CN114149984A - 一种番茄抗逆基因及其编码的蛋白质和应用 - Google Patents
一种番茄抗逆基因及其编码的蛋白质和应用 Download PDFInfo
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Abstract
本发明公开了一种番茄抗逆基因及其编码的蛋白质和应用，其在低温、盐和干旱三种胁迫处理中，抵抗干旱能力的表型最为明显，抗盐次之，抵抗低温能力也比较明显。过表达SlCIPK8转基因植株可以通过提高抗氧化酶的活性提高植株的耐低温、抗盐性和抗旱性能力，因此本发明对于番茄的遗传改良及抗性品种选育具有重要的意义，具有很强的应用价值。
Description
技术领域
本发明属于植物育种技术领域，特别涉及一种抗逆基因及其编码的蛋白质和应用，尤其涉及一种番茄抗逆基因及其编码的蛋白质和应用。
背景技术
番茄(Solanum lycopersicum)即西红柿，是管状花目、茄科、番茄属的一种一年生或多年生草本植物，是世界性的经济作物，在全球的蔬菜作物栽培中有着重要的地位。中国作为世界上番茄种植面积最大，产量最高的国家，在番茄种植过程中会受到各种逆境胁迫的影响，如：低温、盐、干旱等。这些非生物胁迫的影响会导致番茄受到伤害，严重的情况下甚至会导致番茄死亡，影响番茄作物的产量和品质，从而造成严重的经济损失。因此，提高番茄抗低温、盐、干旱等非生物胁迫具有十分重要的意义。
钙离子是广泛存在的第二信使，在细胞内信号转导途径当中有着非常重要的作用。钙信号通过几种钙离子调节的蛋白质传感器来感知，类钙调磷酸酶B(CBL)蛋白作为其中一种蛋白质传感器通过与CBL结合蛋白激酶(CIPKs)的协同作用参与各种应激反应。CBLs-CIPKs信号通路在不同胁迫下表现出多样性，受细胞内复杂机制的调控，已经发现其在响应非生物胁迫中有着重要的作用。目前已经在多种植物中鉴定出了CIPK基因，例如，在拟南芥的CIPK基因家族被鉴定出来有25个成员，辣椒的CIPK基因家族被鉴定出来有21个成员，茄子的CIPK基因家族被鉴定出来有15个成员。虽然已经在番茄中鉴定出22个CIPK基因家族成员，但在番茄中还没有CIPK8在低温、盐和干旱中行使的功能的作用鉴定及其在实际育种中的应用。
发明内容
本发明的目的是提供一种番茄抗逆基因及其编码的蛋白质和应用，其在低温、盐和干旱三种胁迫处理中，抵抗干旱能力的表型最为明显，抗盐次之，抵抗低温能力也比较明显。
为此，本发明技术方案如下：
第一方面，本发明提供一种蛋白质，所述蛋白质由序列表中序列1所示的氨基酸序列组成。序列1(SEQ ID No.1)：
MSSSTSTSSPFSRSRTRLGKYELGKTLGEGSFAKVKYAKNIQTGENVAIKIINRDRVLRQNIMEQIKREISTMKLIRHPNVVRIFEVMASKTKIYIVLEYVHGGELFDEIARHGRLKEDEARRYFQQLINAVDYCHSRGVFHRDLKPENLLLDSFGILKVSDFGLSALSKQVRDDGLLHTACGTPNYVAPEVLTDKGYDGTTTDVWSCGVILFVLMAGYLPFDEPNLNALYRKILKATFSLPPWLSSSSKNLINRILDPDPLTRITIPEILEDEWFKKDYKPPPFEQDEDVNLDDIDAIFNGSDDHLVTERKEKPASVNAFELISRSKSFNLENLFEKQALVKRETQFTSRSPANEIISKIEETARPLGFSVQKKNYKMKLQGDRTGRKGHLAVATEVFEVAPSVHLVELRKTGGDTLEFHKFYKNFSSGLKDIVWTTEQTTEQPSEEKGS*
第二方面，本发明提供编码如第一方面所述蛋白质的基因。
优选的，所述基因为序列表中序列2所示的DNA分子。序列2(SEQ ID No.2)：ATCACATTCGGTTTTCGAGGATATCTACAAGAACTCATACGATAATGAGTAGCTCAACGTCAACGTCATCGCCATTTAGTCGTAGCAGGACACGTTTAGGGAAATATGAATTGGGTAAGACATTAGGTGAGGGAAGTTTTGCTAAGGTAAAGTATGCCAAAAATATACAAACAGGTGAAAATGTAGCCATCAAAATCATCAATCGTGATCGCGTCCTACGACAAAACATTATGGAACAGATTAAAAGAGAAATATCAACGATGAAGTTGATCAGACATCCAAATGTCGTGAGGATCTTTGAGGTTATGGCTAGTAAGACGAAGATCTATATTGTCCTCGAGTATGTACATGGAGGAGAGCTCTTTGATGAAATTGCTAGACATGGAAGACTCAAAGAAGACGAAGCTAGAAGATACTTTCAACAGCTTATTAATGCCGTTGATTACTGTCATAGTAGAGGTGTTTTCCATCGAGACTTGAAGCCTGAGAATCTATTGCTGGATTCATTTGGTATTCTGAAAGTTTCAGACTTCGGATTGAGTGCACTTTCCAAGCAAGTTCGAGACGATGGACTGCTTCATACTGCTTGTGGCACACCAAACTATGTCGCGCCTGAGGTGCTAACTGACAAAGGCTATGATGGTACAACAACTGATGTCTGGTCCTGTGGAGTCATTCTCTTTGTTCTAATGGCTGGATACTTACCTTTTGATGAGCCAAACCTAAATGCTCTATACCGAAAAATCCTGAAGGCTACGTTTTCACTTCCACCATGGTTGTCCTCCAGCTCAAAGAATCTGATCAACCGTATTCTTGACCCGGATCCACTCACAAGGATCACTATTCCAGAGATCCTCGAAGATGAGTGGTTCAAGAAAGATTACAAGCCGCCTCCTTTTGAGCAAGACGAAGATGTCAATCTTGATGACATTGATGCCATATTCAATGGCTCGGACGATCATCTCGTTACAGAAAGAAAGGAAAAACCTGCTTCTGTCAATGCATTCGAGCTCATTTCAAGGTCAAAAAGTTTCAACCTAGAAAATTTGTTCGAAAAACAGGCCCTTGTCAAGAGAGAAACACAATTTACTTCGCGGTCGCCTGCTAATGAGATTATCTCCAAAATTGAGGAAACTGCAAGACCATTGGGATTCAGTGTCCAGAAGAAAAATTACAAGATGAAGTTACAAGGCGACAGGACCGGAAGGAAAGGCCACCTTGCTGTAGCAACAGAGGTGTTTGAAGTAGCACCCTCAGTGCATTTGGTTGAGCTCCGAAAAACTGGTGGTGATACATTAGAGTTTCATAAGTTTTACAAGAACTTCTCTTCAGGATTAAAAGATATCGTCTGGACAACAGAACAAACAACCGAACAACCCTCTGAAGAAAAAGGGTCTTAA
第三方面，本发明提供含有如第二方面所述基因的重组表达载体、表达盒或重组菌。
第四方面，本发明提供如第二方面所述基因的应用。
优选的，所述基因的应用至少包括以下步骤：将如第二方面所述的基因导入目的番茄，使目的番茄加强对于干旱的抵抗能力。
在本发明中，测量了在SlCIPK8转基因植株中的三个胁迫相关响应基因SlAREB、SlDREB2A和SlRD29A，干旱胁迫条件下的表达量与先前研究中已被证明增强干旱胁迫调控的基因出现了一致的结果，比野生型植物更高的表达量。盐和干旱胁迫均会引起ABA(脱落酸)水平的升高，ABA水平也会部分地介导盐和干旱胁迫反应，包括基因诱导。因此推测SlCIPK8是一个积极调控干旱胁迫的基因，调控方式不是只通过ABA依赖途径，做出反应的方式也通过ABA非依赖两种途径来进行，两种方式共同作用来调控番茄在干旱胁迫下生长和发育。
优选的，所述基因的应用至少包括以下步骤：将如第二方面所述的基因导入目的番茄，使目的番茄加强对于高盐的抵抗能力。
为了探究SlCIPK8耐盐性增强可能的分子机制，SlAEEB、SlDREB2A、SlRD29A作为三个与盐胁迫相关的基因，在盐胁迫下测量了这三个基因在SlCIPK8转基因番茄植株和野生植株的表达量，测量后发现在SlCIPK8转基因植株中发现这三个胁迫响应基因SlAREB、SlDREB2A和SlRD29，在盐胁迫条件下表现出比野生型番茄更高的表达量。因此，本发明说明SlCIPK8是一个对盐胁迫调控有正向意义的基因，并且不仅通过ABA依赖来做出反应，还通过ABA非依赖两种途径对盐胁迫作出反应。实验初步验证了SlCIPK8基因在番茄盐胁迫耐响应ABA调控相关。
优选的，所述基因的应用至少包括以下步骤：将如第二方面所述的基因导入目的番茄，使目的番茄加强对于低温的抵抗能力。
