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Lesson Objective: Give overall outline of photosynthetic process that leads to the production of glucose.

PHOTOSYNTHESIS
Green plants use sunlight as an energy source, CO2 & water as raw materials for photosynthesis. The light energy trapped by greens plant is converted to chemical energy & stored in the bonds of organic molecules such as carbohydrates. O2 is released as a by product. 6CO2 + 12H20
Light energy Absorb by chlorophyll

C6H1206 + 6O2 + 6H2O

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6.1: Photosynthesis requires:


1. Chlorophyll pigments in chloroplasts. 2. CO2 3. Water 4. Optimum temperature (photosynthesis is a biochemical reaction that involves many enzyme). 5. Light
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Lesson Objective: List and explain the photosynthetic pigments involved in photosynthesis

Chlorophyll
Thylakoid membranes contain several kinds of pigments. Different pigments absorb light of different wavelength. Chlorophyll, the main pigment of photosynthesis, absorb light primarily in blue & red regions of visible spectrum.

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TOPIC 6: PHOTOSYNTHESIS

Chlorophyll a is the most abundant pigment in plants. It absorbs light mainly in the blue-violet (430nm) & red (662nm) region.

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Chlorophyll b is absorbs light of about 453nm & 642nm. Chlorophyll b helps to increase the range of light a plant can utilize for photosynthesis. Carotenoid absorb light maximally in the blueviolet region (460-550nm).

The most common carotene is carotene, an orange pigment found for example in carrots. Carotenoids act as accessory or antenna pigments. These pigments pass the light energy to chlorophyll a in the reaction centre. The carotenoid also protect the chlorophyll from oxidation.
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The mechanism of photosynthesis


The photosynthetic process involves 2 stages; 1. The light dependent reaction which requires light energy & is biochemical in nature. 2. Light independent reaction which consist of a series enzymatic biochemical reactions that involve the bonding of CO2 to complex organic compounds.

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Lesson Objective: Explain the photoactivation of chlorophyll resulting in the conversion of light energy into ATP and reduced NADPH+.

6.3: Light Dependent Reaction


These stages occur in the chloroplast thylakoids which contains complex photosynthetic pigments. The photosynthetic process involves the following process; i. Photoactivation ii. Photolysis of water iii. Photophosphorylation (produced ATP & NADPH).
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i. Photoactivation

= when a molecule of chlorophyll

absorbs a photon, one of the molecules electrons is elevated to an orbital where it has more potential energy, the pigment molecule is said to be in an excited state photo-activation process.
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The energy of an absorbed photon is converted to the potential energy of an electron raised from the ground state to an exited state. The chlorophyll & accessory pigments are group into 2 photosystem; a) Photosystem I (PSI) b) Photosystem II (PSII
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Photosystem I has a reaction center chlorophyll, the P700 center, that has an absorption peak at 700nm. Photosystem II has a reaction center with a peak at 680nm. These two photosystems work together to use light energy to generate ATP and NADPH.

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The phosystems are embedded in the thylakoid membranes of the chloroplast. A photosystem contains a complex of about 200-300 pigment molecules (consisting of a cluster of a few hundred chlorophyll a, chlorophyll b & carotenoid molecules), primary electron acceptor, electron transfer system & enzymes.
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Each photosystem has an antenna of a few hundred pigment molecules. When a photon strikes a pigment molecule, the energy is passed from molecule to molecule until it reached the reaction center. At the reaction center, the energy drives an oxidationreduction reaction. An excited electron from the reaction-center chlorophyll is captured by a specialised molecule called the primary electron acceptor.
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ii. Photolysis of Water

The Hill Reaction

Photolysis of water is a process in which water is split during the light reaction of photosynthesis. This process occurs in the space of the thylakoid & catalysed by a water splitting enzyme. The enzyme passes the electron from water to the reaction center of PS II & forming O2, which is liberated. H2O O2 + 2H+ + 2e H+ ions combine with NADP+ NADPH + H+
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iii. Photophosphorylation
= The process of generating ATP from

ADP + Pi during the light reactions of photosynthesis. Light reaction consist of; a. non-cyclic photophosphorylation b. cyclic photophosphorylation
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a. Non-cyclic Photophosphorylation
Non-cyclic photophosphorylation involves 2 PS; - PS I - PS II Light energy is absorbed by antenna pigments of PS II. The energy is transferred from one antenna molecule to another molecules then transferred to the reaction center in PS680 in the PS II.

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The P680 become photoactivated & high energy electron released ( exited ) & captured by the primary electron acceptor.
photoactivation

Chlorophyll
(Reduced form)

chlorophyll + e
(oxidised form)

This creates an electron deficiency in PS II. Each photoexited e passes from the primary e acceptor of PS II to PS I via an e transport chain. This chain is very similar to the one that functions in cellular respiration.
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Electron from the primary electron acceptor, then transferred through plastoquinone (Pq) & cytochrome complex. The e are then passed to plastocyanin (Pc) & then to PS I. The energy released in the passage from Pq to cytochrome is used by the cell to make ATP from ADP & Pi.
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When an e- reached the bottom of the etransport chain, it fills an e- hole in P700 (the chlorophyll a molecule in the reaction center of PS I). This replace the e- that light energy drives from the chlorophyll to the primary eacceptor of PS I. The primary e- acceptor of PS I passes the photoexited e- to ferredoxin (Fd).
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An enzyme called NADP+ reductase then transfers the e- from Fd to NADP+. This is the redox reaction that stores the high-energy e- in NADPH, the molecules that will provide reducing power for the synthesis of sugar in the Calvin cycle. 2H+ + 2e- + NADP+ NADPH + H+

The H+ ions formed in water splitting is used in formation of NADPH.


