BIN2 (BRASSINOSTEROID-INSENSITIVE 2; also ASK7/ASK-eta, AtSK21, DWF12, UCU1; At4g18710) is a group-II GSK3/SHAGGY-like serine/threonine kinase and the central negative regulator of brassinosteroid (BR) signalling. In the absence of BR, active BIN2 (requiring autophosphorylation of Tyr200 in its activation loop) phosphorylates the BZR1 and BES1/BZR2 transcription factors, which inhibits their DNA binding, promotes 14-3-3 binding and cytoplasmic retention/nuclear export, and targets them for degradation. BR perception by BRI1-BAK1 leads, via BSK1 and CDG1, to activation of the BSU1 phosphatase, which dephosphorylates BIN2 pTyr200 and inactivates it; BR also promotes KIB1-mediated ubiquitination and proteasomal degradation of BIN2. Gain-of-function bin2 alleles (e.g. bin2-1, in the conserved TREE motif) are semidominant BR-insensitive dwarfs. BIN2 is found in the cytoplasm and nucleus and can be recruited to the plasma membrane by scaffolds and receptors (OPS, POLAR/BASL, TDR). It also phosphorylates many other substrates, including YDA and SPCH (stomatal development), ARF2/ARF7 (auxin signalling), EGL3/TTG1 (root epidermal cell fate) and RAPTOR1B (TOR/autophagy), making it a hub that integrates BR status with other developmental and stress pathways. BIN2 acts redundantly with its paralogs BIL1/ASK-iota and BIL2/ASK-theta.
| GO Term | Evidence | Action | Reason |
|---|---|---|---|
| GO:0004672 protein kinase activity | IEA GO_REF:0000002 | ACCEPT | Summary: InterPro protein kinase activity. Reason: Correct (parent of Ser/Thr kinase activity). Supporting Evidence: PMID:11847343 Here we show that the Arabidopsis BRASSINOSTEROID-INSENSITIVE 2 (BIN2) gene encodes a GSK3/SHAGGY-like kinase. |
| GO:0004672 protein kinase activity | TAS PMID:11820813 The UCU1 Arabidopsis gene encodes a SHAGGY/GSK3-like kinase ... | ACCEPT | Summary: Traceable author statement that UCU1/BIN2 is a kinase. Reason: Correct. Supporting Evidence: PMID:11820813 which encodes an AtSK protein involved in the cross-talk between auxin and brassinosteroid signaling pathways |
| GO:0004674 protein serine/threonine kinase activity | HDA PMID:21477822 Autophosphorylation profiling of Arabidopsis protein kinases... | ACCEPT | Summary: Ser/Thr autophosphorylation activity in a high-throughput screen. Reason: Consistent with focused biochemistry. Supporting Evidence: PMID:21477822 Luminescent analysis revealed that 179 of the 759 PKs had autophosphorylation activity. |
| GO:0004674 protein serine/threonine kinase activity | IBA GO_REF:0000033 | ACCEPT | Summary: Phylogenetic inference of Ser/Thr kinase activity for GSK3 family. Reason: Ancestral GSK3 activity retained in BIN2. Supporting Evidence: PMID:11847343 Here we show that the Arabidopsis BRASSINOSTEROID-INSENSITIVE 2 (BIN2) gene encodes a GSK3/SHAGGY-like kinase. |
| GO:0004674 protein serine/threonine kinase activity | IDA PMID:24771765 Brassinosteroids control root epidermal cell fate via direct... | ACCEPT | Summary: BIN2 phosphorylates EGL3/TTG1 in vitro. Reason: Direct evidence. Supporting Evidence: PMID:24771765 One of the GSK3-like kinases, BIN2, interacted with and phosphorylated EGL3 |
| GO:0004674 protein serine/threonine kinase activity | IEA GO_REF:0000120 | ACCEPT | Summary: Automated Ser/Thr kinase activity. Reason: Consistent with direct evidence. Supporting Evidence: PMID:11847343 Here we show that the Arabidopsis BRASSINOSTEROID-INSENSITIVE 2 (BIN2) gene encodes a GSK3/SHAGGY-like kinase. |
| GO:0005515 protein binding | IPI PMID:12007405 BES1 accumulates in the nucleus in response to brassinostero... | REMOVE | Summary: IPI with BES1, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:12007405 BES1 is phosphorylated and appears to be destabilized by the glycogen synthase kinase-3 (GSK-3) BIN2, a negative regulator of the BR pathway. |
| GO:0005515 protein binding | IPI PMID:17681130 An essential role for 14-3-3 proteins in brassinosteroid sig... | REMOVE | Summary: IPI with BZR1, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:12114546 BIN2 directly interacts with BZR1 in yeast two-hybrid assays, phosphorylates BZR1 in vitro, and negatively regulates BZR1 protein accumulation in vivo. |
| GO:0005515 protein binding | IPI PMID:18599455 Integration of auxin and brassinosteroid pathways by Auxin R... | REMOVE | Summary: IPI with ARF2, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:18599455 Phosphorylation by BIN2 results in loss of ARF2 DNA binding and repression activities. |
| GO:0005515 protein binding | IPI PMID:22307275 Brassinosteroid regulates stomatal development by GSK3-media... | REMOVE | Summary: IPI with BZR1, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:12114546 BIN2 directly interacts with BZR1 in yeast two-hybrid assays, phosphorylates BZR1 in vitro, and negatively regulates BZR1 protein accumulation in vivo. |