植物的冷应激反应是由一个复杂的调控网络控制的，为了探索SlCIPK8是如何增强抗寒性的，测定了在抗寒性中的作用调控几个重要抗寒基因的表达SlCBF1、SlCBF2、SlDREB2A和SlRD29A，根据结果推测SlCIPK8可能参与了一种信号转导级联反应，其中应激信号首先被传递到激活CBFs/DREBs表达的因子，这些因子又与RD29A基因中的CRE/DREs结合，从而激活它们的转录。冷胁迫处理后SlCIPK8转基因番茄株的SlDREB2A显著高于野生型植株，SlDREB2A与启动子中的DRE/CRT元件结合，以非ABA依赖的方式调节基因表达，因此推测SlCIPK8调控了一条不依赖ABA的冷胁迫途径。
优选的，所述基因的应用至少包括以下步骤：将如第二方面所述的基因导入目的番茄，使目的番茄加强对于干旱、高盐和低温的抵抗能力。
第五方面，本发明提供一种番茄育种方法，将第四方面所述的应用获得的番茄作为育种材料用于番茄育种。
与现有技术相比，本发明以SlCIPK8基因为基础，将SlCIPK8基因所编码的蛋白质进行了功能鉴定，实验表明SlCIPK8基因的基因序列号为位于5号染色体上，基因长度为1356bp。SlCIPK8蛋白由451个氨基酸编码构成，为碱性蛋白、非跨膜蛋白、亲水性蛋白，预测含有65个磷酸化位点。SlCIPK8基因顺式作用元件中存在与激素、调控、胁迫相关的元件。推测SlCIPK8蛋白与多个CBL蛋白存在互作关系，以及几个非CBL蛋白也可能与存其在互作关系。
另外，SlCIPK8蛋白定位于细胞膜上，SlCIPK8基因在叶、茎、根组织中表达量较高。在低温、盐和干旱三种胁迫处理中，抵抗干旱能力的表型最为明显，抗盐次之，抵抗低温能力也比较明显。过表达SlCIPK8转基因植株可以通过提高抗氧化酶的活性提高植株的耐低温、抗盐性和抗旱性能力，因此本发明对于番茄的遗传改良及抗性品种选育具有重要的意义，具有很强的应用价值。
附图说明
图1为干旱胁迫下转基因株系和野生型株系的表型。其中，A为干旱胁迫处理3h前，B为干旱胁迫处理3h后。C为干旱胁迫处理9h前，D为干旱胁迫处理9h后。
图2为干旱胁迫下基因SlDREB2A表达量变化图。
图3为干旱胁迫下基因SlRD29A表达量变化图。
图4为干旱胁迫下基因SlABRE表达量变化图。
图5为盐处理胁迫下转基因株系和野生型株系的表型。其中，A为盐处理3h前，B为盐处理3h后。C为盐处理9h前，D为盐处理9h后。
图6为盐胁迫下基因SlDREB2A表达量变化图。
图7为盐胁迫下基因SlRD29A表达量变化图。
图8为盐胁迫下基因SlABRE表达量变化图。
图9为低温胁迫下转基因株系和野生型株系的表型。其中，A为4℃处理9h前，B为4℃处理9h后。C为4℃处理12h前，D为4℃处理12h后。
图10为低温胁迫下基因SlCBF1表达量变化图。
图11为低温胁迫下基因SlCBF2表达量变化图。
图12为低温胁迫下基因SlDREB2A表达量变化图。
图13为低温胁迫下基因SlRD29A表达量变化图。
具体实施方式
下面详细描述本发明的实施例，所述实施例的示例在附图中示出，其中自始至终相同或类似的标号表示相同或类似的元件或具有相同或类似功能的元件。下面通过参考附图描述的实施例是示例性的，旨在用于解释本发明，而不能理解为对本发明的限制。
在本说明书的描述中，参考术语“一个实施例”、“一些实施例”、“示例”、“具体示例”、或“一些示例”等的描述意指结合该实施例或示例描述的具体特征、结构、材料或者特点包含于本发明的至少一个实施例或示例中。在本说明书中，对上述术语的示意性表述不必须针对的是相同的实施例或示例。而且，描述的具体特征、结构、材料或者特点可以在任何的一个或多个实施例或示例中以合适的方式结合。此外，本领域的技术人员可以将本说明书中描述的不同实施例或示例进行接合和组合。
试验材料
番茄品种AC(Alisa Craig)的种子由发明人保存，播种后长至五叶一心。烟草品种为本生烟草(Nicotiana benthamiana)由发明人保存，培养至一月苗龄。
大肠杆菌Trans5α的化学感受态细胞(北京全式金生物)。农杆菌GV3101的感受态(上海唯地生物技术公司)。质粒psuper1300myc-GFP载体、帮助菌P19均来自于发明人实验室-80℃冰箱保存。
试验中所用的限制性核酸内切酶、DNA Maker和TransZol Up(购自北京全式金生物)。RNA逆转录试剂盒为ReverTra Ace
(东洋纺公司)。Solution I连接酶(购自宝日医生物技术(北京)有限公司)，其余试剂均为国产分析纯。
SICIPK8基因过表达载体的构建
