青稞HvERF039基因的克隆及功能研究

doi:10.3724/SP.J.1006.2025.51011

作物学报 ›› 2025, Vol. 51 ›› Issue (9): 2341-2357.doi: 10.3724/SP.J.1006.2025.51011

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

青稞HvERF039基因的克隆及功能研究

马娟娥(), 姚有华, 姚晓华, 吴昆仑^*(), 崔永梅^*()

青海大学农林科学院 / 青海大学省部共建三江源生态与高原农牧业国家重点实验室 / 青藏高原种质资源研究与利用实验室 / 青海省青稞遗传育种重点实验室, 青海西宁 810016

收稿日期:2025-01-24 接受日期:2025-06-01 出版日期:2025-09-12 网络出版日期:2025-06-19
通讯作者: ^*吴昆仑, E-mail: wklqaaf@sina.com; 崔永梅, E-mail: 15251892177@163.com
作者简介:E-mail: 18393567953@163.com
基金资助:
本研究由省部共建三江源生态与高原农牧业国家重点实验室自主研究项目(2023-ZZ-01);青海大学青年科研基金项目(2022-QNY-3);青海省农林科学院创新基金项目(2022-NKY-04);青藏高原种质资源研究与利用实验室项目(2025)

Cloning and functional analysis of HvERF039 gene in Qingke (Hordeum vulgare L. var. nudum Hook. f)

MA Juan-E(), YAO You-Hua, YAO Xiao-Hua, WU Kun-Lun^*(), CUI Yong-Mei^*()

Academy of Agriculture and Forestry Sciences, Qinghai University / State Key Laboratory of Plateau Ecology and Agriculture, Qinghai University / Laboratory for Research and Utilization of Qinghai Tibet Plateau Germplasm Resources / Qinghai Key Laboratory of Hulless Barley Genetics and Breeding, Xining 810016, Qinghai, China

Received:2025-01-24 Accepted:2025-06-01 Published:2025-09-12 Published online:2025-06-19
Contact: ^*E-mail: wklqaaf@sina.com; E-mail: 15251892177@163.com
Supported by:
Open Project of State Key Laboratory of Plateau Ecology and Agriculture(2023-ZZ-01);Qinghai University Natural Science Foundation for Young Scholars(2022-QNY-3);Innovation Fund of Qinghai Academy of Agricultural and Forestry Sciences(2022-NKY-04);Laboratory for Research and Utilization of Qinghai Tibet Plateau Germplasm Resources(2025)

摘要/Abstract

摘要：

AP2/ERF (APETALA2/ethylene responsive factor)转录因子在植物非生物胁迫响应中起重要作用, 但青稞中该类基因的功能研究较少。本研究克隆了青稞HvERF039基因, 并通过生物信息学、qRT-PCR、拟南芥异源过表达等方法研究了其在低温胁迫中的功能。生物信息学分析结果显示, HvERF039是具有典型AP2保守结构域的亲水性不稳定蛋白, 其启动子区域含有与光响应、植物激素响应、低温胁迫响应等相关的顺式作用元件。亚细胞定位和转录自激活活性试验表明, HvERF039是一个定位于质膜和细胞核上、具有转录激活活性的转录因子。qRT-PCR分析表明, HvERF039在低温胁迫下显著上调表达, 其在不同组织中均有表达, 且在叶中的表达量最高。低温胁迫下的功能验证结果显示, 拟南芥HvERF039过表达株系的萌发率和存活率显著高于野生型, 且抗逆生理指标离子渗漏率、H₂O₂含量、MDA含量和CAT活性优于野生型。此外, HvERF039与多个逆境胁迫相关蛋白之间存在物理互作关系。本研究表明, HvERF039在青稞低温胁迫响应中具有正调控作用, 为青稞耐低温品种改良提供了新的基因资源。

关键词: 青稞, HvERF039, 转录因子, 异源表达, 低温胁迫

Abstract:

AP2/ERF (APETALA2/ethylene responsive factor) transcription factors play crucial roles in responses to abiotic stresses, but their functions remain poorly understood in Qingke (hulless barley). In this study, we cloned the HvERF039 gene and investigated its role in cold stress response through bioinformatics analysis, qRT-PCR, and heterologous expression in Arabidopsis thaliana. HvERF039 encoded a hydrophilic, unstable protein containing a typical AP2 conserved domain. The promoter region of HvERF039 contained various cis-acting elements, including those responsive to light, hormones and cold stress. Subcellular localization and transactivation assays revealed that HvERF039 protein was localized in both the membrane and nucleus and possessed transcriptional activation activity. qRT-PCR analysis showed that HvERF039 expression was significantly upregulated under cold treatment, with the highest expression observed in leaves and detectable levels in most other tissues. Functional analysis demonstrated that transgenic Arabidopsis plants overexpressing HvERF039 exhibited significantly higher germination and survival rates under cold stress compared to wild-type plants. These transgenic lines also showed enhanced physiological resistance, including lower ion leakage, reduced H₂O₂ and MDA contents, and increased CAT activity. Furthermore, HvERF039 was found to physically interact with multiple stress-related proteins. Collectively, these findings suggest that HvERF039 plays a positive regulatory role in cold stress tolerance and represents a promising genetic resource for improving cold resistance in Qingke cultivars.

Key words: Qingke, HvERF039, transcription factors, heterologous overexpression, cold stress

马娟娥, 姚有华, 姚晓华, 吴昆仑, 崔永梅. 青稞HvERF039基因的克隆及功能研究[J]. 作物学报, 2025, 51(9): 2341-2357.

MA Juan-E, YAO You-Hua, YAO Xiao-Hua, WU Kun-Lun, CUI Yong-Mei. Cloning and functional analysis of HvERF039 gene in Qingke (Hordeum vulgare L. var. nudum Hook. f)[J]. Acta Agronomica Sinica, 2025, 51(9): 2341-2357.

图/表 13

表1

本试验所用的引物"

引物名称 Primer name	引物序列 Primer sequence (5′-3′)	目的 Purpose
AT3G18780-F AT3G18780-R	CTCCTGAAGAGCACCCTG CCCTCGTAGATTGGCACA	拟南芥扩增内参引物 Internal reference primer for Arabidopsis
XM_045099138.1-F XM_045099138.1-R	GCGGCGTGAGATGAACCT CGTGGAGCAAGTCGTGTAAGA	青稞扩增内参引物 Internal reference primer for barley
ERF039-F ERF039-R	ATGGACGACATCGCCGAGCCGA TCAGTACTCCCACAGGAGTG	ERF039 CDS序列扩增引物 Primers for clone HvERF039 CDS sequence
ERF039-RT-F ERF039-RT-R	ATGGACGACATCGCCGAGC CAGATGCGCGACTTCTTGC	荧光定量扩增引物 Primers for qRT-PCR
ERF039-gateway-F ERF039-gateway-R	GGGGACAAGTTTGTACAAAAAAGCAGGCTTCATGGACGACATCGCCGAGC GGGGACCACTTTGTACAAGAAAGCTGGGTCTCAGTACTCCCACAGGAGT	ERF039拟南芥过表达和亚细胞定位载体构建引物 Primers for Arabidopsis overexpression gernation and subcellular localization assay
P2YN-ERF039-F P2YN-ERF039-R	CATTTACGAACGATAGTTAATTAATGTCTCGTGCGCACCTTCT CACTGCCACCTCCTCCACTAGTCTTGGCCATCATGACCTTGA	BiFC载体构建引物 Primers for BiFC assays
P2YC-CRK2-F P2YC-CRK2-R	CATTTACGAACGATAGTTAATTAAATGGGGCAGTGCTAC CACTGCCACCTCCTCCACTAGT ATGTCGTCTTGTGTTTG	BiFC载体构建引物 Primers for BiFC assays
P2YC-CRK3-F P2YC-CRK3-R	CATTTACGAACGATAGTTAATTAAATGGGGGGCTGCCAC CACTGCCACCTCCTCCACTAGTTCTCATCTTTGACAG	BiFC载体构建引物 Primers for BiFC assays
P2YC-CRK4-F P2YC-CRK4-R	CATTTACGAACGATAGTTAATTAAATGGGTCTCTGTCACG CACTGCCACCTCCTCCACTAGTGGCTTTCGGGATCGAC	BiFC载体构建引物 Primers for BiFC assays
P2YC-MYB15-F P2YC-MYB15-R	CATTTACGAACGATAGTTAATTAA ATGGGGAGGGCGCCGTGC CACTGCCACCTCCTCCACTAGTCTGTGGCAAGTCTAGCGCGC	BiFC载体构建引物 Primers for BiFC assays
P2YN-CAMTA4-F P2YN-CAMTA4-R	CATTTACGAACGATAGTTAATTAAATGAGCCAGAGTTTTGACA CACTGCCACCTCCTCCACTAGTGCACGTGAATTTGTCGATTT	BiFC载体构建引物 Primers for BiFC assays
P2YN-SOS3-F P2YN-SOS3-R	CATTTACGAACGATAGTTAATTAA ATGGGCTGCGTGTCGTCGT CACTGCCACCTCCTCCACTAGTTTTGCTGATTCCGCTGTAATC	BiFC载体构建引物 Primers for BiFC assays
P2YN-HKT1;5-F P2YN-HKT1;5-R	CATTTACGAACGATAGTTAATTAA ATGGGTTCTTTGCATGTC CACTGCCACCTCCTCCACTAGTCACTATCCTCCATGCCTG	BiFC载体构建引物 Primers for BiFC assays
ERF039-BD-F ERF039-BD-R	ATGGCCATGGAGGCCGAATTCATGGACGACATCGCCGAGCCGA CCGCTGCAGGTCGACGGATCCTCAGTACTCCCACAGGAGTG	转录自激活活性验证载体构建引物 Primers for transactivation activity assays

