CN114214341B - 番茄SlSERAT1;1基因或其片段在植物发育过程中的应用 - Google Patents
番茄SlSERAT1;1基因或其片段在植物发育过程中的应用 Download PDFInfo
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Abstract
本发明属于基因工程技术领域，具体涉及番茄SlSERAT1；1基因或其片段在植物发育过程中的应用。为改良番茄中含硫氨基酸含量，提高番茄的经济价值，本发明提供一种番茄SlSERAT1；1基因或其片段在植物发育过程中的应用，其核苷酸序列如SEQ ID No.1所示，片段为番茄SlSERAT1；1基因的cDNA序列，其核苷酸序列如SEQ ID No.7所示。番茄SlSERAT1；1基因属SERAT(丝氨酸乙酰转移酶)家族成员，而SERAT作为半胱氨酸生物合成过程中的限速酶，参与植物许多发育以及对环境的适应过程，具有较好的潜在应用价值。
Description
技术领域
本发明属于基因工程技术领域，具体涉及番茄SlSERAT1；1基因或其片段在植物发育过程中的应用。
背景技术
硫是植物生长发育中的必需元素之一，对植物的生长代谢有重要作用。半胱氨酸(Cysteine，Cys)是植物将无机硫转变为有机硫的过程中产生的第一个含硫有机物，在植物代谢过程中占据重要位置。丝氨酸乙酰转移酶(Serine acetyltransferase，SERAT)催化半胱氨酸合成的第一步反应，是半胱氨酸合成过程中的限速步骤。
有研究发现，L-半胱氨酸作为巯基化合物，具有抗褐变、抗氧化的作用，在多酚的氧化过程中，通过竞争结合位点使半胱氨酸酶促褐变减慢，采摘后将其用于荔枝果实的保存可有效减缓果皮的褐变。此外，研究发现，L-半胱氨酸可以诱导果实对环境压力的耐受性，在龙眼、黄瓜和其他水果蔬菜中使用它可以增加水果的抗氧化活性，并延缓水果腐败变质。当然，半胱氨酸作为第一种有机含硫化合物，在植物代谢中占据中心位置，其重要性不仅取决于它作为蛋白质中氨基酸的作用，还取决于它是还原硫供体分子，是植物生命活动所必须的多种含硫代谢产物的前体，例如甲硫氨酸、谷胱甘肽、酶辅助因子、一些重要维生素、硫酯化合物和其他含硫衍生物等。其中研究较多的是谷胱甘肽，它是一种抗氧化剂，由γ谷酰基-半胱氨酰-谷氨酸构成的三肽化合物，被认为是细胞氧化还原稳态的主要决定因素。通过谷胱甘肽衍生的多肽植物螯合素可以实现重金属解毒，谷胱甘肽还可在谷胱甘肽硫转移酶等的作用下发挥解毒作用。在所有这些生物分子中，它们作为官能团的含硫部分都来源于半胱氨酸，这表明它们的生物合成途径是紧密相连的。另一种与半胱氨酸密切相关的代谢物是激素乙烯，它参与植物种子萌发、根毛发育、叶片和花瓣脱落、幼苗生长、跃变型果实成熟、以及逆境反应调节等许多方面。此外，半胱氨酸本身及其衍生分子还在各种细胞区室发生的氧化还原信号过程中起作用。在细胞质中，半胱氨酸在植物免疫中起着重要作用；在线粒体中，这种分子在氰化物解毒中起着重要作用，氰化物对根毛发育和植物对病原体的反应至关重要。因此，半胱氨酸在植物的初级代谢和次生代谢中都占据着中心位置。
除了半胱氨酸外，含硫氨基酸(SAA)还包括甲硫氨酸(Methionine，Met) 和胱氨酸(Cystine)。其中，甲硫氨酸是人体的必需氨基酸之一，仅能通过食物获取，在动物中亦是如此。在植物中，半胱氨酸是甲硫氨酸的前体，甲硫氨酸生物合成的最后一步是，同型半胱氨酸在甲硫氨酸合成酶的作用下合成甲硫氨酸。
在日常生活中，含硫氨基酸不仅对动物的生长发育有重要作用，比如提供营养和参与机体免疫等。同时，还可在多方面提高食品的品质，包括营养价值、感观感受(比如抑制褐变)和生理作用(比如抗氧化)等。含硫氨基酸的应用前景十分广阔，不仅可以应用于食品开发，而且可以运用于医药保健以及动物养殖等多方面。因此，掌握在植物中提高半胱氨酸产量的方法具有重要的应用价值。
番茄(Solanum lycopersicum)是重要的经济效益蔬菜作物之一，全球年总产量达1.7亿吨，在蔬菜作物中位居首位。在番茄中提高半胱氨酸产量，一方面可以加强番茄的质量，例如：半胱氨酸的抗氧化作用可以保护番茄中的维生素等营养物质不被氧化间接提高营养价值；同时半胱氨酸在抑制酶促褐变时还会产生水果的清香味，在感官上增强吸引力；此外，半胱氨酸能与黄曲霉毒素、脱氢丙氨酸等作用进而发挥解毒功能等。另一方面，在番茄中提高半胱氨酸产量，可以提高番茄的经济价值，即半胱氨酸强烈的抗氧化、抗褐化作用可以延缓番茄腐坏变质，增长其保鲜期，进而带来良好的经济效益。总之，对番茄中含硫氨基酸含量的改良，与我们的的健康和生活息息相关。本发明通过调节番茄中的半胱氨酸合成相关基因SlSERAT1；1的表达进而提高了番茄中的半胱氨酸产量，开发了基于调节SlSERAT1；1的表达提高半胱氨酸含量的手段，同时获得了相关转基因植株。
发明内容
针对上述问题本发明提供了番茄SlSERAT1；1基因或其片段在植物发育过程中的应用，以及番茄SlSERAT1；1基因编码的SlSERAT1；1蛋白，含有番茄 SlSERAT1；1基因或其片段的载体，含有所述载体的宿主细胞。
为了达到上述目的，本发明采用了下列技术方案：
番茄SlSERAT1；1基因或其片段在植物发育过程中的应用，所述番茄 SlSERAT1；1基因的核苷酸序列如SEQ ID No.1所示，所述片段为番茄SlSERAT1；1 基因的cDNA序列，其核苷酸序列如SEQ ID No.7所示。
进一步，所述番茄SlSERAT1；1基因或其片段影响番茄植株半胱氨酸的合成。
