part directly in the light-dependent reactions which convert solar energy into chemical energy.
▪ It is found in all photosynthetic organisms except photosynthetic bacteria.
▪ Chlorophyll a is blue-green.
▪ Chlorophyll-a itself exists in several forms differing slightly in their red absorbing peaks e.g. at 670, 680, 690,
700 nm.

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Chlorophyll-b:

▪ Chlorophyll b is found along with chlorophyll a in all green plants (embryophytes) and green algae.
▪ Chlorophyll b is yellow-green.

In Eukaryotes:

▪ Chlorophyll a, b, c and d are found in eukaryotic photosynthetic plants and algae.

In Prokaryotes:

▪ Chlorophylls found in photosynthetic bacteria are known as bacteriochlorophylls.

Light Absorption by Chlorophyll “a”:

Reason of Appearing Plant as Green

▪ Chlorophylls absorb mainly violet-blue and orange-red wave lengths.
▪ Green, yellow and indigo wave lengths are least absorbed by chlorophylls and are transmitted or reflected.
▪ The yellows are often masked by darker green color.
▪ Hence plants appear green, unless masked by other pigments.

Structure of Chlorophyll Molecule:

A chlorophyll molecule has two main parts:

1. Head:
▪ Head is flat, square, light absorbing hydrophilic region.
▪ The head is complex porphyrin ring which is made up of 4 joined smaller pyrrole rings composed of carbon and
nitrogen atoms.
▪ An atom of magnesium is present in the center of porphyrin ring and is coordinated with the nitrogen of each
pyrrole ring.
▪ That is why magnesium deficiency causes yellowing in plants.
▪ Haem portion of hemoglobin is also a porphyrin ring but containing an iron atom instead of magnesium atom
in the center.
2. Tail:
▪ The tail is long, anchoring, hydrophobic hydrocarbon.
▪ Long hydrocarbon tail which is attached to one of the pyrrole rings is phytol (C20 H39).
▪ The chlorophyll molecule is embedded in the hydrophobic core of thylakoid membrane by this tail.

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Fig: A chlorophyll molecule

Differentiate between Chlorophyll a and chlorophyll b:

Sr # Chlorophyll a Chlorophyll b
1 Molecular formula of chlorophyll a is Molecular formula of chlorophyll b is C55
C55 H72 05 N4 Mg. H70 06 N4 Mg.
2 The functional group bounded to The functional group bounded to
porphyrin ring is methyl group. (-CH3) porphyrin ring is carbonyl group. (-CHO)
3 It most abundant of all photosynthetic It is less abundant.
pigments.
4 It takes part directly in light reaction. It transfers its energy to chlorophyll a.
5 It is blue-green. It is yellow-green.

The molecular formulae for chlorophyll a and b are:

Sr # Chlorophyll a Chlorophyll b
1 Molecular formula of chlorophyll a is Molecular formula of chlorophyll b is C55
C55 H72 05 N4 Mg. H70 06 N4 Mg.

Name of pigment Absorption of light Color of Pigment

Chlorophyll-a Violet-blue, Orange-red Green (Blue-Green)
Chlorophyll-b Violet-Blue Yellow (Yellow-Green)
Carotenoids Violet-Blue Yellow/Orange-Red
(Tetra-terpenoids)

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Carotenes Blue-Green Orange-Red
(Hydrocarbons)
Xanthophylls Blue-Green Yellow
(Oxygen)
▪ Blue light is maximum absorbed by photosynthetic pigments.
▪ Red light has maximum effectiveness in photosynthesis.

Carotenoids-Accessory Pigments
▪ Carotenoids are yellow and red to orange pigments that absorb strongly the blue-violet range.
▪ There absorption range has different wave lengths than the chlorophyll absorbs.
▪ So they broaden the spectrum of light that provides energy for photosynthesis.
▪ These and chlorophyll b are called accessory pigments because they absorb light and transfer the energy to
chlorophyll a, which then initiates the light reactions.
▪ It is generally believed that the order of transfer of energy is:

Carotenoids →Chlorophyll b → Chlorophyll a

Importance of Carotenoids:
1. Some carotenoids protect chlorophyll from intense light by absorbing and dissipating excessive light energy,
rather than transferring energy to chlorophyll.
2. Similar carotenoids may be protecting human eye.

