PHOTOSYNTHES

IS AND
PHOTORESPIRAT
ION
Dr M.S. Mafa
CARBON DIOXIDE FIXATION AND THE PHOTOSYNTHETIC
CARBON REDUCTION CYCLE
 NADPH and ATP, produced during the light dependent reaction is used in the Calvin-Benson cycle

 This cycle consists of 13 enzymatic reactions that take place in the stroma

 Calvin-Benson cycle is divided in to three phases:

 (i) carboxylation, in which CO2 is added to ribulose-1,5- bisphosphate (RuBP), and produce 3-

phosphoglycerate (3-PGA)
 (ii) Reduction, glyceraldehyde 3-phosphate (GAP)is produced from each 3-PGA

 (iii) Regeneration, which results in the formation of more acceptor molecules

 Two molecules each of ATP and NADPH per atom of carbon fixed are used in the reductive phase of the cycle

 The first stable products of CO2 fixation are glyceric acid mono- and bis-phosphates and phosphorylated trioses.

 Calvin–Benson cycle is often referred to as the C3 pathway of photosynthesis

RUBISCO CATALYZES THE FIRST REACTION IN THE CALVIN–
BENSON CYCLE
 The first reaction in the Calvin–Benson cycle is carboxylation of ribulose-1,5-bisphosphate (RuBP) by

CO2,
 This reaction is catalyzed by the enzyme RuBP carboxylase-oxygenase (aka rubisco);

 The product of this reaction is 3-phosphoglycerate (3-PGA).

 “The reaction catalyzed by rubisco proceeds by a five-step mechanism, during which a transient six-

carbon intermediate is formed and then hydrolyzed to yield two molecules of the 3-PGA product.”

 Rubisco is one of the most abundant enzymes on Earth and it account for more than half the total protein

nitrogen of leaves

 Rubisco breakdown during senescence and is used as source of amino acid if it does not participate in the

photosynthesis

 The products of the rubisco reaction enter the reduction phase of the cycle, followed by a series of
RUBISCO Reactions

Carboxylation: Rubisco (CO2/O2)

• C5 + C1  C6 + H2O  2 C3

• If O2 is used  Photorespiration,
forms 1 C3 and 1 C2
• Rubisco has 2 subunits (Large and
Small).
• L carries the catalytic site while S is
for conformation.
THE TWO-STEP REDUCTION PHASE OF THE CALVIN–
BENSON CYCLE USES ATP AND NADPH
 3-PGA, is converted to the triose phosphate glyceraldehyde-3-phosphate (GAP) in two enzymatic steps

 Phosphoglycerate kinase catalyzes phosphorylation of 3-PGA to form 1,3-bisphosphoglycerate

 Which in turn is reduced to GAP by NADPH-dependent GAP dehydrogenase.

 This phase of the cycle uses 1 mole each of ATP and NADPH per CO 2 fixed.

 GAP can be use as a precursor for other

During the regeneration phase of the Calvin-Benson cycle, ten
enzyme reactions convert five 3-carbon to three 5-carbon
intermediates
 Regeneration phase comprises 10 of the 13 enzymes of the Calvin–Benson cycle

 Sedoheptulose-1,7-bisphosphatase dephosphorylates a C7 diphosphosugar to yield a monophosphosugar

 Phosphoribulokinase phosphorylate ribulose-5-phosphate to form RuBP

 Only one triose molecule represents net gain of carbon from every 6 triose phosphate molecules produced by the reduction

phase

 The remaining five regenerate three molecules of RuBP that re-enter the cycle

1. Two molecules of triose phosphate (C3) combine to make fructose-1,6,-bisphosphate (C6).

2. Dephosphorylated C6 sugar phosphate reacts with another triose phosphate to make one C5 and one C4 product.

3. C4 sugar phosphate with another triose phosphate molecule produces a C7 sugar phosphate

4. dephosphorylated C7 reacts with C3 to yield two C5 sugar phosphates

5. The final step in the regeneration cycle is the conversion of ribulose-5-phosphate to RuBP
PHOTOSYNTHESIS IS DEPENDENT ON THE EXCHANGE OF
METABOLITES ACROSS THE CHLOROPLAST ENVELOPE
THE CALVIN–BENSON CYCLE PROVIDES THE PRECURSORS OF
CARBOHYDRATES FOR TRANSLOCATION AND STORAGE
 Triose is (GAP) used for sucrose or starch synthesis

 Sucrose synthesis takes place in the cytosol

 Enzymes used are similar to the C3-CR except that they are cytosolic

 F-1,6-BPase reaction is irreversible and SPS are important control points

 Sucrose phosphate phosphatase also regulate sucrose synthesis

 F-1,6-BPase is modulated by FB-2,6-P (which is sensitive to Pi)

 If photosynthesis is high, there will be high Pi, therefore FB-2,6Pase will be low and F-1,6-BPase is

active high sucrose synthesis

 When Triose-P are low, FB-2,6-Pase builds up and inhibit FBPase  Carbon is diverted to glycolysis
 If supply of sucrose to the metabolic sinks exceeds the

demand  Carbon is diverted away from sucrose to be

stored as starch

 In some plants when this happens, sucrose is stored in

the vacuole of photosynthetic leaves.

 High concentration of sucrose in vacuoles drive

synthesis of Fructans (soluble fructose polymers-
branched or linear).

 Fructans are short term storage carbohydrates.