提取番茄品种AC(Alisa Craig)的RNA，逆转录合成cDNA，然后以cDNA为模板，扩增SICIPK8全场片段，扩增引物分别如序列3(SEQ ID No.3，酶切位点XbaⅠ)和序列4(SEQ IDNo.4，酶切位点KpnⅠ)所示：
序列3(5′–3′)：CTCTAGAATGAGTAGCTCAACGTCAACGTC
序列4(5′–3′)：GGGTACCTTAAGACCCTTTTTCTTCAGAGG
PCR反应程序：
循环1:(1X)
步骤1:98.0℃，00:30min
循环2:(35X)
步骤1:98.0℃，00:10min
步骤2:58.0℃，00:05min
步骤3:68.0℃，00:05min
循环3:(1X)
步骤1:68.0℃，5:00min
经过PCR产物纯化，纯化产物和psupermyc1300-GFP载体的双酶切，酶切产物的胶回收，连接产物转化大肠杆菌感受态，鉴定重组子，提取大肠杆菌质粒。
SICIPK8基因过表达载体的遗传转化
经过预培养(消毒、诱导无菌苗、外植体制备)，转化(摇菌、预培养和今染、共培养)，延筛，愈伤的诱导及筛选，分化培养，生根培养，对生根植物进行转基因验证，继续培养验证成功植物至开花结果，收取种子。
转基因番茄的筛选鉴定
CTAB法提取植物基因组DNA，然后分别经过转基因番茄的PCR鉴定和qRT-PCR检测转基因株系的表达量鉴定。将各转基因株系的植株和野生型的DNA用引物(SEQ ID No.5和SEQ ID No.6)进行PCR，出现目的条带为阳性植株。
SEQ ID No.5：TGTCTGGTCCTGTGGAGTCA
SEQ ID No.6：CCTTTCCTTCCGGTCCTGTC
荧光定量检测抗逆相关基因的表达量
在相同的环境下种植转基因株系和野生型植株，取用种植6周后形态一致的番茄植株进行试验处理。样品均采集植株形态学上端第二片完全展开的功能叶，每个处理取三次生物学重复。处理时间点以前人CIPK8的VIGS实验为参考，选取酶活性差异较大的2个时间点。
(1)低温胁迫处理在4℃下，取样的时间点分别为4℃处理后的0h、9h、12h。
(2)干旱胁迫试验中需要把番茄幼苗先从土中移出，进行炼苗处理，清洗根系移至1/4Hoagland培养液中，适应培养液条件后进行干旱胁迫处理。干旱胁迫处理条件为20％PEG6000的1/4Hoagland培养液，取样的时间点分别为干旱胁迫处理后的0h、3h、9h后取样。
(3)盐胁迫试验中需要把番茄幼苗先从土中移出，进行炼苗处理，清洗根系移至1/4Hoagland培养液中，适应培养液条件后进行盐胁迫处理。盐胁迫处理条件为200mMNaCl的1/4Hoagland培养液，取样的时间点分别为干旱胁迫处理后的0h、3h、9h后取样。
利用荧光定量检测抗逆相关基因的表达量，其中，冷胁迫处理下检测SlCBF1、SlCBF3、SlDREB2A和SlRD29A基因表达量，干旱和盐胁迫处理下检测SlDREB2A、SlRD29A和SlABRE基因表达量。荧光定量PCR分析引物如下所示：
SlCBF1(5’–3’)引物:
SEQ ID No.7：CGGCTGAAATGGCGGCTAGAG
SEQ ID No.8：GGCAGCCTCCAAGCAGAATCAG
SlCBF3(5’–3’)引物:
SEQ ID No.9：TCTGCTTGGAGGCTGCCTACTC
SEQ ID No.10：GAAGATTTCGGCGGCCTGAGC
SlDREB2A(5’–3’)引物:
SEQ ID No.11：CCGCCGTAAAAGCCGCTCAC
SEQ ID No.12：CCGCCGTAAAAGCCGCTCAC
SlRD29A(5’–3’)引物:
SEQ ID No.13：AGGAGTTAGCCCGCTTGTTC
SEQ ID No.14：TGTGGATTGCTCGGATTCATA
SlAREB(5’–3’)引物:
SEQ ID No.15：GCTTTGCCCTATGGTGCTC
SEQ ID No.16：CCACTGGCTCCTAAACCTACC
Actin(5’–3’)引物:
SEQ ID No.17：TGTCCCTATTTACGAGGGTTATGC
SEQ ID No.18：CAGTTAAATCACGACCAGCAAGAT
一、抗旱性测试
过表达SlCIPK8基因提高了转基因番茄的抗旱性
选取6周大的番茄苗，对其进行干旱处理3h、9h后发现，在进行干旱处理3h时，过表达SlCIPK8的转基因株系和野生型均开始萎蔫，未出现明显差异(图1A、B)。在进行干旱处理9h时，野生型茎完全无法直立，过表达SlCIPK8的转基因株系叶片萎蔫但仍保持直立(图1C、D)。从图1可以看出SlCIPK8的转基因株系相对于野生型更耐干旱。(注：A为干旱胁迫处理3h前，B为干旱胁迫处理3h后。C为干旱胁迫处理9h前，D为干旱胁迫处理9h后。其中，左一为野生型，左二至四为过表达SlCIPK8的转基因株系的三个重复。)
