Photosynthesis 1

Photosynthesis:
Photosynthesis is the biochemical process in autotrophs in which energy poor
inorganic substances i.e. water and carbon dioxide are reduced energy rich organic
substance carbohydrate (sugar-glucose) using light energy that is absorbed and converted
into chemical energy by chlorophyll and some other photosynthetic pigments. The
process can be summarized as follows.

6CO2 + 12H2O + light Chlorophyll C6H12O6 + 6O2 + 6H2O

Photosynthesis vs. Respiration:

Look at the reactions of both phenomena. You will observe that both are reverse to each
other.
Photosynthetic reaction:
6CO2 + 12H2O + light Chlorophyll C6H12O6 + 6O2 + 6H2O

Respiration:
C6H12O6 + 6O2 + 6H2O 6CO2 + 12H2O + Energy

 They are almost opposite to each other.

 Photosynthesis uses the products of respiration and respiration uses the products
of photosynthesis.
 Photosynthesis occurs only during daytime, whereas respiration goes on day and
night.
 Photosynthesis acts as an energy-capturing while respiration as an energy
releasing process.

Compensation point:
The compensation point is the amount of light intensity on the light curve
where the rate of photosynthesis exactly matches the rate of respiration. At this point, the
uptake of CO2 through photosynthetic pathways is exactly matched to the respiratory
release of carbon dioxide, and the uptake of O2 by respiration is exactly matched to the
photosynthetic release of oxygen.
This point is reached during early mornings and late evenings. Respiration is relatively
constant whereas photosynthesis depends on the amount of sunlight. When the rate of
photosynthesis equals the rate of respiration, compensation point occurs.
At the compensation point, the rate of photosynthesis is balanced to the rate of respiration
so that the plant is neither consuming nor building biomass. For aquatic plants where the
level of light at any given depth is roughly constant for most of the day, the compensation
point is the depth at which light penetrating the water creates the same balanced effect.

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Photosynthesis 2
Water and photosynthesis:
Water is the source of oxygen in photosynthetic phenomena. This was proposed by Van
Neil in 1931. He discovered that the source of oxygen in photosynthesis was from the
water……. Not the CO2 gas
Later on scientists in 1940s proved this by performing an experiment.
Two groups’ plants were used in the experiment. In the 1st group of plants water was
supplied with the heavy-oxygen isotope (O18) and in the 2nd group of plants
Carbon dioxide was provided with the heavy-oxygen isotope. It was observed that first
group produced O18 but the 2nd group did not.

Group-1 plants: CO2 + 2H2O18 CH2O + H2O + O218

Group-1 plants: CO218 + 2H2O CH2O18 + H2O + O2

SITE OF PHOTOSYNTHESIS:
Chloroplast is the site of photosynthesis in plants.
Leave is the major site of chloroplasts. Chloroplasts are present in very large number,
about half million per square millimeter of leaf surface. Chloroplasts are mainly present
in the cells of messophyll tissues inside the leaf. Each mesophyll cell has about 20-100
chloroplasts.
Structure of chloroplasts:
Chloroplast has double membrane envelope that encloses dense fluid-filled region, the
stroma which contains most of the enzymes required to produce carbohydrate molecules.
Another system of membrane is suspended in ht 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. In some places, thylakoid sacs are stacked in
columns called grana (singular granam). Chlorophyll (and other photosynthetic
pigments) are found embedded in the thylakoid membranes and give green color to the
plant. Electron acceptors of “Electron Transport Chain” are also parts of these
membranes. Thylakoid membranes are thus involved in ATP synthesis by
Chemiosmosis.
Chlorophyll and other pigments absorb 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.

REACTIONS OF PHOTOSYNTHESIS:

By M Awais Shakir
Photosynthesis 3
The reactions of photosynthesis consist of two parts.
1. The light-dependent reaction (light reactions)
2. The light independent reactions (dark reactions)

LIGHT-DEPENDENT REACTIONS:
Light reaction occurs whenever light energy is provided. Light energy is absorbed by
photosynthetic pigments which drives the process of photosynthesis. Photosynthetic
pigments are organized into clusters, called photosystems, for efficient absorption and
utilization of solar energy in thylakoid membranes.
There are two photosystems, photosystem I (PS I) and Photosystem II (PS II). They are
named so in order of their discovery. Photosystem I has chlorophyll “a” molecule which
absorbs maximum light of 700nm and is called P700, whereas photosystem II absorbs
wavelength of 680nm thus called P680.
Each photosystem consists of a light gathering antenna complex and a reaction center.
The antenna complex has many molecules of chlorophyll a, chlorophyll b and
carotenoids, most of them channeling the energy to reaction center. Reaction center has
one or more molecules of chlorophyll a along with 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. 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.

Phosphorylation (Cyclic and Non-Cyclic):

Photosynthetic phosphorylation or photophosphorylation is the process of phosphate
group transfer into ADP to synthesize energy rich ATP molecule making use of light as
external energy source. According to chemi osmotic hypothesis (Mitchell 1961) the ATP
is synthesized on ATPase complexes located on the non appressed portions of thylakoid
membranes particularly towards margins.

Non- Cyclic Phosphorylation (Z- scheme):

In this type the electron passes through both the photosystems. This is the predominant
type of phosphorylation in photosynthesis.
Following steps are taken to complete this stage.

1. Light Absorption:
Photosystem II absorbs light, an electron excited to a higher energy level in the
reaction center chlorophyll P680, Which is captured by the primary electron acceptor
of PSII. The oxidized chlorophyll is now a very strong oxidizing agent; its electron
hole must be filled.
2. Photolysis
This hole is filled by the electrons which are extracted, by an enzyme, from water.
This reaction splits water into two hydrogen ions and an oxygen atom, which
immediately combines with another oxygen atom to form O2. This water splitting
step of photosynthesis that releases oxygen is called photolysis. The oxygen produced
during photolysis is the main source of atmospheric oxygen.

