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Pepper ( Capsicum annuum ) is an economically important vegetable and heat stress can severely impair pepper growth, development, and productivity. The molecular mechanisms underlying pepper thermotolerance are therefore important to understand but remain elusive. In the present study, we characterized the function of  CaHSL1 , encoding a HAESA-LIKE (HSL) receptor-like protein kinase (RLK), during the response of pepper to high temperature and high humidity (HTHH). CaHSL1 exhibits the typical structural features of an arginine-aspartate RLK. Transient overexpression of CaHSL1 in the mesophyll cells of  Nicotiana benthamiana  showed that CaHSL1 localizes throughout the cell, including the plasma membrane, cytoplasm, and the nucleus.  CaHSL1  was significantly upregulated by HTHH or the exogenous application of abscisic acid but not by  R. solanacearum  inoculation. However,  CaHSL1  was downregulated by exogenously applied salicylic acid, methyl jasmonate, or ethephon. Silencing of  CaHSL1  by virus-induced gene silencing significantly was reduced tolerance to HTHH and downregulated transcript levels of an associated gene  CaHSP24 . In contrast, transient overexpression of  CaHSL1  enhanced the transcript abundance of  CaHSP24  and increased tolerance to HTHH, as manifested by enhanced optimal/maximal photochemical efficiency of photosystem II in the dark (Fv/Fm) and actual photochemical efficiency of photosystem II in the light. In addition, CaWRKY40 targeted the promoter of  CaHSL1  and induced transcription during HTHH but not in response to  R. solanacearum . All of these results suggest that  CaHSL1  is directly modulated at the transcriptional level by CaWRKY40 and functions as a positive regulator in the response of pepper to HTHH.

CaHSL1 Acts as a Positive Regulator of Pepper Thermotolerance Under High Humidity and Is Transcriptionally Modulated by CaWRKY40