Non-transcriptional regulatory activity of SMAX1 and SMXL2 mediates karrikin-regulated seedling response to red light in Arabidopsis

Karrikins and strigolactones govern plant development and environmental responses through closely related signaling pathways.The transcriptional repressor proteins SUPPRESSOR OF MAX21(SMAX1),SMAX1-like2(SMXL2),and D53-like SMXLs mediate karrikin and strigolactone signaling by directly binding downstream genes or byinhibiting the activities of transcription factors.In this study,we characterized the non-transcriptional regulatory activities of SMXL proteins in Arabidopsis.We discovered that SMAX1 and SMXL2 with mutations in their ethylene-responsefactor-associated amphiphilic repression(EAR)motif had undetectable or weak transcriptional repression activities but still partially rescued the hypocotyl elongation defects and fully reversed the cotyledon epinasty defects of the smax1 smxl2 mutant.SMAX1 and SMXL2 directly interact with PHYTOCHROME INTERACTION FACTOR4(PIF4)and PIF5 to enhance their protein stability by interacting with phytochrome B(phyB)and suppressing the association of phyB with PIF4 and PIF5.The karrikin-responsive genes were then identified by treatment with GR24ent-ssa,GR24 analog showing karrikin activity.Interestingly,INDOLE-3-ACETIC ACID INDUCIBLE 29(IAA29)expression was repressed by GR24^(ent-5D)streatment in a PIF4-and PIF5-dependent and EARindependent manner,whereas KARRIKIN UPREGULATED F-BOX 1(KUF1)expression was induced in a PIF4-and PIF5-independent and EAR-dependent manner.Furthermore,the non-transcriptional regulatory activity of SMAX1,which is independent of the EAR motif,had a global effect on gene expression.Taken together,these results indicate that non-transcriptional regulatory activities of SMAX1 and SMXL2 mediate karrikin-regulated seedling response to red light.

Auxin signaling:Research advances over the past 30 years

Auxin,one of the first identified and most widely studied phytohormones,has been and will remain a hot topic in plant biology.After more than a century of passionate exploration,the mysteries of its synthesis,transport,signaling,and metabolism have largely been unlocked.Due to the rapid development of new technologies,new methods,and new genetic materials,the study of auxin has entered the fast lane over the past 30 years.Here,we highlight advances in understanding auxin signaling,including auxin perception,rapid auxin responses,TRANSPORT INHIBITOR RESPONSE 1 and AUXIN SIGNALING F-boxes(TIR1/AFBs)-mediated transcriptional and non-transcriptional branches,and the epigenetic regulation of auxin signaling.We also focus on feedback inhibition mechanisms that prevent the over-amplification of auxin signals.In addition,we cover the TRANSMEMBRANE KINASE-mediated non-canonical signaling,which converges with TIR1/AFBs-mediated transcriptional regulation to coordinate plant growth and development.The identification of additional auxin signaling components and their regulation will continue to open new avenues of research in this field,leading to an increasingly deeper,more comprehensive understanding of how auxin signals are interpreted at the cellular level to regulate plant growth and development.