LIGHT-THE DRIVING ENERGY

▪ Light is a form of energy called electromagnetic energy or radiation.
▪ Light behaves as waves as well as sort of particles called photons.
▪ The radiations most important to life are the visible light that ranges from about 380 to 750 nm in wave
length.
▪ It is the sunlight energy that is absorbed by chlorophyll, converted into chemical energy, and drives the
photosynthetic process.
▪ Not all the light falling on the leaves is absorbed.
▪ Only about one percent of the light falling on the leaf surface is absorbed, the rest is reflected or transmitted.

Absorption spectrum of Chlorophylls:

▪ Absorption spectrum for chlorophylls indicates that absorption is maximum in blue and red parts of the
spectrum.
▪ They have two absorption peaks being at around 430 nm and 670 nm respectively.

Absorption Spectrum of Carotenoids:

▪ Absorption peaks of carotenoids are different from those of chlorophylls.

▪ Different wavelengths are not only differently absorbed by photosynthetic pigments but are also differently
effective in photosynthesis.

Absorption Spectrum:

▪ A graph plotting absorption of light of different wave lengths by a pigment is called absorption spectrum of the
pigment.
▪ Peaks in absorption spectrum are narrow.

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▪ Valley is broader and deep.

Action Spectrum:

▪ Graph showing relative effectiveness of different wavelengths of light in driving photosynthesis is called action
spectrum of photosynthesis.
▪ Peaks in action spectrum are broader.
▪ Valley is narrower and not deep.

Important Information:
The first action spectrum was obtained by German biologist, T.W. Engelmann in 1883. He worked on Spirogyra.

Differentiate between absorption and action spectrum:

Sr# Absorption Spectrum Action Spectrum

1 Spectrum which shows the relative The graph which shows the relative
absorption of different wavelengths of light effectiveness of different wavelengths of light
by photosynthetic pigments is called in photosynthesis is called action spectrum.
absorption spectrum.
2 In absorption spectrum, peaks are narrower. In action spectrum, peaks are broader.
3 Valleys are broader and deep. Valleys are narrower and not deep.

Procedure to Obtain Action Spectrum:

Action spectrum can be obtained by following ways:

▪ Action spectrum can be obtained by illuminating plant with light of different wavelengths (or colors).
▪ Then estimating relative C02 consumption or oxygen release during photosynthesis.

Fig: Engel man illuminated a filament of Spirogyra with light that had been passed through a prism. Aerobic bacteria
moved toward the portions of the algal filament emitting the most oxygen, along the cells in blue and red portion of
the spectrum.

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Fig: (a) Absorption spectrum of chlorophyll and carotenoids. (b) Action spectrum for photosynthesis.

Some Facts by Comparing Action and Absorption Spectrums:

▪ Action spectrum of photosynthesis corresponds to absorption spectrum of chlorophyll.
▪ The same two peaks and the valley are obtained for absorption of light as well as for CO2 consumption.
▪ This also shows that chlorophyll is the photosynthetic pigment.
▪ However, the action spectrum of photosynthesis does not parallel the absorption spectrum of chlorophyll
exactly.
▪ Photosynthesis in the most absorbed range is more than the absorption itself.
▪ Likewise, photosynthesis in 500-600 nm (including green light) is more than the absorption of green light by
the chlorophyll.
▪ This difference occurs because the accessory pigments, the carotenoids, absorb light in this zone and pass on
some of the absorbed light to chlorophylls which then convert light energy to chemical energy.
▪ When equal intensities of light are given, there is more photosynthesis in red than in blue part of spectrum.

ROLE OF CARBON DIOXIDE:

A PHOTOSYNTHETIC REACTANT

▪ Sugar is formed during light - independent reactions of photosynthesis by the reduction of CO2 using ATP and
NADH.
▪ Obviously photosynthesis does not occur in the absence of CO2.

Occurrence of Photosynthesis:

▪ About 10 percent of total photosynthesis is carried out by terrestrial plants.

▪ The rest of photosynthesis occurs in oceans, lakes and ponds.

Source of Carbon Dioxide for Photosynthesis:

▪ Air contains about 0.03 - 0.04 percent CO2.

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▪ Photosynthesis occurring on land utilizes this atmospheric C02.
▪ Aquatic photosynthetic organisms use dissolved CO2, bicarbonates and soluble carbonates that are present in
water as carbon source.

Entry of Carbon Dioxide:

▪ Carbon dioxide enters the leaves through stomata and gets dissolved in the water absorbed bythe cell walls of
mesophyll cells.
▪ Stomata are found in a large number in a leaf.
▪ Their number being proportional to the amount of gas diffusing into the leaf.
▪ Stomata cover only 1 - 2 percent of the leaf surface but they allow proportionally much more gas to diffuse.
▪ The entry of C02 into the leaves depends upon the opening of stomata.