以拟南芥SERAT(SERAT1；1)的蛋白质序列作为查询条件，对翻译后的番茄核苷酸数据库进行tblastn搜索进而找到目的基因，SlSERAT1；1相似性最高，为89.93％，具有如SEQ ID No.1所示的核苷酸序列。在此过程中，我们还发现了另外3个具有较高相似性水平(E值<e-10)的SERAT编码基因。
番茄SlSERAT1；1基因编码的SlSERAT1；1蛋白，其为由SEQ ID No.2所示的氨基酸序列组成的蛋白，或为SEQ ID No.2所示的氨基酸序列经取代、缺失和/ 或增加一个或多个氨基酸且具有同等活性的衍生的蛋白。
通过系统发育分析，我们将发现的4个番茄SERATs划分在三个不同的分支中，另参照拟南芥命名方法，将这4个番茄SERAT基因命名为SlSERAT1；1、 SlSERAT2；1、SlSERAT2；2、SlSERAT3；1。此外，将这些SlSERAT的蛋白序列与拟南芥和大豆的SERAT蛋白进行了比较。多重蛋白序列比对表明：SlSERATs 中N端α-螺旋和C端β-折叠是保守的。
应当理解，在研究番茄SlSERAT1；1基因或其片段在植物发育过程中的作用时，本领域技术人员可根据本发明公开的氨基酸序列，在不影响其活性的前提下，取代、缺失和/或增加一个或几个氨基酸，得到所述蛋白的突变序列。因此，本发明的番茄SlSERAT1；1蛋白还包括SEQ ID No.2所示氨基酸序列经取代、替换和/或增加一个或几个氨基酸，具有番茄SlSERAT1；1蛋白同等活性的由番茄 SlSERAT1；1蛋白衍生得到的蛋白质。在研究番茄SlSERAT1；1基因或其片段在植物发育过程中的作用时，本发明基因包括编码所述蛋白的核酸序列。此外，应理解，考虑到密码子的简并性以及不同物种密码子的偏爱性，本领域技术人员可以根据需要使用适合特定物种表达的密码子。
在研究番茄SlSERAT1；1基因或其片段在植物发育过程中的作用时，本发明还提供含有上述番茄SlSERAT1；1基因或其片段的载体，以及含有该载体的宿主细胞；所述载体为所述番茄SlSERAT1；1基因或其片段的克隆载体或各类表达载体；所述片段是指番茄SlSERAT1；1基因的cDNA序列，其核苷酸序列如SEQ ID No.7所示。
具体地说，为了研究番茄SlSERAT1；1基因或其片段在植物发育过程中的作用，本发明将番茄SlSERAT1；1基因的cDNA序列(882bp，其核苷酸序列如SEQ ID No.7所示)克隆到pCAM2300载体中，并在大肠杆菌DH5α中扩繁。
本发明还通过农杆菌介导转化方法，将构建好的过表达载体成功转化农杆菌EHA105菌株，并进一步获得了番茄转化植株。
与现有技术相比本发明具有以下优点：
在研究番茄SlSERAT1；1基因或其片段在植物发育过程中的作用时，番茄SlSERAT1；1基因属于SERAT家族成员，具有影响番茄植株半胱氨酸合成的潜在应用价值。
附图说明
图1是本发明实施例1的克隆载体pMD-18T；
图2是本发明实施例2的SlSERAT1；1蛋白的系统进化分析，即将拟南芥和番茄中的SERAT蛋白进行系统进化分析。拟南芥OASTL家族蛋白AtOAS-A1 用作组外对照；
图3是本发明实施例3的番茄SlSERAT1；1的基因结构，括号中的数字代表基因CDS的长度；
图4是本发明实施例4的番茄SlSERAT1；1编码蛋白的结构分析，将番茄、拟南芥和大豆中的SERAT蛋白序列进行比较。二级结构显示在序列上方：白色矩形代表α-螺旋；白色箭头代表β-折叠；黑条代表环状。黑色三角形表示C末端的关键氨基酸位点——Ile。黑色背景上的白色字母表示最保守的氨基酸(在所有分析的蛋白质序列中相同)，而逐渐变浅的背景色表示保守性降低；
图5是本发明实施例5的植物过表达载体pCAM2300，其在插入基因的5’端有一个3x Flag的编码序列；
图6是本发明实施例6的SlSERAT1；1过表达植株DNA水平的鉴定(a)和 RNA水平的鉴定(b)。其中，M代表DL2000 DNA Marker；WT代表MicroTom 野生型；CK代表阴性对照；其余泳道标号代表不同的转基因株系；
图7是本发明实施例7的番茄SlSERAT1；1转基因植株中SERAT活性(a) 和半胱氨酸含量(b)分析图；
图8是本发明实施例8番茄SlSERAT1；1基因编码蛋白的亚细胞定位分析；
图9是本发明实施例9番茄SlSERAT1；1基因的表达模式分析图。其中，R，根；SD，幼苗；S，茎；YL，幼叶；OL，老叶；IN，花序；Se，萼片；Pe，花瓣；St，雄蕊；Ca，心皮；PD，花柄；IMG，未成熟果实；BF，露白期果实； YF，黄果期；RF，红果期；
图10是本发明实施例10的蛋白纯化与酶活分析图。(a)SlSERAT1；1-His 融合蛋白的诱导与纯化，其中，M，蛋白质分子量标准；1，未诱导菌液蛋白；2，诱导后的菌液蛋白；3，纯化的蛋白；箭头，纯化的蛋白质条带；红色框架，70KD 的蛋白条带。(b)SlSERAT1；1-His融合蛋白的酶活分析。
具体实施方式
实施例1：番茄SlSERAT1；1基因编码区的克隆
该基因是以拟南芥SERAT(SERAT1；1)的蛋白质序列作为查询条件，对翻译后的番茄核苷酸数据库进行Tblastn搜索进而找到的，其相似性最高，为 89.93％，具有如SEQ IDNo.1所示的核苷酸序列。在此过程中，我们还发现了另外3个具有较高相似性水平(E值<e-10)的SERAT编码基因。然后，以番茄cDNA 为模板，PCR扩增和克隆了番茄SlSERAT1；1基因的CDSs，并对其序列进行了验证。即利用核苷酸序列如SEQ ID No.3所示的正向引物和核苷酸序列如SEQ ID No.4所示的反向引物，从番茄Micro Tom的叶片cDNA中克隆SlSERAT1；1 基因的编码区序列；
