TT2 controls rice thermotolerance through SCT1-dependent alteration of wax biosynthesis

Global warming threatens crop production. G proteins mediate plant responses to multiple abiotic stresses. Here we identified a natural quantitative trait locus, TT2 ( THERMOTOLERANCE 2 ), encoding a Gγ subunit, that confers thermotolerance in rice during both vegetative and reproductive growth without a yield penalty. A natural allele with loss of TT2 function was associated with greater retention of wax at high temperatures and increased thermotolerance. Mechanistically, we found that a transcription factor, SCT1 (Sensing Ca 2+ Transcription factor 1), functions to decode Ca 2+ through Ca 2+ -enhanced interaction with calmodulin and acts as a negative regulator of its target genes (for example, Wax Synthesis Regulatory 2 ( OsWR2 )). The calmodulin–SCT1 interaction was attenuated by reduced heat-triggered Ca 2+ caused by disrupted TT2 , thus explaining the observed heat-induced changes in wax content. Beyond establishing a bridge linking G protein, Ca 2+ sensing and wax metabolism, our study illustrates innovative approaches for developing potentially yield-penalty-free thermotolerant crop varieties.