Control of Opening and Closing of Stomata:

▪ The guard cells guarding the stoma regulate the opening and closing of stomata.
▪ Stomata are adjustable pores that are usually open during the day when CO2 is required for photosynthesis
and partially closed at night when photosynthesis stops.
▪ Daily rhythmic opening and closing of stomata is also due to an internal clock located in the guard cells. Even if
a plant is kept in a dark closet, stomata will continue their daily rhythm of opening and closing.

REACTIONS OF PHOTOSYNTHESIS
Photosynthesis is a ‘redox process’ that can be represented by the following simplified summary equation:

▪ However, it is not a simple, single step process, but is a complex one that is completed by a series of simple
steps or reactions.

These reactions of photosynthesis consist of two parts:

1. Light Reaction:
▪ The light-dependent reactions (light reactions) which use light directly.
▪ Light dependent reactions constitute that phase of photosynthesis during which light energy is absorbed by
chlorophyll and other photosynthetic pigment molecules and converted into chemical energy.
▪ As a result of this energy conversion, reducing and assimilating power in the form of NADPH (NADPH + H+) and
ATP, are formed.
▪ Both temporarily store energy and then to be carried along with H to the light independent reactions.
2. Dark Reaction:
▪ The light-independent reactions (dark reactions) which do not use light directly.
▪ NADPH provides energized electron (and H+) and reducing power.
▪ ATP provides chemical energy for the synthesis of sugar by reducing CO2.
▪ The energy is thus stored in the molecules of sugar.
▪ This phase of photosynthesis is also called dark reactions because these reactions do not use light directly and
can take place equally well both in light and dark provided NADPH2 and ATP of light reactions are available.

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Fig: An overview of photosynthesis: Light - Dependent Reactions (Energy - conversion) and Light - Independent
Reactions (Energy - conservation)

Light Dependent Reactions

(Energy-conversion phase; formation of ATP and NADPH)

Photosystems:
▪ Photosynthetic pigments are organized into clusters called photosystems.

Role:

▪ They are used for efficient absorption and utilization of solar energy in thylakoid membranes

Fig: Photosystem

Components of Photosystem:

Each photosystem consists of following parts:

1. A light-gathering antenna complex.

Composition:

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▪ The antenna complex has many molecules of chlorophyll a, chlorophyll b and carotenoids.

Role:

▪ Most of pigments transfer the energy to reaction center.

2. Reaction Center:

Composition:

▪ Reaction center has one or more molecules of chlorophyll a along with a primary electron acceptor, and
associated electron carriers of ‘electron transport system’.
▪ Chlorophyll a molecules of reaction center and associated proteins are closely linked to the nearby electron
transport system.
▪ A specialized molecule called primary electron acceptor is also associated nearby each reaction center.

Role:

▪ Electron transport system plays role in generation of ATP by chemiosmosis.

▪ Light energy absorbed by the pigment molecules of antenna complex is transferred ultimately to the reaction
center.
▪ There the light energy is converted into chemical energy.

Types of Photosystems:

There are two photosystems:

1. Photosystem I (PS I):

▪ Photosystem I has chlorophyll a molecule which absorbs maximum light of 700 nm and is called P700.
▪ It is discovered earlier so named as photosystem-I.
▪ It comes later non-cyclic photophosphorylation.
2. Photosystem II (PS II):
▪ Photosystem II has chlorophyll a molecule which absorbs maximum light of 680 nm and is called P700.
▪ It is discovered later so named as photosystem-II.
It comes earlier in non-cyclic photophosphorylation.

Differentiate between photosystem-I and photosystem-II:

Sr # Photosystem-I Photosystem-II
1 Photosystem I has chlorophyll a Photosystem II has chlorophyll a molecule
molecule which absorbs maximum light which absorbs maximum light of 680 nm and
of 700 nm and is called P680. is called P700.

2 It is discovered earlier. It is discovered later.

3 It is used later in non-cyclic It is used earlier.
photophosphorylation.

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Non-cyclic Phosphorylation
It has following steps:

