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ABA signaling pathway

Abscisic acid (ABA,)
Abscisic Acid
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.[1][2][3] In Arabidopsis, this pathway involves families of ABA receptors (PYR1/RCAR11, PYL1/RCAR12, PYL2/RCAR14, PYL3/RCAR13, PYL8/RCAR3, PYL9/RCAR1)[4][5][6][7], protein phosphatase 2Cs (ABI1, ABI2, HAB1, HAB2, PP2CA/AHG3)[8][9][10], SNRK2 protein kinases (SnRK2.6/OST1/SRK2E, SNRK2.2/SRK2D and SnRK2.3/SRK2IA).[11][12][13][14]

The core ABA signaling pathway is shown in the figure to the left. In unstimulated cells, the ABA receptor (R) is an unliganded dimer[15][16][17][18] (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[19][20][21]. The activated receptor (R.ABA) binds to a protein phosphatase 2C[22] (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[23] and KAT1[24] in guard cells and stomatal closure; 2) transcription factor ABI5[13] in seeds/seedlings and dormancy/growth arrest; or 3) phosphorylation of transcription factor AREB/ABF [25][14] in vegetative tissue and stress tolerance and growth regulation.

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.

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

Left panel - apo PYL2 3kdh Middle panel - ABA bound to PYL2 3kdi or PYR1 3k3k Right panel - PYL.ABA bound to HAB13ujl

3kdh - apo-Pyl2


3kdh scenes
PYL2 is shown as a monomer. See also Scene 3 of the middle panel, which shows the receptor dimer.
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.

3kdi - ABA bound to PYL2


3kdi scenes
ABA (CPK spheres) binds to a water-filled (not shown) pocket of PYL2.
The gate folds over ABA and interacts with the latch.

3ujl - PYR2-HAB1


3ujl scenes
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.
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).

3k3k - PYR1 dimer


3k3k scene
in which one monomer is bound to ABA. The native form of the receptor is a dimer[26][27].

3klx scene

4dsc scene

Melcher K, Ng LM, Zhou XE, Soon FF, Xu Y, Suino-Powell KM,Park SY, Weiner JJ, Fujii H, Chinnusamy V, Kovach A, Li J, Wang Y, Peterson FC, Jensen DR, Yong EL, Volkman BF,Cutler SR, Zhu JK, Xu HE(2009) A gate-latch-lock mechanism for hormone signalling by abscisic acid receptors.Nature 462, 602–608.[28]
Melcher K, Zhou XE, Xu HE (2010) Thirsty plants and beyond: Structural mechanisms of abscisic acid perception and signaling. Curr. Opin. Struct. Biol. 20, 722–729.[29]
Miyazono K, Miyakawa T, Sawano Y, Kubota K, Kang HJ, Asano A, Miyauchi Y, Takahashi M, Zhi Y, Fujita Y, Yoshida T, Kodaira KS, Yamaguchi-Shinozaki K, Tanokura M (2009) Structural basis of abscisic acid signalling. Nature 462, 609–614[30]
Nishimura N, Hitomi K, Arvai AS, Rambo RP, Hitomi C, Cutler SR, Schroeder JI, Getzoff ED (2009) Structural mechanism of abscisic acid binding and signaling by dimeric PYR1. Science 326, 1373–1379[31]
Santiago J, Rodrigues A, Saez A, Rubio S, Antoni R, Dupeux F, Park SY, Marquez JA, Cutler SR, Rodriguez PL (2009b) Modulation of drought resistance by the abscisic acid receptor PYL5 through inhibition of clade A PP2Cs. Plant J. 60, 575–588[32]
YinP,FanH,HaoQ,YuanX,WuD,PangY,YanC,LiW,WangJ,Yan N (2009) Structural insights into the mechanism of abscisic acid signaling by PYL proteins. Nat. Struct. Mol. Biol. 16, 1230–1236[33]
Structural insights into PYR/PYL/RCAR ABA receptors and PP2Cs.
Santiago J, Dupeux F, Betz K, Antoni R, Gonzalez-Guzman M, Rodriguez L, Márquez JA, Rodriguez PL.Plant Sci. 2012 182:3-11[34]