| GO:0005515 protein binding | IPI PMID:22307275 Brassinosteroid regulates stomatal development by GSK3-media... | MODIFY | Summary: IPI with the protein kinase YDA. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:22307275 BIN2 phosphorylates YDA to inhibit YDA phosphorylation of its substrate MKK4 |
| GO:0005515 protein binding | IPI PMID:23169658 MYBL2 is a substrate of GSK3-like kinase BIN2 and acts as a ... | REMOVE | Summary: IPI with MYBL2, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:23169658 MYBL2 is a substrate of GSK3-like kinase BIN2 |
| GO:0005515 protein binding | IPI PMID:23496207 BSKs are partially redundant positive regulators of brassino... | MODIFY | Summary: IPI with the protein kinase BSK11. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:23496207 The BIN2 and BIL2 GSK3-like kinases, which are negative regulators of BR signaling, interacted in vivo with BSKs and phosphorylated them in vitro, probably at different sites to BRI1. |
| GO:0005515 protein binding | IPI PMID:23496207 BSKs are partially redundant positive regulators of brassino... | MODIFY | Summary: IPI with the protein kinase BSK1. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:23496207 The BIN2 and BIL2 GSK3-like kinases, which are negative regulators of BR signaling, interacted in vivo with BSKs and phosphorylated them in vitro, probably at different sites to BRI1. |
| GO:0005515 protein binding | IPI PMID:23496207 BSKs are partially redundant positive regulators of brassino... | MODIFY | Summary: IPI with the protein kinase BSK8. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:23496207 The BIN2 and BIL2 GSK3-like kinases, which are negative regulators of BR signaling, interacted in vivo with BSKs and phosphorylated them in vitro, probably at different sites to BRI1. |
| GO:0005515 protein binding | IPI PMID:23496207 BSKs are partially redundant positive regulators of brassino... | MODIFY | Summary: IPI with the protein kinase BSK6. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:23496207 The BIN2 and BIL2 GSK3-like kinases, which are negative regulators of BR signaling, interacted in vivo with BSKs and phosphorylated them in vitro, probably at different sites to BRI1. |
| GO:0005515 protein binding | IPI PMID:24362628 A secreted peptide acts on BIN2-mediated phosphorylation of ... | REMOVE | Summary: IPI with ARF7, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:24362628 Here, we show that phosphorylation of ARF7 and ARF19 by BRASSINOSTEROID-insensitive2 (BIN2) can also potentiate auxin signalling output during lateral root organogenesis. |
| GO:0005515 protein binding | IPI PMID:24362628 A secreted peptide acts on BIN2-mediated phosphorylation of ... | MODIFY | Summary: IPI with the protein kinase TDR. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:24362628 TDIF-initiated TDR signalling directly acts on BIN2-mediated ARF phosphorylation, leading to the regulation of auxin signalling during lateral root development. |
| GO:0005515 protein binding | IPI PMID:24662460 Plant GSK3 proteins regulate xylem cell differentiation down... | MODIFY | Summary: IPI with the protein kinase TDR. Reason: Generic protein binding is uninformative; the partner is a protein kinase (substrate, upstream activator or downstream target of BIN2), so protein kinase binding is more informative. Proposed replacements: protein kinase binding Supporting Evidence: PMID:24662460 TDR interacts with GSK3s at the plasma membrane and activates GSK3s in a TDIF-dependent fashion. |
| GO:0005515 protein binding | IPI PMID:24771765 Brassinosteroids control root epidermal cell fate via direct... | REMOVE | Summary: IPI with EGL3, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:24771765 One of the GSK3-like kinases, BIN2, interacted with and phosphorylated EGL3 |
| GO:0005515 protein binding | IPI PMID:24771765 Brassinosteroids control root epidermal cell fate via direct... | REMOVE | Summary: IPI with WER, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:24771765 we conducted GST pull-down and BiFC (biomolecular fluorescence complementation) assays to test their interactions, and found that BIN2 can interact with WER, EGL3, and TTG1 |
| GO:0005515 protein binding | IPI PMID:24771765 Brassinosteroids control root epidermal cell fate via direct... | REMOVE | Summary: IPI with TTG1, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:24771765 BIN2 phosphorylated TTG1 to inhibit the activity of the WER-GL3/EGL3-TTG1 complex. |
| GO:0005515 protein binding | IPI PMID:28575660 The F-box Protein KIB1 Mediates Brassinosteroid-Induced Inac... | REMOVE | Summary: IPI with KIB2 (AGI_LocusCode:AT4G12820). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:28575660 Here we identify KIB1 as an F-box E3 ubiquitin ligase that promotes the degradation of BIN2 while blocking its substrate access. |