1. Photoexcitation of PSII:
▪ When photosystem II absorbs light, an electron from P680 is excited to a higher energy level in the reaction
center.
▪ It is captured by the primary electron acceptor of PS II.
▪ The oxidized chlorophyll is now a very strong oxidizing agent.
▪ Its electron “hole” must be filled.
2. Photolysis of Water:
▪ The hole of PSII is filled by the electrons which are extracted by an enzyme from water.
▪ This reaction splits a water molecule into two hydrogen ions and an oxygen atom.
▪ This oxygen immediately combines with another oxygen atom to form O2.
▪ The oxygen produced during photolysis is the main source of replenishment of atmospheric oxygen.
▪ This water splitting step of photosynthesis that releases oxygen is called photolysis.
3. Transportation of Photoexcited Electron:
▪ Each photoexcited electron passes from the primary electron acceptor of photosystem II to photosystem I via
an electron transport chain.
▪ This chain consists of an electron carrier called plastoquinone (Pq), a complex of two cytochromes and a
copper containing protein called plastocyanin (Pc).
4. Photophosphorylation:
▪ Formation of ATP by movement of photoexcited electron is called photophosphorylation.
▪ As electrons move down the chain, their energy goes on decreasing and is used by thylakoid membrane to
produce ATP.
▪ This ATP synthesis is called photophosphorylation because it is driven by light energy.
▪ Specifically, ATP synthesis during non-cyclic electron flow is called non-cyclic photophosphorylation.
▪ This ATP generated by the light reactions will provide chemical energy for the synthesis of sugar during the
Calvin cycle, the second major stage of photosynthesis.
5. Photoexcitation of PS-I:
▪ The hole is created in P700 by absorbing light.
▪ The electron from PS-II reaches the “bottom” of the electron transport chain and fills an electron “hole” inP700.
6. Formation of NADPH2:
▪ The primary electron acceptor of photosystem I passes the photoexcited electrons to a second electron
transport chain which transmits them to ferredoxin (Fd), an iron containing protein.
▪ An enzyme called NADP reductase then transfers the electrons from Fd to NADP.
▪ This is the redox reaction that stores the high-energy electrons in NADPH.
▪ The NADPH molecule will provide reducing power for the synthesis of sugar in the Calvin cycle.
▪ The path of electrons through the two photosystems during non-cyclic photophosphorylation is known as Z-
scheme from its shape.

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Fig: Non-cyclic electron low during photosynthesis ATP, NADPH and oxygen are generated. The arrows trace the
current of light-driven electrons from water to NADPH. Each photon of light excites single electron, but the diagram
tracts two electrons at a time, the number of electrons required to reduce NADP+. The numbered steps are described
in the text.

Fig: Cyclic flow of electron

Differentiate between non-cyclic and cyclic photophosphorylation:

Sr # Non-Cyclic Cyclic Photophosphorylation

Photophosphorylation
1 Formation of ATP by the movement of Formation of ATP by the movement of
photoexcited electron in non-cyclic way photoexcited electron in cyclic way is called
is called non-cyclic cyclic photophosphorylation.
photophosphorylation.
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2 ATP and NADPH both are formed. Only ATP is formed.
3 Oxygen is released. Oxygen is not released.
4 Photolysis of water occurs. Photolysis of water does not occur.
5 Both photosystems are involved. Only PS-I is involved.
6 It is normally occurring process. It occurs only when system is running at low
ATP.

Chemiosmosis
▪ The process that uses membranes to couple redox reactions to ATP production during which electron
transport chain pumps protons (H+) across the membrane of thylakoids in case of photosynthesis into the
thylakoids space is called chemiosmosis.
▪ The energy used for this pumping comes from the electrons moving through the electron transport chain.
▪ This energy is transformed into potential energy stored in the form of H+ gradient across the membrane.
▪ Next the hydrogen ions move down their gradient through special complexes called ATP synthase which are
built in the thylakoid membrane.
▪ During this diffusion of H+ the energy of electrons is used to make ATP.

Fig: Electron Transport chain and chemiosmosis, coupling of ETC and formation of ATP by chemiosmosis.

Light independent (or Dark) Reactions

Calvin cycle:

▪ Dark reactions are light independent reactions in which there is carbon fixation, reduction and synthesis of
sugar occurs.
▪ The dark reactions take place in the stroma of chloroplast.
▪ These reactions do not require light directly and can occur in the presence or absence of light provided the
assimilatory power in the form of ATP and NADPH, produced during light reactions is available.
▪ Energy of these compounds is used in the formation of carbohydrates from C02.

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These reactions can be summarized as follows:

3CO2 + 6NADPH + 9ATP → (CH 2 O)3 + 6NADP + 9ADP + 9Pi +3H 2O

▪ The details of path of carbon in these reactions were discovered by Melvin Calvin and his colleagues at the
University of California.
▪ Calvin was awarded Nobel Prize in 1961.

Calvin Cycle.
▪ The cyclic series of reactions, catalyzed by respective enzymes, by which the carbon is fixed and reduced
resulting in the synthesis of sugar during the dark reactions of photosynthesis is called Calvin cycle.

