冷胁迫下甘蓝型冬油菜表达蛋白及BnGSTs基因家族的鉴定与分析

doi:10.3724/SP.J.1006.2023.14207

作物学报 ›› 2023, Vol. 49 ›› Issue (1): 153-166.doi: 10.3724/SP.J.1006.2023.14207

• 作物遗传育种·种质资源·分子遗传学 • 上一篇下一篇

冷胁迫下甘蓝型冬油菜表达蛋白及BnGSTs基因家族的鉴定与分析

马骊¹(), 白静², 赵玉红³, 孙柏林³, 侯献飞⁴, 方彦¹, 王旺田¹, 蒲媛媛¹, 刘丽君¹, 徐佳¹, 陶肖蕾¹, 孙万仓¹^,^*(), 武军艳¹^,^*()

¹省部共建干旱生境作物学国家重点实验室 / 甘肃省油菜工程技术研究中心 / 甘肃农业大学农学院, 甘肃兰州 730070
²张掖市农业科学院, 甘肃张掖 734000
³甘肃亚盛农业研究院有限公司, 甘肃兰州 730030
⁴新疆农业科学院经济作物研究所, 新疆乌鲁木齐 830091

收稿日期:2021-11-04 接受日期:2022-05-05 出版日期:2023-01-12 网络出版日期:2022-05-19
通讯作者: 孙万仓,武军艳
作者简介:E-mail: mal@gsau.edu.cn
基金资助:
甘肃省青年科技计划项目(21JR7RA835);省部共建干旱生境作物学国家重点实验室(甘肃农业大学)开放基金课题(GSCS-2020-06);财政部和农业农村部国家现代农业产业技术体系建设专项(CARS-12)

Protein and physiological differences under cold stress, and identification and analysis of BnGSTs in Brassica napus L.

MA Li¹(), BAI Jing², ZHAO Yu-Hong³, SUN Bo-Lin³, HOU Xian-Fei⁴, FANG Yan¹, WANG Wang-Tian¹, PU Yuan-Yuan¹, LIU Li-Jun¹, XU Jia¹, TAO Xiao-Lei¹, SUN Wan-Cang¹^,^*(), WU Jun-Yan¹^,^*()

¹State Key Laboratory of Aridland Crop Science / Gansu Research Center of Rapeseed Engineering and Technology / College of Agronomy, Gansu Agricultural University, Lanzhou 730070, Gansu, China
²Zhangye Academy of Agricultural Sciences, Zhangye 734000, Gansu, China
³Gansu Yasheng Agricultural Research Institute Co. Ltd, Lanzhou 730030, Gansu, China
⁴Institute of Economic Crops, Xinjiang Academy of Agricultural Sciences, Urumqi 830091, Xinjiang, China

Received:2021-11-04 Accepted:2022-05-05 Published:2023-01-12 Published online:2022-05-19
Contact: SUN Wan-Cang,WU Jun-Yan
Supported by:
Gansu Provincial Youth Science & Technology Department(21JR7RA835);Research Program Sponsored by State Key Laboratory of Aridland Crop Science (Gansu Agricultural University)(GSCS-2020-06);China Agriculture Research System of MOF and MARA(CARS-12)

摘要/Abstract

摘要：

谷胱甘肽S-转移酶(glutathione S-transferases, GST)参与调节植物生长、发育和逆境胁迫反应的许多方面。本研究利用双向电泳和质谱技术分析‘16VHNTS309’在冷胁迫下差异表达的蛋白质, 基于GO和KEGG分析鉴定出BE、APX、SOD、GST等参与冷胁迫反应的蛋白质。利用qRT-PCR和生理指标鉴定到响应冷胁迫的关键蛋白质GST, 采用同源克隆法克隆到‘16VHNTS309’的GST基因。该基因CDS长度为642 bp, 编码213个氨基酸, 是一个不稳定蛋白, 属于GST_N_3谷胱甘肽S-转移酶家族, 与甘蓝型油菜‘ZS11’序列相似性为99.22%, 与‘ZS11’和‘Vision’两者的氨基酸序列相比较发现第127位亮氨酸(L)突变为脯氨酸(P)。基因家族分析表明, 在甘蓝型油菜中共鉴定到153个BnGSTs成员, 按其功能主要分为Zeta、Phi、Theta、CHQ、DHAR、Lambda和Tau这七大类型, 大部分的BnGSTs属于Phi和Tau这2种类型。系统进化将BnGSTs分为12个亚家族, 亚家族I和VIII包含了较多的成员, BnGSTs不均匀的分布在18条染色体上, C06染色体上分布BnGSTs基因数量最多, 含有10个保守的蛋白质基序。BnGSTs基因家族中有99对基因存在共线性关系, 131个基因来自基因复制事件, 片段重复事件在BnGSTs基因的进化中起着重要作用。冷胁迫下BnaA02g35760D、BnaC06g20450D、BnaC06g35490D、BnaA02g03230D和BnaA02g35980D在强抗寒品种中的显著高表达, 是弱抗寒品种的7~12倍, 并且强抗寒品种具有较高的生理酶活性。另外, 在冷冻胁迫下鉴定到一些瞬时和持续表达的关键候选基因。这些结果为进一步研究BnGSTs基因在强抗寒甘蓝型冬油菜抗寒分子调控中的作用奠定基础。

