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1. Photosynthesis in Chloroplasts 2. Induction of Electron Transfer 3. Light Reactions: Role of Photosystems 4. Accessory Pigments
1. Photosynthesis in Chloroplasts
Photosynthesis Photosynthesis is the vital process happening in plants where the light energy is used for the synthesis of carbohydrates from water and CO2. For the organisms which give out oxygen, the empirical equation of photosynthesis can be written as follows:
presence of light CO2 + H 2 O ⎯⎯⎯⎯⎯⎯ [CH 2 O] + O 2 →
[CH2O] is represented by a carbohydrate molecule like glucose or fructose which is a six-carbon sugar. Photosynthesis is basically a light dependent reaction in which hydrogen from a suitable oxidizable compound reduces carbon dioxide into carbohydrates. This reaction can be well expressed in the following equation:
presence of light 2H 2 X + CO2 ⎯⎯⎯⎯⎯⎯ 2X + [CH 2O] + H 2O →
Considering the green plants, H2O is the hydrogen donor and it is oxidized into O2. There are some exceptional organisms which do not release oxygen during photosynthesis. For example, in case of purple and green sulphur bacteria, the hydrogen donor is H2S instead of H2O and hence the oxidation product will be Sulphur instead of Oxygen. Thus it is clear that the O2 which is released from the green plants come from the H2O and not from the CO2 molecule. Hence the overall process of photosynthesis can be written as follows:
presence of light 6CO 2 + 12H 2 O ⎯⎯⎯⎯⎯⎯ C 6 H12 O 6 + 6H 2 O + 6O 2 →
C6H12O6 represents glucose, the simplest carbohydrate molecule. It is clear that photosynthesis is not a single step process; it is a multistep process. Chloroplasts: Leaves are the significant parts in plants in which the photosynthesis takes place. The mesophyll cells present in the leaf contains a large number of chloroplasts. The chloroplasts are aligned along the walls of the mesophyll cells so that they can receive a maximum amount of incident light. The chloroplasts consists of three parts namely membranous grana, stroma lamella and the fluid stroma. Each part has its own significance carrying out specific function. Thylakoids are membranous disk-like structures that are stacked together like a stack of coins. Chlorophyll and carotenoid pigments are located inside the membranes of the thylakoids.
Grana trap the light energy and are responsible for the synthesis of NADPH and ATP. These reactions are directly dependent and driven by the light and hence they are called as light reactions. The fluid-filled space surrounding the grana is the stroma.
Many enzymes needed in photosynthesis are found in the stroma. Inside the stroma, the CO2 inhaled by the plants undergoes enzymatic reactions leading to synthesis of sugars, which finally produces starch. These reactions are not directly in requirement of light but are dependent on the products of light reactions like NADPH and ATP. Hence they are known as dark reactions.
2. Induction of Electron Transfer:
Two sets of reactions rule over the light reactions. The reactions which happen in the presence of light which are denoted as ‘Light-Dependent Reactions’. These reactions occur inside the thylakoid membrane of the chloroplasts and are responsible for the production of ATP and NADPH. These two molecules are the major requirement for the production of glucose in the lightindependent reactions (dark reactions).
presence of light 6CO 2 + 12H 2O + energy ⎯⎯⎯⎯⎯→ C6 H12O 6 + 6O 2
This reaction of photosynthesis can be correlated with the following diagram of reactions.
The reactions which do not require light as a significant factor are denoted as ‘LightIndependent Reactions’. Light-independent reactions occur in stroma of the chloroplast in light or dark conditions and their purpose is to reduce CO2 to Glucose.
3. Light Reactions: Role of Photosystems
Light reactions include absorption of light, photolysis (splitting of water), release of oxygen and the formation of high- energy chemical intermediates. Various complexes are involved in this multi – step process and the pigments taking part are organized into two distinct photochemical light harvesting complexes (LHC) within the two photosystems.
