CN107254478A - 番茄sllcd基因及其应用 - Google Patents
番茄sllcd基因及其应用 Download PDFInfo
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- CN107254478A CN107254478A CN201710488701.8A CN201710488701A CN107254478A CN 107254478 A CN107254478 A CN 107254478A CN 201710488701 A CN201710488701 A CN 201710488701A CN 107254478 A CN107254478 A CN 107254478A
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Abstract
本发明提供一种番茄SLLCD编码区基因，以及该基因编码的番茄SLLCD蛋白，该基因的编码区核苷酸序列如SEQ ID No.1所示。本发明的优点在于，本发明的番茄SLLCD编码区基因，其属于内源H2S产生直接相关的酶编码基因，在番茄中抑制SLLCD基因的表达能够明显影响到番茄果实中种子的发育，产生没有种子或种子发育不完全的转基因植株；可用于无籽番茄的获得以及果实中有益物质的保持；而且，在SLLCD转基因植株中，果实发育不受影响，因此可以保持番茄果实的优良性状；说明SLLCD在发育过程中具有良好的应用价值。
Description
技术领域
本发明涉及基因工程领域，具体地说，涉及一种番茄内源H2S产生直接相关的酶编码基因及其在产生无籽番茄过程中的应用。
背景技术
种子形成是植物发育过程中的重要过程，有性繁育的植物经过授粉和受精过程，由胚珠发育而来。同时可以利用植物的繁殖方式和遗传变异特点来保持其优良品种。无籽是一个优良的果实性状。无籽果实品质优良，可食率高，更易吸引消费者购买。无籽果实的产生可以提高果实品质，达到较高的无核率，增加果粒的大小，从而最大程度保持果实中的有益物质，获得优良的蔬果品种，提高市场售价和种植收益，有利于保护遗传改良的作物特别是一年生作物品种不被改变。
番茄营养丰富，含有丰富的胡萝卜素，B族维生素，维生素C和番茄红素，可以生食，煮食，也可以加工成番茄酱，番茄汁等产品。番茄中的茄红素是一种很强的抗氧化剂，具有极强的清除自由基的能力，对防治前列腺癌，乳腺癌等有显著效果，还有预防心脑血管疾病，提高免疫力和延缓衰老等功效，是自然界中最强的抗氧化剂之一。其抗氧化作用是β胡萝卜素的2倍，是维生素E的100倍。同时，番茄无籽果实的可溶性固形物和糖的含量要比有籽品种高，而酸与纤维素的含量则低于有籽品种。
关于无籽番茄的获得，通常是为果实施用一定浓度的植物激素，如在授粉前人工涂抹生长素或者生长素类似物，促进子房发育成果实，即抑制种子发育的同时促进果实发育，同时，其细胞内的遗传物质并没有发生变化。随着分子生物学技术日新月异的发展，基因工程技术也得到了飞速的发展，通过基因编辑方法获得无籽果实已成为可能。H2S较早被证实可以打破种子休眠，促进种子萌发。LCD是植物中以L-Cys为底物，产生内源H2S直接相关的酶编码基因(Papenbrock et al.，2007)，与动物的CSE属一类。番茄编码区基因SLLCD突变体中，果实正常发育，产生无籽表型，且番茄叶和心皮等器官中SLLCD基因表达水平下降，说明抑制SLLCD基因有利于无籽番茄的获得。目前，LCD与无籽果实获得的机理仍未报道，值得被进一步研究。
植物体内H2S的生成酶比动物细胞丰富，定位于细胞质，线粒体和叶绿体等多个亚细胞部位，表达具有时空性。内源产生的H2S参与了植物体生长发育和逆境胁迫响应等过程，同时发现H2S和活性氧，NO，CO，Ca2+，以及脱落酸，茉莉酸，赤霉素，生长素，乙烯等植物激素信号路径存在相互作用。植物体内以Cys为底物产生H2S的酶有多种，目前对这些酶类研究最深入的植物是拟南芥，已经被鉴定出的酶类根据其功能主要分为4大类。即半胱氨酸脱巯基酶，半胱氨酸脱硫酶，β-氰丙氨酸合酶和O-乙酰丝氨酸硫醇裂解酶。鉴于L-Cys是植物体内Cys的主要存在形式，推测以L-Cys为底物的CDes是内源H2S生成的主要贡献者。最新的研究表明，H2S可以对蛋白质的活性Cys残基进行巯基化修饰，这可能是其参与各项生命活动的重要作用方式之一(Pei，2016)。目前，其他内源H2S产生相关的酶基因，在番茄果实成熟和种子发育等过程中的功能还未知。
发明内容
本发明的目的在于提供番茄SLLCD编码基因；
本发明的另一目的在于提供所述基因所编码的番茄SLLCD蛋白；
本发明的目的还在于提供含有所述基因或其片段的载体及其宿主细胞；
本发明的最终目的在于提供所述基因或其片段在调节番茄种子发育中的应用。
为了实现上述目的，本发明提供番茄SLLCD编码区基因，该基因是从番茄MicroTom(以下简称MT)中克隆得到的，属于内源H2S产生直接相关的酶编码基因，具有如SEQ IDNo.1所示的核苷酸序列。
本发明还提供上述基因编码的番茄SLLCD蛋白，其为：
1)由SEQ ID No.2所示的氨基酸序列组成的蛋白；或
2)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蛋白。
