三七RALF基因家族鉴定及表达分析

doi:10.13560/j.cnki.biotech.bull.1985.2025-0585

生物技术通报 ›› 2026, Vol. 42 ›› Issue (2): 278-292.doi: 10.13560/j.cnki.biotech.bull.1985.2025-0585

三七RALF基因家族鉴定及表达分析

杨子涵¹^,²^,³(), 李葵秀¹^,²^,³, 李俊良¹^,²^,³, 吕文慧¹^,²^,³, 何温婷⁴, 刘旭燕⁵, 刘冠泽¹^,²()

^1.云南农业大学西南中药材种质创新与利用国家地方联合工程研究中心，昆明 650201
^2.云南农业大学云南省药用植物生物学重点实验室，昆明 650201
^3.云南农业大学农学与生物技术学院，昆明 650201
^4.云南农业大学烟草学院，昆明 650201
^5.云南农业大学植物保护学院，昆明 650201

收稿日期:2025-06-06 出版日期:2026-02-26 发布日期:2026-03-17
通讯作者: 刘冠泽，男，教授，研究方向：药用植物生物技术与育种；E-mail: guanzeliu@ynau.edu.cn
作者简介:杨子涵，女，硕士研究生，研究方向：三七种质资源评价与遗传改良；E-mail: 1439104670@qq.com
基金资助:
国家自然科学基金项目(32260095);云南省重大科技专项(202205AR070001);云南省兴滇英才计划资助(XDYC-QNRC-2022-0210)

Identification and Expression Analysis of the RALF Gene Family in Panax notoginseng

YANG Zi-han¹^,²^,³(), LI Kui-xiu¹^,²^,³, LI Jun-liang¹^,²^,³, LYU Wen-hui¹^,²^,³, HE Wen-ting⁴, LIU Xu-yan⁵, LIU Guan-ze¹^,²()

^1.National & Local Joint Engineering Research Center of Germplasm Innovation and Utilization of Southwest Chinese Medicinal Materials, Yunnan Agricultural University, Kunming 650201
^2.Yunnan Key Laboratory for Medicinal Plant Biology, Yunnan Agricultural University, Kunming 650201
^3.College of Agronomy and Biotechnology, Yunnan Agricultural University, Kunming 650201
^4.College of Tobacco, Yunnan Agricultural University, Kunming 650201
^5.College of Plant Protection, Yunnan Agricultural University, Kunming 650201

Received:2025-06-06 Published:2026-02-26 Online:2026-03-17

摘要/Abstract

摘要：

目的快速碱化因子（rapid alkalinization factors, RALFs）是一类植物多肽类激素，参与植物免疫应答反应，在三七全基因组水平对RALF基因家族进行鉴定并探究PnoRALFs基因家族在三七免疫反应中的作用。方法利用生物信息学分析方法对PnoRALFs基因家族进行鉴定，对该基因家族染色体分布、蛋白理化性质、系统发育树、保守基序、基因结构、启动子顺式元件和组织特异性进行表达分析；采用4种外源激素水杨酸（SA）、茉莉酸甲酯（MeJA）、脱落酸（ABA）、乙烯（ETH）处理及毁灭柱孢菌（Cylindrocarpon destructans）侵染，分析基因表达模式，并利用烟草瞬时表达验证关键基因的功能。结果三七全基因组中共鉴定到13个PnoRALFs基因，分布于8条染色体，分为Ⅰ亚类和Ⅱ亚类；基因结构和保守基序分析发现，PnoRALFs基因含有保守的RRXL基序和YISY保守基序；顺式元件预测发现PnoRALFs基因启动子区段含有多种激素如ABA、MeJA、SA和赤霉素（GA）等响应元件。组织特异性表达分析发现，13个PnoRALFs基因在不同组织中的表达水平存在显著差异；此外，4种外源激素处理下，PnoRALF2、PnoRALF5、PnoRALF6、PnoRALF9表达量呈现上调趋势；毁灭柱孢侵染后转录组数据及RT-qPCR分析发现，9个PnoRALFs响应毁灭柱孢侵染，其中PnoRALF9表达量显著上调；烟草瞬时表达PnoRALF9后，烟草中PTI途径NbFPK1、NbWRKY29和ROS途径NbMPK6、NbGAPDH表达量上调。结论三七全基因组共鉴定到13个PnoRALFs基因，PnoRALF9响应4种外源激素处理及毁灭柱孢侵染，瞬时过表达后诱导植物免疫相关基因表达，推测PnoRALF9在三七抗病免疫反应中发挥重要作用。

关键词: 三七, RALF基因家族, 激素处理, 毁灭柱孢, 表达分析

Abstract:

Objective Rapid alkalinization factors (RALFs) are plant peptide hormones that participate in immune responses. This study is aimed to identify the RALF gene family members in Panax notoginseng and to investigate the roles of PnoRALFs in P. notoginseng immune responses. Method The RALF gene family in P. notoginseng was identified using bioinformatics tools. Chromosomal localization, protein physicochemical properties, phylogenetic relationships, conserved motifs, gene structures, promoter cis-acting elements, and tissue-specific expression patterns were analyzed. Expression profiles were assessed under four hormone treatments-salicylic acid (SA), methyl jasmonate (MeJA), abscisic acid (ABA), and ethylene (ETH) and after infection with C. destructans. The function of a key gene was further validated through transient expression in Nicotiana benthamiana. Result Thirteen PnoRALFs genes were identified and mapped to eight chromosomes. These genes were classified into two subgroups. Conserved motif analysis showed the presence of RRXL and YISY motifs. Promoter regions contained multiple hormone-responsive cis-elements, including those responsive to ABA, MeJA, SA, and gibberellins (GA). Tissue-specific expression analysis revealed considerable variation in gene expressions among different tissues. The expressions of PnoRALF2, PnoRALF5, PnoRALF6, and PnoRALF9 were upregulated by hormone treatments. Transcriptome and RT-qPCR analyses indicated that nine PnoRALFs genes responded to C. destructans infection, with PnoRALF9 showing a marked increase in expression. Transient overexpression of PnoRALF9 in N. benthamiana increased the expressions of immunity-related genes such as NbFPK1 and NbWRKY29 in the pattern-triggered immunity (PTI) pathway and NbMPK6 and NbGAPDH in the reactive oxygen species (ROS) signaling pathway. Conclusion Thirteen PnoRALFs genes are identified in the P. notoginseng genome. Among them, PnoRALF9 responds to multiple hormone treatments and C. destructans infection. Transient overexpression of PnoRALF9 induces plant immunity-related gene expression, suggesting that PnoRALF9 plays a critical role in the immune response in P. notoginseng to disease.

Key words: Panax notoginseng, RALF gene family, hormone treatment, Cylindrocarpon destructans, expression analysis

杨子涵, 李葵秀, 李俊良, 吕文慧, 何温婷, 刘旭燕, 刘冠泽. 三七RALF基因家族鉴定及表达分析[J]. 生物技术通报, 2026, 42(2): 278-292.

YANG Zi-han, LI Kui-xiu, LI Jun-liang, LYU Wen-hui, HE Wen-ting, LIU Xu-yan, LIU Guan-ze. Identification and Expression Analysis of the RALF Gene Family in Panax notoginseng[J]. Biotechnology Bulletin, 2026, 42(2): 278-292.

图/表 15

表1 PnoRALFs基因家族实时荧光定量引物

Table 1 Primers for the RT-qPCR analysis of PnoRALFs

基因 Gene	上游引物Forward primer （5′-3′）	下游引物序列Reverse primer （5′-3′）
PnACT2	GATTGATCTTGGCACCGGGA	TGACCAACCCACGACCTTTC
PnoRALF1	GCTACATGCAACGGTCTGATC	TGCAGTTGTAGTAGGAGGCTC
PnoRALF2	CCACCTGAACTCTATTGGGT	CGCACCGTAGCTGATGTA
PnoRALF3	AGTGATGCTGATGGACTCGG	CGTCTGTAGGGATTGGCTCT
PnoRALF4	CCACCTGAACTCTATTGGGT	GGGAGTACGGATTTGCCTG
PnoRALF5	CGACATGGATTCAGAGAGC	ACGGTTGTAAGGGTTAGCTT
PnoRALF6	CGCCTCCTACTACAACTGTA	ACCAGCATCCATCACAAATC
PnoRALF7	GCAAACAGTCGGTACATCAG	GGGAGTACGGATTTGCCTG
PnoRALF8	GATTCATGGTCGTTTGTCAC	CTGGCCTCGTTTGTTACATG
PnoRALF9	TGATCTTGCTCCCATTTCCA	GGCTGATCTCCGAATCCATA
PnoRALF10	TGTTTCGACGGTGGTGTTTC	CACACACCTGCTCGTTTCTC
PnoRALF11	ATGATGTTGGATTCGGAGGC	CGTCTGTAGGGATTGGCTCT
PnoRALF12	AGTCCTAATCCTGCTGTGTTTC	CGGTTTATCTCCGAATCCATCA
PnoRALF13	CGGAGGTTTCTTGCAGCAAA	CGGAGGTTTCTTGCAGCAAA
pEAQ-HT-PnoRALF9	cccaaattcgcgaccggtATGGGAAAG TCTACTGGTTTTCTCG	atgaaaccagagttaaaggcctcgagACGCC GGCACCGAGTGAT

