中草药

连翘bHLH转录因子家族成员的鉴定与表达分析

PDF下载

闫雪娇 ¹,,², , 原梦 ¹,,², , 李文文 ¹,,², , 张力予 ¹,,², , 许高龙 ¹,,², , 杨崇 ¹,,², , 吕淑芳 ¹,,², , 马占强 ¹,,², , 张红晓 ¹,,², , 王婷 ¹,,², , 侯典云 ¹,,²,

中草药 | 药材与资源 2026,57(16): 6521-6531

收起

中草药 |药材与资源 2026 , 57 (16) : 6521 -6531

连翘bHLH转录因子家族成员的鉴定与表达分析

全屏

闫雪娇¹,,²,, 原梦¹,,²,, 李文文¹,,²,, 张力予¹,,²,, 许高龙¹,,²,, 杨崇¹,,²,, 吕淑芳¹,,²,, 马占强¹,,²,, 张红晓¹,,²,, 王婷¹,,²,, 侯典云¹,,²,

作者信息

^1.河南科技大学农学院, 河南洛阳 471023

^2.河南省药食兼用资源评价与创新利用工程研究中心, 河南洛阳 471023

^3.河南科技大学农学院, 河南洛阳 471023

^4.河南省药食兼用资源评价与创新利用工程研究中心, 河南洛阳 471023

通讯作者:

侯典云

作者简介:

闫雪娇: 闫雪娇(2001—),硕士研究生,主要从事药用植物分子生物学研究。E-mail:yuki10242025@163.com 原梦: 原梦(2000—),博士研究生,主要从事药用植物分子生物学研究。E-mail:ym107107@163.com

Identification and expression analysis of bHLH transcription factors family members in Forsythia suspensa

Affiliations

出版时间: 2026-08-28 doi: 10.7501/j.issn.0253-2670.2026.16.024

文章导航

摘要

收起

目的基于连翘Forsythia suspensa基因组数据筛选鉴定bHLH转录因子家族成员并进行特征与功能分析。方法利用连翘基因组数据筛选鉴定连翘bHLH转录因子家族，分析其蛋白理化性质、系统进化关系，并对bHLH基因进行染色体定位、基因结构、功能注释、顺式作用元件分析等特征分析。此外，采用外源茉莉酸甲酯（methyl jasmonate，MeJA）处理连翘叶片，通过qRT-PCR检测FsbHLHs在处理后48 h内的表达模式，并结合连翘脂素含量变化进行相关性分析。结果筛选出170条具有完整HLH保守结构域的bHLH转录因子，分布在14条染色体上，编码氨基酸长度在67～885个aa，相对分子质量在7 910.58～98 854.78，理论等电点（pI）为4.71～10.44，系统发育分析将其划分为13个亚家族，其中第III亚家族成员最多。通过顺式作用元件分析，FsbHLHs具有多种光调控、激素响应和胁迫响应相关的元件。对筛选获得的8个FsbHLH基因在MeJA处理下的表达模式与连翘脂素含量变化进行相关性分析，结果表明FsbHLH26和FsbHLH139 2个基因极有可能在连翘脂素的合成与积累中发挥关键调控作用。结论筛选鉴定了170个FsbHLH基因家族成员，对其进行系统的生物信息学分析和MeJA处理下基因表达模式分析，筛选到FsbHLH26和FsbHLH139可能是参与到连翘脂素生物合成的关键转录因子，为探究连翘脂素生物合成的分子机制奠定基础。