PCR程序：(1)94℃，2分钟；(2)94℃，30秒；55℃，30秒；72℃，1kb/1 min cDNA；重复38次；(3)72℃，7分钟。
采用回收试剂盒(OMEGA)对上述PCR产物进行纯化，然后将纯化后的 PCR产物直接克隆连接到pMD-18T载体上(如图1所示)；连接产物转化大肠杆菌DH5α，并在其中扩繁，阳性克隆经过测序筛选获得该序列；其核苷酸序列如SEQ ID No.1所示，由其编码的蛋白质的氨基酸序列如SEQ ID No.2所示。
实施例2：番茄SlSERAT1；1蛋白的系统进化分析
系统发育分析使用MEGA 7.0软件，拟南芥的OASTL蛋白质家族OAS-A1 被用作组外对照。从系统发育上看，番茄的SERATs被划分在三个不同的分支中，另参照拟南芥命名方法，将这4个番茄SERAT基因命名为SlSERAT1；1、 SlSERAT2；1、SlSERAT2；2、SlSERAT3；1，其编码区长度分别为882bp、1098bp、 1080bp、1071bp(如图2所示)。
实施例3：番茄SlSERAT1；1基因结构分析
基于CDS和基因组序列之间的比对结果，构建基因结构。SlSERAT1；1的基因结构如图3所示，有1个内含子。
实施例4：番茄SlSERAT1；1蛋白的序列结构分析
为了进一步研究番茄SERAT基因编码的蛋白序列，本实验将这些SlSERAT 的蛋白序列与拟南芥和大豆的SERAT蛋白进行了比较。多重蛋白序列比对表明： SlSERATs中N端α-螺旋和C端β-折叠是保守的，结果如图4所示。
实施例5：番茄SlSERAT1；1基因过表达载体和转基因植株
过表达载体构建采用双酶切方法，过表达载体是pCAM2300，SlSERAT1；1 选用BamHI和SalI作为酶切位点。载体构建过程中SlSERAT1；1的CDS区以实施例1构建好的重组质粒为模板，以核苷酸序列如SEQ ID No.5所示的正向引物和核苷酸序列如SEQ ID No.6所示的反向引物成功扩增出SlSERAT1；1目的基因，利用双酶切法将其连接到pCAM2300载体上，转化大肠杆菌DH5α，挑选有明亮单一条带的菌液提取重组质粒，然后对其双酶切验证，将酶切出正确大小目的条带的质粒送公司测序。
构建好的过表达载体转化农杆菌EHA105菌株，以番茄种子无菌播种后的子叶为受体，获得番茄转化植株，植物中的筛选标记为卡那霉素。农杆菌介导转化方法和步骤如下：
农杆菌转化番茄的方法和步骤如下：
1.种子准备：将1.5g番茄种子依次经95％乙醇洗和20％次氯酸钠消毒和无菌水清洗后，将种子均匀放置于1/2MSO培养基表面，在24℃培养间光照(16h 光照，8h黑暗)培养6天。
2.外植体准备：当子叶从种皮中长出后，用锋利的解剖刀将子叶切成25mm2 大小，并将子叶近轴面向上放置在盛有滤纸的D1培养基的表面(注意无菌操作)；于24℃培养间长光照(16h光照，8h黑暗)培养2天。
3.农杆菌准备：固体培养基上活化-70℃存放的农杆菌(Agrobacteriumtumefaciens，C58C1)，挑单克隆至相应抗性的液体培养基中培养至OD600处于 0.6～0.7之间，收集菌体，MSO清洗悬浮并加入乙酰丁香酮(AS)制备成侵染液。
4.共培养：用上一步制成的侵染液侵染子叶，侵染完成后将子叶远轴端朝上培养于新的盛有滤纸和D1培养基的新培养皿中，在24℃培养间放置2天(16h 光照，8h黑暗)。
5.选择转化愈伤：将子叶转移至2Z培养基上(没有滤纸)，约10天更换一次培养基，直至有芽出现后转移至1Z选择性培养基上继续培养，约两周更换一次培养基。
6.生根：当再生苗至少有2cm长并包含至少一个生长点的时候，可从外植体上切下再生苗(不包括愈伤组织)，将再生苗放入MMSV培养基中进行生根培养。
生根后的植株生长到足够大时，就可转移至装有跖石和营养土的培养钵中，于一般生长箱中生长。这些植株即为T0代植株。
其中，使用到的培养基配方为：
120℃高压灭菌20分钟后加入以下成分，
120℃高压灭菌20分钟后加入以下成分，
120℃高压灭菌20分钟后加入以下成分，
特美汀 200mg/L
卡那霉素 50mg/L
叶酸 0.5mg/L
实施例6：番茄SlSERAT1；1过表达转基因植株分子水平阳性鉴定
首先从实施例5中获得的4株番茄过表达转化植株叶片中提取DNA，并在 DNA水平利用PCR检测进行鉴定。结果如图6中a所示。在DNA水平上，共得到4株SlSERAT1；1过表达植株阳性苗。
提取这些阳性转化株的RNA，反转录成cDNA之后，通过荧光定量PCR(以 ACTIN基因为内参)检测SlSERAT1；1基因的表达水平。结果如图6中b所示，与野生型相比，我们得到2株SlSERAT1；1高表达的转基因株系，其与WT相比，有极显著的差异。
实施例7：番茄SlSERAT1；1转基因植株表型分析以及SERAT活性和半胱氨酸含量分析
我们发现同野生型相比，SlSERAT1；1过表达植株在表型上并无明显变化，因此进一步分析了SERAT活性和半胱氨酸含量，以期解析SlSERAT1；1在体内的功能(图7中a和b)。结果显示，在过表达突变体中，其产生OAS和半胱氨酸的能力均得到了提高，但是由于共阻遏效应，与OE-SlSERAT1；1-6相比较， OE-SlSERAT1；1-3反而具有更高的SERAT活性和半胱氨酸含量。同时在OE-SlSERAT1；1-6中，两个指标其实并没有明显变化。
实施例8：SlSERAT1；1蛋白的亚细胞定位分析