| GO:0005515 protein binding | IPI PMID:28575660 The F-box Protein KIB1 Mediates Brassinosteroid-Induced Inac... | REMOVE | Summary: IPI with KIB2 (UniProtKB:Q9SU04). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:28575660 Here we identify KIB1 as an F-box E3 ubiquitin ligase that promotes the degradation of BIN2 while blocking its substrate access. |
| GO:0005515 protein binding | IPI PMID:28575660 The F-box Protein KIB1 Mediates Brassinosteroid-Induced Inac... | REMOVE | Summary: IPI with KIB1 (UniProtKB:Q9SU05). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:28575660 Here we identify KIB1 as an F-box E3 ubiquitin ligase that promotes the degradation of BIN2 while blocking its substrate access. |
| GO:0005515 protein binding | IPI PMID:28576847 Arabidopsis WRKY46, WRKY54, and WRKY70 Transcription Factors... | REMOVE | Summary: IPI with WRKY54 (UniProtKB:Q93WU8). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:28576847 In addition, WRKY54 is phosphorylated and destabilized by GSK3-like kinase BR-INSENSITIVE2, a negative regulator in the BR pathway. |
| GO:0005515 protein binding | IPI PMID:28576847 Arabidopsis WRKY46, WRKY54, and WRKY70 Transcription Factors... | REMOVE | Summary: IPI with WRKY70 (UniProtKB:Q9LY00). Reason: Generic protein binding is uninformative; WRKY70 is a group III WRKY transcription factor and a candidate BIN2 substrate. The cached abstract reports BIN2 phosphorylation of WRKY54 only; the WRKY70 interaction data are in the full text (not cached), and the row is removed on term-informativeness grounds, not because the interaction is doubted. Supporting Evidence: PMID:28576847 In addition, WRKY54 is phosphorylated and destabilized by GSK3-like kinase BR-INSENSITIVE2, a negative regulator in the BR pathway. |
| GO:0005515 protein binding | IPI PMID:28576847 Arabidopsis WRKY46, WRKY54, and WRKY70 Transcription Factors... | REMOVE | Summary: IPI with WRKY46 (UniProtKB:Q9SKD9). Reason: Generic protein binding is uninformative; WRKY46 is a group III WRKY transcription factor and a candidate BIN2 substrate. The cached abstract reports BIN2 phosphorylation of WRKY54 only; the WRKY46 interaction data are in the full text (not cached), and the row is removed on term-informativeness grounds, not because the interaction is doubted. Supporting Evidence: PMID:28576847 In addition, WRKY54 is phosphorylated and destabilized by GSK3-like kinase BR-INSENSITIVE2, a negative regulator in the BR pathway. |
| GO:0005515 protein binding | IPI PMID:30429609 POLAR-guided signalling complex assembly and localization dr... | REMOVE | Summary: IPI with BASL (UniProtKB:Q5BPF3). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:30429609 Here we identify the plant-specific protein POLAR as a stomatal lineage scaffold for a subset of GSK3-like kinases that confines them to the cytosol and subsequently transiently polarizes them within the cell |
| GO:0005515 protein binding | IPI PMID:30429609 POLAR-guided signalling complex assembly and localization dr... | REMOVE | Summary: IPI with POLAR (UniProtKB:Q6NQ99). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:30429609 Here we identify the plant-specific protein POLAR as a stomatal lineage scaffold for a subset of GSK3-like kinases that confines them to the cytosol and subsequently transiently polarizes them within the cell |
| GO:0005515 protein binding | IPI PMID:32612234 Extensive signal integration by the phytohormone protein net... | REMOVE | Summary: IPI with ZHD3 (UniProtKB:O64722). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:32612234 we experimentally generated a systems-level map of the Arabidopsis phytohormone signalling network, consisting of more than 2,000 binary protein-protein interactions |
| GO:0005515 protein binding | IPI PMID:32612234 Extensive signal integration by the phytohormone protein net... | REMOVE | Summary: IPI with GRF5 (UniProtKB:Q8L8A6). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:32612234 we experimentally generated a systems-level map of the Arabidopsis phytohormone signalling network, consisting of more than 2,000 binary protein-protein interactions |
| GO:0005515 protein binding | IPI PMID:32612234 Extensive signal integration by the phytohormone protein net... | REMOVE | Summary: IPI with ARF2, a BIN2 phosphorylation substrate. Reason: Generic protein binding is uninformative; the interaction is a kinase-substrate interaction already captured by BIN2's kinase activity annotations. Removal does not imply the interaction is false. Supporting Evidence: PMID:18599455 Phosphorylation by BIN2 results in loss of ARF2 DNA binding and repression activities. |
| GO:0005515 protein binding | IPI PMID:32612234 Extensive signal integration by the phytohormone protein net... | REMOVE | Summary: IPI with LBD13 (UniProtKB:Q9AT61). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:32612234 we experimentally generated a systems-level map of the Arabidopsis phytohormone signalling network, consisting of more than 2,000 binary protein-protein interactions |
