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Photosynthesis¶
Part of Module 5: Communication, homeostasis and energy.
Photosynthesis is the process by which light energy is used to drive the endothermic synthesis of organic molecules from inorganic precursors — carbon dioxide and water. It underpins virtually all food chains on Earth and is responsible for atmospheric oxygen. At A level, the focus shifts from the overall equation to the two-stage mechanism: a light-dependent series of reactions in the thylakoid membranes, and a light-independent cycle in the stroma.
What You Need to Learn¶
Further detail: AS Biology A (H020) and A Level Biology A (H420).
On this page you'll learn about overview, chloroplast structure and adaptation, stage 1, and the light-dependent reactions. You'll also cover stage 2, and the light-independent reactions (Calvin cycle) and factors affecting the rate of photosynthesis. The notes bring these ideas together into one clear overview of photosynthesis.
Overview¶
6CO₂ + 6H₂O → C₆H₁₂O₆ + 6O₂ (in the presence of light energy)
This summary equation conceals a complex sequence of reactions. The energy input is light; the light energy is converted first into ATP and reduced NADP, then used to fix CO₂ into organic molecules. Oxygen is a waste product of water splitting.
The process takes place in the chloroplast, an organelle present in the mesophyll cells of leaves.
Chloroplast Structure and Adaptation¶
| Structure | Description | Role in photosynthesis |
|---|---|---|
| Outer and inner membranes | Double membrane; inner membrane is impermeable to many ions | Maintain internal environment suitable for enzymes |
| Thylakoids | Flattened membranous sacs | Site of light-dependent reactions; membranes contain photosynthetic pigments |
| Grana (singular: granum) | Stacks of thylakoids | Increase membrane surface area for light absorption |
| Stroma | Fluid surrounding grana | Site of light-independent reactions; contains Calvin cycle enzymes, DNA and ribosomes |
| Intergranal lamellae | Membranes connecting adjacent grana | Connect thylakoid compartments |
Thylakoid membranes contain photosynthetic pigments arranged into photosystems (PS I and PS II). Photosystems are antenna complexes: accessory pigments (carotenoids, xanthophylls) absorb light of various wavelengths and funnel the energy to a central reaction centre chlorophyll (P700 in PS I, P680 in PS II) where it is used to excite electrons.
Photosynthetic Pigments and Chromatography¶
The main photosynthetic pigments are:
- Chlorophyll a: primary pigment; absorbs red and blue-violet light, reflects green (hence the colour of most leaves); present in all photosynthetic organisms
- Chlorophyll b: accessory pigment; broadens the range of absorbed wavelengths; found alongside chlorophyll a in the light-harvesting complexes
- Xanthophylls and carotenoids: absorb wavelengths not absorbed by chlorophyll, further broadening the usable spectrum
Chromatography can separate these pigments. Each pigment travels a different distance up chromatography paper. The Rf value identifies each pigment:
Rf = distance travelled by pigment from origin
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distance travelled by solvent from origin
Each pigment has a characteristic Rf value (carotene travels farthest; chlorophyll b travels least far).
Stage 1: The Light-Dependent Reactions¶
The light-dependent reactions occur on the thylakoid membranes. Their products are ATP, reduced NADP (NADPH) and oxygen.
