Key Terms
Photophosphorylation |
Synthesis of ATP from ADP and and Pi (inorganic phosphate) using light energy. |
Transducers |
Change energy from one form into another |
Absorption Spectrum |
A graph that shows how much light energy is absorbed at different wavelengths. |
Action Spectrum |
A graph that shows the rate of photosynthesis at different wavelength. |
Antenna Complex |
An array of protein and pigment molecules in the thylakoid membranes with chlorophyll a at the reaction centre. It transfers energy from light of a range of wavelengths to chlorophyll a. |
Limiting Factor |
A factor that limits the rate of a physical process by being in short supply. |
Overview of Photosynthesis
The overall equation for photosynthesis is: |
6CO2 + 6H2) -> C6H12O6 + 6O2 |
Photosynthesis involves two stages: |
1. Light-dependent stage where light energy is converted into chemical energy as the photolysis of water releases protons and electrons which produce ATP via photophosphorylation and reduce the co-enzyme NADP. |
2. Light-independent stage or Calvin cycle where ATP and NADPH from the light-dependent reaction reduce carbon dioxide to produce glucose. |
Structure of the leaf
The leaf is adapted for gas exchange and photosynthesis by having a large surface area allowing the leaf to capture light, and having pores called stomata through which gases diffuse. |
Air spaces between cells allow for carbon dioxide to diffuse to the photosynthesising cells. |
The highest concentration of chloroplasts is found in the palisade mesophyll on the leaf's upper surface. |
The palisade cells are arranged vertically, which allows more light to be absorbed by chloroplasts than if they were stacked horizontally, as light only has to pass through the cuticle, epidermal cells and one palisade cell wall. |
Chloroplasts as transducers
The site of photosynthesis was detected by Engelmann in 1887. In his experiment, he shone a light through a prism to separate the different wavelengths of light, and exposed this to a suspension of algae with evenly distributed, motile, aerobic bacteria.** |
After a period of time, he noticed that the bacteria congregated around the algae exposed to blue and red wavelengths. |
This was because this algae photosynthesised more and so produced more oxygen, attracting the motile bacteria. |
Photosynthetic pigments
In flowering plants there are 2 main types of pigments: |
1. Chlorophylls which absorb red and blue-violet regions of the spectrum, e.g. chlorophyll a and chlorophyll b. |
2. Carotenoids which absorb light energy from the blue-violet region of the spectrum, e.g. B-carotene and xanthophylls, and act as accessory pigments. |
The presence of several pigments allows the plant to absorb a wider range of wavelengths of light than a single pigment. |
Absorption and action spectra
The absorption spectrum shows how much light energy a particular pigment absorbs at different wavelengths, for example chlorophyll a which absorbs red and blue-violet regions of the spectrum. |
It does not indicate whether the particular wavelength is used in photosynthesis. |
An action spectrum shows the rate of photosynthesis at different wavelengths, by measuring the mass of carbohydrate synthesised by plants. |
There is a close correlation between the 2. |
Absorption and action spectra
Light Harvesting
The chlorophylls and accessory pigments are found lying in the thylakoid membranes, grouped into structures called antenna complexes. |
With the aid of special proteins associated with these pigments, light energy (photons) is funnelled towards the reaction centre at the base, containing chlorophyll a. |
There are 2 types of reaction centre: |
1. Photosystem 1 (PS1) chlorophyll a, with an absorption peak of 700nm, also called P700. |
2. Photosystem 11 (PS11) chlorophyll a, with an absorption peak of 680nm also called P680. |
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Identifying different photo pigments from chloro
Pigments can be extracted by grinding plant material in a suitable solvent, e.g. propanone, and separated by paper chromatography. |
By dividing the distance travelled by the pigment by the distance travelled by the solvent front, the Rf value can be calculated. |
Calculating the Rf value
The light-dependent stage of photosynthesis
Occurs on the thylakoid membranes. |
Photophosphorylation occurs via 2 pathways: |
1. Non-cyclic photophosphorylation, which involves both photosystems 1 and 11, generating 2 ATP molecules and reduced NADP. |
Photolysis generates oxygen. |
The electrons take a linear pathway which is referred to as the 'Z scheme'. |
Non-cyclic photophosphorylation
Non-cyclic photophosphorylation
1. Light energy (photons) strikes chlorophyll (PS11) exciting its electrons, boosting them to a higher energy level. |
2. Electrons are accepted by an electron carrier in the thylakoid membrane. |
3. The oxidised chlorophyll removes electrons from water, producing protons and oxygen (photolysis). This occurs in the thylakoid space. |
4. As electrons pass from carrier to carrier, electron energy is lost, which pumps protons from the stroma into the thylakoid space. As protons flow back through the stalked particle, ADP is phosphorylated; 2 ATP are made in total. |
5. Electrons enter photosystem 1 where light excites them, boosting them to an even higher energy level. |
6. Electrons enter a final electron carrier. |
7. Electrons and protons reduce NADP to reduced NADP which pass to the Calvin Cycle with the 2 ATP made. |
ATP production in the chloroplast
Cyclic Phosphorylation
Involves only photosystem 1, producing 1 ATP molecule only. |
As photolysis does not occur, no oxygen is released. |
Electrons take a cyclical pathway. |
If there is no NADP available, then electrons fall back into the electron transport chain (at an intermediate energy level) and generate 1 ATP. |
This cycle continues until NADP is available. |
The ATP produced can be used in the Calvin Cycle, in the stomatal opening mechanism, or for other cellular processes. |
