Photosystem II

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<applet load="1s5l" size="400" frame="true" align="right" />
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<StructureSection load='3a0b' size='350' caption='Photosystem II, [[3a0b]]' scene='' >
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[[Image:1s5l.gif|250px|left]]
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==Background==
==Background==
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This structure of Photosystem II was crystallized from the cyanobacteria, ''Thermosynechococcus elongatus'', at 3.0Å <ref>Loll, B., Kern, J., Saenger, W., Zouni, A., Biesiadka, J. "Towards complete cofactor arrangement in the 3.0 A resolution structure of photosystem II." Nature 2005, 438: 1040-1044. PMID: [http://www.ncbi.nlm.nih.gov/pubmed/16355230 16355230]</ref> and at 3.50 Å <ref>Ferreira, K.N., Iverson, T.M., Maghlaoui, K., Barber, J., Iwata, S. "Architecture of the photosynthetic oxygen-evolving center." Science, March 19, 2004, 303 (5665), 1831-8. PMID:[http://www.ncbi.nlm.nih.gov/pubmed/14764885 14764885]</ref>. PDB codes are [http://www.proteopedia.org/wiki/index.php/2axt 2AXT] and [http://www.proteopedia.org/wiki/index.php/1s5l 1S5L], respectively. Cyanobacteria and plants both contain Photosystem II while photosynthetic bacteria contain the bacterial reaction center. This photosynthetic protein complex is associated with a variety of functional ligands. It is a <scene name='Photosystem_II/Psii_dimer/1'>dimer</scene> composed mainly of alpha-helices. Nineteen <scene name='Photosystem_II/Protein_only/1'>subunits</scene> are in each monomer, with multiple extrinsic subunits associated with the oxygen evolving complex missing from this crystallization. Photosystem II is a membrane bound protein complex that in plants is associated with the thylakoid membrane of chloroplasts. <scene name='Photosystem_II/Hydrophobic_polar/1'>Polar and hydrophobic</scene> regions correlate with membrane associated nature of the protein. '''<FONT COLOR="#616D7E">Hydrophobic</FONT>''' helices make up the transmembranal portion, while '''<FONT COLOR="#C031C7">polar</FONT>''' residues are concentrated externally on either side of the membrane.
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This structure of '''Photosystem II''' was crystallized from the cyanobacteria, ''Thermosynechococcus elongatus'', at 3.0Å <ref>PMID: 16355230</ref> and at 3.50 Å <ref name="Archit">PMID: 14764885</ref>. PDB codes are [[2axt]] and [[1s5l]], respectively. Cyanobacteria and plants both contain Photosystem II while photosynthetic bacteria contain the bacterial reaction center. This photosynthetic protein complex is associated with a variety of functional ligands. It is a <scene name='Photosystem_II/Psii_dimer/1'>dimer</scene> composed mainly of alpha-helices. Nineteen <scene name='Photosystem_II/Protein_only/1'>subunits</scene> are in each monomer, with multiple extrinsic subunits associated with the oxygen evolving complex missing from this crystallization. Photosystem II is a membrane bound protein complex that in plants is associated with the thylakoid membrane of chloroplasts. <scene name='Photosystem_II/Hydrophobic_polar/1'>Polar and hydrophobic</scene> regions correlate with membrane associated nature of the protein. '''<span style="color:gray;background-color:black;font-weight:bold;">Hydrophobic</span>''' helices make up the transmembranal portion, while '''<FONT COLOR="#C031C7">polar</FONT>''' residues are concentrated externally on either side of the membrane.
==Photosynthesis==
==Photosynthesis==
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Photosystem II is an integral part of photosynthesis, the conversion of light energy into chemical energy by living organisms. Photosystem II is linked to a variety of other proteins, including Photosytem I. These proteins ultimately produce NADPH and ATP that power the Calvin cycle. Using this energy, glucose is synthesized from carbon dioxide and water.
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Photosystem II is an integral part of photosynthesis, the conversion of light energy into chemical energy by living organisms. Photosystem II is linked to a variety of other proteins, including Photosytem I. These proteins ultimately produce NADPH and ATP that power the [[Calvin cycle]]. Using this energy, glucose is synthesized from carbon dioxide and water. See also [[Photosynthesis]].
==Electron Transfer==
==Electron Transfer==
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<applet load="1s5l" size="400" frame="true" align="right" />
 
