Lec. 19 Photosynthesis
Lec. 19 Photosynthesis
Photosynthesis 1
Photosynthesis is a series of chemical reactions initiated by the absorption of
sunlight that causes the flow of electrons which is coupled to the synthesis of ATP,
Light absorption and photophosphorylation NADPH and ultimately reduced organic compounds.
The phosphorylation of ADP to form ATP using the energy of sunlight is called
photophosphorylation.
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The capture of solar energy by photosynthetic organisms and its conversion to
chemical energy of reduced organic compounds is the ultimate source of nearly Photosynthesis encompasses two processes;
biological energy on earth. Light-dependent reactions, or light reactions:
Photosynthetic and heterotrophic organisms live in a balanced steady state in the It occur only when light is present. Chlorophyll and other pigments absorb light
biosphere. energy and conserve it as ATP and NADPH and also evolve O2. occur in thylakoid
Solar energy is the deriving force for the continuous cycling of CO2 and O2 membrane.
through biosphere. Carbon-assimilation reactions or carbon-fixation reaction: Driven by products of
light reactions. ATP and NADPH are used to reduce CO2 to carbohydrates. Occur in
stroma of chloroplast.
Photosynthesis 5 Photosynthesis 6
Photosynthesis takes place in chloroplast: Chloroplast also contains three spaces;
Chloroplast has three membranes Intermembrane space, stroma and thylakoid space (lumen) which is continuous
Outer and inner membrane that together form the chloroplast envelope. connected all the thylakoids lumen.
Thylakoid membrane is highly folded into numerous local stacks of flattened The thylakoids membrane contain two large membrane proteins complexes, called
vesicles called grana, interconnected by nonstacked stroma thylakoids. photosystems. Two other proteins complexes, cytochrome b6-f complex and
chloroplast ATP synthase complex are also present in thylakoid membrane
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Light absorption: Light is a kind of energy that travels in the form of
h is planck’s constant (6.626 × 10-34 J.s), v is the frequency of light in cycles/s, c is
electromagnetic waves of different wavelengths.
the speed of light (3.00 × 108 m/s), and λ is the wavelength in meters.
Visible light has wavelengths 400 to 700 nm which a small part of electromagnetic
The energy of a photon of visible light ranges form 150 kJ/einstein for red light to
spectrum ranging from violet to red.
~300 kJ/einstein for violet light.
The energy E, in a single photon (a quantum of light) of visible light is given by the
An einstein of light is Avogdro’s number (6.022 × 1023) of photons
Planck equation: E=hv=hc/λ
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Photosynthesis 10 Photosynthesis 11
Thus the energy of one einstein of photons at 700 nm is given by
Chlorophyll has a long hydrophobic tail that make it behave like a lipid, plus a
(2.84 × 10-19 J/photon)(6.022 × 1023 photons/einstein)=17.1 × 104 J/einstein or 171
porphyrin ring that has a central Mg atom and extensive system of delocalized
kJ/einstein or 40.87 kcal/einstein.
electrons in conjugated double bonds.
Photosystems in the thylakoid membrane are multiprotein assemblies containing
Absorption of a quantum of sunlight (photon) by chlorophyll molecule causes one
large numbers of bound chlorophyll molecules, in addition of cofactors (heme, iron-
of these electrons to move from a low-energy molecular orbital to higher energy
sulfur cluster, and quinones).
orbital.
Photosynthesis 12 Photosynthesis 13
A molecule that has absorbed a photon is in an exited state, which is generally 2. By transferring the energy not the electron, to a neighboring chlorophyll
unstable. molecule by resonance transfer.
The exited electron in a chlorophyll molecule quickly return to it ground state in one 3. By transferring the excited electron with its negative charge to another
of the three ways. nearby molecule, an electron acceptor, and the positive chlorophyll take
1. The absorbed quantum is released in the form of heat or combination of and electron from other molecule (water).
heat and light of longer wavelength (fluorescence). It occurs when
chlorophyll molecule is isolated in solution.
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Photosynthesis 14 Photosystems: These are large multiprotein assemblies in thylakoid membrane
The later two mechanisms occur when chlorophylls are attached to protein in a containing many bound chlorophylls (200), carotenoids (50) and cofactors.
chlorophyll-protein complex. There are two parts of each photosystems;
The light-induced transfer of an electron from a chlorophyll to a mobile electron A photochemical reaction center, which is a transmembrane chlorophyll complex
carrier is the central charge-separation step in photosynthesis in which the chlorophyll and have the special pair of chlorophyll a molecule. It transduce light into chemical
becomes positively changed and an electron carrier becomes negatively charged. energy.