| GO:0005515 protein binding | IPI PMID:32612234 Extensive signal integration by the phytohormone protein net... | REMOVE | Summary: IPI with TCP15 (UniProtKB:Q9C9L2). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:32612234 we experimentally generated a systems-level map of the Arabidopsis phytohormone signalling network, consisting of more than 2,000 binary protein-protein interactions |
| GO:0005515 protein binding | IPI PMID:32612234 Extensive signal integration by the phytohormone protein net... | REMOVE | Summary: IPI with TCP17 (UniProtKB:Q9LEZ9). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:32612234 we experimentally generated a systems-level map of the Arabidopsis phytohormone signalling network, consisting of more than 2,000 binary protein-protein interactions |
| GO:0005515 protein binding | IPI PMID:36648110 Spatiotemporal formation of the large vacuole regulated by t... | REMOVE | Summary: IPI with VLG (AGI_LocusCode:AT2G17740). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:36648110 Further investigation indicated that BIN2 interacted with and phosphorylated VLG, which stabilized VLG, thereby facilitating the large vacuole formation in the FG. |
| GO:0005515 protein binding | IPI PMID:37335918 Plant-specific BLISTER interacts with kinase BIN2 and BRASSI... | REMOVE | Summary: IPI with BLISTER (AGI_LocusCode:AT3G23980). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:37335918 BRASSINOSTEROID INSENSITIVE2 interacts with and phosphorylates BLI at 4 phosphorylation sites (Ser70, Ser146, Thr256, and Ser267) for degradation |
| GO:0005515 protein binding | IPI PMID:37926922 Comparative analysis of BZR1/BES1 family transcription facto... | REMOVE | Summary: IPI with BEH1/BEH2 (AGI_LocusCode:AT3G50750). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:37926922 We also compared their subcellular localizations regulated by the phosphorylation status, interaction with GSK3-like kinases, and heterodimeric combination. |
| GO:0005515 protein binding | IPI PMID:37986925 Regulation of adaptive growth decisions via phosphorylation ... | REMOVE | Summary: IPI with TRS120 (AGI_LocusCode:AT5G11040). Reason: Generic protein binding is uninformative; BIN2 binds many regulators, scaffolds (e.g. POLAR, BASL, KIB1, OPS-like proteins) and substrates. Removal does not imply the interaction is false. Supporting Evidence: PMID:37986925 Kinase assays and pharmacological inhibition provided in vitro and in vivo evidence that AtSKs target the TRAPPII-specific subunit AtTRS120/TRAPPC9. |
| GO:0005524 ATP binding | IEA GO_REF:0000002 | KEEP AS NON CORE | Summary: ATP binding from the kinase domain. Reason: True but uninformative. Supporting Evidence: PMID:11847343 Here we show that the Arabidopsis BRASSINOSTEROID-INSENSITIVE 2 (BIN2) gene encodes a GSK3/SHAGGY-like kinase. |
| GO:0005634 nucleus | IBA GO_REF:0000033 | ACCEPT | Summary: BIN2 is present in the nucleus, where it phosphorylates BZR1/BES1 and other transcription factors. Reason: Consistent with direct localization data. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005634 nucleus | IDA PMID:28652362 Phosphosite charge rather than shootward localization determ... | ACCEPT | Summary: BIN2 is present in the nucleus, where it phosphorylates BZR1/BES1 and other transcription factors. Reason: Consistent with direct localization data. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005634 nucleus | IDA PMID:30429609 POLAR-guided signalling complex assembly and localization dr... | ACCEPT | Summary: BIN2 is present in the nucleus, where it phosphorylates BZR1/BES1 and other transcription factors. Reason: BIN2 nuclear localization is shown directly elsewhere (mRFP-BIN2, PMID:28652362). The localization data from this paper (in which POLAR sequesters a subset of GSK3-like kinases in the cytosol of stomatal lineage cells) are in the full text, which is not cached; deferred to the curator. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005634 nucleus | IEA GO_REF:0000044 | ACCEPT | Summary: BIN2 is present in the nucleus, where it phosphorylates BZR1/BES1 and other transcription factors. Reason: Consistent with direct localization data. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005634 nucleus | ISM GO_REF:0000122 | ACCEPT | Summary: BIN2 is present in the nucleus, where it phosphorylates BZR1/BES1 and other transcription factors. Reason: Consistent with direct localization data. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005737 cytoplasm | EXP PMID:30429609 POLAR-guided signalling complex assembly and localization dr... | ACCEPT | Summary: BIN2 is present in the cytoplasm. Reason: Consistent with direct localization data; POLAR confines BIN2 to the cytosol in stomatal lineage cells. Supporting Evidence: PMID:30429609 Here we identify the plant-specific protein POLAR as a stomatal lineage scaffold for a subset of GSK3-like kinases that confines them to the cytosol and subsequently transiently polarizes them within the cell |