关键词

连翘 / bHLH转录因子 / 连翘脂素 / 生物信息学分析 / 激素处理

Abstract

收起

Objective To screen and identify members of the bHLH transcription factor family based on genomic data from Forsythia suspensa and characterize their features and functions. Methods Using the genomic data of F. suspensa, the bHLH transcription factor family was screened and identified, analysing their physicochemical properties and phylogenetic relationships, and characterizing the bHLH genes by analysing their chromosomal locations, gene structures, functional annotations, and cis-acting elements. In addition, F. suspensa leaves were treated with exogenous methyl jasmonate (MeJA) and expression pattern of FsbHLHs within 48 h after treatment was detected by qRT-PCR. Correlation analysis was conducted in combination with the changes in phillygenin content. Results A total of 170 bHLH transcription factors with complete HLH conserved domains were identified, and these factors were distributed across 14 chromosomes. Their amino acid lengths ranged from 67 to 885 aa, relative molecular masses from 7 910.58 to 98 854.78, and theoretical isoelectric points (pI) from 4.71 to 10.44. Based on phylogenetic analysis, FsbHLHs were grouped into 13 subfamilies, with subfamily III being the largest. Through cis-acting element analysis, it was found that FsbHLHs contained multiple elements associated with light regulation, hormone response, and stress response. Subsequently, the correlation between MeJA-induced expression of eight FsbHLH genes and phillygenin content was analysed, and these results suggested that FsbHLH26 and FsbHLH139 were potentially critical regulators of phillygenin biosynthesis and accumulation. Conclusion This study identified 170 members of the FsbHLH gene family. Through systematic bioinformatics analysis and gene expression pattern analysis following MeJA treatment, FsbHLH26 and FsbHLH139 may be the key transcription factors involved in the biosynthesis of phillygenin, laying the foundation for future research into the molecular mechanisms underlying phillygenin biosynthesis.

Key words

Forsythia suspensa(Thunb.) Vahl / bHLH transcription factors / phillygenin / bioinformatics analysis / hormone treatment

引用本文

闫雪娇, 原梦, 李文文, 张力予, 许高龙, 杨崇, 吕淑芳, 马占强, 张红晓, 王婷, 侯典云. 连翘bHLH转录因子家族成员的鉴定与表达分析. 中草药, 2026 , 57 (16) : 6521 -6531 . DOI: 10.7501/j.issn.0253-2670.2026.16.024

YAN Xuejiao, YUAN Meng, LI Wenwen, ZHANG Liyu, XU Gaolong, YANG Chong, LYU Shufang, MA Zhanqiang, ZHANG Hongxiao, WANG Ting, HOU Dianyun. Identification and expression analysis of bHLH transcription factors family members in Forsythia suspensa[J]. Chinese Traditional and Herbal Drugs, 2026 , 57 (16) : 6521 -6531 . DOI: 10.7501/j.issn.0253-2670.2026.16.024

基金

收起

河南省中央引导地方科技发展资金项目
中央本级重大增减支项目“名贵中药资源可持续利用能力建设项目”
河南省科技攻关项目
河南省中药材产业科技特派员服务团项目

参考文献

收起

文献

收起

参考文献引证文献

排序方式：

陈佳茜, 郭广洋, 谭新杰, 等. 连翘多胺氧化酶基因家族的鉴定及干旱和盐胁迫下的表达分析[J]. 中草药, 2024, 55(9):3077-3084.

郭佳佳, 张雷廷, 车建芳, 等. 太行山南段连翘群落结构及其影响因子研究[J]. 生态学报, 2021, 41(21): 8589-8601.

牛群. 连翘的丰产栽培技术及其药用价值分析[J]. 农业与技术, 2016, 36(12): 101.

李文文, 闫雪娇, 许家齐, 等. 连翘FsUGTs基因家族鉴定与分析[J]. 中草药, 2026, 57(5): 1853-1863.

杨雪艳, 刘成伦. 连翘的研究现状与展望[J]. 贵州农业科学, 2012(9): 33-36.

白映伟, 胡宏. 连翘的多种利用[J]. 甘肃科技, 2012, 28(9): 147-148.

吴永娜, 薛怡萌, 李亚鹏, 等. 连翘化学成分及抗肿瘤活性研究进展[J]. 中国现代中药, 2025, 27(7): 1354-1364.

齐丽娜, 陈炫好, 金华, 等. 中药连翘化学成分及药理活性研究进展[J]. 天津中医药大学学报, 2021, 40(2): 168-175.

安昌, 陆琳, 沈梦千, 等. 植物bHLH基因家族研究进展及在药用植物中的应用前景[J]. 生物技术通报, 2023, 39(10): 1-16.