图1 PnoRALFs基因家族染色体定位

Fig. 1 Chromosomal localization of the PnoRALFs gene family

表2 PnoRALFs基因家族基本理化性质

Table 2 Basic physicochemical properties of PnoRALFs gene family

基因	基因ID	氨基酸数	分子量	等电点	亲水性	亚细胞定位	信号肽
Gene name	Gene ID	Number of amino acids	Molecular weight （kD）	pl	GRAVY	Subcellular localization	Signal peptide
PnoRALF1	Pno01G005926.t1	128	14.06	9.03	-0.30	细胞外Extracellular	Yes
PnoRALF2	Pno03G001289.t1	126	13.74	8.91	-0.11	细胞外Extracellular	Yes
PnoRALF3	Pno03G005478.t1	131	14.39	0.24	-0.22	细胞外Extracellular	Yes
PnoRALF4	Pno03G005479.t1	125	13.75	9.54	-0.44	细胞外Extracellular	Yes
PnoRALF5	Pno06G000518.t1	114	12.33	8.60	-0.20	细胞外Extracellular	Yes
PnoRALF6	Pno07G004032.t1	158	17.28	9.15	0.24	质膜 Plasma membrane	Yes
PnoRALF7	Pno09G001048.t1	109	12.07	7.50	-0.35	细胞外Extracellular	Yes
PnoRALF8	Pno10G000839.t1	123	13.77	9.83	-0.19	细胞外Extracellular	No
PnoRALF9	Pno11G003888.t1	133	14.33	8.99	-0.08	细胞外Extracellular	Yes
PnoRALF10	Pno12G001525.t1	110	12.31	7.54	-0.07	细胞外Extracellular	Yes
PnoRALF11	Pno284G000001.t1	125	12.75	9.54	-0.44	细胞外Extracellular	Yes
PnoRALF12	Pno606G000002.t1	110	12.30	7.54	-0.07	细胞外Extracellular	Yes
PnoRALF13	Pno606G000003.t1	113	12.46	8.69	-0.29	细胞外Extracellular	Yes

图2 三七与拟南芥RALF基因家族的系统发育树

Fig. 2 Phylogenetic tree of RALF gene family in P. notoginseng and Arabidopsisthaliana

图3 PnoRALFs家族保守motif与基因结构分析（A）、motif标识（B）

Fig. 3 Analysis of PnoRALFs family conserved motifs and gene structure(A), motif logo of PnoRALFs (B)

图4 PnoRALF蛋白与已知功能AtRALF蛋白的多序列比对

Fig. 4 Multiple sequence alignment of PnoRALF protein and AtRALF protein with known function

图5 PnoRALFs基因顺式作用元件分析

Fig. 5 Analysis of cis-acting elements of gene PnoRALFs

图6 PnoRALFs基因在不同组织表达模式分析A：PnoRALFs 基因在不同组织的转录组数据；B：PnoRALFs基因在不同组织的相对表达量。*P<0.05，**P<0.01，下同

Fig. 6 Expression profiling of PnoRALFs gene in different tissuesA: Transcriptome expression data of PnoRALFs gene in different tissues. B: Relative expressions of PnoRALFs gene in different tissues. *P<0.05, **P<0.01. The same below

图7 PnoRALFs基因在MeJA不同时间点处理下的表达模式分析

Fig. 7 Analysis of the expression pattern of PnoRALFs gene under different time of MeJA treatment

图8 PnoRALFs基因在SA不同时间点处理下的表达模式分析

Fig. 8 Analysis of the expression pattern of PnoRALFs gene under different time of SA treatment

图9 PnoRALFs基因在ABA不同时间点处理下的表达模式分析

Fig. 9 Analysis of the expression pattern of PnoRALFs gene under different time of ABA treatment

图10 PnoRALFs基因在ETH不同时间点处理下的表达模式分析

Fig. 10 Analysis of the expression pattern of PnoRALFs gene under different time of ETH treatment

图11 PnoRALFs基因响应毁灭柱孢侵染表达模式分析A：PnoRALFs基因响应毁灭柱孢侵染的转录组数据；B：PnoRALFs基因响应毁灭柱孢侵染的相对表达量

Fig. 11 Expression profiling of PnoRALFs gene in response to C. destructans infectionA: Transcriptomic data of PnoRALFs gene in response to C. destructans infection.B: Relative expression of PnoRALFs gene reponding to C. destructans infection

图12 瞬时转化PnoRALF9后烟草表型及相关基因表达分析A：瞬时转化PnoRLAF9后GFP基因表达的烟草表型；B：瞬时转化后PnoRLAF9基因表达量分析；C：烟草中免疫相关基因表达分析

Fig. 12 Analysis of phenotypic and gene expression in N. benthamiana following transient transformation of PnoRALF9A: Phenotype of GFP expression in N. benthamiana following transient expression of PnoRALF9. B: Analysis of PnoRALF9 expression after transient transformation. C: Expression analysis of immune-related genes in N. benthamiana

图13 PnoRALFs关键基因的选择及PnoRALF9瞬时表达模式图

Fig. 13 Selection of key PnoRALFs gene and transient expression patterns of PnoRALF9

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三七RALF基因家族鉴定及表达分析

Identification and Expression Analysis of the RALF Gene Family in Panax notoginseng

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