| GO:0005737 cytoplasm | IBA GO_REF:0000033 | ACCEPT | Summary: BIN2 is present in the cytoplasm. Reason: Consistent with direct localization data; POLAR confines BIN2 to the cytosol in stomatal lineage cells. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005737 cytoplasm | IDA PMID:28652362 Phosphosite charge rather than shootward localization determ... | ACCEPT | Summary: BIN2 is present in the cytoplasm. Reason: Consistent with direct localization data; POLAR confines BIN2 to the cytosol in stomatal lineage cells. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005737 cytoplasm | IEA GO_REF:0000044 | ACCEPT | Summary: BIN2 is present in the cytoplasm. Reason: Consistent with direct localization data; POLAR confines BIN2 to the cytosol in stomatal lineage cells. Supporting Evidence: PMID:28652362 an mRFP-BIN2 fusion protein displayed spotty cytoplasmic as well as nuclear localization |
| GO:0005886 plasma membrane | EXP PMID:30429609 POLAR-guided signalling complex assembly and localization dr... | ACCEPT | Summary: BIN2 can be recruited to the plasma membrane (by OPS, POLAR/BASL, TDR). Reason: Experimentally observed; BIN2 is regulated by plasma-membrane scaffolds and receptors. Supporting Evidence: PMID:30429609 Here we identify the plant-specific protein POLAR as a stomatal lineage scaffold for a subset of GSK3-like kinases that confines them to the cytosol and subsequently transiently polarizes them within the cell |
| GO:0005886 plasma membrane | IDA PMID:28652362 Phosphosite charge rather than shootward localization determ... | ACCEPT | Summary: BIN2 can be recruited to the plasma membrane (by OPS, POLAR/BASL, TDR). Reason: Experimentally observed; BIN2 is regulated by plasma-membrane scaffolds and receptors. Supporting Evidence: PMID:28652362 coexpression of OPS and BIN2 fusion proteins resulted in recruitment of BIN2 to the plasma membrane |
| GO:0005886 plasma membrane | IEA GO_REF:0000044 | ACCEPT | Summary: BIN2 can be recruited to the plasma membrane (by OPS, POLAR/BASL, TDR). Reason: Experimentally observed; BIN2 is regulated by plasma-membrane scaffolds and receptors. Supporting Evidence: PMID:28652362 coexpression of OPS and BIN2 fusion proteins resulted in recruitment of BIN2 to the plasma membrane |
| GO:0005938 cell cortex | IDA PMID:30429609 POLAR-guided signalling complex assembly and localization dr... | KEEP AS NON CORE | Summary: BIN2 polarized at the cell periphery with POLAR/BASL in stomatal lineage cells. Reason: Cell-type-specific polarized localization. Supporting Evidence: PMID:30429609 Here we identify the plant-specific protein POLAR as a stomatal lineage scaffold for a subset of GSK3-like kinases that confines them to the cytosol and subsequently transiently polarizes them within the cell |
| GO:0005938 cell cortex | IEA GO_REF:0000044 | KEEP AS NON CORE | Summary: BIN2 polarized at the cell periphery with POLAR/BASL in stomatal lineage cells. Reason: Cell-type-specific polarized localization. Supporting Evidence: PMID:30429609 Here we identify the plant-specific protein POLAR as a stomatal lineage scaffold for a subset of GSK3-like kinases that confines them to the cytosol and subsequently transiently polarizes them within the cell |
| GO:0006468 protein phosphorylation | IDA PMID:24771765 Brassinosteroids control root epidermal cell fate via direct... | ACCEPT | Summary: BIN2 phosphorylates EGL3 and TTG1. Reason: Direct kinase-substrate evidence; protein phosphorylation is the process BIN2 catalyses. Supporting Evidence: PMID:24771765 One of the GSK3-like kinases, BIN2, interacted with and phosphorylated EGL3 PMID:24771765 BIN2 phosphorylated TTG1 to inhibit the activity of the WER-GL3/EGL3-TTG1 complex. |
| GO:0006468 protein phosphorylation | IPI PMID:12114546 The GSK3-like kinase BIN2 phosphorylates and destabilizes BZ... | ACCEPT | Summary: BIN2 binds and phosphorylates BZR1. Reason: Direct evidence that BIN2 phosphorylates BZR1. Supporting Evidence: PMID:12114546 BIN2 directly interacts with BZR1 in yeast two-hybrid assays, phosphorylates BZR1 in vitro, and negatively regulates BZR1 protein accumulation in vivo. |
| GO:0007033 vacuole organization | IMP PMID:36648110 Spatiotemporal formation of the large vacuole regulated by t... | KEEP AS NON CORE | Summary: BIN2 regulates VLG abundance and large-vacuole formation in the female gametophyte. Reason: Context-specific developmental role. Supporting Evidence: PMID:36648110 gain-of-function mutation of BRASSINOSTEROID INSENSITIVE2 (BIN2) (bin2-1) increases VLG abundance to induce the vacuole formation at stage FG1, and leads to abortion of FG |