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理化性质 Physicochemical property	HvERF039	理化性质 Physicochemical property	HvERF039
分子质量 Molecular weight (Da)	22,527.19	脂溶指数 Aliphatic index	71.56
总原子数 Total number of atoms	3134	α 螺旋比例 Proportions of α helix (%)	18.87
分子式 Formula	C₉₉₃H₁₅₄₆N₂₈₀O₃₁₀S₅	延伸链比例 Proportions of extended strand (%)	7.08
亲水系数 GRAVY	-0.375	无规则卷曲比例 Proportions of random coil (%)	74.06
理论等电点 Theoretical pI	5.49	跨膜结构 Transmembrane structures	无 NO
不稳定指数 Instability index (II)	55.40	亚细胞定位预测 Prediction of subcellular localization	细胞膜, 细胞核 Cell membrane, nucleus

元件名称 Site name	序列 Sequence	功能 Function	数量 Number
GATA-motif	GATAGGA / CCATATCCAAT / GGATAAGGTG / CCACGTGGC	光响应元件 Part of a light responsive element	7
ARE	AAACCA	对厌氧诱导调节顺作用元件 Cis-acting regulatory element essential for the anaerobic induction	4
TGACG-motif	TGACG / CGTCA / CGTCA	参与MeJA反应性的顺式作用调节元件 Cis-acting regulatory element involved in the MeJA-responsiveness	6
TCA-element	CCATCTTTTT	参与水杨酸反应性的顺式作用元件 Cis-acting element involved in salicylic acid responsiveness	1
CAAT-box	CAAAT / CCCAATTT / CCAAT	启动子和增强子区域中常见的顺式作用元件 Common cis-acting element in promoter and enhancer regions	9
GC-motif	CCCCCG	参与缺氧特异性诱导的增强子样元件 Enhancer-like element involved in anoxic specific inducibility	2
LTR	CCGAAA	参与低温响应性的顺式作用元件 Cis-acting element involved in low-temperature responsiveness	1
G-Box	CACGTG / GCCACGTGGA	参与光响应的顺式作用调节元件 Cis-acting regulatory element involved in light responsiveness	8
TATA-box	ATATAT / TATAAAA	启动子核心元件 Core promoter element around -30 of transcription start	31
A-box	CCGTCC	顺式作用调节元件 Cis-acting regulatory element	2
ATCT-motif	AATCTAATCC	参与光响应的保守 DNA 模块的一部分 Part of a conserved DNA module involved in light responsiveness	1
ABRE	CACGTG / ACGTG / CACGTG	参与脱落酸反应的顺式作用元件 Cis-acting element involved in the abscisic acid responsiveness	5

青稞HvERF039基因的克隆及功能研究

Cloning and functional analysis of HvERF039 gene in Qingke (Hordeum vulgare L. var. nudum Hook. f)

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