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作物学报 ›› 2021, Vol. 47 ›› Issue (1): 94-103.doi: 10.3724/SP.J.1006.2021.04156

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蔗ScCRT1基因克隆及其应答SCMV侵染分子机制的研究

张海(), 程光远, 杨宗桃, 王彤, 刘淑娴, 商贺阳, 赵贺, 徐景升*()   

  1. 福建农林大学国家甘蔗工程技术研究中心 / 农业农村部福建甘蔗生物学与遗传育种重点实验室 / 教育部作物遗传育种与综合利用重点实验室, 福建福州 350002
  • 收稿日期:2020-07-14 接受日期:2020-09-13 出版日期:2021-01-12 网络出版日期:2020-09-22
  • 通讯作者: 徐景升
  • 作者简介:E-mail: zhanghai940410@163.com
  • 基金资助:
    国家自然科学基金项目(31971991);福建农林大学科技创新基金项目(CXZX2018026)

Cloning of sugarcane ScCRT1 gene and its response to SCMV infection

ZHANG Hai(), CHENG Guang-Yuan, YANG Zong-Tao, WANG Tong, LIU Shu-Xian, SHANG He-Yang, ZHAO He, XU Jing-Sheng*()   

  1. National Engineering Research Center for Sugarcane / Key Laboratory of Sugarcane Biology and Genetic Breeding, Ministry of Agriculture and Rural Affairs / Key Laboratory of Ministry of Education for Genetics, Breeding and Multiple Utilization of Crops, Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
  • Received:2020-07-14 Accepted:2020-09-13 Published:2021-01-12 Published online:2020-09-22
  • Contact: XU Jing-Sheng
  • Supported by:
    National Natural Science Foundation of China(31971991);Science and Technology Innovation Project of Fujian Agriculture and Forestry University(CXZX2018026)

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摘要：

钙网蛋白(calreticulin, CRT)在真核生物中广泛表达, 是重要的分子伴侣和钙离子结合蛋白, 参与调控Ca2+稳态、钙依赖信号、内质网质量控制、植物生长发育、免疫反应和逆境应答等多种生物学过程。甘蔗(Saccharum spp. hybrid)中CRT应答甘蔗花叶病毒(Sugarcane mosaic virus, SCMV)侵染尚未见报道。本研究从热带种Badila (S. officinarum)中克隆了1个CRT1/CRT2亚型的CRT编码基因, 命名为ScCRT1。该基因开放读码框(open reading frame, ORF)长度为1281 bp, 编码长度为426 aa的蛋白。生物信息学分析表明, ScCRT1具有典型的CRT蛋白结构域, 为稳定的亲水性蛋白, 其N端有一个信号肽, 具有典型的跨膜结构域, C端有典型的内质网定位信号; 二级结构多为无规则卷曲; 系统进化树分析表明, 该蛋白是典型的CRT蛋白, 在单子叶和双子叶植物中具有明显的分化。亚细胞定位表明ScCRT1定位于内质网。实时荧光定量PCR分析发现, ScCRT1基因在甘蔗各组织中都有表达, 在第8节间中的表达量最低, 在心叶中的表达量较高; 该基因在SCMV侵染早期表达量上调, 后期下调表达。酵母双杂交(yeast two hybrid, Y2H)和双分子荧光互补(bimolecular fluorescence complementation, BiFC)试验表明, ScCRT1与SCMV-6K2蛋白互作。推测SCMV-6K2通过与ScCRT1互作调控钙离子稳态进而便于SCMV侵染。

关键词: 甘蔗, 钙网蛋白, SCMV, 6K2

Abstract:

Calreticulin (CRT) is widely expressed in eukaryotes. As a molecular chaperone and a Ca2+ binding protein, CRT is involved in many biological pathways such as the regulation of calcium homeostasis, calcium-dependent signaling, endoplasmic reticulum quality control, plant growth and development, immunity and response to stress. However, the response of CRT of sugarcane (Saccharum spp. hybrid) challenged by Sugarcane mosaic virus (SCMV) has not been reported. In this study, a CRT gene was cloned from the noble cane cultivar Badila (S. officinarum) and designed as ScCRT1. ScCRT1 had an open reading frame (ORF) length of 1281 bp and encoded 426 amino acids. Bioinformatics analysis showed that ScCRT1 was a stable hydrophilic protein and possesses a signal peptide at the N-terminal, a typical transmembrane domain, and a typical endoplasmic reticulum location signal at the C-terminal. The secondary structure of ScCRT1 was composed of mostly random coils. Phylogenetic tree analysis indicated that ScCRT1 belonged to the CRT1/CRT2 subtype and was divergent between monocotyledons and dicotyledons. Subcellular location assays showed that ScCRT1 was mainly located in the endoplasmic reticulum. Real-time quantitative PCR analysis showed that ScCRT1 gene was extensively expressed in different tissues of sugarcane, with the highest expression in leaf roll and the lowest expression in the 8th internode. ScCRT1 gene was up regulated in the early stage of SCMV infection, but down regulated with time going. ScCRT1 interacted with the 6K2 from SCMV as confirmed by yeast two hybrid and bimolecular fluorescence complementation assays. Based on these foundlings, we speculated SCMV interfered the calcium homeostasis by the interaction of 6K2 with ScCRT1, thereby facilitating viral infection of sugarcane.