Fu N N, Wang L, Sun Q L, et al. Genome-wide identification of the bHLH transcription factor family and the regulatory roles of PbbHLH74 in response to drought stress in Phoebe bournei [J]. Int J Biol Macromol, 2024, 283: 137760.

Zuo Z F, Lee H Y, Kang H G. Basic helix-loop-helix transcription factors: Regulators for plant growth development and abiotic stress responses [J]. Int J Mol Sci, 2023, 24(2): 1419.

Sharma A, Shahzad B, Rehman A, et al. Response of phenylpropanoid pathway and the role of polyphenols in plants under abiotic stress [J]. Molecules, 2019, 24(13): 2452.

Hao Y Q, Zong X M, Ren P, et al. Basic helix-loop-helix (bHLH) transcription factors regulate a wide range of functions in Arabidopsis [J]. Int J Mol Sci, 2021, 22(13): 7152.

Arshad K T, Li C H, Li L S, et al. Genome-wide identification and expression profiling of bHLH transcription factors associated with ferulic acid biosynthesis in Angelica sinensis [J]. Front Plant Sci, 2025, 16: 1718585.

Wang X P, You Y Q, Wang J M, et al. Metabolomic and transcriptomic analyses provide insights into the role of MiMYB114 and MiMYC₂ in the MeJA-induced regulatory network for mango pericarp coloration [J]. Int J Biol Macromol, 2025, 311: 143713.

陶娟, 宋小锋, 周圆圆. MeJA对连翘叶片光合作用及次生代谢的影响[J]. 河南农业科学, 2021, 50(6): 54-60.

Xu H, Park N I, Li X H, et al. Molecular cloning and characterization of phenylalanine ammonia-lyase, cinnamate 4-hydroxylase and genes involved in flavone biosynthesis in Scutellaria baicalensis [J]. Bioresour Technol, 2010, 101(24): 9715-9722.

Shabani L, Ehsanpour A A, Asghari G, et al. Glycyrrhizin production by in vitro cultured Glycyrrhiza glabra elicited by methyl Jasmonate and salicylic acid [J]. Russ J Plant Physiol, 2009, 56(5): 621-626.

莫仁连, 洪杰生, 闫晋强, 等. 冬瓜蜡质合成相关基因BhCER1的克隆与表达分析[J]. 中国瓜菜, 2025, 38(2): 32-39.

Letunic I, Bork P. Interactive Tree of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation [J]. Nucleic Acids Res, 2021, 49(W1): W293-W296.

Gao F, Dubos C. The Arabidopsis bHLH transcription factor family [J]. Trends Plant Sci, 2024, 29(6): 668-680.

Khan A, Kanwal F, Ullah S, et al. Plant secondary metabolites: Central regulators against abiotic and biotic stresses [J]. Metabolites, 2025, 15(4): 276.

Li H W, Chen N N, Zhang H B, et al. Multidimensional regulation of transcription factors: Decoding the comprehensive signals of plant secondary metabolism [J]. Front Plant Sci, 2025, 16: 1522278.

An J P, Xu R R, Wang X N, et al. MdbHLH162 connects the gibberellin and jasmonic acid signals to regulate anthocyanin biosynthesis in apple [J]. J Integr Plant Biol, 2024, 66(2): 265-284.

Xiang L E, Jian D Q, Zhang F Y, et al. The cold-induced transcription factor bHLH112 promotes artemisinin biosynthesis indirectly via ERF1 in Artemisia annua [J]. J Exp Bot, 2019, 70(18): 4835-4848.

Xue X S, Li L, Wang D H, et al. SmJAZ1/8 inhibits the stimulation of SmbHLH59, which limits the accumulation of salvianolic acids and tanshinones in Salvia miltiorrhiza [J]. Int J Biol Macromol, 2025, 285: 138348.

Li Q, Wang G, Zhang L, et al. AcbHLH144 transcription factor negatively regulates phenolic biosynthesis to modulate pineapple internal browning [J]. Hortic Res, 2023, 10(10): uhad185.