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b. Cyclic Photophosphorylation
Under certain condition, photoexcited etake an alternative path called cyclic eflow, which uses PS I but not PS II. Light energy absorbed by antenna molecules is transferred to P700 in PS I. Electron from P700 is photoexcited & passed on to a primary e- acceptor. Primary e- acceptor passes the e- to ferredoxin (Fd).
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In cyclic photophosphorylation, an alternative route is used. The e- is passed from Fd to cytochrome complex. Energy is released during the e- flow & this is used in chemiosmosis to generate more ATP. The e- then transferred to plastocyanin (Pc) & is passed back to PS I.
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Lesson Objective: Describe the outline of Calvin cycle involving the light-independent fixation of CO2.

6.4. Light Independent Reaction/ Calvin Cycle


Consist of 4 main stages: a. Carbon dioxide fixation b. Reduction phase c. Regeneration of CO2 acceptor d. Product synthesis phase

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a. Carbon dioxide Fixation


CO2 from the atmosphere diffuse through the stomata into the intercellular spaces of the leaf. It then diffuse into the stroma in the chloroplast of the palisade & spongy mesophyll cells. A five-carbon acceptor, ribulose biphosphate (RuBP) combines with molecule of CO2 to form an unstable six-carbon sugar.\ The process is catalysed by the enzyme RuBP carboxylase. The six-carbon compound immediately splits into 2 molecules of glycerate-3-phosphate (PGA) (3-carbon compound).
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b.Reduction Phase
Glycerate 3-phosphate receives an additional phosphate group from ATP to become glycerate 1,3-diphosphate. Glycerate 1,3-diphosphate combines with hydrogen atoms from reduces dinucleotide phosphate (NADPH + H+) & is converted into glyceraldehyde-3-phosphate (PGAL, triose phosphate).
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TOPIC 6: PHOTOSYNTHESIS

c. Regeneration of CO2 acceptor Some of the PGAL molecules are rearranged in a series of complex reactions to regenerate ribulose biphosphate, this process requires ATP.

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d. Product synthesis phase The rest of the glyceraldehyde-3phosphate is used to assimilate organic molecules such as glucose, amino acids, proteins & lipids.

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PHOTORESPIRATION
RuBP carboxylase (Rubisco), the enzyme that catalysed the carboxylation of ribulose biphosphate, can also catalyze the oxidation of RuBP by molecular O2. CO2 & O2 are alternative substrates that compete with each other for the same active sites on the enzyme.
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When the concentration of CO2 is high & that of O2 is low, CO2 occupies the active sites, carboxylation is favored & carbohydrate synthesis by the Calvin cycle proceeds. But when the concentration of CO2 is low & that of O2 is high, the site is occupied by O2, oxidation is favored or photorespiration occurs. This occurs on hot dry days, which cause water stress in plants. As a result of water stress, plants close their stoma ( help the plant to conserve water)
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Once the stoma close, O2 produced during photosynthesis accumulates in the chloroplast. So rubisco binds RuBP to O2 instead of CO2.
(2 carbon compound)

Phosphoglycolate + glycerate-3-phosphate Two phosphoglycolate molecules then undergo a series of reactions requiring O2 & ATP to produce one molecule of glycerate-3-phosphate & the release

O2 + RuBP

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Photorespiration is wasteful because organic carbon (phosphoglycolate) is converted into CO2 with no net production of ATP or other energy rich metabolites. Photorespiration can reduce the potential photosynthetic yield from between 30-40%. This degradation process is called photorespiration because; i. it occurs during the daylight ii. it requires O2 (like aerobic respiration) iii. It produces CO2 & H2O (like aerobic respiration)
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6.5: Hatch-Slack Pathway (C4 plants)


Eg. of C4 plants: - maize plant (Zea mays) - sugar cane (Saccharum officinale) - sorghum (Sorghum bicolar) - sunflower (Helianthus spp)