Key words: sugarcane, SCMV, calreticulin, 6K2

表1

本研究使用的引物"

引物名称
Primer name		 引物序列
Primer sequence (5′-3′)		 用途
Strategy
ScCRT1-F ATGGCGATCCTCGAGAGG 基因克隆
ScCRT1-R CTAGAGCTCATCATGTTTA Gene cloning
BD-ScCRT1-F ATTAACAAGGCCATTACGGCCATGGCGATCCTCGAGAGGTC 酵母双杂交诱饵载体构建
BD-ScCRT1-R AACTGATTGGCCGAGGCGGCCCCGAGCTCATCATGTTTAGCATC Vector generation for Y2H
221-ScCRT1-F GGGGACAAGTTTGTACAAAAAAGCAGGCTTCATGGCGATCCTCGAGAGGTC 亚细胞定位
221-ScCRT1-R GGGGACCACTTTGTACAAGAAAGCTGGGTCGAGCTCATCATGTTTAGCAT Subcellular localization
ScCRT1-qF CGCCAAGAAGTTAGCAGAGGAGAC 定量PCR
ScCRT1-qR CCTTGTCATCGTCCGCATCATCC Real-time-qPCR
GAPDH-F CACGGCCACTGGAAGCA 内参基因
GAPDH-R TCCTCAG GGTTCCTGATGCC Reference gene
eEF-1α-F TTTCACACTTGGAGTGAAGCAGAT 内参基因
eEF-1α-R GACTTCCTTCACAATCTCATCATAA Reference gene
SCMV-CP-F TACAGAGAGACACACAGCTG SCMV检测
SCMV-CP-R ACGCTACACCAGAAGACACT Detection of SCMV

图1

甘蔗钙网蛋白(ScCRT1)氨基酸序列比对与结构域 高粱: SbCRT1 (XP_021307591.1); 玉米: ZmCRT1 (XP_008670080.1); 谷子: SiCRT1 (XP_004981159.1); 二穗短柄草: BdCRT1 (XP_003563166.1); 水稻: OsCRT1 (XP_015628738.1); 拟南芥: AtCRT1 (NP_176030.1)。黑色背景中的蛋白残基为相同的部分。箭头表示3个结构域(N, P和C)的起始位置。预测的信号肽、保守的CRT家族基序(CRT motif 1: KLDCGGGYVKLL; CRT motif 2: IMFGPDICG)、3个重复的残基(A: PXXIXDPXX KKPEXWDD; B: GXWXAXXIXNPXYK, X为任意氨基酸)[1]和内质网滞留信号HDEL用黑色下划线标记。假定的核靶向序列(PPKKIKDPE)用红色下画线标记。"

图2

ScCRT1与其他物种CRT 蛋白的系统进化树分析"

图3

ScCRT1-YFP在本氏烟表皮细胞中的定位 A: ScCRT1-YFP荧光信号; B: 明场; C: 合并。标尺为 25 μm。"

图4

ScCRT1基因在甘蔗不同组织中的表达模式 误差线为每组处理的标准误差(n = 3)。Root: 根; Leaf roll: 心叶; 1st leaf: 正一叶; 4th leaf: 正四叶; 4th internode: 第四节间; 8th internode: 第八节间。柱上不同的小写字母表示在P < 0.05时显著性的差异。"

图5

ScCRT1基因应答SCMV侵染的表达模式 误差线为每组处理的标准误差(n = 3)。柱上不同的小写字母表示在P < 0.05时显著性的差异。"

图6

Y2H检测ScCRT1与SCMV-6K2的互作 pNubG-Fe65和pTSU2-APP组合作为阳性对照, pNubG-Fe65和pPR3-N组合作为阴性对照。DDO+X-Gal: 添加了5-溴-4-氯-3-吲哚-β-D-半乳糖苷的缺少亮氨酸(Leu)和色氨酸(Trp)的酵母合成限定基本培养基; QDO+X-Gal: 添加了X-Gal的缺少亮氨酸(Leu)、色氨酸(Trp)、组氨酸(His)和腺嘌呤(Ade)的酵母合成限定基本培养基。"

图7

BiFC检测ScCRT1与SCMV-6K2的互作 ScCRT1融合于YFP的C末端, SCMV-6K2融合于YFP的N末端, 在本氏烟叶片中瞬时表达, 48 h后激光共聚焦观察(标尺为25 μm)。"

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