Yu Q B, Huang C, Yang Z N. Nuclear-encoded factors associated with the chloroplast transcription machinery of higher plants [J]. Front Plant Sci, 2014, 5: 316.

Chen X Y, Yao C M, Liu J H, et al. Basic helix-loop-helix (bHLH) gene family in rye (Secale cereale L.): Genome-wide identification, phylogeny, evolutionary expansion and expression analyses [J]. BMC Genom, 2024, 25(1): 67.

Kodama Y, Sano H. Evolution of a basic helix-loop-helix protein from a transcriptional repressor to a plastid-resident regulatory factor [J]. J Biol Chem, 2006, 281(46): 35369-35380.

张诗颖, 周靓, 毛甲绪, 等. 猕猴桃bHLH基因家族鉴定及其在果实贮藏条件下的表达分析[J]. 农业生物技术学报, 2026, 34(1): 47-65.

张建军, 吴京乡, 方昊, 等. 小果白刺bHLH基因家族鉴定与表达分析[J]. 南京林业大学学报: 自然科学版, 2025, 49(6): 102-114.

Liu D Q, Zhang P, Zhou T T, et al. Genome-wide characterization and expression analysis of the bHLH gene family in response to abiotic stresses in Zingiber officinale Roscoe [J]. BMC Genom, 2025, 26(1): 143.

Dong B M, Lang S Y, Gu Y M, et al. Identification of the bHLH gene family and functional analysis of ChMYC₂ in drought stress of Cerasus humilis [J]. Plant Physiol Biochem, 2025, 220: 109519.

Zhu X Y, Chen J Y, Xie Z K, et al. Jasmonic acid promotes degreening via MYC₂/3/4- and ANAC019/055/072-mediated regulation of major chlorophyll catabolic genes [J]. Plant J, 2015, 84(3): 597-610.

Chen Q, Sun J Q, Zhai Q Z, et al. The basic helix-loop-helix transcription factor MYC₂ directly represses PLETHORA expression during jasmonate-mediated modulation of the root stem cell niche in Arabidopsis [J]. Plant Cell, 2011, 23(9): 3335-3352.

Schweizer F, Fernández-Calvo P, Zander M, et al. Arabidopsis basic helix-loop-helix transcription factors MYC₂, MYC3, and MYC4 regulate glucosinolate biosynthesis, insect performance, and feeding behavior [J]. Plant Cell, 2013, 25(8): 3117-3132.

Mai H J, Pateyron S, Bauer P. Iron homeostasis in Arabidopsis thaliana: Transcriptomic analyses reveal novel FIT-regulated genes, iron deficiency marker genes and functional gene networks [J]. BMC Plant Biol, 2016, 16(1): 211.

