Transcriptomic analysis reveals the GRAS family genes respond to gibberellin in Salvia miltiorrhiza hairy roots

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Transcriptomic analysis reveals the GRAS family genes respond to gibberellin in Salvia miltiorrhiza hairy roots Wenrui Li1,2, Chuangfeng Liu3, Jingling Liu3, Zhenqing Bai3 and Zongsuo Liang1,4*

Abstract Background: Salvia miltiorrhiza is one of the most important traditional Chinese medicinal plants with high medicinal value. Gibberellins are growth-promoting phytohormones that regulate numerous growth and developmental processes in plants. However, their role on the secondary metabolism regulation has not been investigated. Results: In this study, we found that gibberellic acid (GA) can promote hairy roots growth and increase the contents of tanshinones and phenolic acids. Transcriptomic sequencing revealed that many genes involved in the secondary metabolism pathway were the GA-responsive. After further analysis of GA signaling pathway genes, which their expression profiles have significantly changed, it was found that the GRAS transcription factor family had a significant response to GA. We identified 35 SmGRAS genes in S. miltiorrhiza, which can be divided into 10 subfamilies. Thereafter, members of the same subfamily showed similar conserved motifs and gene structures, suggesting possible conserved functions. Conclusions: Most SmGRAS genes were significantly responsive to GA, indicating that they may play an important role in the GA signaling pathway, also participating in the GA regulation of root growth and secondary metabolism in S. miltiorrhiza. Keywords: Transcriptome, GRAS family, Gibberellin, Salvia miltiorrhiza hairy roots, Secondary metabolism

Background Salvia miltiorrhiza Bunge (Danshen) is a well-known traditional Chinese medicine with high medicinal and economic value. It is mainly used to treat cardiovascular and cerebrovascular diseases [1]. The Chinese pharmacopeia stipulates that the medicinal part of S. miltiorrhiza is its dried root. There are two major bioactive components of S. miltiorrhiza, lipophilic tanshinones and hydrophilic phenolic acids [2]. More than 40 * Correspondence: [email protected] 1 Institute of Soil and Water Conservation, Chinese Academy of Sciences and Ministry of Water Resources, Yangling 712100, China 4 College of Life Sciences and Medicine, The Key Laboratory of Plant Secondary Metabolism and Regulation of Zhejiang Province, Zhejiang Sci-Tech University, Hangzhou 310018, China Full list of author information is available at the end of the article

tanshinones and 20 hydrophilic phenolic acids have been isolated and identified from S. miltiorrhiza [3]. The tanshinones, including dihydrotanshinone I (DT-I), cryptotanshinone (CT), tanshinone I (T-I) and tanshinone IIA (T-IIA), are biosynthesized via the mevalonic acid (MVA) and 2-C-methyl-D-erythritol-4-phosphate (MEP) pathways [4, 5]. The phenolic acids, including salvianolic acid B (Sal B) and rosmarinic acid (RA), are biosynthesized through the phenylpropanoid and tyrosine-derived pathways [6, 7]. Most of the key biosynthetic enzyme genes of those pathways have been