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作物学报 ›› 2019, Vol. 45 ›› Issue (7): 1002-1016.doi: 10.3724/SP.J.1006.2019.84143

• 作物遗传育种·种质资源·分子遗传学 • 上一篇    下一篇

甘蔗脂氧合酶基因ScLOX1的克隆与表达分析

孙婷婷1,王文举1,娄文月1,刘峰1,张旭1,王玲1,陈玉凤1,阙友雄1,2,许莉萍1,2,李大妹1,2,苏亚春1,2,*()   

  1. 1 福建农林大学 / 农业部福建甘蔗生物学与遗传育种重点实验室, 福建福州 350002
    2 福建农林大学 / 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:2018-11-05 接受日期:2019-01-29 出版日期:2019-07-12 网络出版日期:2019-03-22
  • 通讯作者: 苏亚春
  • 作者简介:E-mail: sunting3221@163.com
  • 基金资助:
    本研究由国家自然科学基金项目(31501363);福建省高校杰出青年科研人才计划项目(苏亚春-2017)(SYC-2017);福建农林大学杰出青年基金(xjq201630);国家现代农业产业技术体系建设专项(CARS-17)

Cloning and expression analysis of sugarcane lipoxygenase gene ScLOX1

SUN Ting-Ting1,WANG Wen-Ju1,LOU Wen-Yue1,LIU Feng1,ZHANG Xu1,WANG Ling1,CHEN Yu-Feng1,QUE You-Xiong1,2,XU Li-Ping1,2,LI Da-Mei1,2,SU Ya-Chun1,2,*()   

  1. 1 Key Laboratory of Sugarcane Biology and Genetic Breeding (Fujian), Ministry of Agriculture / Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    2 Key Laboratory of Crop Genetics and Breeding and Comprehensive Utilization, Ministry of Education / Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China;
  • Received:2018-11-05 Accepted:2019-01-29 Published:2019-07-12 Published online:2019-03-22
  • Contact: Ya-Chun SU
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(31501363);the Research Funds for Distinguished Young Scientists in Fujian Provincial Department of Education(SYC-2017);the Research Funds for Distinguished Young Scientists in Fujian Agriculture and Forestry University(xjq201630);the China Agriculture Research System(CARS-17)

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摘要：

LOX属于脂氧合酶超家族(lipoxygenase superfamily), 是脂肪氧化途径的重要因子, 广泛参与植物生长发育的调节和对外界刺激的抵御。本研究基于甘蔗(Saccharum spp.)转录组数据库, 通过RT-PCR技术, 首次从新台糖22号(ROC22)蔗芽中克隆获得ScLOX1基因(GenBank登录号为MK106188)的cDNA全长序列。生物信息学分析显示, ScLOX1基因cDNA序列长度为2813 bp, 开放读码框全长2664 bp, 编码887个氨基酸, 其编码蛋白的理论等电点为6.23, 不稳定系数为39.77, 亲水性平均值为-0.437, 无信号肽和跨膜结构, 但含有PLAT_LH2和Lipoxygenase活性位点, 与高粱(Sorghum bicolor) LOX (XP_002466613.1)的氨基酸序列相似性高达95.96%。预测ScLOX1基因的编码蛋白为酸性稳定亲水性非分泌蛋白, 属于type I类非传统9-LOX。qPT-PCR分析结果显示, ScLOX1基因在蔗芽组织中特异性表达。接种甘蔗黑穗病菌(Sporisorium scitamineum)后, ScLOX1基因的表达量在抗病品种崖城05-179中短暂上升, 但在感病品种ROC22中显著下降。分别对瞬时表达ScLOX1基因的本氏烟(Nicotiana benthamiana)植株叶片接种烟草茄病镰刀菌蓝色变种(Fusarium solani var. coeruleum)和青枯菌(Ralstonia solanacearum), 表型观察、3,3’-二氨基联苯胺(3,3’-diaminobenzidine, DAB)染色和烟草免疫相关基因的表达情况分析显示, ScLOX1基因的过表达能够增强本氏烟对烟草茄病镰刀菌蓝色变种的防御, 但对烟草青枯菌的作用与对照相比无明显差异。研究还发现, ScLOX1基因的表达受茉莉酸甲酯和水杨酸抑制下调, 但受脱落酸、氯化钠和聚乙二醇诱导上调。以上结果为深入研究甘蔗ScLOX1基因的功能提供参考资料。