陈佳茜, 郭广洋, 谭新杰, 等. 连翘多胺氧化酶基因家族的鉴定及干旱和盐胁迫下的表达分析[J]. 中草药, 2024, 55(9):3077-3084.
郭佳佳, 张雷廷, 车建芳, 等. 太行山南段连翘群落结构及其影响因子研究[J]. 生态学报, 2021, 41(21): 8589-8601.
牛群. 连翘的丰产栽培技术及其药用价值分析[J]. 农业与技术, 2016, 36(12): 101.
李文文, 闫雪娇, 许家齐, 等. 连翘FsUGTs基因家族鉴定与分析[J]. 中草药, 2026, 57(5): 1853-1863.
杨雪艳, 刘成伦. 连翘的研究现状与展望[J]. 贵州农业科学, 2012(9): 33-36.
白映伟, 胡宏. 连翘的多种利用[J]. 甘肃科技, 2012, 28(9): 147-148.
吴永娜, 薛怡萌, 李亚鹏, 等. 连翘化学成分及抗肿瘤活性研究进展[J]. 中国现代中药, 2025, 27(7): 1354-1364.
齐丽娜, 陈炫好, 金华, 等. 中药连翘化学成分及药理活性研究进展[J]. 天津中医药大学学报, 2021, 40(2): 168-175.
安昌, 陆琳, 沈梦千, 等. 植物bHLH基因家族研究进展及在药用植物中的应用前景[J]. 生物技术通报, 2023, 39(10): 1-16.
Fu N N, Wang L, Sun Q L, et al. Genome-wide identification of the bHLH transcription factor family and the regulatory roles of PbbHLH74 in response to drought stress in Phoebe bournei [J]. Int J Biol Macromol, 2024, 283: 137760.
Zuo Z F, Lee H Y, Kang H G. Basic helix-loop-helix transcription factors: Regulators for plant growth development and abiotic stress responses [J]. Int J Mol Sci, 2023, 24(2): 1419.
Sharma A, Shahzad B, Rehman A, et al. Response of phenylpropanoid pathway and the role of polyphenols in plants under abiotic stress [J]. Molecules, 2019, 24(13): 2452.
Hao Y Q, Zong X M, Ren P, et al. Basic helix-loop-helix (bHLH) transcription factors regulate a wide range of functions in Arabidopsis [J]. Int J Mol Sci, 2021, 22(13): 7152.
Arshad K T, Li C H, Li L S, et al. Genome-wide identification and expression profiling of bHLH transcription factors associated with ferulic acid biosynthesis in Angelica sinensis [J]. Front Plant Sci, 2025, 16: 1718585.
Wang X P, You Y Q, Wang J M, et al. Metabolomic and transcriptomic analyses provide insights into the role of MiMYB114 and MiMYC₂ in the MeJA-induced regulatory network for mango pericarp coloration [J]. Int J Biol Macromol, 2025, 311: 143713.
陶娟, 宋小锋, 周圆圆. MeJA对连翘叶片光合作用及次生代谢的影响[J]. 河南农业科学, 2021, 50(6): 54-60.
Xu H, Park N I, Li X H, et al. Molecular cloning and characterization of phenylalanine ammonia-lyase, cinnamate 4-hydroxylase and genes involved in flavone biosynthesis in Scutellaria baicalensis [J]. Bioresour Technol, 2010, 101(24): 9715-9722.
Shabani L, Ehsanpour A A, Asghari G, et al. Glycyrrhizin production by in vitro cultured Glycyrrhiza glabra elicited by methyl Jasmonate and salicylic acid [J]. Russ J Plant Physiol, 2009, 56(5): 621-626.
莫仁连, 洪杰生, 闫晋强, 等. 冬瓜蜡质合成相关基因BhCER1的克隆与表达分析[J]. 中国瓜菜, 2025, 38(2): 32-39.
Letunic I, Bork P. Interactive Tree of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation [J]. Nucleic Acids Res, 2021, 49(W1): W293-W296.
Gao F, Dubos C. The Arabidopsis bHLH transcription factor family [J]. Trends Plant Sci, 2024, 29(6): 668-680.
Khan A, Kanwal F, Ullah S, et al. Plant secondary metabolites: Central regulators against abiotic and biotic stresses [J]. Metabolites, 2025, 15(4): 276.
Li H W, Chen N N, Zhang H B, et al. Multidimensional regulation of transcription factors: Decoding the comprehensive signals of plant secondary metabolism [J]. Front Plant Sci, 2025, 16: 1522278.
An J P, Xu R R, Wang X N, et al. MdbHLH162 connects the gibberellin and jasmonic acid signals to regulate anthocyanin biosynthesis in apple [J]. J Integr Plant Biol, 2024, 66(2): 265-284.