ATP is produced in the chloroplast when protons are pumped across the thylakoid membrane using energy from the electrons and accumulate with protons generated from photolysis of water in the thylakoid space generating an electrochemical (proton) gradient. |
The H+ ions diffuse back into the stroma through stalked particles generating ATP. |
The protons and electrons reduce NADP, which removes H+ ions from the stroma, further contributing to the H+ gradient. |
The movement of protons is referred to as chemiosmosis. |
Calvin Cycle - light indep stage of photosynthesis
This stage occurs in the stroma. |
ATP and reduced NADP from the light-dependent reaction are used to fix carbon from carbon dioxide with the help of the enzyme RuBisCO. |
The sequence was first worked out by Calvin and his team using a radioactive isotope of carbon (14C) present in hydrogen carbonate. |
At regular intervals, Calvin removed samples into hot methanol to kill the chlorella algae used and to stop all enzyme reactions. |
He then performed chromatography to identify the products. |
He exposed his chromatogram to piece of x-ray film which would detect radiation emitted from 14C used. |
This identified products containing 14C in the order they were produced: first was hydrogen carbonate ions, then glycerate 3-phosphate (GP), triose phosphate (TP), ribulose bisphosphate (RuBP) and finally glucose. |
Calvin's lollipop apparatus
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Stages in the Calvin Cycle
1. CO2 diffuses into leaf via stomata, dissolving in the water surrounding palisade mesophyll cells before diffusing into the cells. |
2. CO2 combines with the 5 carbon compound ribulose bisphosphate (RuBP) using the enzyme RuBisCO to form an unstable 6C compound. |
3. Unstable 6C compound immediately breaks down into 2 molecules of glycerate 3-phosphate (GP). |
4. Using one ATP molecule from the light reaction, GP is reduced to triose phosphate (TP) using hydrogen atoms from reduced HADP. |
5. Triose phosphate molecules combine in pairs to form hexose sugars. |
6. 5 out of every 6 triose phosphate molecules produced are used to regenerate RuBP (via the intermediate ribulose phosphate) using ATP from the light-dependent reaction to supply energy and phosphate. This allows the cycle to continue. |
Stages in the Calvin Cycle
1. CO2 diffuses into leaf via stomata, dissolving in the water surrounding palisade mesophyll cells before diffusing into the cells. |
2. CO2 combines with the 5 carbon compound ribulose bisphosphate (RuBP) using the enzyme RuBisCO to form an unstable 6C compound. |
3. Unstable 6C compound immediately breaks down into 2 molecules of glycerate 3-phosphate (GP). |
4. Using one ATP molecule from the light reaction, GP is reduced to triose phosphate (TP) using hydrogen atoms from reduced HADP. |
5. Triose phosphate molecules combine in pairs to form hexose sugars. |
6. 5 out of every 6 triose phosphate molecules produced are used to regenerate RuBP (via the intermediate ribulose phosphate) using ATP from the light-dependent reaction to supply energy and phosphate. This allows the cycle to continue. |
Product Synthesis
Plants must produce all the carbohydrates, fats and proteins they need from the products of the Calvin Cycle. |
Fructose phosphate formed from the 2 molecules of triose phosphate can be converted to glucose, or combined with glucose to produce sucrose. |
Sucrose is then translocated in the phloem to the growing regions of the plant. |
Some a glucose is stored as starch, B glucose forms cellulose in cell walls. |
Fatty acids can be formed from glycerate 3-phosphate, and glycerol from triose phosphate, the building blocks of triglycerides. |
Proteins can be formed from glycerate 3-phosphate, but the amino group requires nitrogen from nitrate ions. |
Other compounds, e.g. chlorophyll, require additional ions e.g. Mg2+, and the middle lamella of cell walls needs Ca2+. |
A lack of nitrogen results in stunted growth in plants, as plants cannot synthesise proteins due to lack of nitrogen, whereas a lack of magnesium causes chlorosis, the yellowing of the leaves, as chlorophyll cannot be synthesised. |
This can be shown experimentally by placing plants in soils with different nutrient contents and observing growth. |
Limiting factors in photosynthesis
The rate of photosynthesis is controlled by a number of factors including the concentration of CO2, light intensity, and temperature. |
The limiting factor is the one which is in shortest supply which controls the rate-limiting step, and therefore an increase in it increases the rate of photosynthesis. |
Limiting Factors Table
Factor |
Explanation |
Carbon Dioxide |
At low concentrations, CO2 concentration is limiting, but above 0.5%, the rate plateaus, showing that something else must be limiting. Above 1% the stomata close, preventing the uptake of carbon dioxide. |
Light intensity |
As light intensity increases the rate of photosynthesis increases up to about 10,000 lux (SI unit if illuminance) when sone other factor becomes limiting. At very high light intensities the rate decreases as chloroplast pigments become bleached. Different plants have evolved to be most efficient at light intensities found in their environment, e.g. sun and shade plants. |
Temperature |
Temperature increases the kinetic energy of the reactants and enzymes involved in photosynthesis. Unlike other factors, a plateau is not reached as enzymes, e.g. RuBisCO, begin to denature so the rate of photosynthesis decreases above the optimum temperature. This will be higher in species adapted to hot, dry environments. |
Measuring the rate of photosynthesis
Aquatic plants are a good subject to use when investigating how different factors affect photosynthesis. |
Temperature and CO2 concentration are more easily controlled than with terrestrial plants, by using a water bath and controlling hydrogen carbonate concentration. |
It is also easy to collect and accurately measure the oxygen produced in a capillary tube. |
The volume of the bubble collected is calculated by the formula: |
Volume = pi r2 x length of bubble |
Where pi = 3.14 and r = radius or diameter/2 |
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