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==ABA signaling pathway== | ==ABA signaling pathway== | ||
Abscisic acid (ABA,)[[Image:Abscisic acid.svg.png|thumb| Abscisic Acid]] is a plant hormone that regulates seed dormancy and germination, stomatal closure, and responses to drought stress, and its core signaling pathway has recently been identified.<ref>PMID:15012233</ref><ref>PMID:12045268</ref><ref>PMID:20192755</ref> In Arabidopsis, this pathway involves | Abscisic acid (ABA,)[[Image:Abscisic acid.svg.png|thumb| Abscisic Acid]] is a plant hormone that regulates seed development, dormancy and germination, stomatal closure, and responses to drought stress, and its core signaling pathway has recently been identified.<ref>PMID:15012233</ref><ref>PMID:12045268</ref><ref>PMID:20192755</ref> In Arabidopsis, this pathway involves families of ABA receptors (PYR1/RCAR11, PYL1/RCAR12, PYL2/RCAR14, PYL3/RCAR13, PYL8/RCAR3, PYL9/RCAR1)<ref>PMID:19407143</ref><ref>PMID:19407142</ref><ref>"PMID:22579247</ref><ref>PMID:23370718</ref>, protein phosphatase 2Cs (ABI1, ABI2, HAB1, HAB2, PP2CA/AHG3)<ref>PMID:19407143</ref><ref>PMID:19407142</ref><ref>PMID:16339800</ref>, SNRK2 protein kinases ([[SnRK2.6/OST1/SRK2E]], SNRK2.2/SRK2D and SnRK2.3/SRK2IA).<ref name = "Mustilli2002">PMID:12468729</ref><ref>PMID:12514244</ref><ref name = "Nakashima2009">PMID:19541597</ref><ref name = "Fujii2007">PMID:17307925</ref> | ||
The core ABA signaling pathway is shown in the figure to the left. In unstimulated cells, the ABA receptor (R) is an unliganded dimer<ref>PMID:19933100</ref><ref>PMID:23265948</ref> (monomer shown) in the cytosol and nucleus , and the SNRK2 protein kinase (K) is bound to a protein phosphatase 2C (P) in a complex (K-P) in which the kinase is dephosphorylated and inactivated by the phosphatase. When the ABA level rises ABA binds to the ABA receptor<ref>PMID:19893533</ref><ref>PMID:19407143</ref><ref>PMID:19407142</ref>. The activated receptor (R<sup>.</sup>ABA) binds to a protein phosphatase 2C<ref>PMID:19624469</ref> (R<sup>.</sup>ABA-P), and frees the protein kinase to be activated by autophosphorylation or phosphorylation by another protein kinase. Activation of SnRK2.6/OST1/SRK2E leads to phosphorylation of: 1) ion channels SLAC1<ref>PMID: 19955405</ref> and KAT1<ref>PMID:19785574</ref> in guard cells and stomatal closure; 2) transcription factor ABI5<ref name = "Nakashima2009"/> in seeds/seedlings and dormancy/growth arrest; or 3) phosphorylation of transcription factor AREB/ABF <ref>PMID: 16446457</ref><ref name = "Fujii2007"/> in vegetative tissue and stress tolerance and growth regulation. | The core ABA signaling pathway is shown in the figure to the left. In unstimulated cells, the ABA receptor (R) is an unliganded dimer<ref>PMID:19898420</ref><ref>PMID:19933100</ref><ref>PMID:23265948</ref><ref>"PMID:22579247</ref> (monomer shown) in the cytosol and nucleus, and the SNRK2 protein kinase (K) is bound to a protein phosphatase 2C (P) in a complex (K-P) in which the kinase is dephosphorylated and inactivated by the phosphatase. When the ABA level rises ABA binds to the ABA receptor<ref>PMID:19893533</ref><ref>PMID:19407143</ref><ref>PMID:19407142</ref>. The activated receptor (R<sup>.