SMXL7

UniProt ID: O80875
Organism: Arabidopsis thaliana
Review Status: COMPLETE
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Gene Description

SMAX1-LIKE7 (SMXL7; D53-like 1) is a nuclear Clp-ATPase-related (double Clp-N / P-loop NTPase domain) protein and the most influential member of the SMXL6/7/8 clade of strigolactone (SL)-regulated transcriptional co-repressors. Through an EAR motif it recruits TOPLESS/TPR co-repressors and represses target genes such as BRANCHED1 (BRC1), and it promotes PIN1 accumulation and auxin transport in the stem, allowing axillary bud outgrowth. SL perceived by the receptor D14 triggers SL-dependent association of nuclear SMXL7 with D14 and SCF(MAX2), followed by rapid polyubiquitination and proteasomal degradation; deletion of the conserved RGKT (P-loop) motif stabilises SMXL7 and confers a highly branched phenotype. SMXL7 functions in the nucleus, and part of its activity is independent of the EAR motif. It acts redundantly with SMXL6 and SMXL8 and also influences leaf shape, lateral root density and drought tolerance.

Existing Annotations Review

GO Term Evidence Action Reason
GO:0005515 protein binding
IPI
PMID:26546446
Strigolactone Signaling in Arabidopsis Regulates Shoot Devel...
REMOVE
Summary: SMXL7 interacts with the SL receptor D14 in a GR24-enhanced manner (Y2H/co-IP).
Reason: The interaction is real and central to SL signalling (SMXLs are the receptor's targets), but protein binding conveys no functional information about SMXL7 itself; its activity is represented by transcription corepressor activity. Removal does not imply the interaction is false.
Supporting Evidence:
PMID:26546446
In response to rac -GR24 treatment, SMXL6 and SMXL7 directly interacted with D14 in yeast cells in a concentration-dependent manner
GO:0005515 protein binding
IPI
PMID:37759806
SUPPRESSOR OF MAX2 LIKE 6, 7, and 8 Interact with DDB1 BINDI...
REMOVE
Summary: SMXL6/7/8 interact with DWA1, a CUL4-based E3 substrate receptor.
Reason: Protein binding is uninformative; the DWA1 association reflects regulation of SMXL stability rather than an activity of the SMXL protein. Removal does not imply the interaction is false.
Supporting Evidence:
PMID:37759806
physically interacted with SMXL6,7,8
GO:0005524 ATP binding
IEA
GO_REF:0000002
MARK AS OVER ANNOTATED
Summary: InterPro (AAA ATPase core) mapping to ATP binding.
Reason: SMXL proteins share Clp-ATPase-like nucleotide-binding domains, but nucleotide binding has not been demonstrated and the conserved P-loop region (RGKT) functions as the SL-triggered degron. The domain-based inference overstates the evidence.
GO:0005634 nucleus
EXP
PMID:26546446
Strigolactone Signaling in Arabidopsis Regulates Shoot Devel...
ACCEPT
Summary: GFP-SMXL7 co-localises with a nuclear marker in protoplasts.
Reason: Direct localisation; SMXL co-repressors act on target promoters in the nucleus.
Supporting Evidence:
PMID:26546446
the GFP-SMXL6, GFP-SMXL7, and GFP-SMXL8 proteins colocalized with SV40NLS-mCherry
PMID:27317673
Here, we show that SMXL7 functions in the nucleus, as does the SL receptor, DWARF14 (D14).
GO:0005634 nucleus
IBA
GO_REF:0000033
ACCEPT
Summary: Phylogenetic inference of nuclear activity for the SMXL clade.
Reason: Consistent with experimental nuclear localisation and nuclear transcriptional function.
GO:0005634 nucleus
IEA
GO_REF:0000044
ACCEPT
Summary: Electronic mapping to nucleus.
Reason: Consistent with experimental nuclear localisation.
GO:0005737 cytoplasm
ISM
GO_REF:0000122
MARK AS OVER ANNOTATED
Summary: AtSubP prediction of cytoplasm.
Reason: Sequence-based prediction only; experimental data place SMXL6/7/8 in the nucleus, and SMXL7 activity requires nuclear localisation.
GO:0006457 protein folding
IEA
GO_REF:0000108
REMOVE
Summary: Protein folding inferred (GO_REF:0000108) from the IBA chaperone annotation.
Reason: Derived from the protein folding chaperone IBA, which is not supported for the SMXL clade.
GO:0009414 response to water deprivation
IEP
PMID:32295207
Negative Roles of Strigolactone-Related SMXL6, 7 and 8 Prote...
KEEP AS NON CORE
Summary: smxl6,7,8 triple mutants are more drought resistant than wild type.
Reason: A downstream physiological role of SL signalling; the evidence is from the triple mutant, so the contribution of the individual paralog is inferred.
Supporting Evidence:
PMID:32295207
Our findings together indicate that the SMXL6, 7 and 8 act as negative regulators of drought resistance
GO:0016887 ATP hydrolysis activity
IEA
GO_REF:0000002
MARK AS OVER ANNOTATED
Summary: InterPro (AAA ATPase core) mapping to ATP hydrolysis activity.
Reason: No ATPase activity has been demonstrated for SMXL6/7/8; their characterised activity is transcriptional co-repression. Domain similarity to Clp ATPases is insufficient to assert catalytic function.
GO:0044183 protein folding chaperone
IBA
GO_REF:0000033
REMOVE
Summary: IBA transfer of protein folding chaperone activity from a deep Clp-ATPase node seeded by bacterial ClpB/ClpC (P9WPC9).
Reason: The SMAX1/SMXL clade has diverged from Clp chaperones into hormone-regulated transcriptional co-repressors; there is no evidence of chaperone or disaggregase activity for any SMXL protein, which lack the ClpB/ClpC middle domain architecture. The ancestral chaperone function should not propagate into this clade.
Propagation Review
Root cause: PROPAGATION BAD
Failure modes: FUNCTIONAL DIVERGENCE
Sources checked:
PANTHER:PTN002461480 Β· Clp ATPase ancestral node SUPPORTS SOURCE BUT NOT TARGET
Chaperone activity is well grounded for bacterial ClpB/ClpC (P9WPC9) but the plant SMXL clade is functionally diverged.
Supporting Evidence:
PMID:26546446
D53-like SMXLs exhibit TPR2-dependent transcriptional repression activity and repress the expression of BRANCHED1.
GO:0003714 transcription corepressor activity
IDA
PMID:26546446
Strigolactone Signaling in Arabidopsis Regulates Shoot Devel...
NEW
Summary: SMXL7 fused to GAL4-BD represses reporter transcription in an EAR-motif- and TPR2-dependent manner.
Reason: The characterised molecular activity of the SMXL6/7/8 clade is EAR-motif-dependent co-repression via TOPLESS-related proteins; GOA has no informative MF for this protein.
Supporting Evidence:
PMID:26546446
Compared with GAL4 alone, the relative luciferase activities were dramatically reduced when GAL4 was fused with SMXL6, SMXL7, or SMXL8, and this was dependent on the EAR motif
PMID:26546446
D53-like SMXLs exhibit TPR2-dependent transcriptional repression activity and repress the expression of BRANCHED1.
GO:2000032 regulation of secondary shoot formation
IMP
PMID:26546446
Strigolactone Signaling in Arabidopsis Regulates Shoot Devel...
NEW
Summary: smxl6/7/8 loss suppresses the bushy max2/max3 phenotypes; SMXL6/7/8 promote axillary bud outgrowth.
Reason: SMXL6/7/8 repress BRC1 and are the direct effectors whose SL-triggered removal inhibits bud outgrowth; GOA lacks a branching term for this gene.
Supporting Evidence:
PMID:26546446
the triple mutant smxl6 / 7 / 8 completely restored max3-9 and displayed a significant decrease in secondary cauline branch number, suggesting that SMXL6 , SMXL7 , and SMXL8 function redundantly in promoting the outgrowth of axillary buds
PMID:26546447
SMXL6, SMXL7, and SMXL8 are co-orthologs of rice D53 that promote shoot branching
GO:1902348 cellular response to strigolactone
IDA
PMID:26546446
Strigolactone Signaling in Arabidopsis Regulates Shoot Devel...
NEW
Summary: SMXL7 is ubiquitinated and degraded within minutes of SL treatment, in a D14/MAX2-dependent manner.
Reason: SMXL proteins are the repressor components of the SL signalling pathway, whose regulated degradation is the signalling step.
Supporting Evidence:
PMID:26546446
Exogenous application of the SL analog rac-GR24 causes ubiquitination and degradation of SMXL6, 7, and 8; this requires D14 and MAX2.

