Gene Symbol: BIN2
Description: Protein kinase superfamily protein
Alias: ATSK21, BRASSINOSTEROID-INSENSITIVE 2, DWARF 12, DWF12, F28A21.120, F28A21_120, SHAGGY-LIKE KINASE 21, SK21, UCU1, ULTRACURVATA 1, Protein kinase superfamily protein
Species: thale cress

Top Publications

  1. Kim T, Michniewicz M, Bergmann D, Wang Z. Brassinosteroid regulates stomatal development by GSK3-mediated inhibition of a MAPK pathway. Nature. 2012;482:419-22 pubmed publisher
    ..brassinosteroid signalling inhibits stomatal development through the glycogen synthase kinase 3 (GSK3)-like kinase BIN2, and BIN2 acts upstream of YDA but downstream of the ERECTA family of receptor kinases...
  2. Yin Y, Wang Z, Mora Garcia S, Li J, Yoshida S, Asami T, et al. BES1 accumulates in the nucleus in response to brassinosteroids to regulate gene expression and promote stem elongation. Cell. 2002;109:181-91 pubmed
    ..BES1 is phosphorylated and appears to be destabilized by the glycogen synthase kinase-3 (GSK-3) BIN2, a negative regulator of the BR pathway...
  3. Tang W, Yuan M, Wang R, Yang Y, Wang C, Oses Prieto J, et al. PP2A activates brassinosteroid-responsive gene expression and plant growth by dephosphorylating BZR1. Nat Cell Biol. 2011;13:124-31 pubmed publisher
    When brassinosteroid levels are low, the GSK3-like kinase BIN2 phosphorylates and inactivates the BZR1 transcription factor to inhibit growth in plants...
  4. Peng P, Yan Z, Zhu Y, Li J. Regulation of the Arabidopsis GSK3-like kinase BRASSINOSTEROID-INSENSITIVE 2 through proteasome-mediated protein degradation. Mol Plant. 2008;1:338-46 pubmed publisher
    ..b>BIN2 is an Arabidopsis GSK3-like kinase that negatively regulates brassinosteroid (BR) signaling...
  5. Gampala S, Kim T, He J, Tang W, Deng Z, Bai M, et al. An essential role for 14-3-3 proteins in brassinosteroid signal transduction in Arabidopsis. Dev Cell. 2007;13:177-89 pubmed
    ..through a signal transduction pathway that involves cell-surface receptors (BRI1 and BAK1) and a GSK3 kinase (BIN2). How BR-regulated phosphorylation controls the activities of BZR1/BZR2 is not fully understood...
  6. Ryu H, Kim K, Cho H, Hwang I. Predominant actions of cytosolic BSU1 and nuclear BIN2 regulate subcellular localization of BES1 in brassinosteroid signaling. Mol Cells. 2010;29:291-6 pubmed publisher
    During brassinosteroid (BR) signaling in Arabidopsis, BSU1 (bri1 SUPPRESSOR1) phosphatase and BIN2 (BRASSINOSTEROID INSENSITIVE2) kinase regulate the signal intensity by determining the phosphorylation status of the transcription factors ..
  7. Li J, Nam K. Regulation of brassinosteroid signaling by a GSK3/SHAGGY-like kinase. Science. 2002;295:1299-301 pubmed
    ..Here we show that the Arabidopsis BRASSINOSTEROID-INSENSITIVE 2 (BIN2) gene encodes a GSK3/SHAGGY-like kinase...
  8. He J, Gendron J, Yang Y, Li J, Wang Z. The GSK3-like kinase BIN2 phosphorylates and destabilizes BZR1, a positive regulator of the brassinosteroid signaling pathway in Arabidopsis. Proc Natl Acad Sci U S A. 2002;99:10185-90 pubmed
    ..BR signaling involves the cell-surface receptor BRI1, the glycogen synthase kinase-3-like kinase BIN2 as a negative regulator, and nuclear proteins BZR1 and BZR2/BES1 as positive regulators...
  9. Ryu H, Kim K, Cho H, Park J, Choe S, Hwang I. Nucleocytoplasmic shuttling of BZR1 mediated by phosphorylation is essential in Arabidopsis brassinosteroid signaling. Plant Cell. 2007;19:2749-62 pubmed

More Information


  1. Yan Z, Zhao J, Peng P, Chihara R, Li J. BIN2 functions redundantly with other Arabidopsis GSK3-like kinases to regulate brassinosteroid signaling. Plant Physiol. 2009;150:710-21 pubmed publisher
    ..have so far uncovered eight mutants, all of which carry gain-of-function mutations in BRASSINOSTEROID-INSENSITIVE2 (BIN2), one of the three members in group II...
  2. Perez Perez J, Ponce M, Micol J. The UCU1 Arabidopsis gene encodes a SHAGGY/GSK3-like kinase required for cell expansion along the proximodistal axis. Dev Biol. 2002;242:161-73 pubmed
    ..Here, we describe the characterization of mutant alleles of the Arabidopsis ULTRACURVATA1 (UCU1) gene, the two strongest of which dramatically reduce cell expansion along the proximodistal axis, dwarfing the ..