关键词: 甘蔗, 脂氧合酶, 生物信息学, 实时荧光定量PCR, 生物和非生物胁迫

Abstract:

LOX, which belongs to the lipoxygenase superfamily, is an important factor for fat oxidation and widely involved in the regulation of plant growth and development and the resistance to external stimuli. In this study, based on sugarcane (Saccharum spp.) transcriptome database, we first cloned a full-length cDNA sequence of ScLOX1 gene (GenBank accession number: MK106188) from ROC22 bud by RT-PCR. Bioinformatics analysis showed that the cDNA length of ScLOX1 gene was 2813 bp which had a 2664 bp length of open reading frame, encoding 887 amino acids. The theoretical isoelectric point, instability coefficient, and average hydrophilicity of the ScLOX1 protein were 6.23, 39.77, and -0.437, respectively. There were no signal peptide and transmembrane structure, but the PLAT_LH2 and lipoxygenase active sites in ScLOX1 protein. The similarity of amino acid sequences between ScLOX1 and Sorghum bicolor LOX (XP_002466613.1) was 95.96%. The protein encoded by ScLOX1 gene was predicted to be an acid-stable, hydrophilic, and non-secreted protein which belongs to the type I non-traditional 9-LOX. qRT-PCR results showed that ScLOX1 was specifically expressed in sugarcane bud tissue. The expression level of ScLOX1 gene was transiently increased in the smut-resistant sugarcane variety Yacheng05-179 but significantly decreased in the smut-susceptible sugarcane variety ROC22 after inoculated with Sporisorium scitamineum. When leaves of Nicotiana benthamiana were transiently overexpressed ScLOX1 gene and inoculated with tobacco pathogens Fusarium solani var. coeruleum and Ralstonia solanacearum, respectively, the results of phenotypic observation, 3,3’-diaminobenzidine (DAB) staining and expression analysis of tobacco immune-related genes revealed that the overexpression of ScLOX1 gene could enhance the defense of N. benthamiana to the F. solani var. coeruleum, but had no significant difference with the control on the defense effect against R. solanacearum. In addition, the expression level of ScLOX1 was down-regulated by methyl jasmine and salicylic acid, but up-regulated by abscisic acid, sodium chloride and polyethylene glycol. The above results provide references for further study on the function of sugarcane ScLOX1 gene.

Key words: sugarcane, lipoxygenase, bioinformatics, real-time flourescent quantitative PCR, biotic and abiotic stresses

表1

本研究所用引物序列及用途"