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Photosynthesis 4
3. Transfer of energized electron to PS I:

Each photoexcited electron passes from the primary electron acceptor of PS II to PS I

via electron transport chain. This chain consists of an electron carrier called
plastoquinone (Pq), a complex of two cytochromes and a copper containing protein
called plastocynin (Pc).

4. De- Excitation of electrons:

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. Filling of Hole in P700.

The electron reaches the bottom of the electron transport chain of and fills an electron
hole in P700, the chlorophyll a molecules in the reaction center of PS I. this hole is
created when light energy is absorbed by molecules of P700 and drives an electron
from P700 to the primary electron acceptor of PS I.

6. Formation of NADPH.

The primary electron acceptor of PS 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 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
phosphorylation is known as Z-scheme from its shape.
The diagram of Z-scheme is shown as follows.

By M Awais Shakir
Photosynthesis 5

Cyclic Phosphorylation:
Under certain conditions, photoexcited electrons take an alternative path called cyclic
electron flow. This path uses PS I only. Possibly it happens when the chloroplast runs
low on ATP for Calvin Cycle, the cycle slows down and NADPH accumulate in the
chloroplast. This rise in NADPH may stimulate a temporary shift from non-cyclic to
cyclic electron flow until ATP supply meets the demand. The cyclic flow is short circuit.
The electrons cycle back from primary electron acceptor to ferredoxin (Fd) to the
cytochromes complex and from there continue on to the P700 chlorophyll. ATP is
generated by coupling ETC by Chemiosmosis. There is no production of NADPH and no
release of oxygen. Cyclic flow continues to generate ATP. This is called cyclic
phosphorylation.

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Photosynthesis 6

CHEMIOSMOSIS

Chemiosmosis is the diffusion of ions across a selectively-permeable membrane. More

specifically, it relates to the generation of ATP by the movement of hydrogen ions across
a membrane during cellular respiration.

An Ion gradient has potential energy and can be used

to power chemical reactions when the ions pass
through a channel.

Hydrogen ions (protons) will diffuse from an area

of high proton concentration to an area of lower
proton concentration. Peter Mitchell proposed that
an electrochemical concentration gradient of
protons across a membrane could be harnessed to
make ATP. He likened this process to osmosis, the
diffusion of water across a membrane, which is why
it is called chemiosmosis.

ATP synthase is the enzyme that makes ATP by chemiosmosis. It allows protons to pass
through the membrane using the kinetic energy to phosphorylate ADP making ATP. The
generation of ATP by chemiosmosis occurs in chloroplasts and mitochondria as well as
in some bacteria.

LIGHT INDEPENDENT OR DARK REACTIONS:

Dark reactions take place in the stroma of chloroplast. These reactions don’t require light
directly and can occur in the presence or absence of light provided assimilatory power in

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Photosynthesis 7
the form of ATP and NADPH is available. Energy of these compounds is used in the
formation of carbohydrates from CO2, and thus stored their in. these reactions can be
summarized as follows.

C6H12O6 +12NADP +18ADP +18Pi

12H2O + 12NADP + 18ADP +18Pi +6CO2
+6H2O +6CO2

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 perspective enzymes, by which the carbon is
fixed and reduced, resulting in the synthesis of sugar during dark reactions of
photosynthesis, is called Calvin Cycle.
The Calvin cycle can be divided into three phases.
1. Carbon Fixation.
2. Reduction.
3. Regeneration of CO2 acceptor (RUBP)

PHASE I. Carbon Fixation:

Carbon fixation refers to the initial incorporation of CO2 into organic material. 3
molecules of CO2 are required to produce one molecule of carbohydrate, a triose). The
Calvin cycle begins when a molecule of CO2 reacts with a highly reactive
phosphorylated five-carbon sugar named ribulose biphosphate (RUBP). This reaction is
catalyzed by the enzyme ribulose biphosphate carboxylase, also known as Rubisco.
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 part of CO2
molecule is now a part of an organic molecule: the carbon has bee. “Fixed”. Because the
product of initial carbon fixation is a three-carbon compound, the Calvin cycle is known
as C3 pathway.

PHASE II. Reduction:

Each molecule of (PGA) receives an additional phosphate from ATP of light reaction,
forming 1, 3-biphosphoglycerate as the product. 1, 3-biphosphoglycerate is reduced to
glyceraldehyde 3-phosphate (G3P) by receiving a pair of electrons donated by NADPH
of light reactions. In this way fixed carbon is reduced to G3P with energy and reducing
power of ATP and NADPH (both the products of light-phase), having the energy stored
in it. Actually G3p and not glucose is the carbohydrate produced directly from Calvin
cycle. For every three molecules of CO2 entering the cycle and combining with 3
molecules of five-carbon RUBP, six molecules of G3P (containing 18 carbon in all) are
produced. But only one molecule of G3P and not glucose 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,

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Photosynthesis 8
and other five molecules are recycled to regenerate the three molecules of five-carbon
RUBP, the CO2 acceptor.

PAHSE III. Regeneration of CO2 acceptor, RUBP:

Through a complex series of reactions, the carbon skeleton of five molecules of three-
carbon G3P is rearranged into three molecules of five carbon Ribulose phosphate (RuP).
Each RuP is phosphorylated to ribulose biphosphate (RUBP), the very five-carbon CO2
acceptor with which the cycle started. Again three molecules of ATP of light phase are
used for this phosphorylation of three RuP molecules. These RUBP are now prepared to
receive CO2 again, and the cycle continues.

THANK YOU
Awais Shakir
Batch 6
BSc (Hons)
Environmental Sciences

By M Awais Shakir