  3. Tsugama D, Liu S, Takano T. Arabidopsis heterotrimeric G protein ? subunit, AGB1, regulates brassinosteroid signalling independently of BZR1. J Exp Bot. 2013;64:3213-23 pubmed publisher
    ..GSKs), which are known components of BR signalling, the interaction between AGB1 and one of the Arabidopsis GSKs, BIN2, was examined...
  4. Li J, Nam K, Vafeados D, Chory J. BIN2, a new brassinosteroid-insensitive locus in Arabidopsis. Plant Physiol. 2001;127:14-22 pubmed
    ..Genetic analysis of the bin2 (BR-INSENSITIVE 2) mutant indicated that the BR-insensitive dwarf phenotype was due to a semidominant mutation in ..
  5. Zhao J, Peng P, Schmitz R, Decker A, Tax F, Li J. Two putative BIN2 substrates are nuclear components of brassinosteroid signaling. Plant Physiol. 2002;130:1221-9 pubmed
    ..b>BIN2 is a GSK3-like kinase in Arabidopsis that functions as a negative regulator of brassinosteroid (BR) signaling...
  6. Hu Y, Yu D. BRASSINOSTEROID INSENSITIVE2 interacts with ABSCISIC ACID INSENSITIVE5 to mediate the antagonism of brassinosteroids to abscisic acid during seed germination in Arabidopsis. Plant Cell. 2014;26:4394-408 pubmed publisher
    ..Here, we show that the Glycogen Synthase Kinase 3-like kinase BRASSINOSTEROID INSENSITIVE2 (BIN2), a critical repressor of BR signaling, positively regulates ABA responses during seed germination and ..
  7. Kondo Y, Ito T, Nakagami H, Hirakawa Y, Saito M, Tamaki T, et al. Plant GSK3 proteins regulate xylem cell differentiation downstream of TDIF-TDR signalling. Nat Commun. 2014;5:3504 pubmed publisher
    ..Our findings provide insight into the regulation of cell fate determination in meristem maintenance. ..
  8. Gudesblat G, Schneider Pizoń J, Betti C, Mayerhofer J, Vanhoutte I, Van Dongen W, et al. SPEECHLESS integrates brassinosteroid and stomata signalling pathways. Nat Cell Biol. 2012;14:548-54 pubmed publisher
    ..The GSK3/SHAGGY-like kinase BIN2 (BR INSENSITIVE2) phosphorylates residues overlapping those targeted by the MAPKs, as well as four residues in the ..
  9. Sreeramulu S, Mostizky Y, Sunitha S, Shani E, Nahum H, Salomon D, et al. BSKs are partially redundant positive regulators of brassinosteroid signaling in Arabidopsis. Plant J. 2013;74:905-19 pubmed publisher
    ..The BIN2 and BIL2 GSK3-like kinases, which are negative regulators of BR signaling, interacted in vivo with BSKs and ..
  10. Peng P, Zhao J, Zhu Y, Asami T, Li J. A direct docking mechanism for a plant GSK3-like kinase to phosphorylate its substrates. J Biol Chem. 2010;285:24646-53 pubmed publisher
    ..Brassinosteroid-INsensitive 2 (BIN2) is an Arabidopsis GSK3-like kinase that negatively regulates brassinosteroid (BR) signaling by phosphorylating ..
  11. Xiong F, Zhang R, Meng Z, Deng K, Que Y, Zhuo F, et al. Brassinosteriod Insensitive 2 (BIN2) acts as a downstream effector of the Target of Rapamycin (TOR) signaling pathway to regulate photoautotrophic growth in Arabidopsis. New Phytol. 2017;213:233-249 pubmed publisher
    ..Systems, genetic, and biochemical analyses revealed that Brassinosteriod Insensitive 2 (BIN2) acts as a novel downstream effector of S6K2, and the phosphorylation of BIN2 depends on TOR-S6K2 signaling in ..
  12. Youn J, Kim T, Kim E, Bu S, Kim S, Wang Z, et al. Structural and functional characterization of Arabidopsis GSK3-like kinase AtSK12. Mol Cells. 2013;36:564-70 pubmed publisher
    ..Only one of these Arabidopsis GSK3s, BIN2 (also named AtSK21), has been characterized by biochemical and genetic studies...
  13. Zhu J, Li Y, Cao D, Yang H, Oh E, Bi Y, et al. The F-box Protein KIB1 Mediates Brassinosteroid-Induced Inactivation and Degradation of GSK3-like Kinases in Arabidopsis. Mol Cell. 2017;66:648-657.e4 pubmed publisher
    ..In Arabidopsis, the GSK3-like kinase BIN2 phosphorylates a range of proteins to control broad developmental processes, and BIN2 is degraded through unknown ..
