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==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 ~~at least four~~ ABA receptors (PYR1/RCAR11, PYL1/RCAR12, PYL2/RCAR14, PYL9/RCAR1)<ref>PMID:19407143</ref><ref>PMID:19407142</ref>, a protein phosphatase ~~2C in Clade C~~ (ABI1, ABI2, ~~and~~ HAB1)<ref>PMID:19407143</ref><ref>PMID:19407142</ref>, ~~and a~~ protein ~~kinase in subclass III of SnRK2s~~ ([[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>

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.ABA) binds to a protein phosphatase 2C<ref>PMID:19624469</ref> (R.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.ABA) binds to a protein phosphatase 2C<ref>PMID:19624469</ref> (R.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 ~~PYR2~~, [[3kdh]]; R.ABA, ~~PYR2~~.ABA, [[3kdi]]; K-P; SnRK2.6-HAB1, [[3ujg]]; R.ABA-P, ~~PYR2~~.ABA-ABI2, [[3ujl]]; K, SnRK2.6,[[3uc4]].

Structures in the figure are: R, apo PYL2, [[3kdh]]; R.ABA, PYL2.ABA, [[3kdi]]; K-P; SnRK2.6-HAB1, [[3ujg]]; R.ABA-P, PYL2.ABA-ABI2, [[3ujl]]; K, SnRK2.6,[[3uc4]].

~~==Pyr~~/~~Pyl~~/RCAR family of ABA receptors==

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~~

~~PYR1, PYL1, PYL2, and PYL3 show ABA-dependent inhibition~~ of ~~PP2Cs such as ABI1, HYPERSENSITIVE TO ABA 1 (HAB1), HAB2, and~~ ABA~~-HYPERSENSITIVE GERMINATION 3 (AHG3, also named PP2CA)<ref>PMID:23265948</ref>~~

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|-

|<applet load='3k3k' size='300' frame='true' align='left' caption='3k3k - PYR1 dimer' scene = '56/564059/Pyr1dimer/2'/> '''3k3k scene''' <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'/> '''3klx scene''' <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'/> '''4dsc scene''' <scene name='56/564059/Pylabatransdimer/2'>PYL3.ABA ''trans'' dimer'</scene>

|}

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[[3kay]], apo AtPYL1

[[3kdh]], [[3kaz]], [[3kl1]] apo AtPYL2

[[3klx]], ~~apo~~ AtPYL3

[[3klx]], Apo AtPYL3

[[4jdl]], Apo AtPYL5

[[3rt2]], [[3uqh]] apo AtPYL10

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 ==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 at least four ABA receptors (PYR1/RCAR11, PYL1/RCAR12, PYL2/RCAR14, PYL9/RCAR1)<ref>PMID:19407143</ref><ref>PMID:19407142</ref>, a protein phosphatase 2C in Clade C (ABI1, ABI2, and HAB1)<ref>PMID:19407143</ref><ref>PMID:19407142</ref>, and a protein kinase in subclass III of SnRK2s ([[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>
+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 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]].
+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]].
-==Pyr/Pyl/RCAR family of ABA receptors==
+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
-PYR1, PYL1, PYL2, and PYL3 show ABA-dependent inhibition of PP2Cs such as ABI1, HYPERSENSITIVE TO ABA 1 (HAB1), HAB2, and ABA-HYPERSENSITIVE GERMINATION 3 (AHG3, also named PP2CA)<ref>PMID:23265948</ref>
@@ Line 28: / Line 25: @@
 |-
 |<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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 [[3kay]], apo AtPYL1 <br>
 [[3kdh]], [[3kaz]], [[3kl1]] apo AtPYL2<br>
-[[3klx]], apo AtPYL3<br>
+[[3klx]], Apo AtPYL3<br>
 [[4jdl]], Apo AtPYL5<br>
 [[3rt2]], [[3uqh]] apo AtPYL10<br>