序列分析结果显示，番茄SLLCD基因属于L型半胱氨酸脱巯基酶(以下简称LCD)编码基因，由该基因所编码的蛋白可以以L型半胱氨酸为底物产生内源H2S。系统进化分析显示，由该基因所编码的蛋白同拟南芥中LCD的相似性为63％；番茄SLLCD基因在萼片、花瓣、雄蕊等器官中表达较弱，在叶片中表达较强，通过精细分析后发现，该基因在心皮中的表达最强。
应当理解，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。本发明的番茄SLLCD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SLLCD蛋白同等活性的由番茄SLLCD蛋白衍生得到的蛋白。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。
本发明还提供含有上述番茄SLLCD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SLLCD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SLLCD基因cDNA的一段3’端序列，其核苷酸序列如SEQ ID No.5所示。
具体地说，本发明将番茄SLLCD基因cDNA的一段3’端序列(400bp，其核苷酸序列如SEQ ID No.5所示)首先反向构建到中间载体pKANNIBAL中，最终构建到双元载体pART27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pART27携带的SLLCD编码区基因cDNA的3’端序列转入番茄，获得番茄转化植株。
本发明的番茄SLLCD编码区基因在调节番茄种子发育中的应用。所述应用是指通过抑制番茄SLLCD编码区基因的表达，产生没有种子或种子发育不完全的转基因植株，从而保护各种番茄作物的编码区基因优势以及番茄果实的优良性状。
本发明的优点在于，本发明的番茄SLLCD基因，其属于内源H2S产生直接相关的酶编码基因，在番茄中抑制SLLCD编码区基因的表达能够明显影响到番茄种子的发育，产生没有种子或种子发育不完全的转基因植株；可用于保护各种番茄作物的编码区基因优势；而且，在SLLCD转基因植株中，果实发育不受影响，因此可以保持番茄果实的优良性状；说明SLLCD在发育过程中具有良好的应用价值。
附图说明
图1是本发明的番茄SLLCD编码蛋白的结构分析；
图2是本发明实施例1的克隆载体pET28a；
图3是本发明实施例3的克隆中间载体pKANNIBAL；
图4是本发明实施例3的植物表达载体pART27；
图5是本发明实施例4的番茄SLLCD基因影响番茄种子发育图；
图6是本发明实施例4的番茄SLLCD转基因植株中的无籽表型图；
图7是本发明实施例5的番茄SLLCD基因的表达谱；其中，SD，幼苗；R，根；S，茎；L，叶；Se，萼片；Pe，花瓣；Sta，雄蕊；Ca，心皮；IMG，未成熟绿果；MG，成熟绿果；B，破果期果实，Rf，成熟期果实。
图8是本发明实施例6的番茄SLLCD基因在转基因植株中的表达水平；其中，1，2，4代表不同的转基因株系；
图9是本发明实施例7的番茄SLLCD基因在转基因植株中的表达水平；其中，1，2，4，5，6，7代表不同的转基因株系。
具体实施方式
以下结合附图和实施例，进一步详细说明本发明。以下实施例用于说明本发明，但不用来限制本发明的范围。
实施例1番茄SLLCD编码区基因的克隆
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MT的心皮cDNA中克隆SLLCD基因的编码区序列；
PCR程序：94℃，5分钟；94℃，30秒；55℃，30秒；72℃，1分30秒；重复35次；72℃，10分钟。
将上述PCR产物通过双酶切的方法连接到克隆载体pET28a上(如图2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番茄SLLCD基因的编码区cDNA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 IDNo.2所示。
实施例2番茄SLLCD蛋白的序列结构分析
番茄SLLCD具有产生内源H2S的功能，同拟南芥中LCD编码区蛋白序列的相似性为63％。结果如图1所示。
实施例3番茄SLLCD基因表达载体及转基因植株
番茄内源H2S产生酶相关基因SLLCD的一段3’端序列(400bp，其核苷酸序列如SEQID No.5所示)，反向构建在图4所示的中间载体pART27中，用于利用RNA干扰的原理在番茄中抑制SLLCD的表达水平，进而研究其功能。
通过农杆菌介导转化方法，将pART27携带的SLLCD基因cDNA的3’端序列转入番茄，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％乙醇清洗，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℃培养间长光照(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(Whatman Filter Paper)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-80℃存放的农杆菌(Agrobacterium tumefaciens，EHA105)，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CI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SI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，就可转移至装有蛭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：