</sup>ABA) binds to a protein phosphatase 2C<ref>PMID:19624469</ref> (R<sup>.</sup>ABA-P), and this sequestration of the phosphatase frees the protein kinase to be activated by autophosphorylation or phosphorylation by another protein kinase. Activation of SnRK2.6/OST1/SRK2E leads to phosphorylation of: 1) ion channels SLAC1<ref>PMID: 19955405</ref> and KAT1<ref>PMID:19785574</ref> in guard cells and stomatal closure; 2) transcription factor ABI5<ref name = "Nakashima2009"/> in seeds/seedlings and dormancy/growth arrest; or 3) phosphorylation of transcription factor AREB/ABF <ref>PMID: 16446457</ref><ref name = "Fujii2007"/> in vegetative tissue and stress tolerance and growth regulation. | ||
Structures in the figure are: R, apo | Structures in the figure are: R, apo PYL2, [[3kdh]]; R<sup>.</sup>ABA, PYL2<sup>.</sup>ABA, [[3kdi]]; K-P; SnRK2.6-HAB1, [[3ujg]]; R<sup>.</sup>ABA-P, PYL2<sup>.</sup>ABA-ABI2, [[3ujl]]; K, SnRK2.6,[[3uc4]]. | ||
For further information about proteins involved in this pathway see [[PYR/PYL/RCAR family of ABA receptors]], [[ABA-regulated Protein Phosphatase 2C]], [[ABA-regulated SNRK2 Protein Kinase]]. | |||
==PYR/PYL/RCAR family of ABA receptors== | |||
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|- | |- | ||
|<applet load='3k3k' size='300' frame='true' align='left' caption='3k3k - PYR1 dimer' scene = '56/564059/Pyr1dimer/2'/><br clear='both'>'''3k3k scene'''<Br><scene name='56/564059/Pyr1dimer/2'>3. PYR1 dimer</scene> in which one monomer is bound to ABA. The native form of the receptor is a dimer<ref>PMID:19933100</ref><ref>PMID:23265948</ref>. | |<applet load='3k3k' size='300' frame='true' align='left' caption='3k3k - PYR1 dimer' scene = '56/564059/Pyr1dimer/2'/><br clear='both'>'''3k3k scene'''<Br><scene name='56/564059/Pyr1dimer/2'>3. PYR1 dimer</scene> in which one monomer is bound to ABA. The native form of the receptor is a dimer<ref>PMID:19933100</ref><ref>PMID:23265948</ref>. | ||
|<applet load='3klx' size='300' frame='true' align='left' caption='3klx - PYL3 ''cis'' dimer' scene = '56/564059/Apopyl3cisdimer/2'/><br clear='both'>'''3klx scene'''<Br><scene name='56/564059/Apopyl3cisdimer/2'>Apo PYL3 ''cis'' dimer </scene> | |||
|<applet load='4dsc' size='300' frame='true' align='left' caption='4dsc - PYL3.ABA ''trans'' dimer' scene = '56/564059/Pylabatransdimer/2'/><br clear='both'>'''4dsc scene'''<Br><scene name='56/564059/Pylabatransdimer/2'>PYL3.ABA ''trans'' dimer'</scene> | |||
|} | |} | ||
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Santiago J, Dupeux F, Betz K, Antoni R, Gonzalez-Guzman M, Rodriguez L, Márquez JA, Rodriguez PL.Plant Sci. 2012 182:3-11<ref>PMID:22118610</ref> | Santiago J, Dupeux F, Betz K, Antoni R, Gonzalez-Guzman M, Rodriguez L, Márquez JA, Rodriguez PL.Plant Sci. 2012 182:3-11<ref>PMID:22118610</ref> | ||
=== | ===PYR/PYL/RCAR structures=== | ||
At is ''Arabidopsis thaliana'' | |||
'''Apo structures'''<br> | '''Apo structures'''<br> | ||
