Heterologous expression of heat stress-responsive AtPLC9 confers heat tolerance in transgenic rice

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RESEARCH ARTICLE

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Heterologous expression of heat stressresponsive AtPLC9 confers heat tolerance in transgenic rice Yuliang Liu1†, Xinye Liu1†, Xue Wang1, Kang Gao1, Weiwei Qi1, Huimin Ren1, Haorui Hu1,2, Daye Sun1, Jiaoteng Bai1,3* and Shuzhi Zheng1,3*

Abstract Background: As global warming becomes increasingly severe, it is urgent that we enhance the heat tolerance of crops. We previously reported that Arabidopsis thaliana PHOSPHOINOSITIDE-SPECIFIC PHOSPHOLIPASE C9 (AtPLC9) promotes heat tolerance. Results: In this study, we ectopically expressed AtPLC9 in rice to examine its potential to improve heat tolerance in this important crop. Whereas AtPLC9 did not improve rice tolerance to salt, drought or cold, transgenic rice did exhibit greater heat tolerance than the wild type. High-throughput RNA-seq revealed extensive and dynamic transcriptome reprofiling in transgenic plants after heat stress. Moreover, the expression of some transcription factors and calcium ion-related genes showed specific upregulation in transgenic rice after heat stress, which might contribute to the enhanced heat tolerance. Conclusions: This study provides preliminary guidance for using AtPLC9 to improve heat tolerance in cereal crops and, more broadly, highlights that heterologous transformation can assist with molecular breeding. Keywords: AtPLC9, Ca2 + , Heterologous expression, Rice, Thermotolerance

Background Plants must cope with a complicated and varied environment. The primary abiotic stresses plants facing include heat, cold, drought and salt. Temperature affects plant growth, development, and geographical distribution, and extreme temperatures can adversely affect crop quality and productivity. As temperatures increase above the optimum, plants begin to experience heat stress (HS). Plants perceive and transmit HS signals through a complicated pathway. Heat stress initially increases the fluidity of * Correspondence: [email protected]; [email protected] † Yuliang Liu and Xinye Liu contributed equally to this work. 1 Ministry of Education Key Laboratory of Molecular and Cellular Biology, Hebei Collaboration Innovation Center for Cell Signaling, Hebei Key Laboratory of Molecular and Cellular Biology, College of Life Sciences, Hebei Normal University, Shijiazhuang 050024, China Full list of author information is available at the end of the article

the plasma membrane, and researchers have proposed that HS activates cyclic nucleotide-gated channel family proteins through these changes in membrane fluidity [1, 2]. While it is still not clear what the HS signal transduction comprises, a number of HS signal components capable of regulating thermotolerance in plants have been identified, including Ca2+, calcium-dependent protein kinases (CDPKs), 1,4,5-inositol triphosphate (1,4,5-IP3), cyclic AMP (cAMP) and mitogen-activated protein kinases (MPK) [3, 4]. Moreover, gene expression changes are critical for plant adaptation to high-temperature stress. Recent research reports that transcription factors are important for pla