The Calvin cycle can be divided into three phases:

Phase-1: Carbon Fixation:

▪ Carbon Fixation refers to the initial incorporation of CO2 into organic material.
▪ Following three molecules of CO2 through the reaction one molecule of carbohydrate, a triose is produced.
▪ The Calvin cycle begins when a molecule of CO2 reacts with a highly reactive phosphorylated five – carbon
sugar named ribulose bisphosphate (RuBP).
▪ This reaction is catalyzed by the enzyme ribulosebisphosphate carboxylase, also known as Rubisco which is the
most abundant protein in chloroplasts, and probably the most abundant protein on Earth.
▪ The product of this reaction is a highly unstable, six - carbon intermediate that immediately breaks into two
molecules of three - carbon compound called 3 - phosphoglycerate (phosphoglyceric acid-PGA).
▪ The carbon that was originally part of CO2 molecule is now a part of an organic molecule so the carbon has
been “fixed”.

Calvin Cycle as C3 Pathway:

▪ Because the product of initial carbon fixation is a three - carbon compound, the Calvin cycle is also known as
C3pathway.

Phase-2: Reduction:

▪ Each molecule of (PGA) receives an additional phosphate from ATP of light reaction forming 1,3 -
bisphosphoglycerate as the product.
▪ 1,3 bisphosphoglycerate is reduced to glyceraldehyde 3-phosphate(G3P) by receiving a pair of electrons
donated from NADPH of light reactions.
▪ G3P is the same three-carbon sugar which is formed in glycolysis (first phase of cellular respiration) by the
splitting of glucose.
▪ In this way fixed carbon is reduced to energy rich G3P with the energy and reducing power of ATP and NADPH
(both the products of light-dependent reactions), having the energy stored in it.
▪ Actually G3P, and not glucose, is the carbohydrate produced directly from the Calvin cycle.
▪ For every three molecules of CO2 entering the cycle and combining with 3molecules of five-carbon RuBP, six
molecules of G3P (containing 18 carbon in all) are produced.
▪ But only one molecule of G3P can be counted as a net gain of carbohydrate.
▪ Out of every six molecules of G3P formed, only one molecule leaves the cycle to be used by the plant for
making glucose, sucrose starch or other carbohydrates, and other organic compounds.
▪ The other five molecules are recycled to regenerate the three molecules of five-carbon RuBP, the CO2
acceptor.

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Phase-3: Regeneration of CO2 Acceptor, RuBP:

▪ Through a complex series of reactions, the carbon skeletons of five molecules of three-carbon G3P are
rearranged into three molecules of five-carbon ribulose phosphate (RuP).
▪ Each RuP is phosphorylated to ribulose bisphosphate (RuBP), the very five-carbon C02 acceptor with which the
cycle started.
▪ Again, three more molecules of ATP of lightreactions are used for this phosphorylation of three RuP molecules.
▪ These RuBP are now prepared to receive C02 again, and the cycle continues.

Fig: The Calvin cycle occurs in stroma of chloroplast. Carbon is fixed and reduced to sugar.

RESPIRATION
▪ Living organisms need energy to carry on their vital activities.
▪ This energy is provided from within the cells by the phenomenon of respiration.
▪ Respiration is the universal process by which organism breakdown complex compounds containing carbon in a
way that allows the cells to harvest a maximum of usable energy.

In biology the term respiration is used in two following ways:

1. External Respiration:
▪ The exchange of respiratory gases (CO2 and O2) between the organism and its environment is called external
respiration.
2. Cellular Respiration:

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▪ The cellular respiration is the process by which energy is made available to cells in a step by step breakdown of
C-chain molecules in the cells.

Aerobic and Anaerobic Respiration:

▪ The most common fuel used by the cell to provide energy by cellular respiration is glucose.
▪ The way glucose is metabolized depends on the availability of oxygen.
▪ Prior to entering a mitochondrion, the glucose molecule is split to form two molecules of pyruvic acid.
▪ This reaction is called glycolysis (glycolysis literally means splitting of sugar), and occurs in the cytosol and is
represented by the equation:

▪ This reaction occurs in all the cells and biologists believe that an identical reaction may have occurred in the
first cell that was organized on earth.
▪ The next step in cellular respiration varies depending on the type of the cell and the prevailing conditions.