| GO:0009414 response to water deprivation | IMP PMID:36508461 A role for brassinosteroid signalling in decision-making pro... | KEEP AS NON CORE | Summary: bin2 alleles alter differential organ growth when light and water are withdrawn. Reason: Phenotypic, downstream consequence of altered BR signalling output in a growth-allocation screen. Supporting Evidence: PMID:36508461 This identified BRASSINOSTEROID INSENSITIVE 2 (BIN2) alleles as decision mutants |
| GO:0009416 response to light stimulus | IMP PMID:36508461 A role for brassinosteroid signalling in decision-making pro... | KEEP AS NON CORE | Summary: bin2 alleles alter seedling growth responses under light withdrawal. Reason: Phenotypic, downstream consequence of BR signalling output. Supporting Evidence: PMID:36508461 This identified BRASSINOSTEROID INSENSITIVE 2 (BIN2) alleles as decision mutants |
| GO:0009553 embryo sac development | IMP PMID:36648110 Spatiotemporal formation of the large vacuole regulated by t... | KEEP AS NON CORE | Summary: bin2-1 causes female gametophyte abortion. Reason: Context-specific developmental role. Supporting Evidence: PMID:36648110 gain-of-function mutation of BRASSINOSTEROID INSENSITIVE2 (BIN2) (bin2-1) increases VLG abundance to induce the vacuole formation at stage FG1, and leads to abortion of FG |
| GO:0009729 detection of brassinosteroid stimulus | IMP PMID:11553730 BIN2, a new brassinosteroid-insensitive locus in Arabidopsis... | REMOVE | Summary: bin2 gain-of-function mutants are BR-insensitive. Reason: Detection of brassinosteroid stimulus is the receptor (BRI1) step. BIN2 is a cytoplasmic/nuclear GSK3-like kinase that negatively regulates the pathway downstream of the receptor; the insensitivity phenotype of semidominant bin2 alleles reflects constitutive negative regulation, not BR detection. The correct process, GO:1900458 negative regulation of brassinosteroid mediated signaling pathway, is already carried by BIN2 (accepted IMP row), so this row is removed rather than modified onto a duplicate. Supporting Evidence: PMID:11553730 bin2 mutants exhibited BR insensitivity when assayed for root growth inhibition and feedback inhibition of CPD gene expression PMID:11847343 Thus, BIN2 acts as a negative regulator to control steroid signaling in plants. |
| GO:0009733 response to auxin | IMP PMID:11820813 The UCU1 Arabidopsis gene encodes a SHAGGY/GSK3-like kinase ... | KEEP AS NON CORE | Summary: ucu1 (bin2) alleles show altered auxin responses. Reason: BIN2 links BR and auxin signalling (e.g. via ARF2/ARF7 phosphorylation) but this is a secondary role. Supporting Evidence: PMID:11820813 which encodes an AtSK protein involved in the cross-talk between auxin and brassinosteroid signaling pathways PMID:18599455 Phosphorylation by BIN2 results in loss of ARF2 DNA binding and repression activities. |
| GO:0009742 brassinosteroid mediated signaling pathway | IBA GO_REF:0000033 | ACCEPT | Summary: Phylogenetic inference of BR signalling for the plant group-II GSK3 clade. Reason: BIN2 is an experimental source for this node; BIL1/BIL2 share the role. Supporting Evidence: PMID:11847343 Thus, BIN2 acts as a negative regulator to control steroid signaling in plants. |
| GO:0009742 brassinosteroid mediated signaling pathway | IMP PMID:11553730 BIN2, a new brassinosteroid-insensitive locus in Arabidopsis... | ACCEPT | Summary: BIN2 is a core component (negative regulator) of BR signalling. Reason: Well established; BIN2 is the central negative regulator between the receptor complex and BZR1/BES1. Supporting Evidence: PMID:11553730 bin2 mutants exhibited BR insensitivity when assayed for root growth inhibition and feedback inhibition of CPD gene expression PMID:11847343 Thus, BIN2 acts as a negative regulator to control steroid signaling in plants. |
| GO:0009825 multidimensional cell growth | IMP PMID:11820813 The UCU1 Arabidopsis gene encodes a SHAGGY/GSK3-like kinase ... | KEEP AS NON CORE | Summary: ucu1 alleles reduce proximodistal cell expansion. Reason: Downstream growth output of BR signalling. Supporting Evidence: PMID:11820813 which encodes an AtSK protein involved in the cross-talk between auxin and brassinosteroid signaling pathways |
| GO:0009965 leaf morphogenesis | IMP PMID:11820813 The UCU1 Arabidopsis gene encodes a SHAGGY/GSK3-like kinase ... | KEEP AS NON CORE | Summary: ucu1 alleles produce spirally rolled leaves. Reason: Downstream developmental output. Supporting Evidence: PMID:11820813 which encodes an AtSK protein involved in the cross-talk between auxin and brassinosteroid signaling pathways |
| GO:0010082 regulation of root meristem growth | IMP PMID:36508461 A role for brassinosteroid signalling in decision-making pro... | KEEP AS NON CORE | Summary: bin2 alleles alter root growth regulation. Reason: Downstream developmental output of BR signalling. Supporting Evidence: PMID:36508461 This identified BRASSINOSTEROID INSENSITIVE 2 (BIN2) alleles as decision mutants |