Photosystem II (P680)¶
- Light energy is absorbed by PS II antenna pigments; energy is transferred to the P680 reaction centre
- Energy excites electrons to a higher energy level; the excited electrons leave P680
- These electrons are accepted by the primary electron acceptor and passed to the electron transport chain in the thylakoid membrane
- P680 is left with a deficit — it has a very positive redox potential; it obtains replacement electrons from the photolysis of water:
2H₂O → 4H⁺ + 4e⁻ + O₂
- Oxygen is released as a waste product; protons (H⁺) accumulate in the thylakoid lumen
Electron Transport Chain and ATP Synthesis (Photophosphorylation)¶
- Electrons pass through a series of electron carriers in the thylakoid membrane (plastoquinone, cytochrome b₆f complex, plastocyanin)
- As electrons move to lower energy levels, energy is released and used to pump H⁺ ions from the stroma into the thylakoid lumen (chemiosmosis)
- H⁺ ions accumulate in the thylakoid lumen, creating a proton gradient (high H⁺ inside thylakoid)
- H⁺ ions flow back down the gradient through ATP synthase (located in the thylakoid membrane)
- This drives the synthesis of ATP from ADP and Pᵢ — called photophosphorylation
Photosystem I (P700)¶
- Electrons arriving at the end of the transport chain are re-energised by PS I (absorbs light at 700 nm)
- The re-excited electrons are passed to the final electron acceptor ferredoxin
- Ferredoxin reduces NADP using the electrons and the H⁺ from the lumen:
NADP + 2e⁻ + H⁺ → NADPH (reduced NADP)
Cyclic and Non-Cyclic Photophosphorylation¶
| Type | Photosystems involved | Oxygen produced? | NADPH produced? | ATP produced? |
|---|---|---|---|---|
| Non-cyclic | PS II and PS I | Yes (from photolysis) | Yes | Yes |
| Cyclic | PS I only | No | No | Yes (supplementary ATP) |
In cyclic photophosphorylation, electrons from PS I return to the electron transport chain via ferredoxin → cytochrome b₆f complex → plastocyanin → PS I. No NADPH is produced, but extra ATP is generated to meet the demand of the Calvin cycle.
Summary of Light-Dependent Reaction Products¶
| Product | Where it comes from | Where it goes |
|---|---|---|
| ATP | Photophosphorylation | Calvin cycle (light-independent stage) |
| Reduced NADP (NADPH) | Reduction of NADP by PS I | Calvin cycle |
| Oxygen (O₂) | Photolysis of water | Released as waste; diffuses out |
Stage 2: The Light-Independent Reactions (Calvin Cycle)¶
The Calvin cycle occurs in the stroma of the chloroplast. It uses ATP and NADPH from the light-dependent stage to fix CO₂ into organic molecules. It does not directly require light, but it is dependent on the continuous supply of ATP and NADPH.
The Calvin Cycle Steps¶
Step 1 — Carbon fixation:
- CO₂ combines with the 5-carbon acceptor molecule ribulose bisphosphate (RuBP) in a reaction catalysed by the enzyme RuBisCO (ribulose-1,5-bisphosphate carboxylase/oxygenase)
- This produces an unstable 6-carbon intermediate that immediately splits into two molecules of glycerate-3-phosphate (GP) — a 3-carbon compound
CO₂ + RuBP (C5) → 2 × GP (C3)
Step 2 — Reduction:
- GP is reduced using ATP and NADPH from the light-dependent reactions
- GP is converted to triose phosphate (TP) — a 3-carbon sugar phosphate (specifically glyceraldehyde-3-phosphate, G3P)
GP + ATP + NADPH → TP + ADP + NADP + Pᵢ
Step 3 — Regeneration of RuBP:
- Most TP molecules (5 out of every 6) are used to regenerate RuBP using ATP
- This step maintains the cycle by ensuring the CO₂ acceptor is continuously available
Step 4 — Production of organic molecules:
- The remaining TP (1 in 6 molecules) is used to synthesise glucose and other organic compounds
- Glucose is then converted to starch, sucrose, lipids, amino acids, nucleotides or cellulose
Fate of Triose Phosphate¶
| Product | How it is made from TP |
|---|---|
| Glucose / sucrose | Two TP condensed, then dephosphorylated |
| Starch | Polymerisation of glucose |
| Cellulose | Polymerisation of glucose (different glycosidic linkage) |
| Lipids | Glycerol (from TP) + fatty acids (from acetyl CoA derived from pyruvate) |
| Amino acids | TP → pyruvate or oxaloacetate; combined with NH₃ (from nitrate reduction) |
| Nucleotides | Ribose phosphate derived from TP intermediates |
Worked Example: Effect of Changing Light on Calvin Cycle Intermediates¶
If light is suddenly removed:
- Light-dependent reactions stop; no more ATP or NADPH produced
- GP can no longer be reduced to TP → GP accumulates
- TP is no longer available to regenerate RuBP → TP and RuBP levels fall
If CO₂ concentration is suddenly reduced:
- Less CO₂ fixation occurs; less GP is produced → GP levels fall
- TP continues to be used to regenerate RuBP for a while → RuBP accumulates initially
Factors Affecting the Rate of Photosynthesis¶
The rate of photosynthesis is determined by whichever factor is in shortest supply — this is the limiting factor.