关键词: 甘蓝型冬油菜, 冷胁迫, 蛋白质谱, GST基因家族, 表达模式

Abstract:

Glutathione S-transferases (GST) are involved in regulating many aspects of plant growth, development, and response to adversity stress. In this study, the differentially expressed proteins of ‘16VHNTS309’ under cold stress were analyzed using bidirectional electrophoresis and mass spectrometry, and proteins involved in the cold stress response, such as BE, APX, SOD, and GST, were identified based on GO and KEGG analyses. The key protein GST was identified using qRT-PCR and physiological indicators in response to cold stress, and the GST gene of ‘16VHNTS309’ was cloned using homologous cloning. This gene CDS length was 642 bp, and encoded 213 amino acids, which was an unstable protein and belonged to the GST_N_3 glutathione S-transferase family. The sequence similarity with Brassica napus ‘ZS11’ was 99.22%, and a comparison of the amino acid sequences of both ‘ZS11’ and ‘Vision’ revealed a mutation from leucine (L) to proline (P) at position 127. The gene family analysis showed that 153 BnGSTs members were identified in Brassica napus and were classified into seven main types according to their functions: Zeta, Phi, Theta, CHQ, DHAR, Lambda, and Tau. Most BnGSTs belonged to Phi and Tau types. Phylogeny divided BnGSTs into 12 subfamilies, subfamilies I and VIII contained more members. BnGSTs were unevenly distributed on 18 chromosomes, and the number of BNGSTs genes on C06 chromosome was the largest, it contained 10 conserved protein motifs. There were 99 pairs of genes in the BnGSTs gene family that were colinearly related, 131 genes from gene duplication events, and segmental duplication events played an important role in the evolution of BnGSTs genes. Significantly higher expression of BnaA02g35760D, BnaC06g20450D, BnaC06g35490D, BnaA02g03230D, and BnaA02g35980D in strong cold-resistant varieties was 7-12 times higher than that in weak cold-resistant varieties under cold stress. And the strong cold-resistant varieties had higher physiological enzyme activities. In addition, a number of key candidate genes were identified for transient and sustained expression under freezing stress. This result lays the foundation for further studies on the molecular regulation of BnGSTs genes for cold resistance in strong cold resistant for Brassica napus.

Key words: winter Brassica napus L., cold stress, protein mass spectrometry, GST gene family, expression pattern

马骊, 白静, 赵玉红, 孙柏林, 侯献飞, 方彦, 王旺田, 蒲媛媛, 刘丽君, 徐佳, 陶肖蕾, 孙万仓, 武军艳. 冷胁迫下甘蓝型冬油菜表达蛋白及BnGSTs基因家族的鉴定与分析[J]. 作物学报, 2023, 49(1): 153-166.

MA Li, BAI Jing, ZHAO Yu-Hong, SUN Bo-Lin, HOU Xian-Fei, FANG Yan, WANG Wang-Tian, PU Yuan-Yuan, LIU Li-Jun, XU Jia, TAO Xiao-Lei, SUN Wan-Cang, WU Jun-Yan. Protein and physiological differences under cold stress, and identification and analysis of BnGSTs in Brassica napus L.[J]. Acta Agronomica Sinica, 2023, 49(1): 153-166.

