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PHOTOSYNTHESIS

Essentially all free energy utilized by biological systems arises from solar energy that is
trapped by the process of photosynthesis.
The basic equation of photosynthesis is :
Light
CO2 + H2O -----------------à (CH2O) + O2

(CH2O) represents primarily sucrose and starch.

Photosynthesis in green plant takes place in chloroplasts.

The energy of light captured by pigment molecules, called chlorophylls, is used to


generate high-energy electrons with greater reducing potential. These electrons are used
to produce NADPH as well as ATP in a series of reactions called the light reaction.
NADPH and ATP formed by the action of light then reduced CO2 and convert it to
hexose sugar through 3-phosphoglycerate by a series of reaction called the Calvin cycle
or the dark reaction.
• The primary events of photosynthesis namely, synthesis of NADPH and ATP,
take place in thylakoid membranes.
• The stroma contains the soluble enzymes that utilize the NADPH and ATP to
convert CO2 into sugar.
The light reaction:
• In the photosynthesis the first event is the absorption of light.
• The principal photoreceptor in the chloroplast of most green plants is
chlorophyll.
• The peak molar absorption coefficient of chlorophyll a is higher than 10 5 M-1 cm
–1
, among the highest observed for organic compounds.
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• The excited chlorophyll transfer its electron to a neighboring molecule through


resonance energy transfer.
• This process results in the formation of a positive charge on the initial molecule
and a negative charge on the acceptor and is, hence, referred to as photoinduced
charge separation.
• The site where the separational charge occurs is called the reaction center.

• The electron, extracted from its initial site by absorption of light, can reduce other
species to store the light energy in chemical forms.
• Photosynthesis of green plants is mediated by two kinds of membrane bound,
light-sensitive complexes – photosystem I (PS I) and photosystem II (PS II).

• PS I : 13 polypeptide chains, >60 chlorophyll mol., a quinone, and 3 4Fe-4S


clusters.
• PS II : at least 10 polypeptide chains, >30 chlorophyll mol., a nonheme iron ion,
and 4 Mn ions.

• In PS II, excitation of P680, a special pair of chlorophyll molecules, leads to


electron transfer to plastoquinone.
• The electrons are replenished by the extraction of electrons from water at a center
containing 4 Mn ions resulting in generation of O2.
• The plastoquinol, formed from plastoquinone, is reoxidized by the cytochrome bf
complex, which transfer the electron to plastocyanine.
• From plastocyanine the electron enters the PS I.
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The +Ve charge left at P700 is neutralized by electron transferred from reduced
plastocyanin.

• In PS I, excitation of special pair P700 releases electrons that flow to ferredoxin, a


powerful reductant.
• Ferredoxin-NADP+ reductase, located on the stromal side of the membrane, then
catalyzes the formation of NADPH.
• A proton gradient across the thylakoid membrane is generated as electrons pass
through PS II, through cytochrome bf complex, and through ferredoxin-NADP+
reductase, which resulted the formation of PMF.

• The PMF used by the ATP synthase to form the ATP.


• The ATP synthase of chloroplasts (also called CF0-CF1) closely resembles the
ATP-syhthesizing assemblies of bacteria and mitochondria (F0-F1).
• If the NADPH:NADP+ ratio is high, electrons transferred to ferredoxin by PS I
can reenter the cytochrome bf complex. This process, called cyclic
photophosphorylation, leads to the generation of a proton gradient by the
cytochrome bf complex without the formation of NADPH.
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Figure: Pathway of electron flow from water to NADP+ in photosynthesis (endergonic


reaction).
Symbols: Ph, pheophytin; QA & QB, plastoquinone-binding proteins; Pc, plastocyanin;
Ao & A1, acceptors of electrons from P700; Fd, ferredoxin; Mn, Manganese.

The standard free energy for oxidation of reduced plastocyanin by oxidized ferredoxin is
+18.9 kCal/ mol. This uphill reaction is driven by the absorption of a 700 nm photon
which has an energy of 40.9 kcal/mol. The cooperation between PSI and II creates
electron flow from water to NADP+. The pathway of electron flow is called Z-scheme of
photosynthesis as the redox diagram looks like the letter Z.

Comparison of photosynthesis and oxidative phosphorylation:


• The light induced electron transfer in photosynthesis drives protons into thylakoid
lumen.The excess protons flow out of the lumen through ATP synthase to
generate ATP in the stroma.
• In oxidative phosphorylation, electron flow down the electron transport chain
pumps protons out of the mitochondrial matrix. Excess protons from the
intermembrane space flow into the matrix through ATP synthase to generate ATP
in the matrix.
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• The Dark Reaction (CALVIN CYCLE):

• Calvin cycle takes place in the stroma of chloroplast.


The Calvin cycle synthesizes hexoses from carbon dioxide and water.
The Calvin cycle comprises 3 stages :
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1. The fixation of CO2 by ribulose 1,5-biphosphate to form 2 molecules of 3-


phosphoglycerate.

2. The reduction of 3-phosphoglycerate to form hexose sugars.


3. The regeneration of ribulose 1,5-biphosphate so that more CO2 can be fixed.

The initial incorporation of CO2 into


3-phosphoglycerate is highly exergonic reaction catalyzed by ribulose 1,5-biphosphate
carboxylase/oxygenase (rubisco).

Rubisco also catalyzes a wasteful oxygenase reaction


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• Phosphoglycolate is not a versatile metabolite.


• A salvage pathway recover part of its carbon skeleton. This salvage pathway
serves to recycle 3 of the 4 C-atoms of 2 molecules of glycolate. However, 1 C-
atom is lost as CO2. This process is called photorespiration because O2 is
consumed and CO2 is released.

• Photorespiration is wasteful because organic carbon is converted into CO2


without the production of ATP, NADPH or other E rich metabolites. The
oxygenase activity increases more rapidly with temperature than carboxylase
activity. This wasteful side reaction is minimized in tropical plants through an
accessory pathway-called C4 pathway –for concentrating CO2 at the site of
Calvin cycle.

Hexdose phosphates are made from phosphoglycerate :


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Starch in chloroplasts and sucrose in the cytosol are the major carbohydrate stores in
plants.