Light Intensity¶
- Higher light intensity → more photons available → more PS II reaction centre activation → more photophosphorylation → more ATP and NADPH → faster Calvin cycle
- If light is limiting: GP rises (CO₂ still being fixed but less ATP/NADPH to reduce it); TP and RuBP fall
CO₂ Concentration¶
- Higher CO₂ → more carbon fixation by RuBisCO → more GP → more TP → more glucose
- If CO₂ is limiting: RuBP accumulates (not being used for fixation); GP falls
Temperature¶
- Temperature affects enzyme activity: RuBisCO and Calvin cycle enzymes have an optimal temperature
- At low temperatures, enzyme kinetic energy is low, reactions are slow, rate of photosynthesis limited
- At very high temperatures, enzymes denature
- The light-dependent reactions are less temperature-sensitive (photochemical processes, not enzyme-controlled), so temperature mainly limits the Calvin cycle
Saturation Point¶
The saturation point is the value of a limiting factor at which it is no longer limiting the rate of photosynthesis — another factor has become limiting instead. At and beyond the saturation point, increasing the original factor produces no further increase in the rate of photosynthesis.
Combined Effect: Compensation Point¶
The compensation point is the light intensity at which the rate of photosynthesis equals the rate of respiration — there is no net gas exchange. Below the compensation point, the plant consumes more O₂ (respiration) than it produces. Shade-adapted plants have lower compensation points.
Key Terms¶
- Chloroplast: organelle in plant and algal cells where photosynthesis takes place.
- Photosystem II: pigment-protein complex that absorbs light and drives photolysis and electron excitation.
- Photolysis: splitting of water by light.
- Photophosphorylation: synthesis of ATP using light energy during the light-dependent stage.
- Chemiosmosis: ATP production driven by proton flow down an electrochemical gradient through ATP synthase.
- Reduced NADP: electron-carrying molecule formed in the light-dependent stage and used in the Calvin cycle.
- Photosystem I: pigment-protein complex that re-excites electrons so NADP can be reduced.
- Cyclic photophosphorylation: pathway in which electrons cycle back to PSI and generate ATP but not reduced NADP.
- RuBisCO: enzyme that catalyses fixation of CO2 to RuBP in the Calvin cycle.
- GP: glycerate 3-phosphate, the first stable product of carbon fixation in the Calvin cycle.
- TP: triose phosphate, the reduced three-carbon product used to make carbohydrates and regenerate RuBP.
- RuBP: ribulose bisphosphate, the five-carbon carbon acceptor in the Calvin cycle.
- Rf value: the ratio of pigment travel distance to solvent travel distance.
- Saturation point: the value of a limiting factor beyond which it no longer limits the rate of photosynthesis.
- Compensation point: light intensity at which photosynthesis and respiration occur at equal rates.
Connected Pages¶
- 5.2.2 Respiration (complementary energy processes; ATP)
- 2.1.3 Nucleotides and nucleic acids (ATP structure)
- 2.1.2 Biological molecules (products of Calvin cycle — glucose, lipids)
- 6.3.1 Ecosystems (gross and net primary productivity)
- Photosynthesis vs respiration
- Module 5: Communication, homeostasis and energy