引物名称
Primer name		 引物序列
Primer sequence (5'-3')		 用途
Purpose
ScLOX1-cDNAF CCATCCATCCACCAACCA RT-PCR
ScLOX1-cDNAR ACGGCACAGCACAACATTTA RT-PCR
GAPDH-QF CACGGCCACTGGAAGCA qRT-PCR
GAPDH-QR TCCTCAGGGTTCCTGATGCC qRT-PCR
ScLOX1-QF ATCATTGAGGCTGTTCTT qRT-PCR
ScLOX1-QR TCTGCTATTAGTGGATTGT qRT-PCR
ScLOX1-Gate-F GGGGACAAGTTTGTACAAAAAAGCAGGCTTCATGTTCTGGCACGGGGTCGC Transient overexpression vector construction
ScLOX1-Gate-R GGGGACCACTTTGTACAAGAAAGCTGGGTCTATGGAGATGCTGTTGGGAA Transient overexpression vector construction
M13-F TGTAAAACGACGGCCAGT Transient overexpression vector construction
M13-R CAGGAAACAGCTATGACC Transient overexpression vector construction
NtHSR201-F CAGCAGTCCTTTGGCGTTGTC qRT-PCR after transient overexpression
引物名称
Primer name		 引物序列
Primer sequence (5'-3')		 用途
Purpose
NtHSR201-R GCTCAGTTTAGCCGCAGTTGTG qRT-PCR after transient overexpression
NtHSR203-F TGGCTCAACGATTACGCA qRT-PCR after transient overexpression
NtHSR203-R GCACGAAACCTGGATGG qRT-PCR after transient overexpression
NtHSR515-F TTGGGCAGAATAGATGGGTA qRT-PCR after transient overexpression
NtHSR515-R TTTGGTGAAAGTCTTGGCTC qRT-PCR after transient overexpression
NtPR-1a/c-F AACCTTTGACCTGGGACGAC qRT-PCR after transient overexpression
NtPR-1a/c-R GCACATCCAACACGAACCGA qRT-PCR after transient overexpression
NtPR2-F TGATGCCCTTTTGGATTCTATG qRT-PCR after transient overexpression
NtPR2-R AGTTCCTGCCCCGCTTT qRT-PCR after transient overexpression
NtPR3-F CAGGAGGGTATTGCTTTGTTAGG qRT-PCR after transient overexpression
NtPR3-R CGTGGGAAGATGGCTTGTTGTC qRT-PCR after transient overexpression
NtEFE26-F CGGACGCTGGTGGCATAAT qRT-PCR after transient overexpression
NtEFE26-R CAACAAGAGCTGGTGCTGGATA qRT-PCR after transient overexpression
NtAccdeaminase-F TCTGAGGTTACTGATTTGGATTGG qRT-PCR after transient overexpression
NtAccdeaminase-R TGGACATGGTGGATAGTTGCT qRT-PCR after transient overexpression
NtEF-1α-F TGCTGCTGTAACAAGATGGATGC qRT-PCR after transient overexpression
NtEF-1α-R GAGATGGGGACAAAGGGGATT qRT-PCR after transient overexpression

图1

ScLOX1基因核苷酸序列及其推导的氨基酸序列 下画线部分为克隆引物; *表示终止密码子。"

图2

ScLOX1蛋白三级结构预测"

图3

ScLOX1氨基酸序列的保守结构域"

图4

ScLOX1与其他植物LOX蛋白构建的系统发育树 ScLOX1用▲标识。玉米LOXs: ZmLOX1 (AF271894)、ZmLOX3 (AF329371)、ZmLOX6 (DQ335764)、ZmLOX10 (DQ335768)、ZmLOX11 (DQ335769); 水稻LOXs: OsLOX1 (DQ389164)、r9-LOX1 (AB099850); 番茄LOXs: TomLOXA (U09026)、TomLOXB (U09025)、TomLOXC (U37839)、TomLOXD (U37840)、TomLOXE (AY008278)、TomLOXF (FJ617476); 拟南芥LOXs: AtLOX1 (NM_104376)、AtLOX2 (AY062611)、AtLOX3 (AJ249794)、AtLOX4 (NM_105911); 烟草LOXs: NaLOX1 (X84040)、NaLOX2 (AY254348)、NaLOX3 (AY254349); 马铃薯LOXs: StLOX1 (X95513)、StLOX2 (X96405)、StLOX3 (X96406); 大豆LOXs: GmLOX9 (EU003576)、GmLOX10 (EU003577); 花生LOX: PnLOX2 (DQ068249); 豌豆LOX: LOXN2 (AJ749702); 桃LOXs: PpLOX1 (EU883638)、PpLOX2 (FJ029110)、PpLOX3 (FJ032015)、PpLOX4 (EF568783); 葡萄LOXs: VvLOXA (FJ858255)、VvLOXC (FJ858256)、VvLOXO (FJ858257); 橄榄LOXs: OeLOX (EU678670)、Oep1LOX2 (EU513352)、Oep2LOX2 (EU513353); 辣椒LOXs: CaLOX1 (FJ377708)、CaLOX2 (JQ219046); 杨树LOXs: PdLOX1 (DQ131178)、PdLOX2 (DQ131179); 杏LOXs: LOX1:Pd:1 (AJ404331)、LOX1:Pd:2 (AJ418043); 菜豆LOX: PvLOX6 (EF196866); 甘蓝LOX: BoLOX (EF123056); 鬼箭锦鸡儿LOX: CjLOX (EF530043); 茶树LOX: CasLOX1 (EU195885); 榛子LOX: CaLOX (AJ417975)。"

