菜心减数分裂相关基因BrASY2参与植株耐热性的功能分析

doi:10.3864/j.issn.0578-1752.2025.11.012

摘要/Abstract

摘要：

【目的】菜心属于不耐热、喜冷凉的蔬菜，高温是夏秋菜心生产中面临的主要问题之一。实验室前期在菜心高温胁迫转录组数据库中挖掘到一个可能与高温胁迫响应相关的基因片段，将其命名为减数分裂相关基因BrASY2。通过进一步研究其理化性质、蛋白结构、系统进化及表达特性，并探究其在耐热性调控中的功能，以期为耐热菜心分子育种提供参考。【方法】采用全基因合成的方法获得BrASY2全长，利用ExPASy ProtParam、ProtScale、SignalP-5.0、SOPMA、Swiss-Model和PlantCARE对其理化性质、信号肽、蛋白结构、顺式作用元件进行分析，使用MEGA 6.0构建系统发育树，采用实时荧光定量PCR（qRT-PCR）技术分析其在高温胁迫下的表达特性。分别构建酵母异源超表达载体pYES2-NTB-BrASY2和基因沉默表达载体pK7GWIWG2（I）-BrASY2，获得转基因酵母和沉默表达菜心植株，并进行耐热性功能分析。采用花药压片法对BrASY2沉默表达植株细胞减数分裂染色体行为进行细胞学观察。【结果】 BrASY2长度为852 bp，编码283个氨基酸，蛋白相对分子量为69.16 kDa，理论等电点为5.09。基因结构分析表明，其含有丰富的α-螺旋和无规则卷曲。信号肽预测发现，BrASY2为无信号肽的亲水性蛋白。顺式作用元件分析显示，BrASY2含有9个高温胁迫响应元件CAAT-box。系统进化分析表明，BrASY2与白菜、甘蓝型油菜同源性最高，亲缘关系更近。转录组数据分析表明，在高温胁迫前（0 h），BrASY2在热敏自交系CX7-3和耐热自交系CX1-7中的转录表达水平均较低；在高温胁迫6 h后，BrASY2在CX1-7中的转录表达水平均迅速上升，而在CX7-3中的表达量变化不明显。qRT-PCR结果显示，随着高温胁迫处理时间的延长，耐热自交系CX1-7叶片中BrASY2的表达量呈上调趋势，在处理后24 h时表达量最高，而在热敏自交系CX7-3中始终处于低水平表达状态，表达量无明显变化。经酵母异源超表达分析，超表达酵母菌pYES2-NTB-BrASY2更耐高温，能够帮助酵母菌度过高温逆境。BrASY2沉默表达分析表明，高温胁迫后，与对照植株相比，沉默表达植株的BrHsfA2、BrHsp90和BrMn-SOD的表达均受到抑制，植株叶片颜色变浅，叶绿素含量降低，相对电导率和H₂O₂含量升高，SOD活性下降。减数分裂细胞学观察显示，与对照植株相比，BrASY2沉默植株减数分裂各时期的染色体行为均表现正常，减数分裂后期的均衡分裂细胞和四分体占比，均未表现出明显差异。【结论】成功获得菜心BrASY2，BrASY2在菜心响应高温胁迫中发挥正向调控作用。

关键词: 菜心, BrASY2, 耐热性, 基因超表达, 基因沉默

Abstract:

【Objective】 Chinese flowering cabbage (Brassica campestris L. ssp. chinensis var. utilis Tsen et Lee) was a heat-resistant and enjoy cold vegetable, high temperature is one of the main problems faced in the production of summer and autumn vegetable production. In the early stage, we discovered a gene fragment that may be closely related to the response to heat stress in the transcriptome database of Chinese flowering cabbage, named BrASY2. By studying its physicochemical properties, protein structure, systematic evolution, and expression characteristics, and exploring its function in heat tolerance regulation, this study provides a reference for molecular breeding of heat-resistant. 【Method】 The BrASY2 gene sequence was obtained by whole gene synthesis, the physicochemical properties, signal peptides, protein structure, and cis acting elements of BrASY2 were analyzed using ExPASy ProtParam, ProtScale, SignalP-5.0, SOPMA, Swiss Model, and PlantCARE, respectively. Construct phylogenetic tree using MEGA 6.0, and the expression characteristics under heat stress were analyzed using real-time fluorescence quantitative PCR (qRT-PCR) method. Construct yeast heterologous overexpression vector pYES2-NTB-BrASY2 and gene silencing vector pK7GWIWG2(I)-BrASY2, respectively, to obtain transgenic yeast and silenced expression cabbage plants, and perform heat tolerance functional analysis.Cytological observation of meiotic chromosome behavior in BrASY2 silenced plant cells using anther pressing method.【Result】BrASY2 with an ORF of 852 bp was obtained, encoding 283 amino acids, the relative molecular weight was 69.16 kDa with a theoretical isoelectric point being 5.09. The results of gene structure analysis indicate that it contains abundant alpha helices and random coi. Signal peptide prediction revealed that BrASY2 was a hydrophilic protein without signal peptides. Evolutionary analysis revealed that BrASY2 had the closest genetic relationship with Chinese cabbage and Brassica napus. Transcriptome data showed that the BrASY2 gene expression levels were low in both the heat sensitive material CX7-3 and the heat-resistant material CX1-7 before high temperature stress (0 h), and the expression level was rapidly increased in CX1-7 after 6 h of heat stress, however, there was no significant change in CX7-3. qRT-PCR results showed that the expression level of BrASY2 gene in the leaves of heat-resistant inbred line CX1-7 showed an upregulation trend with the prolongation of heat stress treatment time, the highest expression level was occurred at 24 h. However, in the thermosensitive inbred line CX7-3, the relative expression level remained in a low-level expression state with no significant change. Analysis of heterologous overexpression of yeast revealed that overexpression of yeast pYES2-NTB-BrASY2 was tolerant to high temperatures and can help the yeast overcome heat stress.The silencing results showed that the expression of BrHsfA2, BrHsp90, and BrMn-SOD in the silenced plants were inhibited after heat stress compared with the control plants, and the plant leaves color became lighter, chlorophyll content decreased, relative conductivity and H₂O₂ content increased, and SOD activity decreased. The cytological observation results of meiosis showed that, compared with the control plants, the chromosome behavior of BrASY2 silenced plants at all stages was normal, and there was no significant difference in the proportion of balanced splits cells and quaternary inmeiosis anaphase.【Conclusion】 This study successfully obtained the BrASY2 gene in Chinese flowering cabbage, and it plays a positive regulatory role in response to heat stress.

Key words: Chinese flowering cabbage, BrASY2, heat resistance, gene overexpression, gene silencing expression

庞强强, 孙晓东, 陈贻诵, 周曼, 王亚强, 施国宾. 菜心减数分裂相关基因BrASY2参与植株耐热性的功能分析[J]. 中国农业科学, 2025, 58(11): 2239-2252.

PANG QiangQiang, SUN XiaoDong, CHEN YiSong, ZHOU Man, WANG YaQiang, SHI GuoBin. Heat Resistance Analysis of Meiosis Related Gene BrASY2 in Chinese Flowering Cabbage[J]. Scientia Agricultura Sinica, 2025, 58(11): 2239-2252.

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引物名称Primer name	引物序列Primer sequence (5′-3′)	引物用途Primer purpose
BrASY2-F	GGCAGATGGGTTCAGATTCG	qRT-PCR
BrASY2-R	AAGAGGCGTGCTCATACTCA
BrHsfA2-F	GACCATTGCTCCAAGACACC
BrHsfA2-R	GGAAGAGACGGTGACCTACG
BrHsp90-F	CTGCAACAGTCTCACCCTCT
BrHsp90-R	CTGAGAGCAAGTGGTGGAAC
BrMn-SOD-F	AGATGAGTGCTGAAGGTGCT
BrMn-SOD-R	CTGATTGGCGGTTGTGTCAA
BrCKⅠ-F	GTCTAACTTCCTCGCCACCT	内参基因Reference gene
BrCKⅠ-R	GATTCGACGACAAGCCAGAC	内参基因Reference gene

材料 Material	后期ⅠAnaphaseⅠ			后期ⅡAnaphaseⅡ
材料 Material	观察细胞数 Observe the No. of cells	均衡分裂数 No. of balanced splits	比例 Proportion (%)	二分体 Binary	三分体 Ternary	四分体 Quaternary	四分体比例 Quaternary proportion (%)
BrASY2沉默植株 BrASY2 silent plants	51	48	94.12	2	0	26	92.86
对照植株 Control plant	49	46	93.88	2	0	24	92.31