A photosystem is a molecular assemblage consisting of several hundred pigment molecules and bounded proteins. Photosystems are an adaptation involved in the absorption of light energy and the production of high energy electrons in photosynthesis. Each photosystem consists of the two major parts: antenna molecules and reaction centre. Antenna molecules: An antenna molecule is a light absorbing accessory pigment molecule which assists in making the photosynthesis process more efficient. They are known as antenna because they absorb light energy analogously to how a radio antenna absorbs radio waves. An antenna molecule consists of a carotenoid pigment, absorbing blue and green light or a phycocyanin absorbing green and yellow. Reaction centre: The reaction center consists of one of the several different chlorophyll molecules. The reaction centre is different for both of the photosystems. The photosystem II has a reaction center consisting of chlorophyll ‘a’ molecule. In plants there is also a second type of reaction centre in photosystem I and this reaction center has chlorophyll called chlorophyll ‘b’. The reaction centre in photosystem II is designated P680, since chlorophyll ‘a’ has its maximum absorption at 680 nm. The reaction centre in photosystem I is often called P700 since its pigment absorbs its optimum at 700nm.
In the photosystem II, the reaction centre chlorophyll ‘a’ absorbs 680 nm wavelength of red light. This causes the electrons to be excited and jumped into an orbit beyond the atomic nucleus.
These electrons are accepted by an electron accepter which allows them to pass them into an electron transport system (ETC) which consists of cytochromes. These electrons are not consumed when they pass through the electron transport chain, but passed into the pigments of Photosystem I. At the same time, the electrons present in the reaction centre of PS I are also excited when they receive the red light which is of wavelength 700 nm. These excited electrons are then transferred to another accepter molecule having greater redox potential. These electrons are now in a downhill again to the energy –rich NADP+ molecules. The addition of these electrons reduces NADP+ to NADH + H+. This whole system of electron transfer is called Z scheme due to its characteristic shape. It starts from PS II, uphill to the accepter, down the electron transport chain to PS I, excitation of electrons, transfer to another accepter, and finally down hill to NADP+. When all the carriers are placed in a sequence on a redox potential scale, this particular characteristic shape will be formed.
The antennas are represented by a single green circle. The blue circles represent the electron transport system (ETC).
The electron transport system is implanted within the thylakoid membrane and functions in the production of ATP. The system contains membrane-bound electron carriers that pass electrons from one to another.
As a result of gaining an electron (reduction), the first carrier of the electron transport system gains energy. It uses some of the energy to pump H+ into the thylakoid.
The carrier then passes the electron to the next carrier, because it used some energy to pump H+ and has less energy (reducing capability) to pass to the next H+ pump. This carrier uses some of the remainder of the energy to pump more H+ into the thylakoid. The electron is passed to the next carrier which also pumps H+.
The electron transport system functions to create a concentration gradient of H+ inside the thylakoid. The concentration gradient of H+ is used to synthesize ATP.
ATP is produced from ADP and Pi when hydrogen ions pass out of the thylakoid through ATP synthase. This method of synthesizing ATP by using a H+ gradient in the thylakoid is called photophosphorylation.
At this point, the electron has little energy left for reducing and is passed to the P700 antenna. A pigment molecule in the P700 antenna absorbs a photon of solar (light) energy. The energy from that molecule is passed to neighboring molecules within the antenna. The energy is eventually passed to the reaction center of this antenna. As a result of being energized, the P700 reaction center loses the electron to an electron acceptor. The acceptor passes it to NADP+, which becomes reduced to NADPH. According to the following equation, NADP+ has the capacity to carry two electrons. NADP+ + 2e- + H+ → NADPH
The electron transport system and photophosphorylation in the chloroplast is similar to the system found in the mitochondria to produce ATP during cellular respiration.
The electron that was lost from the antenna complex of photosystem I is replaced by splitting water. (Photolysis) Photolysis of water: The electrons which were moved from the PS II are replaced by the available electrons from the photolysis of water. This splitting of water is associated with PS II; water is split into H+, [O] and electrons.
Thus oxygen is created which is one of the important net products of photosynthesis. The electrons needed to replace those which are removed from PS I are now provided by PS II.
2 H 2O ⎯ 4 H + + O2 + 4e − ⎯→
Water splitting complex is associated with the PS II, which is itself actually located on the inner side of the membrane of the thylakoid. In the light reactions, electrons move one way from water to NADPH and the energy of sunlight is used to synthesize ATP.