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[[Image:Chlorophyll_a.svg.png|thumb|170px|left|structure of chlorophyll ''a'']]
[[Image:Chlorophyll_a.svg.png|thumb|170px|left|structure of chlorophyll ''a'']]
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<scene name='Photosystem_II/Chlorophyll_green/4'>Chlorophyll</scene> surround Photosystem II and capture energy from sunlight, exciting electrons. Chlorophyll are highly conjugated and absorb visible light, along with accessory light harvesting pigments such as <scene name='Photosystem_II/Betacarotene/3'>beta carotene</scene>. Beta carotene absorbs visible light of other wavelengths and also protects Photosystem II by destroying reactive oxygen species that result from this photoexcitation. [[Image:b-car.svg.png|b-car.svg.png|thumb|right|400px|structure of beta carotene]] Electrons are passed from chlorophyll to <scene name='Photosystem_II/Pheophytin_purple/5'>pheophytin</scene>. Pheophytin are very similar to chlorophyll except they contain 2 H<sup>+</sup> instead of a Mg<sup>2+</sup> ion. From the pheophytin, electrons transferred to <scene name='Photosystem_II/Quinone_pink/5'>plastoquinones</scene>, which are reduced. Between each pair of quinones, an iron, in red, helps to transfer the electron. These plastoquinones eventually move to a plastoquinone pool which travels to another large protein subunit, cytochrome b <sub>6</sub>/ f. Eventually these electrons reduce NADP<sup>+</sup> to NADPH. The <scene name='Photosystem_II/Electron_pathway/3'>electron pathway</scene> through Photosystem II is shown, with '''<FONT COLOR="#F87217">beta-carotenes</FONT>''', '''<FONT COLOR="#571B7e">pheophytins</FONT>''', '''<FONT COLOR="#E42217">iron</FONT>''' and '''<FONT COLOR="#F535AA">plasotoquinones</FONT>'''.
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<scene name='Photosystem_II/Chlorophyll_green/4'>Chlorophyll</scene> surround Photosystem II and capture energy from sunlight, exciting electrons. Chlorophyll are highly conjugated and absorb visible light, along with accessory light harvesting pigments such as <scene name='Photosystem_II/Betacarotene/3'>beta carotene</scene>. Beta carotene absorbs visible light of other wavelengths and also protects Photosystem II by destroying reactive oxygen species that result from this photoexcitation. Electrons are passed from chlorophyll to <scene name='Photosystem_II/Pheophytin_purple/5'>pheophytin</scene>. Pheophytin are very similar to chlorophyll except they contain 2 H<sup>+</sup> instead of a Mg<sup>2+</sup> ion. From the pheophytin, electrons transferred to <scene name='Photosystem_II/Quinone_pink/5'>plastoquinones</scene>, which are reduced. Located between each pair of quinones, an iron helps to transfer the electron. These plastoquinones eventually move to a plastoquinone pool which travels to another large protein subunit, cytochrome b <sub>6</sub>/ f. Eventually these electrons reduce NADP<sup>+</sup> to NADPH. The <scene name='Photosystem_II/Electron_pathway/3'>electron pathway</scene> through Photosystem II is shown, with '''<span style="color:orange;background-color:black;font-weight:bold;">beta-carotenes</span>''', '''<FONT COLOR="#571B7e">pheophytins</FONT>''', '''<FONT COLOR="#E42217">iron</FONT>''' and '''<FONT COLOR="#F535AA">plasotoquinones</FONT>'''.
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[[Image:plastoquinone.jpg|thumb|300px|right|reduced plastoquinone]]
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{{Clear}}
==Oxygen Evolution==
==Oxygen Evolution==
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Another important facet of Photosystem II is its ability to oxidize water to oxygen with its <scene name='Photosystem_II/Oxygen_evolving_centers/11'>oxygen evolving centers</scene>. These centers are <scene name='Photosystem_II/Oxygen_evolving_centers/3'>cubane-like</scene> structures with 3 '''<FONT COLOR="#8D38C9">manganese</FONT>''', 4 '''<FONT COLOR="#C11B17">oxygen</FONT>''' and a '''<FONT COLOR="#59E817">calcium</FONT>''' linked to a fourth manganese.<ref>Ferreira, K.N., Iverson, T.M., Maghlaoui, K., Barber, J., Iwata, S. "Architecture of the photosynthetic oxygen-evolving center." Science, March 19, 2004, 303 (5665), 1831-8. PMID:[http://www.ncbi.nlm.nih.gov/pubmed/14764885 14764885]</ref> Oxidation of water leaves 2 H <sup>+</sup> on the lumenal side of the membrane, helping to establish the proton gradient essential for ATP synthesis in the CF<sub>1</sub>CF<sub>0</sub>-ATP sythase protein.
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Another important facet of Photosystem II is its ability to oxidize water to oxygen with its <scene name='Photosystem_II/Oxygen_evolving_centers/11'>oxygen evolving centers</scene>. These centers are <scene name='Photosystem_II/Single_oxygen_evolving/1'>cubane-like</scene> structures with 3 '''<FONT COLOR="#8D38C9">manganese</FONT>''', 4 '''<FONT COLOR="#C11B17">oxygen</FONT>''' and a '''<span style="color:lime;background-color:black;font-weight:bold;">calcium</span>''' linked to a fourth manganese<ref name="Archit" />. Oxidation of water leaves 2 H <sup>+</sup> on the lumenal side of the membrane, helping to establish the proton gradient essential for ATP synthesis in the CF<sub>1</sub>CF<sub>0</sub>-ATP sythase protein.
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[[Image:b-car.svg.png|b-car.svg.png|thumb|left|400px|structure of beta carotene]]
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[[Image:plastoquinone.jpg|thumb|300px|left|reduced plastoquinone]]
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==3D structures of photosystem II==
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[[Photosystem II 3D structures]]
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</StructureSection>
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==Additional Resources==
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For additional information, see: [[Photosynthesis]]
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<br />
==References==
==References==
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<references/>
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<references/>
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2. Garrett, R.H., Grisham, C.M. ''Biochemistry, 3rd Edition.'' Belmont, CA: Thomson Brooks/ Cole, 2005.
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[[Category:Topic Page]]

Current revision

Photosystem II, 3a0b

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Additional Resources

For additional information, see: Photosynthesis

References

  1. Loll B, Kern J, Saenger W, Zouni A, Biesiadka J. Towards complete cofactor arrangement in the 3.0 A resolution structure of photosystem II. Nature. 2005 Dec 15;438(7070):1040-4. PMID:16355230 doi:http://dx.doi.org/10.1038/nature04224
  2. 2.0 2.1 Ferreira KN, Iverson TM, Maghlaoui K, Barber J, Iwata S. Architecture of the photosynthetic oxygen-evolving center. Science. 2004 Mar 19;303(5665):1831-8. Epub 2004 Feb 5. PMID:14764885 doi:http://dx.doi.org/10.1126/science.1093087
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