干旱胁迫下过表达SlCIPK8基因中干旱胁迫相关基因表达量检测
为了进一步确认SlCIPK8转基因植株的抗旱能力，对SlCIPK8转基因植株和野生型进行了3h和9h的干旱胁迫处理，检测了SlCIPK8转基因植株和野生型中的干旱胁迫相关基因SlDREB2A、SlRD29A和SlABRE的表达量。如图2-4所示，随着干旱胁迫时间增加，SlDREB2A、SlRD29A和SlABRE的表达量随着胁迫时间的增加，在SlCIPK8转基因植株SlDREB2A、SlRD29A和SlABRE的表达量极显著高于野生型植株。
二、耐盐性测试
过表达SlCIPK8基因提高了转基因番茄的抗盐性
选取了6周大的番茄苗，如图5所示，对其进行了盐处理3h、9h时发现：过表达SlCIPK8转基因番茄株系在低温处理3h时表型与野生型相比没有显著变化；两者在盐处理9h时，野生型株系茎已经萎蔫，过表达SlCIPK8转基因番茄株茎仍挺直。根据图5可以看出，过表达SlCIPK8转基因番茄株系比野生型植株更耐盐。(注：A为盐胁迫处理3h前，B为盐胁迫处理3h后。C为盐胁迫处理9h前，D为盐胁迫处理9h后。)
盐胁迫下过表达SlCIPK8基因中低温胁迫相关基因表达量检测
分别检测盐胁迫3h和9h时的SlCIPK8转基因株系和野生型中的SlDREB2A、SlRD29A和SlABRE的表达量，结果如图6-8所示。随着盐胁迫时间增加，SlDREB2A、SlRD29A和SlABRE的表达量随着胁迫时间的增加，在SlCIPK8转基因株系中SlDREB2A、SlRD29A和SlABRE的表达量极显著高于野生型株系。
三、耐低温测试
低温对过表达SlCIPK8的转基因番茄表型的影响
如图9所示，选取6周大的番茄苗，对其进行了低温处理9h和12h后发现，过表达SlCIPK8转基因番茄株系在低温处理9h时表型与野生型相比没有显著变化，两者在低温处理12h时，野生型叶片卷曲更明显，过表达SlCIPK8转基因番茄株系比野生型植株更耐低温。(注：A为4℃处理9h前，B为4℃处理9h后。C为4℃处理12h前，D为4℃处理12h后。)
低温下过表达SlCIPK8基因中低温胁迫相关基因表达量检测
为了进一步确认SlCIPK8转基因株系的抗低温能力，分别检测了低温胁迫9h和12h的SlCIPK8转基因株系和野生型中的SlCBF1、SlCBF2、SlDREB2A和SlRD29A几个相关基因的表达量(图10-13)。结果发现，随着冷胁迫时间增加，前者在该指标上始终显著高于后者。
综上所述，本发明提供的抗逆基因在低温、盐和干旱三种胁迫处理中，抵抗干旱能力的表型最为明显，抗盐次之，抵抗低温能力也比较明显。过表达SlCIPK8转基因植株可以通过提高抗氧化酶的活性提高植株的耐低温、抗盐性和抗旱性能力，因此本发明对于番茄的遗传改良及抗性品种选育具有重要的意义，具有很强的应用价值。
应该注意到并理解，在不脱离后附的权利要求所要求的本发明的精神和范围的情况下，能够对上述详细描述的本发明做出各种修改和改进。因此，要求保护的技术方案的范围不受所给出的任何特定示范教导的限制。
申请人声明，以上内容是结合具体的优选实施方式对本发明所作的进一步详细说明，不能认定本发明的具体实施只局限于这些说明。对于本发明所属技术领域的普通技术人员来说，在不脱离本发明构思的前提下，还可以做出若干简单推演或替换，都应当视为属于本发明的保护范围。
序列表
<110> 东北农业大学
<120> 一种番茄抗逆基因及其编码的蛋白质和应用
<160> 18
<170> SIPOSequenceListing 1.0
<210> 1
<211> 451
<212> PRT
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 1
Met Ser Ser Ser Thr Ser Thr Ser Ser Pro Phe Ser Arg Ser Arg Thr
1 5 10 15
Arg Leu Gly Lys Tyr Glu Leu Gly Lys Thr Leu Gly Glu Gly Ser Phe
20 25 30
Ala Lys Val Lys Tyr Ala Lys Asn Ile Gln Thr Gly Glu Asn Val Ala
35 40 45