PYR/PYL/RCAR structures

At is Arabidopsis thaliana

Apo structures

3k3k, AtPYR1 dimer, one monomer is bound to ABA and the other unliganded
3kay, apo AtPYL1
3kdh, 3kaz, 3kl1 apo AtPYL2
3klx, Apo AtPYL3
4jdl, Apo AtPYL5
3rt2, 3uqh apo AtPYL10


Structures with (+)-ABA
3k90, AtPYR1.ABA
3k3k, AtPyr1 dimer, one monomer is bound to ABA and the other unliganded
3jrs, AtPYL1.ABA
3kdi, 3kb0 AtPYL2.ABA
4dsb, 4dsc AtPYL3 with ABA
3oqu, AtPYL9.ABA
3r6p, AtPYL10.ABA


Structures with (-)-ABA
4jda, AtPYL3 with (-)-ABA


Structures with pyrabactin
3njo, AtPYR1.Pyrabactin
3nef, 3neg, 3nr4 AtPYL1.pyrabactin
3nj0, 3ns2 AtPYL2.Pyrabactin
3nj1, AtPYL2 V114I mutant.Pyrabactin
3nmh, AtPYL2 in complex with pyrabactin
3nmp, AtPYL2 mutant A93F in complex with pyrabactin
3oji, AtPYL3 with pyrabactin(?)


Structures with (+)-ABA or homolog and a PP2C
3qn1, AtPYR1.ABA - AtHAB1
3zvu, AtPYR1 H60P mutant .ABA - AtHAB1
3kb3, AtPYL1.ABA - HAB1
3jrq, 3kdj AtPYL1.ABA - ABI1
3ujl, AtPYL2.ABA - AtABI2
4lga, 4lgb AtPYL2.ABA mimic - AtHAB1
4ds8, AtPYL3.ABA complex with AtHAB1 3rt0, AtPYL10.ABA - AtHAB1


Structures with pyrabactin or homolog and a PP2C
4la7, 4lg5 AtPYL2.Quinabactin - AtHAB1
3nmn, AtPYL1.pyrabactin in complex with AtABI1
3nmt, AtPYL2 mutant A93F.pyrabactin in complex with type 2C protein phosphatase AtHAB1
3nmv, AtPYL2 mutant A93F.pyrabactin in complex with type 2C protein phosphatase AtABI1


