ABA Signaling Pathway

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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][8], protein phosphatase 2Cs (ABI1, ABI2, HAB1, HAB2, PP2CA/AHG3)[4][6][9][10], and 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[5][15][16][7] (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[4][6][17], and this creates a binding surface for PP2C. The activated receptor (R.ABA) dissociates to monomers which bind to protein phosphatase 2C[9] (R.ABA-P). The sequestration of the protein 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[18] and KAT1[19] 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 [20][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.

Structures of Proteins

For details about the structures of proteins involved in this pathway see PYR/PYL/RCAR family of ABA receptors, ABA-regulated Protein Phosphatase 2C, ABA-regulated SNRK2 Protein Kinase.

References

  1. ↑ Leung J, Giraudat J. ABSCISIC ACID SIGNAL TRANSDUCTION. Annu Rev Plant Physiol Plant Mol Biol. 1998 Jun;49:199-222. PMID:15012233 doi:10.1146/annurev.arplant.49.1.199
  2. ↑ Finkelstein RR, Gampala SS, Rock CD. Abscisic acid signaling in seeds and seedlings. Plant Cell. 2002;14 Suppl:S15-45. PMID:12045268
  3. ↑ Cutler SR, Rodriguez PL, Finkelstein RR, Abrams SR. Abscisic acid: emergence of a core signaling network. Annu Rev Plant Biol. 2010;61:651-79. doi: 10.1146/annurev-arplant-042809-112122. PMID:20192755 doi:10.1146/annurev-arplant-042809-112122
  4. ↑ 4.0 4.1 4.2 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
  5. ↑ 5.0 5.1 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
  6. ↑ 6.0 6.1 6.2 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
  7. ↑ 7.0 7.1 Zhang X, Zhang Q, Xin Q, Yu L, Wang Z, Wu W, Jiang L, Wang G, Tian W, Deng Z, Wang Y, Liu Z, Long J, Gong Z, Chen Z. Complex Structures of the Abscisic Acid Receptor PYL3/RCAR13 Reveal a Unique Regulatory Mechanism. Structure. 2012 May 9;20(5):780-90. PMID:22579247 doi:10.1016/j.str.2012.02.019
  8. ↑ Antoni R, Gonzalez-Guzman M, Rodriguez L, Peirats-Llobet M, Pizzio GA, Fernandez MA, De Winne N, De Jaeger G, Dietrich D, Bennett MJ, Rodriguez PL. PYRABACTIN RESISTANCE1-LIKE8 plays an important role for the regulation of abscisic acid signaling in root. Plant Physiol. 2013 Feb;161(2):931-41. doi: 10.1104/pp.112.208678. Epub 2012 Dec , 14. PMID:23370718 doi:10.1104/pp.112.208678
  9. ↑ 9.0 9.1 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
  10. ↑ Yoshida T, Nishimura N, Kitahata N, Kuromori T, Ito T, Asami T, Shinozaki K, Hirayama T. ABA-hypersensitive germination3 encodes a protein phosphatase 2C (AtPP2CA) that strongly regulates abscisic acid signaling during germination among Arabidopsis protein phosphatase 2Cs. Plant Physiol. 2006 Jan;140(1):115-26. Epub 2005 Dec 9. PMID:16339800 doi:10.1104/pp.105.070128
  11. ↑ Mustilli AC, Merlot S, Vavasseur A, Fenzi F, Giraudat J. Arabidopsis OST1 protein kinase mediates the regulation of stomatal aperture by abscisic acid and acts upstream of reactive oxygen species production. Plant Cell. 2002 Dec;14(12):3089-99. PMID:12468729
  12. ↑ Yoshida R, Hobo T, Ichimura K, Mizoguchi T, Takahashi F, Aronso J, Ecker JR, Shinozaki K. ABA-activated SnRK2 protein kinase is required for dehydration stress signaling in Arabidopsis. Plant Cell Physiol. 2002 Dec;43(12):1473-83. PMID:12514244
  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
  14. ↑ 14.0 14.1 Fujii H, Verslues PE, Zhu JK. Identification of two protein kinases required for abscisic acid regulation of seed germination, root growth, and gene expression in Arabidopsis. Plant Cell. 2007 Feb;19(2):485-94. Epub 2007 Feb 16. PMID:17307925 doi:tpc.106.048538
  15. ↑ 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
  16. ↑ 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
  17. ↑ 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
  18. ↑ 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
  19. ↑ 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
  20. ↑ 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

See Also

[1] Abscisic Acid in Wikipedia

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