Ile Lys Ile Ile Asn Arg Asp Arg Val Leu Arg Gln Asn Ile Met Glu
50 55 60
Gln Ile Lys Arg Glu Ile Ser Thr Met Lys Leu Ile Arg His Pro Asn
65 70 75 80
Val Val Arg Ile Phe Glu Val Met Ala Ser Lys Thr Lys Ile Tyr Ile
85 90 95
Val Leu Glu Tyr Val His Gly Gly Glu Leu Phe Asp Glu Ile Ala Arg
100 105 110
His Gly Arg Leu Lys Glu Asp Glu Ala Arg Arg Tyr Phe Gln Gln Leu
115 120 125
Ile Asn Ala Val Asp Tyr Cys His Ser Arg Gly Val Phe His Arg Asp
130 135 140
Leu Lys Pro Glu Asn Leu Leu Leu Asp Ser Phe Gly Ile Leu Lys Val
145 150 155 160
Ser Asp Phe Gly Leu Ser Ala Leu Ser Lys Gln Val Arg Asp Asp Gly
165 170 175
Leu Leu His Thr Ala Cys Gly Thr Pro Asn Tyr Val Ala Pro Glu Val
180 185 190
Leu Thr Asp Lys Gly Tyr Asp Gly Thr Thr Thr Asp Val Trp Ser Cys
195 200 205
Gly Val Ile Leu Phe Val Leu Met Ala Gly Tyr Leu Pro Phe Asp Glu
210 215 220
Pro Asn Leu Asn Ala Leu Tyr Arg Lys Ile Leu Lys Ala Thr Phe Ser
225 230 235 240
Leu Pro Pro Trp Leu Ser Ser Ser Ser Lys Asn Leu Ile Asn Arg Ile
245 250 255
Leu Asp Pro Asp Pro Leu Thr Arg Ile Thr Ile Pro Glu Ile Leu Glu
260 265 270
Asp Glu Trp Phe Lys Lys Asp Tyr Lys Pro Pro Pro Phe Glu Gln Asp
275 280 285
Glu Asp Val Asn Leu Asp Asp Ile Asp Ala Ile Phe Asn Gly Ser Asp
290 295 300
Asp His Leu Val Thr Glu Arg Lys Glu Lys Pro Ala Ser Val Asn Ala
305 310 315 320
Phe Glu Leu Ile Ser Arg Ser Lys Ser Phe Asn Leu Glu Asn Leu Phe
325 330 335
Glu Lys Gln Ala Leu Val Lys Arg Glu Thr Gln Phe Thr Ser Arg Ser
340 345 350
Pro Ala Asn Glu Ile Ile Ser Lys Ile Glu Glu Thr Ala Arg Pro Leu
355 360 365
Gly Phe Ser Val Gln Lys Lys Asn Tyr Lys Met Lys Leu Gln Gly Asp
370 375 380
Arg Thr Gly Arg Lys Gly His Leu Ala Val Ala Thr Glu Val Phe Glu
385 390 395 400
Val Ala Pro Ser Val His Leu Val Glu Leu Arg Lys Thr Gly Gly Asp
405 410 415
Thr Leu Glu Phe His Lys Phe Tyr Lys Asn Phe Ser Ser Gly Leu Lys
420 425 430
Asp Ile Val Trp Thr Thr Glu Gln Thr Thr Glu Gln Pro Ser Glu Glu
435 440 445
Lys Gly Ser
450
<210> 2
<211> 1400