References

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  5. Park SY, Fung P, Nishimura N, Jensen DR, Fujii H, Zhao Y, Lumba S, Santiago J, Rodrigues A, Chow TF, Alfred SE, Bonetta D, Finkelstein R, Provart NJ, Desveaux D, Rodriguez PL, McCourt P, Zhu JK, Schroeder JI, Volkman BF, Cutler SR. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins. Science. 2009 May 22;324(5930):1068-71. doi: 10.1126/science.1173041. Epub 2009, Apr 30. PMID:19407142 doi:10.1126/science.1173041
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  9. Park SY, Fung P, Nishimura N, Jensen DR, Fujii H, Zhao Y, Lumba S, Santiago J, Rodrigues A, Chow TF, Alfred SE, Bonetta D, Finkelstein R, Provart NJ, Desveaux D, Rodriguez PL, McCourt P, Zhu JK, Schroeder JI, Volkman BF, Cutler SR. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins. Science. 2009 May 22;324(5930):1068-71. doi: 10.1126/science.1173041. Epub 2009, Apr 30. PMID:19407142 doi:10.1126/science.1173041
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  13. 13.0 13.1 Nakashima K, Fujita Y, Kanamori N, Katagiri T, Umezawa T, Kidokoro S, Maruyama K, Yoshida T, Ishiyama K, Kobayashi M, Shinozaki K, Yamaguchi-Shinozaki K. Three Arabidopsis SnRK2 protein kinases, SRK2D/SnRK2.2, SRK2E/SnRK2.6/OST1 and SRK2I/SnRK2.3, involved in ABA signaling are essential for the control of seed development and dormancy. Plant Cell Physiol. 2009 Jul;50(7):1345-63. doi: 10.1093/pcp/pcp083. Epub 2009, Jun 18. PMID:19541597 doi:10.1093/pcp/pcp083
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  15. Melcher K, Ng LM, Zhou XE, Soon FF, Xu Y, Suino-Powell KM, Park SY, Weiner JJ, Fujii H, Chinnusamy V, Kovach A, Li J, Wang Y, Li J, Peterson FC, Jensen DR, Yong EL, Volkman BF, Cutler SR, Zhu JK, Xu HE. A gate-latch-lock mechanism for hormone signalling by abscisic acid receptors. Nature. 2009 Dec 3;462(7273):602-8. PMID:19898420 doi:10.1038/nature08613
  16. Nishimura N, Hitomi K, Arvai AS, Rambo RP, Hitomi C, Cutler SR, Schroeder JI, Getzoff ED. Structural mechanism of abscisic acid binding and signaling by dimeric PYR1. Science. 2009 Dec 4;326(5958):1373-9. Epub 2009 Oct 22. PMID:19933100
  17. Miyakawa T, Fujita Y, Yamaguchi-Shinozaki K, Tanokura M. Structure and function of abscisic acid receptors. Trends Plant Sci. 2013 May;18(5):259-66. doi: 10.1016/j.tplants.2012.11.002. Epub, 2012 Dec 22. PMID:23265948 doi:10.1016/j.tplants.2012.11.002
  18. "PMID:22579247
  19. Yin P, Fan H, Hao Q, Yuan X, Wu D, Pang Y, Yan C, Li W, Wang J, Yan N. Structural insights into the mechanism of abscisic acid signaling by PYL proteins. Nat Struct Mol Biol. 2009 Dec;16(12):1230-6. Epub 2009 Nov 5. PMID:19893533 doi:10.1038/nsmb.1730
  20. Ma Y, Szostkiewicz I, Korte A, Moes D, Yang Y, Christmann A, Grill E. Regulators of PP2C phosphatase activity function as abscisic acid sensors. Science. 2009 May 22;324(5930):1064-8. doi: 10.1126/science.1172408. Epub 2009, Apr 30. PMID:19407143 doi:10.1126/science.1172408
  21. Park SY, Fung P, Nishimura N, Jensen DR, Fujii H, Zhao Y, Lumba S, Santiago J, Rodrigues A, Chow TF, Alfred SE, Bonetta D, Finkelstein R, Provart NJ, Desveaux D, Rodriguez PL, McCourt P, Zhu JK, Schroeder JI, Volkman BF, Cutler SR. Abscisic acid inhibits type 2C protein phosphatases via the PYR/PYL family of START proteins. Science. 2009 May 22;324(5930):1068-71. doi: 10.1126/science.1173041. Epub 2009, Apr 30. PMID:19407142 doi:10.1126/science.1173041
  22. Santiago J, Rodrigues A, Saez A, Rubio S, Antoni R, Dupeux F, Park SY, Marquez JA, Cutler SR, Rodriguez PL. Modulation of drought resistance by the abscisic acid receptor PYL5 through inhibition of clade A PP2Cs. Plant J. 2009 Nov;60(4):575-88. doi: 10.1111/j.1365-313X.2009.03981.x. Epub 2009 , Jul 16. PMID:19624469 doi:10.1111/j.1365-313X.2009.03981.x
