Class XI Biology · Chapter 11

PHOTOSYNTHESIS IN HIGHER PLANTS

Photosynthesis in Higher Plants
Chapter overview: photosynthesis

Photosynthesis is the physico-chemical process by which green plants use light energy to synthesise organic compounds. It is the primary source of food on Earth and releases oxygen into the atmosphere. Chlorophyll, light and carbon dioxide are essential requirements.

Chapter map Early experiments -> chloroplast and pigments -> light reaction -> electron transport -> Calvin cycle -> C4 pathway -> photorespiration -> limiting factors.

Overall equation: 6CO2 + 12H2O -> C6H12O6 + 6H2O + 6O2. The oxygen evolved comes from water, not carbon dioxide.

11.1 Early experiments and requirements

Starch tests on variegated or partially covered leaves show that photosynthesis occurs only in green parts exposed to light. A leaf region enclosed with KOH-soaked cotton does not form starch because KOH absorbs CO2; this proves the requirement of carbon dioxide.

ScientistKey contribution
Joseph PriestleyPlants restore air damaged by breathing animals and burning candles.
Jan IngenhouszSunlight and green parts are necessary; green aquatic parts release oxygen bubbles in light.
Julius von SachsGreen parts form glucose, usually stored as starch; chlorophyll occurs in special bodies later called chloroplasts.
T.W. EngelmannAerobic bacteria accumulated in blue and red regions around illuminated alga, showing action spectrum of photosynthesis.
Cornelius van NielShowed that the hydrogen donor is oxidised; in green plants water supplies hydrogen, so released O2 comes from water.
11.2 Site of photosynthesis: chloroplast

Photosynthesis occurs in green leaves and other green plant parts. Mesophyll cells contain numerous chloroplasts, which can orient along cell walls to optimise incident light. Chloroplasts have a membrane system of grana and stromal lamellae, plus the surrounding stroma.

Chloroplast regionMajor role
Thylakoid/granal membrane systemTraps light energy and synthesises ATP and NADPH through light reactions.
StromaEnzyme-driven carbon reactions synthesise sugars that can form starch.
Terminology Carbon reactions are conventionally called dark reactions because they are not directly light-driven; they do not mean reactions that occur only in darkness, because they use ATP and NADPH formed in light reactions.
11.3 Photosynthetic pigments

Paper chromatography separates four leaf pigments: chlorophyll a (blue-green), chlorophyll b (yellow-green), xanthophylls (yellow) and carotenoids (yellow to yellow-orange). Pigments absorb light at specific wavelengths; most photosynthesis occurs in blue and red regions.

Pigment groupRole
Chlorophyll aChief photosynthetic pigment and reaction-centre pigment.
Chlorophyll b, xanthophylls and carotenoidsAccessory pigments: absorb additional wavelengths, transfer energy to chlorophyll a and protect it from photo-oxidation.
11.4 Light reaction and photosystems

Light reactions include absorption of light, splitting of water, oxygen evolution, electron transport, and formation of ATP and NADPH. Pigment molecules in the light-harvesting complex/antenna funnel energy to a reaction-centre chlorophyll a molecule.

PhotosystemReaction centreKey role
Photosystem II (PS II)P680, absorbs red light at 680 nmReceives electrons from photolysis of water and initiates non-cyclic electron flow.
Photosystem I (PS I)P700, absorbs red light at 700 nmRe-excites electrons and reduces NADP+ to NADPH in non-cyclic flow.

Water splitting is associated with PS II: 2H2O -> 4H+ + O2 + 4e-. The electrons replace those lost by PS II, protons contribute to the gradient, and oxygen is released.

Electron transport and photophosphorylation

In non-cyclic photophosphorylation, electrons move from PS II through carriers to PS I, are re-excited and finally reduce NADP+ to NADPH. This route produces ATP, NADPH and O2.

In cyclic photophosphorylation, electrons excited from PS I cycle back through the electron transport chain to PS I. It produces ATP only: no NADPH is formed and no O2 is evolved. It is useful when the chloroplast needs additional ATP relative to NADPH.

Electron transport builds a proton gradient across the thylakoid membrane. High H+ concentration develops in the lumen through water splitting, proton translocation via electron carriers, and use of stromal H+ in NADPH formation. H+ flows back through CF0-CF1 ATP synthase; this chemiosmotic flow powers ATP formation.