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The other part is antenna complex or light-harvesting complex (LHC), which Chloroplast contains two functionally different but structurally related
absorb and collect a sufficient number of photons and passes the energy from one photosystems.
protein-bound chlorophyll to another by resonance energy transfer until it reaches the Photosystem I absorbs light of 700nm and is called P700. it is confined to the
special pair of the reaction center. unstacked stroma thylakoids.
Without it the process would be slow and inefficient. Photosystem II absorbs light of 680nm and is called P680, which is confined to the
stacked grana thylakoid.
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Photosynthesis 20 Photosynthesis 21
Z scheme or photosynthetic electron-transport chain or light reactions:
The path of electrons through the two photosystems can be described as a Z-like
Photosystem I causes second light-driven charge-separation reaction that leads to
trajectory and is known as the Z scheme.
NADPH production.
The reaction center of photosystem II first withdraws an electron from water. The
Photosystem II: After absorption of photons, the special pair of chlorophyll (P680)
electron passes via an electron-transport chain (composed of electron carriers
rapidly transfer an electron to Pheophytin (the chlorophyll with the 2H+ at its center
plastoquinone, the cytochrome b6-f complex, and protein plastocyanin) to P I.
instead of Mg+).
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The electron is then transferred to a tightly bound plastoquinone QA and then to Photosynthesis 23
the exchangeable pool of plastoquinone QB.
The manganese cluster is very strong oxidant and extracts electrons from H2O and
At the arrival of second electron, the exchangeable plastoquinone extract two
transfer it to P680+. A molecule of O2 is formed for every 2H2O.
protons form water in stroma and is reduced to QH2 leaving positive charge on P680+
2H2O + 4 photons → 4H+ + 4e- + O2
There is manganese cluster linked to the protein towards the lumen side of
This reaction is called photolysis and the protons are released in the thylakoid space
thylakoid.
(lumen) which contribute the proton gradient across the thylakoid membrane.
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All of the oxygen in the earth’s atmosphere has been generated in this way and this Cytochrome b6-f complex: Its structure is homologous to the cytochrome c
protein environment that endows life has remained unchanged through billions of reductase in mitochondria.
years of evolution. It forms the connecting link between PII and PI in the chloroplast ETC.
The exact details of water-oxidation reaction in PII are still not fully understood but It is intermingled both in grana and stroma thylakoid.
the scientists are trying to construct an artificial system that mimic this process. If The reduce plastoquinone (mobile electron carrier) transfer it electrons to
successful, this might provide an endless supply of clean energy to solve the world’s cytochrome b6-f complex.
energy crises.
Photosynthesis 26 Photosynthesis 27
Photosystem I: It receive electrons from plastocyanin in the thylakoid space and
It pumps protons into thylakoid space using the Q cycle and contributes to the transfer them, via a second charge-separation reaction, to the small protein ferredoxin
proton gradient across the thylakoid membrane. on the opposite membrane surface.
It passes electron one at a time to the mobile electron carrier plastocyanin (a small Then in final steps, ferredoxin feeds its electrons to a membrane-associated enzyme
copper-containing electron carrier, a homolog of cy. c) complex, the ferredoxin-NADP+ reductase, which uses the electrons to produce
Plastocyanin transfers electron to PI. NADPH from NADP+
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The chloroplast ATP synthase: The complex is Three small mobile electron carriers—
embedded in the thylakoid membrane and is plastoquinone, plastocyanin, and ferredoxin-
confined to stroma thylakoid. The catalytic participate in this process.
globular head is toward stromal side. Together electron-driven proton pump cy.b6-f
Starting with the withdrawal of electrons from complex, the photosystems generates a large
water, the light-driven charge separation steps proton gradient across the thylakoid membrane
in photosystem II and I enable the energetically The ATP synthase the harness this proton
unfavorable (uphill) flow of electrons from gradient to produce a large amounts of ATP in
water to NADPH. the chloroplast stroma.
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Cyclic electron transport chain:
The linear (non-cyclic) Z scheme can be switched to circular mode of electron
flow through photosystem I and the b6-f complex.
Here, the reduced ferrodoxin diffuses back to cytochrome b6-f complex to
reduce plastoquinone, instead of passing its electrons to the ferrodoxin-NADP +
reductase and thus generates only ATP.
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