图/表 13

表1

图1

附图1

附表1

甘蓝型冬油菜响应冷胁迫的差异蛋白质鉴定"

蛋白点编号 Spot number	基因登录号 Accession number	蛋白质名称 Protein name	分子质量 Molecular weight (kD)	等电点 pI	亚细胞定位 Cellular location	差异倍数 Fold change
2	XP_013688985.1	RNA-binding protein CP29B	31,076	4.82	叶绿体Chloroplast	NDT
3	XP_013656707.1	glycine-rich RNA-binding protein 10-like	16,038	5.24	微体Microbody	1.87±0.06
4	XP_013656707.1	glycine-rich RNA-binding protein 10-like	16,038	5.24	微体Microbody	0.49±0.01
6	XP_013687602.1	bifunctional enolase 2/transcriptional activator	47,556	5.56	内质网 Endoplasmic reticulum	1.99±0.09
8	XP_013713332.1	oxygen-evolving enhancer protein 1-1	35,164	5.55	叶绿体Chloroplast	2.54±0.03
9	XP_013688985.1	RNA-binding protein CP29B	31,076	4.82	叶绿体Chloroplast	NDT
11	XP_013649537.1	cysteine protease inhibitor WSCP	23,891	8.66	细胞外Extracellular	1.9±0.04
12	XP_013673398.1	ATP synthase subunit beta-2	53,800	5.39	细胞质Cytoplasm	2.52±0.06
13	XP_013659948.1	cysteine protease inhibitor WSCP-like	24,956	6.52	细胞外Extracellular	NDC
14	XP_013678556.1	actin-7	41,665	5.31	细胞质Cytoplasm	2.01±0.11
15	XP_013641139.1	L-ascorbate peroxidase 1, cytosolic-like isoform X3	27,594	5.58	微体Microbody	2.63±0.06
16	XP_013697709.1	probable phospholipid hydroperoxide glutathione peroxidase 6	25,260	9.2	线粒体Mitochondrial	2.08±0.18
17	AIF75338.1	chloroplast ribulose 1,5-bisphosphate carboxylase/oxygenase small subunit	19,852	7.59	细胞外Extracellular	2.21±0.05
18	XP_013720130.1	PREDICTED: superoxide dismutase [Cu-Zn] 2	21,343	6.79	细胞外Extracellular	2.75±0.11
19	XP_013649537.1	cysteine protease inhibitor WSCP	23,891	8.66	细胞外Extracellular	3.01±0.05
20	XP_013649391.1	nucleoside diphosphate kinase 1	16,384	6.29	微体Microbody	1.94±0.04
21	XP_013712048.1	CBS domain-containing protein CBSX3	22,650	8.71	线粒体Mitochondrial	NDC
23	AIF75338.1	chloroplast ribulose 1,5-bisphosphate carboxylase/oxygenase small subunit	19,852	7.59	细胞外Extracellular	3.33±0.39
24	XP_013649614.1	kunitz trypsin inhibitor 1-like	23,553	6.6	细胞外Extracellular	NDC
25	XP_013641139.1	L-ascorbate peroxidase 1, cytosolic-like isoform X3	27,594	5.58	微体Microbody	3.41±0.22
26	AIF75338.1	chloroplast ribulose 1,5-bisphosphate carboxylase/oxygenase small subunit	19,852	7.59	细胞外Extracellular	3.11±0.29
27	XP_013641880.1	glutathione S-transferase DHAR1, mitochondrial	23,533	5.76	细胞质Cytoplasm	2.1±0.1
28	XP_013666490.1	malate dehydrogenase 1, cytoplasmic-like	35,695	6.11	细胞外Extracellular	3.12±0.06
29	XP_013666490.1	malate dehydrogenase 1, cytoplasmic-like	35,695	6.11	细胞外Extracellular	2.91±0.21
30	XP_013649614.1	kunitz trypsin inhibitor 1-like	23,553	6.6	细胞外Extracellular	NDC

附表1

图2

图3

图4

附图2

图5

图6

图7

图8

图9

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基因名称 Gene name	引物序列 Primer sequence (5'-3')	产物大小 Product size (bp)
BE	F: GATGGAACCCAGAACGAG; R: GGAATGCCACTGACAACG	110
SPI	F: CCCCTCTTTATCGGACAG; R: GGAACGAACCCAACTCTA	89
APX	F: GAGCGGTGAGAAGGAAGG; R: CTCGTCAGCAGCGTATTT	100
SOD	F: GCCACATTTCAACCCTAACA; R: ACCACAAAGGCTCTTCCAAC	174
GST	F: ATCAACATCTCCGACAAACC; R: CGTCAGAATCAGACACCCAC	100
RBP	F: CGTGTTTATGTCGGCAATC; R: GCCACTGTCCCTGTCGTAG	114
β-Actin	F: TGGGTTTGCTGGTGACGAT; R: TGCCTAGGACGACCAACAATACT	63

冷胁迫下甘蓝型冬油菜表达蛋白及BnGSTs基因家族的鉴定与分析

Protein and physiological differences under cold stress, and identification and analysis of BnGSTs in Brassica napus L.

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