Growth-regulating factors (GRFs) are plant-specific transcription factors that play an important role in plant growth and development. In this study, fifteen GRF gene members containing QLQ and WRC domains were identified in Zanthoxylum armatum. Phylogenetic and collinearity analysis showed that ZaGRFs were closely related to CsGRFs and AtGRFs, and distantly related to OsGRFs. There are a large number of cis-acting elements related to hormone response and stress induction in the GRF gene promoter region of Z. armatum. Tissue-specific expression analysis showed that except for ZaGRF7, all the ZaGRFs were highly expressed in young parts with active growth and development, including terminal buds, seeds, and young flowers, suggesting their key roles in Z. armatum growth and development. Eight ZaGRFs were selected to investigate the transcriptional response to auxin, gibberellin and drought treatments. A total of six ZaGRFs in the NAA treatment, four ZaGRFs in the GA₃ treatment, and six ZaGRFs in the PEG treatment were induced and significantly up-regulated. Overexpression of ZaGRF6 increased branching and chlorophyll content and delayed senescence of transgenic Nicotiana benthamiana. ZaGRF6 increased the expression of CRF2 and suppressed the expression of ARR4 and CKX1, indicating that ZaGRF6 is involved in cytokinin metabolism and signal transduction. These research results lay a foundation for further analysis of the GRF gene function of Z. armatum and provide candidate genes for growth, development, and stress resistance breeding of Z. armatum.

中文翻译：

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花椒DC中生长调节因子家族的全基因组鉴定和分析以及ZaGRF6在叶片大小和寿命调节中的功能分析

生长调节因子（GRFs）是植物特异性转录因子，在植物生长发育中起重要作用。在这项研究中，在花椒中鉴定了 15 个含有 QLQ 和 WRC 结构域的GRF基因成员。系统发育和共线性分析表明，ZaGRFs与CsGRFs和AtGRFs亲缘关系较近，与OsGRFs亲缘关系较远。Z. armatum的GRF基因启动子区存在大量与激素反应和应激诱导相关的顺式作用元件。组织特异性表达分析表明，除ZaGRF7 外，所有ZaGRF在生长发育活跃的年轻部位，包括顶芽、种子和幼花中高度表达，表明它们在Z. armatum生长发育中的关键作用。选择了8个 ZaGRF来研究对生长素、赤霉素和干旱处理的转录反应。NAA处理中总共有6个ZaGRFs、GA ₃处理中的4个ZaGRFs和PEG处理中的6个ZaGRFs被诱导并显着上调。ZaGRF6的过表达增加了转基因烟草的分枝和叶绿素含量并延缓了衰老。ZaGRF6增加CRF2的表达，抑制ARR4和CKX1的表达，表明ZaGRF6参与细胞分裂素代谢和信号转导。这些研究成果为进一步分析毛茛的GRF基因功能奠定了基础，为毛茛的生长发育和抗逆育种提供了候选基因。