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Cooper GM. The Cell: A Molecular Approach. 2nd edition. Sunderland (MA): Sinauer Associates; 2000.
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During photosynthesis, energy from sunlight is harvested and used to drive the synthesis of glucose from CO2 and H2O. By converting the energy of sunlight to a usable form of potential chemical energy, photosynthesis is the ultimate source of metabolic energy for all biological systems. Photosynthesis takes place in two distinct stages. In the light reactions, energy from sunlight drives the synthesis of ATP and NADPH, coupled to the formation of O2 from H2O. In the dark reactions, so named because they do not require sunlight, the ATP and NADPH produced by the light reactions drive glucose synthesis. In eukaryotic cells, both the light and dark reactions of photosynthesis occur within chloroplasts—the light reactions in the thylakoid membrane and the dark reactions within the stroma. This section discusses the light reactions of photosynthesis, which are related to oxidative phosphorylation in mitochondria. The dark reactions were discussed in detail in Chapter 2.
Electron Flow through Photosystems I and II
Sunlight is absorbed by photosynthetic pigments, the most abundant of which in plants are the chlorophylls. Absorption of light excites an electron to a higher energy state, thus converting the energy of sunlight to potential chemical energy. The photosynthetic pigments are organized into photocenters in the thylakoid membrane, each of which contains hundreds of pigment molecules (Figure 10.20). The many pigment molecules in each photocenter act as antennae to absorb light and transfer the energy of their excited electrons to a chlorophyll molecule that serves as a reaction center. The reaction center chlorophyll then transfers its high-energy electron to an acceptor molecule in an electron transport chain. High-energy electrons are then transferred through a series of membrane carriers, coupled to the synthesis of ATP and NADPH.
Organization of a photocenter. Each photocenter consists of hundreds of antenna pigment molecules, which absorb photons and transfer energy to a reaction center chlorophyll. The reaction center chlorophyll then transfers its excited electron to an acceptor (more...)
The best characterized photosynthetic reaction center is that of the bacterium Rhodopseudomonas viridis, the structure of which was determined by Johann Deisenhofer, Hartmut Michel, Robert Huber, and their colleagues in 1985 (Figure 10.21). The reaction center consists of three transmembrane polypeptides, bound to a c-type cytochrome on the exterior side of the membrane. Energy from sunlight is captured by a pair of chlorophyll molecules known as the special pair. Electrons are then transferred from the special pair to another pair of chlorophylls and from there to other prosthetic groups (pheophytins and quinones). From there the electrons are transferred to a cytochrome bc complex in which electron transport is coupled to the generation of a proton gradient. The electrons are then transferred to the reaction center cytochrome and finally returned to the chlorophyll special pair. The reaction center thus converts the energy of sunlight to high-energy electrons, the potential energy of which is converted to a proton gradient by the cytochrome bc complex.
Structure of a photosynthetic reaction center. The reaction center of R. viridis consists of three transmembrane proteins (purple, blue, and beige) and a c-type cytochrome (green). Chlorophylls and other prosthetic groups are colored yellow. (Courtesy (more...)
The proteins involved in the light reactions of photosynthesis in plants are organized into five complexes in the thylakoid membrane (Figure 10.22). Two of these complexes are photosystems (photosystems I and II), in which light is absorbed and transferred to reaction center chlorophylls. High-energy electrons are then transferred through a series of carriers in both photosystems and in a third protein complex, the cytochromebfcomplex. As in mitochondria, these electron transfers are coupled to the transfer of protons into the thylakoid lumen, thereby establishing a proton gradient across the thylakoid membrane. The energy stored in this proton gradient is then harvested by a fourth protein complex in the thylakoid membrane, ATP synthase, which (like the mitochondrial enzyme) couples proton flow back across the membrane to the synthesis of ATP.
Electron transport and ATP synthesis during photosynthesis. Five protein complexes in the thylakoid membrane function in electron transport and the synthesis of ATP and NADPH. Photons are absorbed by complexes of pigment molecules associated with photosystems (more...)