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 2
atcacattcg gttttcgagg atatctacaa gaactcatac gataatgagt agctcaacgt 60
caacgtcatc gccatttagt cgtagcagga cacgtttagg gaaatatgaa ttgggtaaga 120
cattaggtga gggaagtttt gctaaggtaa agtatgccaa aaatatacaa acaggtgaaa 180
atgtagccat caaaatcatc aatcgtgatc gcgtcctacg acaaaacatt atggaacaga 240
ttaaaagaga aatatcaacg atgaagttga tcagacatcc aaatgtcgtg aggatctttg 300
aggttatggc tagtaagacg aagatctata ttgtcctcga gtatgtacat ggaggagagc 360
tctttgatga aattgctaga catggaagac tcaaagaaga cgaagctaga agatactttc 420
aacagcttat taatgccgtt gattactgtc atagtagagg tgttttccat cgagacttga 480
agcctgagaa tctattgctg gattcatttg gtattctgaa agtttcagac ttcggattga 540
gtgcactttc caagcaagtt cgagacgatg gactgcttca tactgcttgt ggcacaccaa 600
actatgtcgc gcctgaggtg ctaactgaca aaggctatga tggtacaaca actgatgtct 660
ggtcctgtgg agtcattctc tttgttctaa tggctggata cttacctttt gatgagccaa 720
acctaaatgc tctataccga aaaatcctga aggctacgtt ttcacttcca ccatggttgt 780
cctccagctc aaagaatctg atcaaccgta ttcttgaccc ggatccactc acaaggatca 840
ctattccaga gatcctcgaa gatgagtggt tcaagaaaga ttacaagccg cctccttttg 900
agcaagacga agatgtcaat cttgatgaca ttgatgccat attcaatggc tcggacgatc 960
atctcgttac agaaagaaag gaaaaacctg cttctgtcaa tgcattcgag ctcatttcaa 1020
ggtcaaaaag tttcaaccta gaaaatttgt tcgaaaaaca ggcccttgtc aagagagaaa 1080
cacaatttac ttcgcggtcg cctgctaatg agattatctc caaaattgag gaaactgcaa 1140
gaccattggg attcagtgtc cagaagaaaa attacaagat gaagttacaa ggcgacagga 1200
ccggaaggaa aggccacctt gctgtagcaa cagaggtgtt tgaagtagca ccctcagtgc 1260
atttggttga gctccgaaaa actggtggtg atacattaga gtttcataag ttttacaaga 1320
acttctcttc aggattaaaa gatatcgtct ggacaacaga acaaacaacc gaacaaccct 1380
ctgaagaaaa agggtcttaa 1400
<210> 3
<211> 30
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 3
ctctagaatg agtagctcaa cgtcaacgtc 30
<210> 4
<211> 30
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 4
gggtacctta agaccctttt tcttcagagg 30
<210> 5
<211> 20