实施例4番茄SLLCD基因对种子发育的影响
将实施例3获得的共7株番茄转化植株与其野生型对照(MT)植株在培养间中同时一起培育(16小时光照，22℃/8小时黑暗，20℃的光周期)，直至完成整个生命周期。7株番茄转化植株共表现出如图5，图6所示的2种表型，其中4株如图5所示，即在番茄中抑制SLLCD基因的表达，导致种子不完全发育甚至消失，(左侧A图为野生型对照(MT)，右侧B图为实施例3获得的转基因植株)。上述结果说明：番茄SLLCD参与果实发育，控制种子的形成。
由于不同转基因植株中SLLCD被抑制的程度不同，所以转基因植株的表型会有程度上的差异(没有种子或种子发育不完全)，这可能与外源基因在番茄基因组上的插入位点有关系，运用农杆菌转化的方法来转化植物，外源基因的插入位点是随机的。同时这种转基因植株的表型方式是应用RNA干扰的方法来沉默基因的一种普遍的表型表现方式，更说明了SLLCD编码区参与种子的发育。
实施例5
通过实时定量荧光PCR(quantitative real time qRT-PCR)测定番茄SLLCD基因在各器官中的表达谱。结果如图7所示。
结果表明：番茄SLLCD基因在心皮中高水平表达。
实施例6
利用RT-PCR测定SLLCD反义转基因株系中SLLCD的表达量，结果如图8所示。
实施例7
利用qRT-PCR测定SLLCD反义转基因株系中SLLCD的表达量，结果如图9所示。
结果表明：番茄中SLLCD基因在SLLCD反义转基因植株中表达量下降。
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。
SEQUENCE LISTING
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Claims (7)
1.番茄SLLCD编码区基因，其核苷酸序列如SEQ ID No.1所示。
2.权利要求1所述基因编码的番茄SLLCD蛋白，其为：
1)由SEQ ID No.2所示的氨基酸序列组成的蛋白；或
2)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蛋白。
3.含有权利要求1所述基因或其片段的载体。
4.如权利要求3所述基因片段的载体，其特征在于，所述片段为番茄SLLCD基因cDNA的一段3’端序列，其核苷酸序列如SEQ ID No.5所示。
5.含有权利要求3所述载体的宿主细胞。
6.如权利要求1所述基因或其片段在调节番茄种子发育中的应用。
7.如权利要求6所述的应用，其特征在于，所述应用是通过抑制番茄SLLCD编码区基因的表达，产生没有种子或种子发育不完全的转基因植株。
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| CN108893481A (zh) * | 2018-06-21 | 2018-11-27 | 山西大学 | 番茄SlOAS7基因及其应用 |
| CN108893481B (zh) * | 2018-06-21 | 2021-07-02 | 山西大学 | 番茄SlOAS7基因及其应用 |
| WO2020120242A1 (en) * | 2018-12-13 | 2020-06-18 | Nunhems B.V. | Solanaceous plant capable of stenospermocarpic fruit formation |
| CN110079537A (zh) * | 2019-05-27 | 2019-08-02 | 河南科技大学 | 葡萄细胞分裂素响应调节因子VvRR基因及其编码蛋白和应用 |
| CN113563439A (zh) * | 2021-07-15 | 2021-10-29 | 浙江省农业科学院 | 一种果形发育相关蛋白及其编码基因与应用 |
| CN113462706A (zh) * | 2021-08-04 | 2021-10-01 | 合肥工业大学 | 一种增加番茄果实重量及心室数目的基因及其调控方法 |
| CN113462706B (zh) * | 2021-08-04 | 2024-04-26 | 合肥工业大学 | 一种增加番茄果实重量及心室数目的基因及其调控方法 |
| CN113430213A (zh) * | 2021-08-13 | 2021-09-24 | 合肥工业大学 | 一种调控番茄侧枝的基因及方法 |
| CN114214341A (zh) * | 2021-12-30 | 2022-03-22 | 山西大学 | 番茄SlSERAT1;1基因或其片段在植物发育过程中的应用 |
| CN114214341B (zh) * | 2021-12-30 | 2023-12-26 | 山西大学 | 番茄SlSERAT1;1基因或其片段在植物发育过程中的应用 |
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