  23. Geiger D, Scherzer S, Mumm P, Stange A, Marten I, Bauer H, Ache P, Matschi S, Liese A, Al-Rasheid KA, Romeis T, Hedrich R. Activity of guard cell anion channel SLAC1 is controlled by drought-stress signaling kinase-phosphatase pair. Proc Natl Acad Sci U S A. 2009 Dec 15;106(50):21425-30. doi:, 10.1073/pnas.0912021106. Epub 2009 Dec 2. PMID:19955405 doi:10.1073/pnas.0912021106
  24. Sato A, Sato Y, Fukao Y, Fujiwara M, Umezawa T, Shinozaki K, Hibi T, Taniguchi M, Miyake H, Goto DB, Uozumi N. Threonine at position 306 of the KAT1 potassium channel is essential for channel activity and is a target site for ABA-activated SnRK2/OST1/SnRK2.6 protein kinase. Biochem J. 2009 Dec 10;424(3):439-48. doi: 10.1042/BJ20091221. PMID:19785574 doi:10.1042/BJ20091221
  25. Furihata T, Maruyama K, Fujita Y, Umezawa T, Yoshida R, Shinozaki K, Yamaguchi-Shinozaki K. Abscisic acid-dependent multisite phosphorylation regulates the activity of a transcription activator AREB1. Proc Natl Acad Sci U S A. 2006 Feb 7;103(6):1988-93. Epub 2006 Jan 30. PMID:16446457 doi:10.1073/pnas.0505667103
  26. Nishimura N, Hitomi K, Arvai AS, Rambo RP, Hitomi C, Cutler SR, Schroeder JI, Getzoff ED. Structural mechanism of abscisic acid binding and signaling by dimeric PYR1. Science. 2009 Dec 4;326(5958):1373-9. Epub 2009 Oct 22. PMID:19933100
  27. Miyakawa T, Fujita Y, Yamaguchi-Shinozaki K, Tanokura M. Structure and function of abscisic acid receptors. Trends Plant Sci. 2013 May;18(5):259-66. doi: 10.1016/j.tplants.2012.11.002. Epub, 2012 Dec 22. PMID:23265948 doi:10.1016/j.tplants.2012.11.002
  28. Melcher K, Ng LM, Zhou XE, Soon FF, Xu Y, Suino-Powell KM, Park SY, Weiner JJ, Fujii H, Chinnusamy V, Kovach A, Li J, Wang Y, Li J, Peterson FC, Jensen DR, Yong EL, Volkman BF, Cutler SR, Zhu JK, Xu HE. A gate-latch-lock mechanism for hormone signalling by abscisic acid receptors. Nature. 2009 Dec 3;462(7273):602-8. PMID:19898420 doi:10.1038/nature08613
  29. Melcher K, Zhou XE, Xu HE. Thirsty plants and beyond: structural mechanisms of abscisic acid perception and signaling. Curr Opin Struct Biol. 2010 Dec;20(6):722-9. doi: 10.1016/j.sbi.2010.09.007. Epub, 2010 Oct 14. PMID:20951573 doi:10.1016/j.sbi.2010.09.007
  30. Miyazono K, Miyakawa T, Sawano Y, Kubota K, Kang HJ, Asano A, Miyauchi Y, Takahashi M, Zhi Y, Fujita Y, Yoshida T, Kodaira KS, Yamaguchi-Shinozaki K, Tanokura M. Structural basis of abscisic acid signalling. Nature. 2009 Dec 3;462(7273):609-14. PMID:19855379 doi:10.1038/nature08583
  31. Nishimura N, Hitomi K, Arvai AS, Rambo RP, Hitomi C, Cutler SR, Schroeder JI, Getzoff ED. Structural mechanism of abscisic acid binding and signaling by dimeric PYR1. Science. 2009 Dec 4;326(5958):1373-9. Epub 2009 Oct 22. PMID:19933100
  32. Santiago J, Rodrigues A, Saez A, Rubio S, Antoni R, Dupeux F, Park SY, Marquez JA, Cutler SR, Rodriguez PL. Modulation of drought resistance by the abscisic acid receptor PYL5 through inhibition of clade A PP2Cs. Plant J. 2009 Nov;60(4):575-88. doi: 10.1111/j.1365-313X.2009.03981.x. Epub 2009 , Jul 16. PMID:19624469 doi:10.1111/j.1365-313X.2009.03981.x
  33. Yin P, Fan H, Hao Q, Yuan X, Wu D, Pang Y, Yan C, Li W, Wang J, Yan N. Structural insights into the mechanism of abscisic acid signaling by PYL proteins. Nat Struct Mol Biol. 2009 Dec;16(12):1230-6. Epub 2009 Nov 5. PMID:19893533 doi:10.1038/nsmb.1730
  34. Santiago J, Dupeux F, Betz K, Antoni R, Gonzalez-Guzman M, Rodriguez L, Marquez JA, Rodriguez PL. Structural insights into PYR/PYL/RCAR ABA receptors and PP2Cs. Plant Sci. 2012 Jan;182:3-11. doi: 10.1016/j.plantsci.2010.11.014. Epub 2010 Dec , 7. PMID:22118610 doi:10.1016/j.plantsci.2010.11.014

See Also

[1] Abscisic Acid in Wikipedia

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Alice Harmon

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