  14. Cai Z, Liu J, Wang H, Yang C, Chen Y, Li Y, et al. GSK3-like kinases positively modulate abscisic acid signaling through phosphorylating subgroup III SnRK2s in Arabidopsis. Proc Natl Acad Sci U S A. 2014;111:9651-6 pubmed publisher
    ..2 and SnRK2.3, that can interact with and be phosphorylated by a GSK3-like kinase, brassinosteroid insensitive 2 (BIN2)...
  15. Cheng Y, Zhu W, Chen Y, Ito S, Asami T, Wang X. Brassinosteroids control root epidermal cell fate via direct regulation of a MYB-bHLH-WD40 complex by GSK3-like kinases. elife. 2014;: pubmed publisher
    ..One of the GSK3-like kinases, BIN2, interacted with and phosphorylated EGL3, and EGL3s mutated at phosphorylation siteswere retained in hair cell ..
  16. Samakovli D, Margaritopoulou T, Prassinos C, Milioni D, Hatzopoulos P. Brassinosteroid nuclear signaling recruits HSP90 activity. New Phytol. 2014;203:743-57 pubmed publisher
    ..GDA treatment promoted the export of BIN2, a regulator of BR signaling, from the nucleus into the cytoplasm, indicating that active HSP90 is required to ..
  17. Anne P, Azzopardi M, Gissot L, Beaubiat S, Hématy K, Palauqui J. OCTOPUS Negatively Regulates BIN2 to Control Phloem Differentiation in Arabidopsis thaliana. Curr Biol. 2015;25:2584-90 pubmed publisher
    ..Directed protein interactions with known BR signaling proteins identified BIN2, a GSK3 protein involved in multiple signaling pathways, as a partner of OPS...
  18. Zhang D, Ye H, Guo H, Johnson A, Lin H, Yin Y. Transcription factors involved in brassinosteroid repressed gene expression and their regulation by BIN2 kinase. Plant Signal Behav. 2014;9:e27849 pubmed
    ..Moreover, BIN2 kinase, a well-established negative regulator in the BR signaling pathway, phosphorylates MYBL2 and HAT1...
  19. Zhang D, Ye H, Guo H, Johnson A, Zhang M, Lin H, et al. Transcription factor HAT1 is phosphorylated by BIN2 kinase and mediates brassinosteroid repressed gene expression in Arabidopsis. Plant J. 2014;77:59-70 pubmed publisher
    ..Furthermore, HAT1 can be phosphorylated and stabilized by GSK3 (GLYCOGEN SYNTHASE KINASE 3)-like kinase BIN2 (BRASSINOSTEROID-INSENSITIVE 2), a well established negative regulator of the BR pathway...
  20. Hao Y, Wang H, Qiao S, Leng L, Wang X. Histone deacetylase HDA6 enhances brassinosteroid signaling by inhibiting the BIN2 kinase. Proc Natl Acad Sci U S A. 2016;113:10418-23 pubmed publisher
    ..The Arabidopsis thaliana GSK3-like kinase BR-INSENSITIVE 2 (BIN2) acts as a key negative regulator in the BR signaling pathway, but the mechanisms regulating BIN2 function remain ..
  21. Khan M, Rozhon W, Bigeard J, Pflieger D, Husar S, Pitzschke A, et al. Brassinosteroid-regulated GSK3/Shaggy-like kinases phosphorylate mitogen-activated protein (MAP) kinase kinases, which control stomata development in Arabidopsis thaliana. J Biol Chem. 2013;288:7519-27 pubmed publisher
    ..thaliana mutants, resulting in clustered stomata. We demonstrate that BIN2 and its homologues, GSK3/Shaggy-like kinases involved in BR signaling, can phosphorylate the MAPK kinases MKK4 and ..
  22. Cho H, Ryu H, Rho S, Hill K, Smith S, Audenaert D, et al. A secreted peptide acts on BIN2-mediated phosphorylation of ARFs to potentiate auxin response during lateral root development. Nat Cell Biol. 2014;16:66-76 pubmed publisher
    ..Here, we show that phosphorylation of ARF7 and ARF19 by BRASSINOSTEROID-insensitive2 (BIN2) can also potentiate auxin signalling output during lateral root organogenesis...
  23. Ye H, Li L, Guo H, Yin Y. MYBL2 is a substrate of GSK3-like kinase BIN2 and acts as a corepressor of BES1 in brassinosteroid signaling pathway in Arabidopsis. Proc Natl Acad Sci U S A. 2012;109:20142-7 pubmed publisher
    ..Moreover, MYBL2 is a substrate of glycogen synthase kinase 3 (GSK3)-like kinase brassinosteroid-insensitive 2 (BIN2), which has been well established as a negative regulator in the BR pathway by phosphorylating and inhibiting the ..
  24. Vert G, Walcher C, Chory J, Nemhauser J. Integration of auxin and brassinosteroid pathways by Auxin Response Factor 2. Proc Natl Acad Sci U S A. 2008;105:9829-34 pubmed publisher
    ..Moreover, we describe a direct connection between the BR-regulated BIN2 kinase and ARF2, a member of the Auxin Response Factor family of transcriptional regulators...