One important difference between electron transport in chloroplasts and that in mitochondria is that the energy derived from sunlight during photosynthesis not only is converted to ATP but also is used to generate the NADPH required for subsequent conversion of CO2 to carbohydrates. This is accomplished by the use of two different photosystems in the light reactions of photosynthesis, one to generate ATP and the other to generate NADPH. Electrons are transferred sequentially between the two photosystems, with photosystem I acting to generate NADPH and photosystem II acting to generate ATP.
The pathway of electron flow starts at photosystem II, which is homologous to the photosynthetic reaction center of R. viridis already described. However, at photosystem II the energy derived from absorption of photons is used to split water molecules to molecular oxygen and protons (see Figure 10.22). This reaction takes place within the thylakoid lumen, so the release of protons from H2O establishes a proton gradient across the thylakoid membrane. The high-energy electrons derived from this process are transferred through a series of carriers to plastoquinone, a lipid-soluble carrier similar to coenzyme Q (ubiquinone) of mitochondria. Plastoquinone carries electrons from photosystem II to the cytochrome bf complex, within which electrons are transferred to plastocyanin and additional protons are pumped into the thylakoid lumen. Electron transport through photosystem II is thus coupled to establishment of a proton gradient, which drives the chemiosmotic synthesis of ATP.
From photosystem II, electrons are carried by plastocyanin (a peripheral membrane protein) to photosystem I, where the absorption of additional photons again generates high-energy electrons. Photosystem I, however, does not act as a proton pump; instead, it uses these high-energy electrons to reduce NADP+ to NADPH. The reaction center chlorophyll of photosystem I transfers its excited electrons through a series of carriers to ferrodoxin, a small protein on the stromal side of the thylakoid membrane. The enzyme NADP reductase then transfers electrons from ferrodoxin to NADP+, generating NADPH. The passage of electrons through photosystems I and II thus generates both ATP and NADPH, which are used by the Calvin cycle enzymes in the chloroplast stroma to convert CO2 to carbohydrates (see Figure 2.39).
Cyclic Electron Flow
A second electron transport pathway, called cyclic electron flow, produces ATP without the synthesis of NADPH, thereby supplying additional ATP for other metabolic processes. In cyclic electron flow, light energy harvested at photosystem I is used for ATP synthesis rather than NADPH synthesis (Figure 10.23). Instead of being transferred to NADP+, high-energy electrons from photosystem I are transferred to the cytochrome bf complex. Electron transfer through the cytochrome bf complex is then coupled, as in photosystem II, to the establishment of a proton gradient across the thylakoid membrane. Plastocyanin then returns these electrons to photosystem I in a lower energy state, completing a cycle of electron transport in which light energy harvested at photosystem I is used to pump protons at the cytochrome bf complex. Electron transfer from photosystem I can thus generate either ATP or NADPH, depending on the metabolic needs of the cell.
The pathway of cyclic electron flow. Light energy absorbed at photosystem I (PS I) is used for ATP synthesis rather than NADPH synthesis. High-energy electrons generated by photon absorption are transferred to the cytochrome bf complex rather than to (more...)
The ATP synthase of the thylakoid membrane is similar to the mitochondrial enzyme. However, the energy stored in the proton gradient across the thylakoid membrane, in contrast to the inner mitochondrial membrane, is almost entirely chemical in nature. This is because the thylakoid membrane, although impermeable to protons, differs from the inner mitochondrial membrane in being permeable to other ions, particularly Mg2+ and Cl-. The free passage of these ions neutralizes the voltage component of the proton gradient, so the energy derived from photosynthesis is conserved mainly as the difference in proton concentration (pH) across the thylakoid membrane. However, because the thylakoid lumen is a closed compartment, this difference in proton concentration can be quite large, corresponding to a differential of more than three pH units between the stroma and the thylakoid lumen. Because of the magnitude of this pH differential, the total free energy stored across the thylakoid membrane is similar to that stored across the inner mitochondrial membrane.
For each pair of electrons transported, two protons are transferred across the thylakoid membrane at photosystem II and two to four protons at the cytochrome bf complex. Since four protons are needed to drive the synthesis of one molecule of ATP, passage of each pair of electrons through photosystems I and II by noncyclic electron flow yields between 1 and 1.5 ATP molecules. Cyclic electron flow has a lower yield, corresponding to between 0.5 and 1 ATP molecules per pair of electrons.
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By agreement with the publisher, this book is accessible by the search feature, but cannot be browsed.