4. Accessory Pigments
Pigments are the substances that have the capability to absorb light, at particular wavelengths of light. In order to convert light energy to chemical energy, the organism must first absorb light energy with the pigments involved in photosynthesis. Hence pigments are significant for photosynthesis. When light is absorbed, the energy stored in it is taken up by some of the electrons in the outer most energy level of the pigment molecules. The electrons jump up to a higher energy level. During photosynthesis the energy stored by these electrons can be harvested in the next steps of photosynthesis. The color seen in the leaves of the plants are not due to the action of a single pigment but due to the result of four major pigments, Chlorophyll a, which depicts bright or blue green in the chromatogram, Chlorophyll b which is yellow green pigment, yellow xanthophylls and yellow –orange carotenoids. The maximum absorption of light is shown by the chlorophyll a pigment and it is the chief pigment of photosynthesis comparing all the rest of the pigments. Chlorophyll ‘a’ is the key photosynthetic pigment in all organisms except bacteria. All the other pigments are called accessory pigments which absorb slightly different wavelengths of light. They do not participate directly in photosynthetic reactions but are able to pass their energy to chlorophyll a. The blend of all of the pigments increases the range of colors that plants can use in photosynthesis. They also protect the chlorophyll a from photo – oxidation. In basic biology laboratories, an instrument called as spectrophotometer is used to determine the percentage of light absorbed or reflected at particular wavelengths of light from pigments extracted from plants or other resources.
Points to Remember:
• • • • • • • • • Photosynthesis is the vital process in plants where light energy is used for the synthesis of carbohydrates from water and CO2. Photosynthesis is basically a light dependent reaction in which hydrogen from a suitable oxidisable compound reduces carbon dioxide into carbohydrates. In case of green plants, H2O is the hydrogen donor and it is oxidized into O2. In case of purple and green sulphur bacteria, the hydrogen donor is H2S instead of H2O and hence the oxidation product will be Sulphur instead of Oxygen. Large number of chloroplasts is present in the mesophyll cells of the leaf. Chloroplasts consists of three parts namely membranous grana, stroma lamella and the fluid stroma. Thylakoids are membranous disk-like structures that are stacked together like a stack of coins. Chlorophyll and carotenoid pigments are located inside the membranes of the thylakoids. Two sets of reactions rule over the light reactions: ‘Light-Dependent Reactions’ which happen in the presence of light and reactions which do not require light as a significant factor are ‘Light-Independent Reactions’. • • • • • Light-independent reactions occur in stroma of the chloroplast in light or dark conditions and their purpose is to reduce CO2 to glucose. Light reactions include absorption of light, photolysis (splitting of water), release of oxygen and the formation of high- energy chemical intermediates. A photosystem is a molecular assemblage consisting of several hundred pigment molecules and bounded proteins. Each photosystem consists of the two major parts: antenna molecules and reaction centre. The reaction centre in photosystem II is designated P680, since chlorophyll a has its maximum absorption at 680 nm, whereas in photosystem I, the reaction centre is often called P700, where its maximum absorption is 700 nm. • Electron transport system is implanted within the thylakoid membrane and functions in the production of ATP.
• • • • • •
ATP is produced from ADP and Pi when hydrogen ions pass out of the thylakoid through ATP synthase. Photophosphorylation is the method of synthesizing ATP by using a H+ gradient in the thylakoid. Splitting of water (Photolysis) is associated with PS II where the water is split into H+, [O] and electrons. Oxygen, one of the important net products of photosynthesis is released during photolysis of water. Pigments are the substances that have the capability to absorb light, at particular wavelengths of light. Chlorophyll a, depicts bright or blue green in the chromatogram, Chlorophyll b is yellow green pigment, xanthophylls are yellow and carotenoids are yellow – orange.
Maximum absorption of light is shown by the chlorophyll a pigment and it is the chief pigment of photosynthesis comparing all the rest of the pigments. Spectrophotometer is used to determine the percentage of light absorbed or reflected at particular wavelengths of light from pigments extracted from plants or other resources.
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