构建SlSERAT1；1-GFP融合基因，并将载有该基因的质粒转化进农杆菌中，然后利用农杆菌介导的瞬时转化将融合基因转化进烟草叶片中。转化细胞中 GFP荧光信号存在的位置指示SlSERAT1；1蛋白在细胞中定位于线粒体中。结果如图8所示。
实施例9：SlSERAT1；1基因的表达模式分析
利用实时定量PCR分析SlSERAT1；1基因在各组织器官的时空表达模式，结果如图9所示。结果显示SlSERAT1；1在番茄的各个组织器官中均匀且高水平地表达，表明其在植物发育过程中起着重要作用。
实施例10：SlSERAT1；1蛋白的酶活分析
构建SlOAS7-pCold原核表达载体，在大肠杆菌BL21中表达SlSERAT1；1-His 融合蛋白并纯化。蛋白纯化后测定其酶活，分析了番茄SlSERAT1；1蛋白催化生成OAS的能力。结果如图10所示。结果显示在没有OASTL蛋白的情况下， SlSERAT1；1蛋白表现出中等的催化OAS生物合成的能力。当将SlOAS6添加到反应中时，SlSERAT1；1的活性显著增加。
虽然，上文中已经用一般性说明及具体实施方式，对本发明作了详尽的描述，但在本发明基础上，可以对之作一些修改或改进，这对本领域技术人员而言是显而易见的。因此，在不偏离本发明精神的基础上所做的这些修改或改进，均属于本发明要求保护的范围。
序列表
SEQ ID No.1
ATGCCAGCCGAAGAACACCGTAACGCCTCGCCGGCGGCGCCGCATCC ACCGACGGACACGGCAGAAGAAGCTATCTGGTTATGGACACAGATCAAAG CAGAAGCTCGGCGGGATGCAGAAGCTGAACCGGCATTAGCAAGCTATTTGTACTCAACTATACTATCTCACTCTTCGCTTGAGCGTTCACTCTCTTTTCATTT GGGGAACAAGCTTTGTTCTTCAACGCTTCTATCTACTCTTCTCTACGATCTGTTCCTCAATAATTTCTCCTCTGATCCTGATCTACGCGCCGCGGCATCCGCTG ACCTACTCGCCGCTCGCTACCGTGACCCAGCCTGTGTTTCCTTCTCTCATTGTTTGCTTAATTACAAAGGCTTCCTTGCTTGTCAGGTTTGTATGTTGTAACTA CTCGTTCCATCTATTTTTACTTATCCAGTATCGACTTACTTGGCCTATCGAGCAATAAATGAAATGACAAGTATAATAAATTTATGTATCGTGAAATGACTGTAG TTAATAATACGATAAACTATTAGTAACTATGTCCTTTTTAGTATATTGTGATACTCTGAAATTGTGAAAAAATAGGGTTTATGTTAAATTTTTCTGGAAAGATGG GAACTTTAGAGTATGAAAAGATGAATTTTTGGGGGGTTTGTTTGAAAAATTGATGATTGAAGTGTGGGAGCTAAGATTTTCGACAAGGGAATTCAAAATATA GAGAAGTAAATGCACAAGGCCTACGGGATTCAGTAACTAGAACAACAACGTACCTAGTGTAATTCCATTTGTGGGGTTTGGGAAGGGCAAGATATATGCAAACGCAAGATCGTATCGTACCTTTGCAGGGTTGAGAGTCTATTTCTAATAGACCATCGGCTTGACATATGAGAAAAGATATATCTACTATTTATGCATGAAGTATA TTATTTACATACAAAATAGTTTTGACCTTGTGCATACTGTGAAATTTTTGGTGATGAATTCCCTTGCAGCATCCTAGCTCCGCCCTGGTCAATGTTCCAATTGT TTCCTTGGAAGTTATACCAAACAAAAGATTGACAGAGGATGTTAGAAGTAT ATGGCATATGCCAATCTACTTGAATGTGAAACTGATAATATGACCTTTTCACTTTGGAATTTGACAATGTTCAAGACTTAGTTGGTGCTATACTACTATGTTATA ACGATAGGTACATTTGTTAAGATATTACTTTAGCAAGGTAGTGGAATTAGATCTTTCTTTGGATGTAGACTCGGTATGAATTAGGATAACTATTGAGTTGAAGT TGAGAGAGTACTTGAAGTTGATGTTGGAAAAAGATAGTGGAAGACTTGAA GTTAGTCTTTATGTTTCGAAATGAAATTCATTTGGAAAGAAAGTTTGTGAATGTTGTGAGTAGAAGTCAAACACTTGAATTGCCTTGCTTCTATAATAAACTTT CAAACCACTACTACAAGATTTTATTTATTGTTCAAATAAGACTTCAAGGCCAAACTACTATTTGTAAATAATTATGTCATGTCTGACCGGGCTAAAATGGTGAA AGTGTGTTCTGCAATGGAGCCAAGGGCTGTGATATTGTTTGCTCAGTAGTC TCTAGTATCTGGTTGGTAGGTATGCTTTGAAAGTAGTATAAAGATAAAAGGCTGCCAGTTTGGTGGTCTATACCATTCCTTGTTGTTTCTTTTCTATCCCAATCC ATTTTGAGATAAGCAGCTAATGAAACATCGACCCCCCCGGATTTTCCTGTTTATTTGTGTCAGAAGGCCCGCACACACCTATATACGCGGATGTAAATGAATG