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 5
tgtctggtcc tgtggagtca 20
<210> 6
<211> 20
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 6
cctttccttc cggtcctgtc 20
<210> 7
<211> 21
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 7
cggctgaaat ggcggctaga g 21
<210> 8
<211> 22
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 8
ggcagcctcc aagcagaatc ag 22
<210> 9
<211> 22
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 9
tctgcttgga ggctgcctac tc 22
<210> 10
<211> 21
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 10
gaagatttcg gcggcctgag c 21
<210> 11
<211> 20
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 11
ccgccgtaaa agccgctcac 20
<210> 12
<211> 20
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 12
ccgccgtaaa agccgctcac 20
<210> 13
<211> 20
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 13
aggagttagc ccgcttgttc 20
<210> 14
<211> 21
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 14
tgtggattgc tcggattcat a 21
<210> 15
<211> 19
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 15
gctttgccct atggtgctc 19
<210> 16
<211> 21
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 16
ccactggctc ctaaacctac c 21
<210> 17
<211> 24
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 17
tgtccctatt tacgagggtt atgc 24
<210> 18
<211> 24
<212> DNA
<213> 2 Ambystoma laterale x Ambystoma jeffersonianum
<400> 18
cagttaaatc acgaccagca agat 24
Claims (10)
1.一种蛋白质，其特征在于，所述蛋白质由序列表中序列1所示的氨基酸序列组成。
2.编码如权利要求1所述蛋白质的基因。
3.如权利要求2所述的基因，其特征在于，所述基因为序列表中序列2所示的DNA分子。
4.含有如权利要求2或3所述基因的重组表达载体、表达盒或重组菌。
5.如权利要求2或3所述基因的应用。
6.如权利要求5所述基因的应用，其特征在于，至少包括以下步骤：将权利要求2或3所述的基因导入目的番茄，使目的番茄加强对于干旱的抵抗能力。
7.如权利要求5所述基因的应用，其特征在于，至少包括以下步骤：将权利要求2或3所述的基因导入目的番茄，使目的番茄加强对于高盐的抵抗能力。
8.如权利要求5所述基因的应用，其特征在于，至少包括以下步骤：将权利要求2或3所述的基因导入目的番茄，使目的番茄加强对于低温的抵抗能力。
9.如权利要求5-8中任一项所述的应用，其特征在于，至少包括以下步骤：将权利要求2或3所述的基因导入目的番茄，使目的番茄加强对于干旱、高盐和低温的抵抗能力。
10.一种番茄育种方法，其特征在于，将权利要求5-8中任一项所述的应用获得的番茄作为育种材料用于番茄育种。
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