11.5 Carbon reactions: Calvin cycle

The Calvin cycle occurs in the stroma of all photosynthetic plants, including C3 and C4 plants. Its primary CO2 acceptor is the 5-carbon ketose ribulose-1,5-bisphosphate (RuBP). RuBisCO carboxylates RuBP, producing two molecules of 3-phosphoglycerate (3-PGA).

StageCore event
CarboxylationCO2 is fixed to RuBP by RuBisCO, yielding two 3-PGA molecules.
ReductionATP and NADPH convert intermediates to carbohydrate/triose phosphate.
RegenerationATP regenerates RuBP so the cycle can continue.

For each CO2 fixed, 3 ATP and 2 NADPH are used. Six turns fix six CO2 to make one glucose, consuming 18 ATP and 12 NADPH.

11.6 C3 pathway vs C4 pathway

All photosynthetic plants use the Calvin cycle to form sugars. C4 plants first concentrate CO2 through the Hatch-Slack pathway, an adaptation of dry tropical plants.

FeatureC3 plantsC4 plants
First stable product of CO2 fixation3-PGA, a 3-carbon acidOxaloacetate (OAA), a 4-carbon acid
Primary acceptorRuBP, 5-carbonPEP, 3-carbon
Initial carboxylationMesophyll cells by RuBisCOMesophyll cells by PEP carboxylase (PEPcase)
Calvin cycleMesophyll cellsBundle sheath cells
Leaf anatomyNo Kranz anatomyKranz anatomy: chloroplast-rich, thick-walled bundle sheath cells with no intercellular spaces
ResponseLower temperature optimum and lower high-light CO2 fixationTolerates high temperature, responds to high light and has greater biomass productivity
PhotorespirationPresentAbsent/negligible

In C4 plants, OAA becomes malate or aspartate in mesophyll cells and is transported to bundle sheath cells. Decarboxylation releases CO2 for Calvin cycle; the 3-carbon compound returns to mesophyll to regenerate PEP.

11.7 Photorespiration

RuBisCO acts as both carboxylase and oxygenase. In C3 plants, O2 can compete with CO2 at the active site. Oxygenation of RuBP produces one phosphoglycerate and one 2-carbon phosphoglycolate, initiating photorespiration.

Why it is wasteful Photorespiration neither synthesises sugars nor ATP or NADPH. It releases CO2 and uses ATP; its biological function remains unknown in NCERT.

C4 plants avoid photorespiration because decarboxylation of C4 acids elevates CO2 around RuBisCO in bundle sheath cells, favouring carboxylase activity over oxygenase activity.

11.8 Factors affecting photosynthesis

Internal factors include leaf number, size, age and orientation; mesophyll cells and chloroplasts; internal CO2; and chlorophyll quantity. External factors are light, temperature, CO2 concentration and water. Blackman's law of limiting factors states that when several factors affect a process, its rate is determined by the factor nearest its minimum value.

FactorNCERT effect
LightCO2 fixation rises linearly at low intensity, then plateaus when other factors limit. Saturation occurs at about 10% of full sunlight; excessive light can break down chlorophyll.
CO2Major limiting factor. Atmospheric level is 0.03-0.04%; increase up to 0.05% raises fixation, but higher long-term levels may be damaging. C4 saturates near 360 microlitres L-1; C3 beyond 450 microlitres L-1.
TemperatureCarbon reactions are strongly temperature-controlled. C4 plants have higher optimum temperature than C3 plants; adaptation to habitat matters.
WaterWater stress closes stomata, reducing CO2; it also causes wilting, reducing leaf area and metabolism.
NCERT revision prompts
  1. State the experiments proving the need for chlorophyll, light and CO2.
  2. Which experiment established that oxygen released during photosynthesis comes from water?
  3. Differentiate the sites and products of light and carbon reactions.
  4. Compare cyclic and non-cyclic photophosphorylation.
  5. Explain the chemiosmotic mechanism of ATP synthesis in chloroplasts.
  6. State the three phases and energy cost of Calvin cycle.
  7. Compare C3 and C4 plants, including Kranz anatomy and the primary CO2 acceptor.
  8. Why is photorespiration negligible in C4 plants?
  9. Apply Blackman's law to light, CO2, temperature and water.
Final recall PS II has P680 and photolyses water; PS I has P700 and reduces NADP+. Calvin cycle uses 3 ATP and 2 NADPH per CO2 fixed.