Processing of Pyruvic Acid:

▪ Cell processes pyruvic acid in three major ways, alcoholic fermentation, lactic acid fermentationand aerobic
respiration as follows:

▪ The first two reactions occur in the absence of oxygen and are referred to as anaerobic (without oxygen).
▪ The complete breakdown of glucose molecule occurs only in the presence of oxygen called aerobic respiration.
▪ During aerobic respiration glucose is oxidized to CO2 and water and energy is released.

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Anaerobic Respiration:
▪ The process by which pyruvic acid is broken down in the absence of oxygen is called anaerobic respiration.
▪ It is also called fermentation.

Fermentation has following two types;

1. Alcoholic Fermentation:
▪ The process by which pyruvic acid is further broken down into alcohol (C2 H5 OH) and CO2is called alcoholic
fermentation.
▪ It occurs in primitive cells and in some eukaryotic cells such as yeast.

2. Lactic acid Fermentation:

▪ In lactic acid fermentation, each pyruvic acid molecule is converted into lactic acid C3 H6 O3 in the absence of
oxygen gas.
▪ This form of anaerobic respiration occurs in muscle cells of humans and other animals during extreme physical
activities, such as sprinting, when oxygen cannot be transported to the cells as rapidly as it is needed.

Differentiate between alcoholic and lactic acid fermentation:

Sr # Alcoholic Fermentation Lactic acid Fermentation

1 Break down of pyruvic acid into alcohol Break down of pyruvic acid into lactic acid in
and carbon dioxide in the absence of the absence of oxygen is called lactic acid
oxygen is called alcoholic fermentation. fermentation.
2 It takes place in some primitive cells and It takes place in muscle cells of human at
some eukaryotic cells like yeasts. extreme stress and some other cells.

Limitations of Fermentation:

▪ Both alcoholic and lactic acid fermentations yield relatively small amounts of energy from glucose molecule.

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▪ Only about 2% of the energy present within the chemical bonds of glucose is converted into adenosine
triphosphate (ATP).

Aerobic respiration:
Role of Mitochondria in Respiration:

▪ Mitochondria are large granular or filamentous organelles that are distributed throughout the cytoplasm of
animal and plant cells.
▪ Each mitochondrion is constructed of an outer enclosing membrane and an inner membrane with elaborate
folds or cristae that extend into the interior of the organelle.
▪ Mitochondria play a part in cellular respiration by transferring the energy of the organic molecules to the
chemical bonds of ATP.
▪ A large “battery” of enzymes and coenzymes slowly release energy from the glucose molecules.
▪ Thus, mitochondria are the “Power houses” that produce energy necessary for many cellular functions.

Adenosine Triphosphate and its Importance:

▪ Adenosine triphosphate, generally abbreviated ‘ATP’ is a compound found in every living cell and is one of the
essential chemicals of life.
▪ It plays the key role in most biological energy transformations.

High Energy Bonds in ATP:

▪ Conventionally, ‘P’ stands for the entire phosphate group.

▪ The second and the third phosphate represent the so called “high energy” bonds.
▪ If these are broken by hydrolysis, far more free energy is released as compared to the other bond in the ATP
molecule.
▪ The breaking of the terminal phosphate of ATP releases about 7.3 K cal. of energy.
▪ The high energy ‘P’ bond enables the cell to accumulate a great quantity of energy in a very small space and
keeps it ready for use as soon as it is needed.

Role of ATP:

▪ The ATP molecule is used by cells as a source of energy for various functions for example, synthesis of more
complex compounds, active transport across the cell membrane, muscular contraction, and nerve conduction,
etc.

Biological Oxidation:
▪ The maintenance of living system requires a continual supply of free energy which is ultimately derived from
various oxidation reduction reactions.

Example:

▪ Cellular respiration is essentially an oxidation process.

Exception in Extraction of Energy:

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▪ Except for photosynthetic and some bacterial chemosynthetic processes, which are themselves oxidation
reduction reactions, all other cells depend ultimately for their supply of free energy on oxidation reactions in
respiratory processes.

Dehydrogenase:
▪ In some cases, biological oxidation involves the removal of hydrogen, a reaction catalyzed by the
dehydrogenases linked to specific coenzymes.

Cellular Respiration:
Cellular respiration may be sub-divided into 4 stages:

1. Glycolysis.
2. Pyruvic acid oxidation.
3. Krebs cycle or citric acid cycle.
4. Respiratory chain
▪ Out of these stages the first occurs in the cytosol for which oxygen is not essential.
▪ Other three occur within the mitochondria where the presence of oxygen is essential.