Xiang L E, Jian D Q, Zhang F Y, et al. The cold-induced transcription factor bHLH112 promotes artemisinin biosynthesis indirectly via ERF1 in Artemisia annua [J]. J Exp Bot, 2019, 70(18): 4835-4848.
Xue X S, Li L, Wang D H, et al. SmJAZ1/8 inhibits the stimulation of SmbHLH59, which limits the accumulation of salvianolic acids and tanshinones in Salvia miltiorrhiza [J]. Int J Biol Macromol, 2025, 285: 138348.
Li Q, Wang G, Zhang L, et al. AcbHLH144 transcription factor negatively regulates phenolic biosynthesis to modulate pineapple internal browning [J]. Hortic Res, 2023, 10(10): uhad185.
Yu Q B, Huang C, Yang Z N. Nuclear-encoded factors associated with the chloroplast transcription machinery of higher plants [J]. Front Plant Sci, 2014, 5: 316.
Chen X Y, Yao C M, Liu J H, et al. Basic helix-loop-helix (bHLH) gene family in rye (Secale cereale L.): Genome-wide identification, phylogeny, evolutionary expansion and expression analyses [J]. BMC Genom, 2024, 25(1): 67.
Kodama Y, Sano H. Evolution of a basic helix-loop-helix protein from a transcriptional repressor to a plastid-resident regulatory factor [J]. J Biol Chem, 2006, 281(46): 35369-35380.
张诗颖, 周靓, 毛甲绪, 等. 猕猴桃bHLH基因家族鉴定及其在果实贮藏条件下的表达分析[J]. 农业生物技术学报, 2026, 34(1): 47-65.
张建军, 吴京乡, 方昊, 等. 小果白刺bHLH基因家族鉴定与表达分析[J]. 南京林业大学学报: 自然科学版, 2025, 49(6): 102-114.
Liu D Q, Zhang P, Zhou T T, et al. Genome-wide characterization and expression analysis of the bHLH gene family in response to abiotic stresses in Zingiber officinale Roscoe [J]. BMC Genom, 2025, 26(1): 143.
Dong B M, Lang S Y, Gu Y M, et al. Identification of the bHLH gene family and functional analysis of ChMYC₂ in drought stress of Cerasus humilis [J]. Plant Physiol Biochem, 2025, 220: 109519.
Zhu X Y, Chen J Y, Xie Z K, et al. Jasmonic acid promotes degreening via MYC₂/3/4- and ANAC019/055/072-mediated regulation of major chlorophyll catabolic genes [J]. Plant J, 2015, 84(3): 597-610.
Chen Q, Sun J Q, Zhai Q Z, et al. The basic helix-loop-helix transcription factor MYC₂ directly represses PLETHORA expression during jasmonate-mediated modulation of the root stem cell niche in Arabidopsis [J]. Plant Cell, 2011, 23(9): 3335-3352.
Schweizer F, Fernández-Calvo P, Zander M, et al. Arabidopsis basic helix-loop-helix transcription factors MYC₂, MYC3, and MYC4 regulate glucosinolate biosynthesis, insect performance, and feeding behavior [J]. Plant Cell, 2013, 25(8): 3117-3132.
Mai H J, Pateyron S, Bauer P. Iron homeostasis in Arabidopsis thaliana: Transcriptomic analyses reveal novel FIT-regulated genes, iron deficiency marker genes and functional gene networks [J]. BMC Plant Biol, 2016, 16(1): 211.

2026年第57卷第16期

PDF下载

引用本文

BibTeX

文章信息

doi: 10.7501/j.issn.0253-2670.2026.16.024

接收时间：2026-02-03
首发时间：2026-10-22
出版时间：2026-08-28

补充材料

相关文章

文章信息

作者

出版历史

收稿日期：2026-02-03

基金

河南省中央引导地方科技发展资金项目

中央本级重大增减支项目“名贵中药资源可持续利用能力建设项目”

河南省科技攻关项目

河南省中药材产业科技特派员服务团项目

作者信息

^1.河南科技大学农学院, 河南洛阳 471023

^2.河南省药食兼用资源评价与创新利用工程研究中心, 河南洛阳 471023

^3.河南科技大学农学院, 河南洛阳 471023

^4.河南省药食兼用资源评价与创新利用工程研究中心, 河南洛阳 471023

通讯作者:

参考文献

分享链接

https://castjournals.cast.org.cn/joweb/zcy/CN/10.7501/j.issn.0253-2670.2026.16.024

分享至

全文二维码

扫描看全文

引用本文

BibTeX

本文的引用情况

2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

关闭全屏

BibTeX
EndNote
RefWorks
TxT