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In C4 plants, there are 2 distinct types of photosynthetic cells: a. bundle-sheath cells b. mesophyll cells. Bundle sheath cells are arranged into tightly packed sheets around the veins of the leaf. Between the bundle sheets & the leaf surface are the more loosely arranged mesophyll cells.
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The arrangement pattern of the cells around the vascular bundle is known as Krantz anatomy.
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CO2 in the atmosphere diffuse into the mesophyll cells of C4 plants where they combine with phosphoenolpyruvate (PEP,3C acceptor) to produce oxaloacetate (4C). This process is catalysed by the enzyme phosphoenol-pyruvate carboxylase (Pepco) which has higher affinity for CO2 at low CO2 concentration.
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Oxaloacetate is reduced to malate (4C). Malate is shunted through plasmodesmata into the bundle sheath cells. Malate is then oxidised to pyruvate (3C) by the removal of hydrogen & CO2. This increases the concentration of CO2 in the bundle cells & the CO2 undergoes fixation as in the normal C3 pathway. High CO2 concentration in the bundle sheath cells inhibits photorespiration & as a result, production of carbohydrate (glucose) increases 20-25%.
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Pyruvate diffuses into the mesophyll cells & phosphorylated to regenerate phosphoenol-pyruvate. In general, C4 plants are found in hot climates (tropical). Under hot conditions, C4 plants attain a higher photosynthetic rate compared to C3 plants.
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Differences between C3 & C4 Plants


Aspect Examples C3 plants C4 plants Tomato,tobacco, Sugar cane, legume, wheat. Sorghum, maize.

CO2 fixation

Once, only in Twice, first in mesophyll cells. mesophyll cells & then bundle sheath cells.
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CO2 acceptor

RuBP ( 5C)

PEP (3C) in mesophyll cells. RuBP in bundle sheath cells.

Enzyme

Rubisco which Pepco which has is inefficient at high affinity for low CO2 CO2 at low concentration. concentration. RuBP which is efficient at high CO2 concentration.
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First product Glycerate 3formed. phosphate (a C3 acid) Photorespiration O2 acts as competitive inhibitor.

Oxaloacetate, (a C4 acid).

Photorespiration is inhibited by high concentration of Photorespiration CO2. occurs.


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Efficiency

Less efficient photosynthesis than C4 plants. Yields usually lower.

In the tropics, light intensity & temperature are higher & O2 is not a competitive inhibitor. Photosynthesis is more efficient. Photorespiration is prevented & yields are usually much higher.

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Leaf anatomy

Krantz
anatomy absent. Only one type of chloroplast in mesophyll cells.

Krantz anatomy present with the columnar palisade cells arranged in a ring around the bundle sheath cells & the vascular tissue.

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Leaf anatomy

There are 2 different forms of chloroplasts. Mesophyll palisade cells contain few small chloroplasts, have many large well-developed grana but not starch grains. Bundle sheath cells have many large chloroplasts with fewer poorly developed grana & contain starch grains.

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CAM Plants = Crassulacean Acid Metabolism


E.g. of CAM plants are succulent plants such as the cacti & pineapples. This succulent (Crassulaceae), avoid water loss in their hot environment by closing their stomata during the day & opening them at night. CAM plants only contain mesophyll cells & no Krantz anatomy.
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Photosynthetic pathway is the same as C4 plants but fixation of CO2 takes place at night when stomata are open. Phosphoenolpyruvate (PEP) 3C combines with CO2 to produce oxaloacetate (OAA) 4C. Malate is stored in the cell vacuole at night to prevent pH changes in the cytoplasm. During daytime, malate is oxidised producing pyruvate & CO2. Concentration of CO2 increases in the mesophyll cell & photorespiration is prevented. CO2 is used in Calvin Cycle producing organic molecules.

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6.6: FACTORS LIMITING THE RATE OF PHOTOSYNTHESIS

a) Light Intensity
At low light intensity there is no photosynthesis

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As in figure, a higher intensity of light increases the rate of photosynthesis, unless influenced by other limiting factors, such as very low CO2 concentration. Under ideal conditions with no other limiting factors, the rate of photosynthesis is directly proportional to the intensity of light. Increase in CO2 concentration increases the rate of photosynthesis until saturation point is again reached.

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At this point, CO2 concentration or another new factor may limit the photosynthetic process. Light intensity only effects the lightdependent reactions & not the lightindependent reaction, which is catalysed by specific enzymes.

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b) Wavelength of Light
Visible light of different wavelengths or different colour affect the rate of photosynthesis differently. Red light of around 650750nm & blue light of 430-500nm are most effective for photosynthesis whereas green light of 540nm is least effective.
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This is because red & blue lights are absorbed by the chlorophyll whereas green light is not absorbed at all. The effects of different wavelengths of light on photosynthesis can be seen from the action spectrum below. The action spectrum is corresponding to the absorption spectrum.
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c) Temperature
The rate of photosynthesis is temperaturedependent because it is a series of reaction that depend on enzymes.

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In general, the rate of photosynthesis is at a maximum between 25oC 35oC. The optimum temperature for photosynthesis is about 30oC. Between 0oC to optimum temp., the temp. quotient (Q10) is equal to 2. This is same as to say for every 10oC increased in temp.,there is corresponding double increased in the rate of photosynthesis. Temp. above 40oC can break up the proteins in enzymes (denatured) & as a result photosynthesis would come to a stop.
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d) CO2 Concentration The rate of photos. is directly proportional to the CO2 concentration, in condition where the light intensity & temp. are not limiting factors. The concentration of CO2 in the atmosphere is only 0.035%. When this value increases, the rate of photos. would increase until a maximum is reached at approximately 1.0%.
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A concentration of CO2 that exceeds 1.0% will stimulate the closing of the stoma & will reduce the rate of photosynthesis.
Rate of photosynthesis

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