Core Functions

Strigolactone-regulated transcriptional co-repressor: SMXL7 recruits TOPLESS-related co-repressors through its EAR motif to repress SL-responsive genes such as BRC1, and is removed by D14/SCF(MAX2)-dependent ubiquitination and proteasomal degradation when SL is perceived, de-repressing bud-outgrowth inhibitors.

Supporting Evidence:
  • PMID:26546446
    D53-like SMXLs exhibit TPR2-dependent transcriptional repression activity and repress the expression of BRANCHED1.
  • PMID:26546446
    Exogenous application of the SL analog rac-GR24 causes ubiquitination and degradation of SMXL6, 7, and 8; this requires D14 and MAX2.
  • PMID:26546446
    the triple mutant smxl6 / 7 / 8 completely restored max3-9 and displayed a significant decrease in secondary cauline branch number, suggesting that SMXL6 , SMXL7 , and SMXL8 function redundantly in promoting the outgrowth of axillary buds
  • PMID:26546447
    SMXL7 is degraded rapidly after treatment with the synthetic strigolactone mixture rac-GR24.
  • PMID:27317673
    Here, we show that SMXL7 functions in the nucleus, as does the SL receptor, DWARF14 (D14).

References

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Suggested Questions for Experts

Q: Does the Clp-ATPase-like region of SMXL proteins bind or hydrolyse ATP, and does nucleotide state influence D14-SCF(MAX2) recognition?

Q: Which DNA-binding transcription factors (e.g. SPL, BES1, TCP partners) tether SMXL6/7/8 to BRC1 and other targets in axillary buds?

Suggested Experiments

Experiment: In vitro ATPase assays with purified SMXL7 (wild type versus Walker-motif mutants), combined with complementation of smxl6/7/8, to test whether ATPase activity contributes to repression.

Type: biochemistry and genetic complementation

πŸ“š Additional Documentation

Notes

(SMXL7-notes.md)

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