Glycolysis:
▪ Glycolysis is the breakdown of glucose upto the formation of pyruvic acid.
▪ Glycolysis can take place both in the absence of oxygen (anaerobic condition) or in the presence of oxygen.
▪ In both, the end product of glucose breakdown is pyruvic acid.
▪ The breakdown of glucose takes place in a series of steps, each catalyzed by a specific enzyme.
▪ All these enzymes are found dissolved in the cytosol.
▪ In addition to the enzymes, ATP and coenzyme NAD (nicotinamide adenine dinucleotide) are also essential.

Glycolysis can be divided into two phases:

1. Preparatory Phase:
▪ In the preparatory phase breakdown of glucose occurs.
▪ Energy is expended.

Steps of preparatory phase are as follows:

Step-1:

▪ The first step in glycolysis is the transfer of a phosphate group from ATP to glucose.
▪ As a result, a molecule of glucose-6 -phosphate is formed.

Step-2:

▪ An enzyme catalyzes the conversion of glucose-6-phosphate to its isomer, fructose-6 - phosphate.

Step-3:

▪ At this stage another ATP molecule transfers a second phosphate group.

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▪ The product is fructose 1,6-bisphosphate.

Step-4 & 5:

▪ The next step in glycolysis is the enzymatic splitting of fructose 1 ,6 -bisphosphate into two fragments.
▪ Each of these molecules contains three carbon atoms.
▪ One is called 3 - phospho- glyceraldehyde, 3-PGAL or Glyceraldehyde 3-phosphate (G3P) while the other is
dihydroxyacetone phosphate.
▪ These two molecules are isomers and in fact, are readily interconverted by yet another enzyme of glycolysis.
2. Oxidative Phase/Payoff Phase:
▪ In the oxidative phase high energy phosphate bonds are formed:
▪ Energy is stored.

Oxidative or payoff phase has following steps:

Step-6:

▪ The next step in glycolysis is crucial to this process.

▪ Two electrons or two hydrogen atoms are removed from the molecule of 3- phosphoglyceraldehyde (PGAL)
and transferred to a molecule of NAD.
▪ This is of course, an oxidation-reduction reaction, with the PGAL being oxidized and the NAD being reduced.
▪ During this reaction, a second phosphate group is donated to the molecule from inorganic phosphate present
in the cell.
▪ The resulting molecule is called 1,3 Bisphosphoglycerate(BPG).
▪ The oxidation of PGAL is an energy yielding process.
▪ Thus a “high energy” phosphate bond is created in this molecule.

Step-7:

▪ At the very next step in glycolysis this phosphate group is transferred to a molecule of adenosine diphosphate
(ADP) converting it into ATP.
▪ The end product of this reaction is 3-phospho glycerate (3-PG).

Step-8:

▪ In the next step 3-PG is converted to 2-Phosphoglycerate (2PG).

Step-9:

▪ From 2PG a molecule of water is removed and the product is phosphoenol pyruvate (PEP).

Step-10:

▪ PEP then gives up its ‘high energy’ phosphate to convert a second molecule of ADP to ATP.
▪ The product is pyruvate, pyruvic acid (C3 H4 O3).
▪ It is equivalent to half glucose molecule that has been oxidized to the extent of losing two electrons (as
hydrogen atoms).

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Fig: Glycolysis

Pyruvic Acid Oxidation:

▪ Pyruvic acid (pyruvate), the end product of glycolysis, does not enter the Krebs cycle directly.
▪ The pyruvate (3- carbon molecule) is first changed into 2-carbonacetic acid molecule.
▪ One carbon is released as CO2 (decarboxylation).
▪ Acetic acid on entering the mitochondrion unites with coenzyme-A (Co A) to form acetyl Co A (active acetate).
▪ In addition, more hydrogen atoms are transferred to NAD.

Krebs Cycle or Citric Acid Cycle:

▪ Acetyl CoA now enters a cyclic series of chemical reactions during which oxidation process is completed.
▪ This series of reactions is called the Krebs cycle (after the name of the biochemist who discovered it), or the
citric acid cycle.

Following steps are involved in Krebs cycle:

Step-1:

▪ The first step in the cycle is the union of acetyl CoA with oxaloacetate to form citrate.
▪ In this process, a molecule of CoA is regenerated and one molecule of water is used.
▪ Oxaloacetate is a 4-carbon acid.
▪ Citrate thus has 6 carbon atoms.

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Step-2:

▪ Citrate is converted into isocitrate.

Step-3:

▪ In isocitrate another NAD- mediated oxidation takes place.

▪ This is accompanied by the removal of a molecule of CO2.
▪ The result is α-ketoglutarate.

