Chang_2025_J.Exp.Bot__

SUPPRESSOR OF MAX2 1 (SMAX1) and SMAX1-LIKE (SMXL) proteins comprise a family of plant growth regulators that includes downstream targets of the karrikin (KAR)/KAI2 ligand (KL) and strigolactone (SL) signaling pathways. Following the perception of KAR/KL or SL signals by alpha/beta hydrolases, some types of SMXL proteins are polyubiquitinated by an E3 ubiquitin ligase complex containing the F-box protein MORE AXILLARY GROWTH2 (MAX2)/DWARF3 (D3), and proteolyzed. Because SMXL proteins interact with TOPLESS (TPL) and TPL-related (TPR) transcriptional corepressors, SMXL degradation initiates changes in gene expression. This simplified model of SMXL regulation and function in plants must now be revised in light of recent discoveries. It has become apparent that SMXL abundance is not regulated by KAR/KL or SL alone, and that some SMXL proteins are not regulated by MAX2/D3 at all. Therefore, SMXL proteins should be considered signaling hubs that integrate multiple cues. Here we review the current knowledge of how SMXL proteins impose transcriptional regulation of plant development and environmental responses. SMXL proteins can bind DNA directly and interact with transcriptional regulators from several protein families. Multiple mechanisms of downstream genetic control by SMXL proteins have been identified recently that do not involve the recruitment of TPL/TPR, expanding the paradigm of SMXL function.