TATGCAATGATCTTTCCTTTTCTGTTAAACAATGTTCATTAGCTATCATGTGTTGTATTGGCTCTCTGCTGACAATGAAATAACTGTACAGATGCATTTGGACA GTTCATATTCTAATTGAAAAGAAAAAGTATATTTTTCCTTTGGTTAACGAGTGTGAGATGCTTTTTTATTTCAATAAAATGAGTAGCTATTAGCATCTCATGGA AATCGCATATTTACAGATTGCCTGTATTTCAATAGCAAGTCAAGGGGCTTACTTTTCACTTCATATCACAGACTCTCTTCAAATGGTAGTATTAGGTTTTAACA ATATAATGTTTCTATTTTCAGGCACATCGAGTAGCTCACAAACTTTGGACTCAATCCCGAAGGCCACTTGCACTTGCACTTCAATCCCGAATCTCTGATGTTT TTGCTGTTGACATTCATCCAGCTGCCAAAATCGGAAAAGGCATCCTCTTCG ACCATGCAACAGGAGTGGTTGTTGGTGAAACTGCAGTTATTGGAAACAATGTGTCGATTCTTCACCATGTAACCTTAGGAGGAACTGGTAAGTTTGGTGGT GACCGACACCCTAAGATTGGTGACGGAGTCCTCATAGGTGCAGGTGCCACGATATTAGGCAACATAAATATCGGTGAGGGAGCCAAGATTGGCGCTGGATCAGTGGTTCTGATTGACGTGCCACCACGAACAACTGCAGTTGGAAATCCAGCTAGGTTGGTGGGAGGGAAAGAACAGCCAACTAAGCACGAGGAATGTCC CGGAGAGAGTATGGACCATACATCTTTCATATCTGGATGGTCTGATTACATCATCTGA
SEQ ID No.2
Met Pro Ala Glu Glu His Arg Asn Ala Ser Pro Ala Ala Pro His Pro ProThr Asp Thr Ala Glu Glu Ala Ile Trp Leu Trp Thr Gln Ile Lys Ala Glu Ala ArgArg Asp Ala Glu Ala Glu Pro Ala Leu Ala Ser Tyr Leu Tyr Ser Thr Ile Leu SerHis Ser Ser Leu Glu Arg Ser Leu Ser Phe His Leu Gly Asn Lys Leu Cys Ser SerThr Leu Leu Ser Thr Leu Leu TyrAsp Leu Phe LeuAsnAsn Phe Ser SerAsp Pro AspLeuArg AlaAla Ala Ser Ala Asp Leu Leu Ala Ala Arg Tyr Arg Asp Pro Ala Cys ValSer Phe Ser His Cys Leu Leu Asn Tyr Lys Gly Phe Leu Ala Cys Gln Ala His ArgVal Ala His Lys Leu Trp Thr Gln Ser Arg Arg Pro Leu Ala Leu Ala Leu Gln SerArg Ile Ser Asp Val Phe Ala Val Asp Ile His Pro AlaAla Lys Ile Gly Lys GlyIle Leu Phe Asp His Ala Thr Gly Val Val Val Gly Glu ThrAla Val Ile Gly AsnAsnVal Ser Ile Leu His His Val Thr Leu Gly Gly Thr Gly Lys Phe Gly Gly Asp ArgHis Pro Lys Ile Gly Asp Gly Val Leu Ile Gly Ala GlyAla Thr Ile Leu GlyAsn IleAsn Ile Gly Glu Gly Ala Lys Ile GlyAla Gly Ser Val Val Leu Ile Asp Val ProPro Arg Thr Thr Ala Val Gly Asn Pro Ala Arg Leu Val Gly Gly Lys Glu Gln ProThr Lys His Glu Glu Cys Pro Gly Glu Ser Met Asp His Thr Ser Phe Ile Ser GlyTrp SerAsp Tyr Ile Ile
SEQ ID No.3
GCTGGATCCATGCCAGCCGAAGAACAC
SEQ ID No.4
GACGTCGACTCAGATGATGTAATCAGACC
SEQ ID No.5
CGCGGATCCATGCCAGCCGAAGAACACCG
SEQ ID No.6
ACGCGTCGACGATGATGTAATCAGACCATC
SEQ ID No.7