Step-4:

▪ It, in turn, undergoes further oxidation (NAD + 2H —> NADH) followed by decarboxylation (CO2) and addition
of a molecule of water.
▪ The product then has one carbon atom and one oxygen atom less.
▪ It is succinate.
▪ The conversion of α-ketoglutarate into succinate is accompanied by a free energy change which is utilized in
the synthesis of an ATP molecule.

Step-5:

▪ The next step in the Krebs cycle is the oxidation of succinate to fumarate.
▪ Once again, two hydrogen atoms are removed, but this time the oxidizing agent is a coenzyme called flavin
adenine dinucleotide(FAD), which is reduced to FADH2.

Step-6:

▪ With the addition of another molecule of water, fumarate is converted to malate.

Step-7:

▪ Another NAD mediated oxidation of malate produces oxaloacetate, the original 4-carbon molecule.
▪ This completes the cycle.
▪ The oxaloacetate may now combine with another molecule of acetyl CoA to enter the cycle and the whole
process is repeated.

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Fig: Krebs Cycle

Respiratory Chain:
▪ In the Krebs cycle NADH and H+ are produced from NAD+.
▪ NADH then transfers the hydrogen atom to the respiratory chain (also called electron transport system) where
electrons are transported in a series of oxidation-reduction steps to react, ultimately, with molecular oxygen.

The oxidation reduction substances which take part in respiratory chain are:

i. A coenzyme called coenzyme Q

ii. A series of cytochrome enzymes (b,c,a,a3)
iii. Molecular oxygen (02)
▪ Cytochromes are electron transport intermediates containing haem of related prosthetic groups that undergo
valency changes of iron atom.
▪ Haem is the same iron containing group that is oxygen carrying pigment in hemoglobin.

The path of electrons in the respiratory chain appears to be as follows:

i. NADH is oxidized by coenzyme Q.

ii. This oxidation yields enough free energy to permit the synthesis of a molecule of ATP from ADP and inorganic
phosphate.
iii. Coenzyme Q is in turn oxidized by cytochrome b:
iv. Cytochrome b is then oxidized by cytochrome c.
v. This step also yields enough energy to permit the synthesis of a molecule of ATP.
vi. Cytochrome c then reduces a complex of two enzymes called cytochrome a and a3. (for convenience the
complex is referred as cytochrome a).
vii. Cytochrome a is oxidized by an atom of oxygen and the electrons arrive at the bottom end of the respiratory
chain.
viii. Oxygen is the most electronegative substance and the final acceptor of the electrons.

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ix. A molecule of water is produced.
x. In addition, this final oxidation provides enough energy for the synthesis of a third molecule of ATP.

Fig: The respiratory electron transport chain and its coupling with oxidative phosphorylation.

Oxidative Phosphorylation:
▪ Synthesis of ATP in the presence of oxygen is called oxidative phosphorylation.
▪ Normally, oxidative phosphorylation is coupled with the respiratory chain.
▪ ATP is formed in three steps of the respiratory chain.

The equation for this process can be expressed as follows:

NADH + H+ + 3ADP + 3Pi + 1/2 O2 →NAD+ + H2O + 3ATP

▪ Where Pi is inorganic phosphate.

▪ The molecular mechanism of oxidative phosphorylation takes place in conjunction with the respiratory chain in
the inner membrane of the mitochondrion.
▪ Here also, as in photosynthesis, the mechanism involved is chemiosmosis by which electron transport chain is
coupled with synthesis of ATP.
▪ In this case, however the pumping/movement of protons (H+) is across the inner membrane of mitochondrion
folded into cristae, between matrix of mitochondrion and mitochondrion’s inter membrane space.
The coupling factors in respiration are also different from those in photosynthesis.

Differentiate between photophosphorylation and oxidative phosphorylation:

Sr # Photophosphorylation Oxidative Phosphorylation

1 Formation of ATP by the movement of Formation of ATP in the presence of oxygen is
photoexcited electron moving through called oxidative phosphorylation.
ETC of photosynthesis is called
photophosphorylation.
2 It occurs during movement of electron It occurs during the movement of electron
through transport chain of light through transport chain of respiration.

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reaction.
3 It occurs in chloroplast. It occurs in mitochondria.
4 Usually one ATP is generated. Usually three ATPs are generated.

Differentiate between photosynthesis and respiration:+

Sr # Photosynthesis Respiration
1 Carbon dioxide is absorbed and oxygen Carbon dioxide is released and oxygen is
is released. absorbed.

2 It occurs only in green parts of plants. It occurs in all living cells.

3 It occurs only at day time. It occurs all the time.
4 It is anabolic process. It is catabolic process.

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