| GO:0010508 positive regulation of autophagy | IMP PMID:36151786 Brassinosteroids modulate autophagy through phosphorylation ... | KEEP AS NON CORE | Summary: BIN2 phosphorylates RAPTOR1B; BR pathway activity suppresses autophagy. Reason: Secondary output linking BR status to TOR/autophagy. Supporting Evidence: PMID:36151786 directly phosphorylated RAPTOR1B (regulatory-associated protein of TOR 1B), a substrate-recruiting subunit in the TOR complex PMID:36151786 Autophagy was constitutively up-regulated upon blocking BR biosynthesis or signaling, and down-regulated by increasing the activity of the BR pathway. |
| GO:0030154 cell differentiation | IBA GO_REF:0000033 | KEEP AS NON CORE | Summary: Phylogenetic inference of cell differentiation for GSK3 family. Reason: Very broad process term; BIN2 does influence stomatal, xylem and root-hair cell fate decisions, but as a regulator downstream of signalling. Supporting Evidence: PMID:24662460 TDR interacts with GSK3s at the plasma membrane and activates GSK3s in a TDIF-dependent fashion. PMID:24771765 One of the GSK3-like kinases, BIN2, interacted with and phosphorylated EGL3 |
| GO:0031982 vesicle | IDA PMID:36648110 Spatiotemporal formation of the large vacuole regulated by t... | KEEP AS NON CORE | Summary: BIN2 in vesicle-like structures in the female gametophyte. Reason: Context-specific localization. Supporting Evidence: PMID:36648110 BRASSINOSTEROID INSENSITIVE2 (BIN2) is expressed in the FG and localizes in vesicular-like structures. |
| GO:0042802 identical protein binding | IPI PMID:18599455 Integration of auxin and brassinosteroid pathways by Auxin R... | MARK AS OVER ANNOTATED | Summary: BIN2 self-interaction detected by IPI. Reason: Self-association of BIN2 has no established functional role and is not a defining molecular function. The cached text of this paper does not describe the self-interaction (it reports BIN2 interaction with and phosphorylation of other partners); the IPI evidence is in data not in the cache, so no supporting quote is given and the curator's observation is not disputed. |
| GO:0042802 identical protein binding | IPI PMID:22307275 Brassinosteroid regulates stomatal development by GSK3-media... | MARK AS OVER ANNOTATED | Summary: BIN2 self-interaction detected by IPI. Reason: Self-association of BIN2 has no established functional role and is not a defining molecular function. The cached text of this paper does not describe the self-interaction (it reports BIN2 interaction with and phosphorylation of other partners); the IPI evidence is in data not in the cache, so no supporting quote is given and the curator's observation is not disputed. |
| GO:0042802 identical protein binding | IPI PMID:22466366 SPEECHLESS integrates brassinosteroid and stomata signalling... | MARK AS OVER ANNOTATED | Summary: BIN2 self-interaction detected by IPI. Reason: Self-association of BIN2 has no established functional role and is not a defining molecular function. The cached text of this paper does not describe the self-interaction (it reports BIN2 interaction with and phosphorylation of other partners); the IPI evidence is in data not in the cache, so no supporting quote is given and the curator's observation is not disputed. |
| GO:0042802 identical protein binding | IPI PMID:23169658 MYBL2 is a substrate of GSK3-like kinase BIN2 and acts as a ... | MARK AS OVER ANNOTATED | Summary: BIN2 self-interaction detected by IPI. Reason: Self-association of BIN2 has no established functional role and is not a defining molecular function. The cached text of this paper does not describe the self-interaction (it reports BIN2 interaction with and phosphorylation of other partners); the IPI evidence is in data not in the cache, so no supporting quote is given and the curator's observation is not disputed. |
| GO:0042802 identical protein binding | IPI PMID:24362628 A secreted peptide acts on BIN2-mediated phosphorylation of ... | MARK AS OVER ANNOTATED | Summary: BIN2 self-interaction detected by IPI. Reason: Self-association of BIN2 has no established functional role and is not a defining molecular function. The cached text of this paper does not describe the self-interaction (it reports BIN2 interaction with and phosphorylation of other partners); the IPI evidence is in data not in the cache, so no supporting quote is given and the curator's observation is not disputed. |
| GO:0046777 protein autophosphorylation | HDA PMID:21477822 Autophosphorylation profiling of Arabidopsis protein kinases... | KEEP AS NON CORE | Summary: High-throughput autophosphorylation screen. Reason: BIN2 autophosphorylates (including at Tyr200, which is required for its activity), but autophosphorylation is a regulatory feature rather than its function. Supporting Evidence: PMID:21477822 Luminescent analysis revealed that 179 of the 759 PKs had autophosphorylation activity. PMID:19734888 indicating that Tyr200 phosphorylation is essential for BIN2 to inhibit BR-dependent plant growth and that dephosphorylation of pTyr200 is sufficient to inactivate BIN2 |
