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==Pyr/Pyl/RCAR family of ABA receptors==
[[Image:ABA model 2 med.jpg|left]]
[[Image:ABA model 2 med.jpg|left]]


==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.<ref>PMID:15012233</ref><ref>PMID:12045268</ref><ref>PMID:20192755</ref> As shown in the left figure, ABA binds to an ABA receptor (R)<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 (R<sup>.</sup>ABA-P), which was formerly in a complex with a SnRK2 protein kinase. The protein kinase is now free 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">PMID:19541597</ref> in seeds/seedlings and dormancy/growth arrest; or 3) phosphorylation of transcription factor AREB/ABF <ref>PMID: 16446457</ref><ref name = "Fujii2007">PMID:17307925</ref> in vegetative tissue and stress tolerance and growth regulation.  
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: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.  


The ABA receptor family includes PYR1, PYR2, PYL1, PYL2, and RCAR. The ABA-responsive protein phosphatase 2Cs are in Clade C and they include ABI1, ABI2, and HAB1. The ABA-activated protein kinases are in subclass III of SnRK2s and they include [[SnRK2.6/OST1/SRK2E]], and SNRK2.2/SRK2D and SnRK2.3/SRK2I
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]].


Structures in the figure are: R, apo PYR2, [[3kdh]];  R<sup>.</sup>ABA, PYR2<sup>.</sup>ABA, [[3kdi]]; K-P; SnRK2.6-HAB1, [[3ujg]]; R<sup>.</sup>ABA-P, PYR2<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]].  


The following Proteopedia pages


crystal structures of
==PYR/PYL/RCAR family of ABA receptors==
PYR1/RCAR11 and PYL9/RCAR1






{|
{|
|'''Left scene''' - apo PYL2 [[3KAZ]]
|'''Left panel''' - apo PYL2 [[3kdh]]
|'''Middle scene''' - ABA bound to PYL2  
|'''Middle panel''' - ABA bound to PYL2 [[3kdi]] or PYR1 [[3k3k]]
|'''Right scene''' - PYL.ABA bound to [[3KB3]]
|'''Right panel''' - PYL<sup>'''.'''</sup>ABA bound to HAB1[[3ujl]]
|-
| <applet load='3kdh' size='300' frame='true' align='left' caption='3kdh - apo-Pyl2' scene = '56/564059/Apopyl2/1'/><br  clear='both'>'''3kdh scenes''' <Br><scene name='56/564059/Apopyl2/1'>1. Default scene </scene> PYL2 is shown as a monomer. See also Scene 3 of the middle panel, which shows the receptor dimer. <br><scene name='56/564059/Apopyl2/3'>2. Open gate</scene> The entrance to binding pocket for ABA is regulated by a "latch" shown in orchid and a "gate" shown in blue. Proline 92 is shown in ball and stick. Here the gate and entrance to the binding site are open.
| <applet load='3kdi' size='300' frame='true' align='left' caption='3kdi - ABA bound to PYL2' scene = '56/564059/Abapyl2/1' /><Br clear='both'>'''3kdi scenes'''<Br><scene name='56/564059/Abapyl2/1'>1. Default scene </scene> ABA (CPK spheres) binds to a water-filled (not shown) pocket of PYL2. <br><scene name='56/564059/Abapyl2/2'>2. Closed gate </scene> The gate folds over ABA and interacts with the latch. <br>
|<applet load='3ujl' size='300' frame='true' align='left' caption='3ujl - PYR2-HAB1' scene = '56/564059/Pyl2hab1/2' /><Br clear='both'>'''3ujl scenes'''<Br><scene name='56/564059/Pyl2hab1/1'>1. Default scene</scene> Complex between PYL2 (blue) with bound ABA (CPK spheres)and HAB1 (gold), a protein phosphatase 2C. Magnesium ions in the active site of HAB1 are shown as green spheres. <br><scene name='56/564059/Pyl2hab1/5'>2. Closed gate locked by interaction with HAB1</scene> Gate residue proline 92 (blue ball and stick) interacts with typtophan 290 (gold ball and stick and residues in a hydrophobic loop (dark gold ball and stick) of HAB1. The gate also interacts with residues surrounding the phosphatase's active site, which is marked by magnesium ions (small green spheres).
|-
|-
| <applet load='3kaz' size='300' frame='true' align='left' caption='3kaz - apo-Pyl2' scene = '55/559985/Aposnrk2_6/6' /><br  clear='both'>'''3kaz scenes''' <Br>
|<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='3ujg' size='300' frame='true' align='left' caption='3ujg - SnRK2.6-HAB1' scene = '55/559985/Aposnrk2_6/2' /><Br clear='both'>'''3ujg scenes'''<Br>
|<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='3ujg' size='300' frame='true' align='left' caption='3ujg - SnRK2.6-HAB1' scene = '55/559985/Aposnrk2_6/2' /><Br clear='both'>'''3ujg scenes'''<Br>
|<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>
[[3k3k]], AtPYR1 dimer, one monomer is bound to ABA and the other unliganded<br>
[[3kay]], apo AtPYL1 <br>
[[3kdh]], [[3kaz]], [[3kl1]] apo AtPYL2<br>
[[3klx]], Apo AtPYL3<br>
[[4jdl]], Apo AtPYL5<br>
[[3rt2]], [[3uqh]] apo AtPYL10<br>
'''Structures with (+)-ABA'''<br>
[[3k90]], AtPYR1.ABA<br>
[[3k3k]], AtPyr1 dimer, one monomer is bound to ABA and the other unliganded<br>
[[3jrs]], AtPYL1.ABA<br>
[[3kdi]], [[3kb0]] AtPYL2.ABA<br>
[[4dsb]], [[4dsc]] AtPYL3 with ABA<br>
[[3oqu]], AtPYL9.ABA<br>
[[3r6p]], AtPYL10.ABA<br>
'''Structures with (-)-ABA'''<br>
[[4jda]], AtPYL3 with (-)-ABA<br>
'''Structures with pyrabactin'''<br>
[[3njo]], AtPYR1.Pyrabactin<br>
[[3nef]], [[3neg]], [[3nr4]] AtPYL1.pyrabactin<br>
[[3nj0]], [[3ns2]] AtPYL2.Pyrabactin<br>
[[3nj1]], AtPYL2 V114I mutant.Pyrabactin <br>
[[3nmh]], AtPYL2 in complex with pyrabactin<br>
[[3nmp]], AtPYL2 mutant A93F in complex with pyrabactin<br>
[[3oji]], AtPYL3 with pyrabactin(?)<br>
'''Structures with (+)-ABA or  homolog and a PP2C'''<br>
[[3qn1]], AtPYR1.ABA - AtHAB1<br>
[[3zvu]], AtPYR1 H60P mutant .ABA - AtHAB1<br>
[[3kb3]], AtPYL1.ABA - HAB1<br>
[[3jrq]], [[3kdj]] AtPYL1.ABA - ABI1<br>
[[3ujl]], AtPYL2.ABA - AtABI2<br>
[[4lga]], [[4lgb]] AtPYL2.ABA mimic - AtHAB1<br>
[[4ds8]], AtPYL3.ABA complex with AtHAB1
[[3rt0]], AtPYL10.ABA - AtHAB1<br>
'''Structures with pyrabactin or homolog and a PP2C'''<br>
[[4la7]], [[4lg5]] AtPYL2.Quinabactin - AtHAB1<br>
[[3nmn]], AtPYL1.pyrabactin in complex with AtABI1<br>
[[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 AtABI1<br>




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