图5

甘蔗ScLOX1与其他植物LOX氨基酸序列比对 文本框表示底物结合结构域(Domain I和II)和C末端保守区域。箭头表示与铁结合有关的保守氨基酸残基。加号为TH(V)/R(K), 是一个对确定加氧位置特异性至关重要的基序。OsLOX1: 水稻(DQ389164); ZmLOX1: 玉米(AF271894); HvLOXC: 大麦(L37358); SbLOX4: 高粱(XP_002466613); SiLOX4: 谷子(XP_ 004982082)。"

图6

ScLOX1基因在甘蔗ROC22不同组织中的表达情况 以甘油醛-3-磷酸脱氢酶基因(glyceraldehyde-3-phosphate dehydrogenase, GAPDH)为内参基因。数据点为平均值±标准误(n = 3)。Leaf: 叶; Stem pith: 蔗肉; Bud: 蔗芽; Epidermis: 蔗皮; Root: 根。"

图7

不同基因型甘蔗与黑穗病菌互作过程中ScLOX1基因的表达量 以甘油醛-3-磷酸脱氢酶基因(glyceraldehyde-3-phosphate dehydrogenase, GAPDH)作为内参。柱上不同的字母代表在P < 0.05时显著性的差异。数据点为平均值±标准误(n = 3)。Yacheng 05-179代表甘蔗抗黑穗病品种; ROC22代表甘蔗感黑穗病品种。"

图8

ScLOX1基因在本氏烟叶片中瞬时表达 A: RT-PCR检测ScLOX1基因在接种携带35S::00和35S::ScLOX1载体的农杆菌菌株GV3101的本氏烟叶片1 d的表达情况。B: 对接种携带35S::00和35S::ScLOX1载体的农杆菌2 d的本氏烟叶片进行DAB染色。(1)和(2)分别代表索尼相机和显微镜拍摄的图像。C: 烟草免疫相关标记基因在接种携带35S::00和35S::ScLOX1载体的农杆菌1 d的本氏烟叶片中的相对表达水平。D和F: 瞬时过表达ScLOX1基因1 d后接种烟草青枯菌和茄病镰刀菌蓝色变种1 d和7 d时的本氏烟叶片发病情况和DAB染色结果。(1)和(2)分别代表索尼相机和显微镜拍摄的图像。E和G: 烟草免疫相关标记基因在瞬时过表达ScLOX1基因的本氏烟叶片接种青枯菌和茄病镰刀菌蓝色变种1 d和7 d时的表达情况分析。烟草免疫相关标记基因包括HR标记基因NtHSR201、NtHSR203和NtHSR515, SA途径相关基因NtPR-1a/c, JA途径相关基因NtPR2和NtPR3, ET合成依赖基因NtEFE26和NtAccdeaminase。内参基因为NtEF-1α基因。柱上不同的字母代表在P < 0.05时显著性的差异。所有的点均为平均值±标准误 (n = 3)。a和b分别表示载体35S::00和35S::ScLOX1。"

图9

ScLOX1基因在不同植物激素胁迫下的相对表达量 内参基因为甘油醛-3-磷酸脱氢酶基因(glyceraldehyde-3-phosphate dehydrogenase, GAPDH)。柱上不同的字母代表在P < 0.05时显著性的差异。数据点为平均值±标准误(n = 3)。"

图10

ScLOX1基因在NaCl和PEG胁迫下的表达情况 内参基因为甘油醛-3-磷酸脱氢酶基因(glyceraldehyde-3-phosphate dehydrogenase, GAPDH)。柱上不同小写字母代表在P < 0.05时显著性的差异。数据点为平均值±标准误(n = 3)。"

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