| GO:0046827 positive regulation of protein export from nucleus | IDA PMID:17873094 Nucleocytoplasmic shuttling of BZR1 mediated by phosphorylat... | KEEP AS NON CORE | Summary: BIN2-mediated phosphorylation promotes nuclear export of BZR1. Reason: Mechanistically real consequence of BZR1 phosphorylation (14-3-3 binding), downstream of BIN2 kinase activity. Supporting Evidence: PMID:17873094 GSK3-like kinases induce the nuclear export of BZR1 by modulating BZR1 interaction with the 14-3-3 proteins |
| GO:0048765 root hair cell differentiation | IGI PMID:24771765 Brassinosteroids control root epidermal cell fate via direct... | KEEP AS NON CORE | Summary: BIN2 and its paralogs regulate root epidermal cell fate. Reason: BIN2 phosphorylates EGL3/TTG1; a context-specific developmental role. Supporting Evidence: PMID:24771765 One of the GSK3-like kinases, BIN2, interacted with and phosphorylated EGL3 PMID:24771765 BIN2 phosphorylated TTG1 to inhibit the activity of the WER-GL3/EGL3-TTG1 complex. |
| GO:0106310 protein serine kinase activity | EXP PMID:11847343 Regulation of brassinosteroid signaling by a GSK3/SHAGGY-lik... | ACCEPT | Summary: BIN2 phosphorylates serine residues of substrates (BZR1/BES1, SPCH). Reason: Core catalytic activity. The cached abstract of this paper establishes that BIN2/DWF12 is a GSK3/SHAGGY-family kinase but does not itself show serine phosphorylation; the curator's EXP evidence is in the full text (not cached). Direct serine phosphorylation of substrates by BIN2 is shown in other cached papers (e.g. Ser70/Ser146/Ser267 of BLISTER). Supporting Evidence: PMID:37335918 BRASSINOSTEROID INSENSITIVE2 interacts with and phosphorylates BLI at 4 phosphorylation sites (Ser70, Ser146, Thr256, and Ser267) for degradation PMID:22466366 The GSK3/SHAGGY-like kinase BIN2 (BR INSENSITIVE2) phosphorylates residues overlapping those targeted by the MAPKs |
| GO:0106310 protein serine kinase activity | EXP PMID:12428015 Arabidopsis brassinosteroid-insensitive dwarf12 mutants are ... | ACCEPT | Summary: BIN2 phosphorylates serine residues of substrates (BZR1/BES1, SPCH). Reason: Core catalytic activity. The cached abstract of this paper establishes that BIN2/DWF12 is a GSK3/SHAGGY-family kinase but does not itself show serine phosphorylation; the curator's EXP evidence is in the full text (not cached). Direct serine phosphorylation of substrates by BIN2 is shown in other cached papers (e.g. Ser70/Ser146/Ser267 of BLISTER). Supporting Evidence: PMID:37335918 BRASSINOSTEROID INSENSITIVE2 interacts with and phosphorylates BLI at 4 phosphorylation sites (Ser70, Ser146, Thr256, and Ser267) for degradation PMID:22466366 The GSK3/SHAGGY-like kinase BIN2 (BR INSENSITIVE2) phosphorylates residues overlapping those targeted by the MAPKs |
| GO:0106310 protein serine kinase activity | EXP PMID:22466366 SPEECHLESS integrates brassinosteroid and stomata signalling... | ACCEPT | Summary: BIN2 phosphorylates serine residues of substrates (BZR1/BES1, SPCH). Reason: Core catalytic activity. Supporting Evidence: PMID:22466366 The GSK3/SHAGGY-like kinase BIN2 (BR INSENSITIVE2) phosphorylates residues overlapping those targeted by the MAPKs |
| GO:0106310 protein serine kinase activity | IEA GO_REF:0000116 | ACCEPT | Summary: Rhea-derived protein serine kinase activity. Reason: Consistent. Supporting Evidence: PMID:11847343 Here we show that the Arabidopsis BRASSINOSTEROID-INSENSITIVE 2 (BIN2) gene encodes a GSK3/SHAGGY-like kinase. |
| GO:1900458 negative regulation of brassinosteroid mediated signaling pathway | IMP PMID:28575660 The F-box Protein KIB1 Mediates Brassinosteroid-Induced Inac... | ACCEPT | Summary: BIN2 negatively regulates BR signalling; its BR-induced degradation by KIB1 is required for BR responses. Reason: Core role of BIN2. Supporting Evidence: PMID:28575660 Here we identify KIB1 as an F-box E3 ubiquitin ligase that promotes the degradation of BIN2 while blocking its substrate access. PMID:11847343 Thus, BIN2 acts as a negative regulator to control steroid signaling in plants. |
Loading supporting content…
Download this section (compressed HTML)Q: Is BSU1-mediated dephosphorylation of pTyr200 or KIB1-mediated degradation the predominant mode of BIN2 inactivation in different tissues and BR concentrations?
Q: How do scaffolds (POLAR, OPS, BASL) and receptors (TDR) partition the BIN2 pool among its many substrates?
Experiment: Quantify BIN2 pTyr200 and total BIN2 protein over a BR dose-time course in bsu1 bsl1 bsl2 bsl3 and kib1 kib2 kib3 backgrounds to separate the contributions of dephosphorylation and degradation.
Loading supporting content…
Download this section (compressed HTML)Loading supporting content…
Download this section (compressed HTML)