ATGCCAGCCGAAGAACACCGTAACGCCTCGCCGGCGGCGCCGCATCC ACCGACGGACACGGCAGAAGAAGCTATCTGGTTATGGACACAGATCAAAGCAGAAGCTCGGCGGGATGCAGAAGCTGAACCGGCATTAGCAAGCTATTTG TACTCAACTATACTATCTCACTCTTCGCTTGAGCGTTCACTCTCTTTTCATTTGGGGAACAAGCTTTGTTCTTCAACGCTTCTATCTACTCTTCTCTACGATCTG TTCCTCAATAATTTCTCCTCTGATCCTGATCTACGCGCCGCGGCATCCGCTGACCTACTCGCCGCTCGCTACCGTGACCCAGCCTGTGTTTCCTTCTCTCATTG TTTGCTTAATTACAAAGGCTTCCTTGCTTGTCAGGCACATCGAGTAGCTCACAAACTTTGGACTCAATCCCGAAGGCCACTTGCACTTGCACTTCAATCCCG AATCTCTGATGTTTTTGCTGTTGACATTCATCCAGCTGCCAAAATCGGAAAAGGCATCCTCTTCGACCATGCAACAGGAGTGGTTGTTGGTGAAACTGCAG TTATTGGAAACAATGTGTCGATTCTTCACCATGTAACCTTAGGAGGAACTGGTAAGTTTGGTGGTGACCGACACCCTAAGATTGGTGACGGAGTCCTCATAG GTGCAGGTGCCACGATATTAGGCAACATAAATATCGGTGAGGGAGCCAAGA TTGGCGCTGGATCAGTGGTTCTGATTGACGTGCCACCACGAACAACTGCAGTTGGAAATCCAGCTAGGTTGGTGGGAGGGAAAGAACAGCCAACTAAGC ACGAGGAATGTCCCGGAGAGAGTATGGACCATACATCTTTCATATCTGGAT GGTCTGATTACATCATCTGA
序列表
<110> 山西大学
<120> 番茄SlSERAT1;1基因或其片段在植物发育过程中的应用
<160> 7
<170> SIPOSequenceListing 1.0
<210> 1
<211> 2632
<212> DNA
<213> 番茄(Solanum lycopersicum)
<400> 1
atgccagccg aagaacaccg taacgcctcg ccggcggcgc cgcatccacc gacggacacg 60
gcagaagaag ctatctggtt atggacacag atcaaagcag aagctcggcg ggatgcagaa 120
gctgaaccgg cattagcaag ctatttgtac tcaactatac tatctcactc ttcgcttgag 180
cgttcactct cttttcattt ggggaacaag ctttgttctt caacgcttct atctactctt 240
ctctacgatc tgttcctcaa taatttctcc tctgatcctg atctacgcgc cgcggcatcc 300
gctgacctac tcgccgctcg ctaccgtgac ccagcctgtg tttccttctc tcattgtttg 360
cttaattaca aaggcttcct tgcttgtcag gtttgtatgt tgtaactact cgttccatct 420
atttttactt atccagtatc gacttacttg gcctatcgag caataaatga aatgacaagt 480
ataataaatt tatgtatcgt gaaatgactg tagttaataa tacgataaac tattagtaac 540
tatgtccttt ttagtatatt gtgatactct gaaattgtga aaaaataggg tttatgttaa 600
atttttctgg aaagatggga actttagagt atgaaaagat gaatttttgg ggggtttgtt 660
tgaaaaattg atgattgaag tgtgggagct aagattttcg acaagggaat tcaaaatata 720
gagaagtaaa tgcacaaggc ctacgggatt cagtaactag aacaacaacg tacctagtgt 780
aattccattt gtggggtttg ggaagggcaa gatatatgca aacgcaagat cgtatcgtac 840
ctttgcaggg ttgagagtct atttctaata gaccatcggc ttgacatatg agaaaagata 900
tatctactat ttatgcatga agtatattat ttacatacaa aatagttttg accttgtgca 960
tactgtgaaa tttttggtga tgaattccct tgcagcatcc tagctccgcc ctggtcaatg 1020
ttccaattgt ttccttggaa gttataccaa acaaaagatt gacagaggat gttagaagta 1080
tatggcatat gccaatctac ttgaatgtga aactgataat atgacctttt cactttggaa 1140
tttgacaatg ttcaagactt agttggtgct atactactat gttataacga taggtacatt 1200
tgttaagata ttactttagc aaggtagtgg aattagatct ttctttggat gtagactcgg 1260
