Lettuce (Lactuca sativa L.) is an important leafy vegetable consumed worldwide. Heatâinduced bolting and flowering greatly limit lettuce production during the summer. Additionally, MADSâbox transcription factors are important for various aspects of plant development and architecture (e.g., flowering and floral patterning). However, there has been no comprehensive study of lettuce MADSâbox family genes. In this study, we identified 82 MADSâbox family genes in lettuce, including 23 type I genes and 59 type II genes. Transcriptome profiling revealed that LsMADS gene expression patterns differ among the various floral stages and organs. Moreover, heatâresponsive cisâelements were detected in the promoter regions of many LsMADS genes. An in situ hybridization assay of 10 homologs of flowerâpatterning genes and a yeast twoâhybrid assay of the encoded proteins revealed that the ABC model is conserved in lettuce. Specifically, the APETALA1 (AP1) homolog in lettuce, LsMADS55, is responsive to heat and is specifically expressed in the inflorescence meristem and pappus bristles. The overexpression of LsMADS55 results in early flowering in Arabidopsis thaliana. Furthermore, we observed that the heat shock factor LsHSFB2Aâ1 can bind to the LsMADS55 promoter in lettuce. Therefore, a model was proposed for the LsMADSâregulated floral organ specification and heatâinduced flowering in lettuce.