AtHAP5A modulates freezing stress resistance in Arabidopsis independent of the CBF pathway

The heme-associated proteins (HAPs, also known as nuclear factor y, subunit A/B/C (NF-YA/B/C)) have been reported to bind specifically to DNA fragments containing CCAAT-box, however, the physiological functions and direct targets of these HAP proteins remain unclear in plants. In our recent study, we found that AtHAP5A and AtXTH21 positively modulated freezing stress resistance, and chromatin immunoprecipitation (ChIP) assay and genetic evidence indicated that AtHAP5A might act in the upstream of AtXTH21 in freezing stress. Moreover, AtHAP5A and AtXTH21 had significant effects on inhibiting cold stress-induced reactive oxygen species (ROS) accumulation and activating ABA-related genes' expression. Thus, a possible model that depicting AtHAP5A-mediated cold stress responses was proposed in this study, and we highlighted that AtHAP5A modulates freezing stress resistance in Arabidopsis through binding to CCAAT motif of AtXTH21, which is independent of the CBF pathway.