tatgaattag gataactatt gagttgaagt tgagagagta cttgaagttg atgttggaaa 1320
aagatagtgg aagacttgaa gttagtcttt atgtttcgaa atgaaattca tttggaaaga 1380
aagtttgtga atgttgtgag tagaagtcaa acacttgaat tgccttgctt ctataataaa 1440
ctttcaaacc actactacaa gattttattt attgttcaaa taagacttca aggccaaact 1500
actatttgta aataattatg tcatgtctga ccgggctaaa atggtgaaag tgtgttctgc 1560
aatggagcca agggctgtga tattgtttgc tcagtagtct ctagtatctg gttggtaggt 1620
atgctttgaa agtagtataa agataaaagg ctgccagttt ggtggtctat accattcctt 1680
gttgtttctt ttctatccca atccattttg agataagcag ctaatgaaac atcgaccccc 1740
ccggattttc ctgtttattt gtgtcagaag gcccgcacac acctatatac gcggatgtaa 1800
atgaatgtat gcaatgatct ttccttttct gttaaacaat gttcattagc tatcatgtgt 1860
tgtattggct ctctgctgac aatgaaataa ctgtacagat gcatttggac agttcatatt 1920
ctaattgaaa agaaaaagta tatttttcct ttggttaacg agtgtgagat gcttttttat 1980
ttcaataaaa tgagtagcta ttagcatctc atggaaatcg catatttaca gattgcctgt 2040
atttcaatag caagtcaagg ggcttacttt tcacttcata tcacagactc tcttcaaatg 2100
gtagtattag gttttaacaa tataatgttt ctattttcag gcacatcgag tagctcacaa 2160
actttggact caatcccgaa ggccacttgc acttgcactt caatcccgaa tctctgatgt 2220
ttttgctgtt gacattcatc cagctgccaa aatcggaaaa ggcatcctct tcgaccatgc 2280
aacaggagtg gttgttggtg aaactgcagt tattggaaac aatgtgtcga ttcttcacca 2340
tgtaacctta ggaggaactg gtaagtttgg tggtgaccga caccctaaga ttggtgacgg 2400
agtcctcata ggtgcaggtg ccacgatatt aggcaacata aatatcggtg agggagccaa 2460
gattggcgct ggatcagtgg ttctgattga cgtgccacca cgaacaactg cagttggaaa 2520
tccagctagg ttggtgggag ggaaagaaca gccaactaag cacgaggaat gtcccggaga 2580
gagtatggac catacatctt tcatatctgg atggtctgat tacatcatct ga 2632
<210> 2
<211> 293
<212> PRT
<213> 番茄(Solanum lycopersicum)
<400> 2
Met Pro Ala Glu Glu His Arg Asn Ala Ser Pro Ala Ala Pro His Pro
1 5 10 15
Pro Thr Asp Thr Ala Glu Glu Ala Ile Trp Leu Trp Thr Gln Ile Lys
20 25 30
Ala Glu Ala Arg Arg Asp Ala Glu Ala Glu Pro Ala Leu Ala Ser Tyr
35 40 45
Leu Tyr Ser Thr Ile Leu Ser His Ser Ser Leu Glu Arg Ser Leu Ser
50 55 60
Phe His Leu Gly Asn Lys Leu Cys Ser Ser Thr Leu Leu Ser Thr Leu
65 70 75 80
Leu Tyr Asp Leu Phe Leu Asn Asn Phe Ser Ser Asp Pro Asp Leu Arg
85 90 95
Ala Ala Ala Ser Ala Asp Leu Leu Ala Ala Arg Tyr Arg Asp Pro Ala
100 105 110
Cys Val Ser Phe Ser His Cys Leu Leu Asn Tyr Lys Gly Phe Leu Ala
115 120 125
Cys Gln Ala His Arg Val Ala His Lys Leu Trp Thr Gln Ser Arg Arg
130 135 140
Pro Leu Ala Leu Ala Leu Gln Ser Arg Ile Ser Asp Val Phe Ala Val