[[3k3k]], | [[3k3k]], AtPYR1 dimer, one monomer is bound to ABA and the other unliganded<br> | ||
[[3kay]], apo AtPYL1 <br> | [[3kay]], apo AtPYL1 <br> | ||
[[3kdh]], [[3kaz]], [[3kl1]] apo AtPYL2<br> | [[3kdh]], [[3kaz]], [[3kl1]] apo AtPYL2<br> | ||
[[3klx]], | [[3klx]], Apo AtPYL3<br> | ||
[[4jdl]], Apo | [[4jdl]], Apo AtPYL5<br> | ||
[[3rt2]], [[3uqh]] apo | [[3rt2]], [[3uqh]] apo AtPYL10<br> | ||
'''Structures with (+)-ABA'''<br> | '''Structures with (+)-ABA'''<br> | ||
[[3k90]], | [[3k90]], AtPYR1.ABA<br> | ||
[[3k3k]], AtPyr1 dimer, one monomer is bound to ABA and the other unliganded<br> | [[3k3k]], AtPyr1 dimer, one monomer is bound to ABA and the other unliganded<br> | ||
[[3jrs]], AtPYL1.ABA<br> | [[3jrs]], AtPYL1.ABA<br> | ||
[[3kdi]], [[3kb0]] AtPYL2.ABA<br> | [[3kdi]], [[3kb0]] AtPYL2.ABA<br> | ||
[[4dsb]], [[4dsc]] AtPYL3 with ABA<br> | [[4dsb]], [[4dsc]] AtPYL3 with ABA<br> | ||
[[3oqu]], | [[3oqu]], AtPYL9.ABA<br> | ||
[[3r6p]], | [[3r6p]], AtPYL10.ABA<br> | ||
'''Structures with (-)-ABA'''<br> | '''Structures with (-)-ABA'''<br> | ||
[[4jda]], | [[4jda]], AtPYL3 with (-)-ABA<br> | ||
'''Structures with pyrabactin'''<br> | '''Structures with pyrabactin'''<br> | ||
[[3njo]], | [[3njo]], AtPYR1.Pyrabactin<br> | ||
[[3nef]], [[3neg]], [[3nr4]] | [[3nef]], [[3neg]], [[3nr4]] AtPYL1.pyrabactin<br> | ||
[[3nj0]], [[3ns2]] | [[3nj0]], [[3ns2]] AtPYL2.Pyrabactin<br> | ||
[[3nj1]], | [[3nj1]], AtPYL2 V114I mutant.Pyrabactin <br> | ||
[[3nmh]], AtPYL2 in complex with pyrabactin<br> | [[3nmh]], AtPYL2 in complex with pyrabactin<br> | ||
[[3nmp]], AtPYL2 mutant A93F in complex with pyrabactin<br> | [[3nmp]], AtPYL2 mutant A93F in complex with pyrabactin<br> | ||
[[3oji]], | [[3oji]], AtPYL3 with pyrabactin(?)<br> | ||
'''Structures with (+)-ABA or homolog and a PP2C'''<br> | '''Structures with (+)-ABA or homolog and a PP2C'''<br> | ||
[[3qn1]], | [[3qn1]], AtPYR1.ABA - AtHAB1<br> | ||
[[3zvu]], | [[3zvu]], AtPYR1 H60P mutant .ABA - AtHAB1<br> | ||
[[3kb3]], AtPYL1.ABA - HAB1<br> | [[3kb3]], AtPYL1.ABA - HAB1<br> | ||
[[3jrq]], [[3kdj]] AtPYL1.ABA - ABI1<br> | [[3jrq]], [[3kdj]] AtPYL1.ABA - ABI1<br> | ||
[[3ujl]], | [[3ujl]], AtPYL2.ABA - AtABI2<br> | ||
[[4lga]], [[4lgb]] | [[4lga]], [[4lgb]] AtPYL2.ABA mimic - AtHAB1<br> | ||
[[4ds8]], AtPYL3.ABA complex with | [[4ds8]], AtPYL3.ABA complex with AtHAB1 | ||
[[3rt0]], | [[3rt0]], AtPYL10.ABA - AtHAB1<br> | ||
'''Structures with pyrabactin or homolog and a PP2C'''<br> | '''Structures with pyrabactin or homolog and a PP2C'''<br> | ||
[[4la7]], [[4lg5]] | [[4la7]], [[4lg5]] AtPYL2.Quinabactin - AtHAB1<br> | ||
[[3nmn]], AtPYL1.pyrabactin in complex with | [[3nmn]], AtPYL1.pyrabactin in complex with AtABI1<br> | ||
[[3nmt]], AtPYL2 mutant A93F.pyrabactin in complex with type 2C protein phosphatase | [[3nmt]], AtPYL2 mutant A93F.pyrabactin in complex with type 2C protein phosphatase AtHAB1<br> | ||
[[3nmv]], AtPYL2 mutant A93F.pyrabactin in complex with type 2C protein phosphatase | [[3nmv]], AtPYL2 mutant A93F.pyrabactin in complex with type 2C protein phosphatase AtABI1<br> | ||