145 150 155 160
Asp Ile His Pro Ala Ala Lys Ile Gly Lys Gly Ile Leu Phe Asp His
165 170 175
Ala Thr Gly Val Val Val Gly Glu Thr Ala Val Ile Gly Asn Asn Val
180 185 190
Ser Ile Leu His His Val Thr Leu Gly Gly Thr Gly Lys Phe Gly Gly
195 200 205
Asp Arg His Pro Lys Ile Gly Asp Gly Val Leu Ile Gly Ala Gly Ala
210 215 220
Thr Ile Leu Gly Asn Ile Asn Ile Gly Glu Gly Ala Lys Ile Gly Ala
225 230 235 240
Gly Ser Val Val Leu Ile Asp Val Pro Pro Arg Thr Thr Ala Val Gly
245 250 255
Asn Pro Ala Arg Leu Val Gly Gly Lys Glu Gln Pro Thr Lys His Glu
260 265 270
Glu Cys Pro Gly Glu Ser Met Asp His Thr Ser Phe Ile Ser Gly Trp
275 280 285
Ser Asp Tyr Ile Ile
290
<210> 3
<211> 27
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 3
gctggatcca tgccagccga agaacac 27
<210> 4
<211> 29
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 4
gacgtcgact cagatgatgt aatcagacc 29
<210> 5
<211> 29
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 5
cgcggatcca tgccagccga agaacaccg 29
<210> 6
<211> 30
<212> DNA
<213> 人工序列(Artificial Sequence)
<400> 6
acgcgtcgac gatgatgtaa tcagaccatc 30
<210> 7
<211> 882
<212> DNA
<213> 番茄(Solanum lycopersicum)
<400> 7
atgccagccg aagaacaccg taacgcctcg ccggcggcgc cgcatccacc gacggacacg 60
gcagaagaag ctatctggtt atggacacag atcaaagcag aagctcggcg ggatgcagaa 120
gctgaaccgg cattagcaag ctatttgtac tcaactatac tatctcactc ttcgcttgag 180
cgttcactct cttttcattt ggggaacaag ctttgttctt caacgcttct atctactctt 240
ctctacgatc tgttcctcaa taatttctcc tctgatcctg atctacgcgc cgcggcatcc 300
gctgacctac tcgccgctcg ctaccgtgac ccagcctgtg tttccttctc tcattgtttg 360
cttaattaca aaggcttcct tgcttgtcag gcacatcgag tagctcacaa actttggact 420
caatcccgaa ggccacttgc acttgcactt caatcccgaa tctctgatgt ttttgctgtt 480
gacattcatc cagctgccaa aatcggaaaa ggcatcctct tcgaccatgc aacaggagtg 540
gttgttggtg aaactgcagt tattggaaac aatgtgtcga ttcttcacca tgtaacctta 600
ggaggaactg gtaagtttgg tggtgaccga caccctaaga ttggtgacgg agtcctcata 660
ggtgcaggtg ccacgatatt aggcaacata aatatcggtg agggagccaa gattggcgct 720
ggatcagtgg ttctgattga cgtgccacca cgaacaactg cagttggaaa tccagctagg 780
ttggtgggag ggaaagaaca gccaactaag cacgaggaat gtcccggaga gagtatggac 840
catacatctt tcatatctgg atggtctgat tacatcatct ga 882
Claims (1)
1.番茄SlSERAT1;1基因或其片段促进番茄植株半胱氨酸合成的应用，其特征在于，所述番茄SlSERAT1;1基因的核苷酸序列如SEQ ID No.1所示，所述片段为番茄SlSERAT1;1基因的cDNA序列，其核苷酸序列如SEQ ID No.7所示。
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