CN114369147A - Bfne基因在番茄株型改良和生物产量提高中的应用 - Google Patents
Bfne基因在番茄株型改良和生物产量提高中的应用 Download PDFInfo
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- CN114369147A CN114369147A CN202111231381.0A CN202111231381A CN114369147A CN 114369147 A CN114369147 A CN 114369147A CN 202111231381 A CN202111231381 A CN 202111231381A CN 114369147 A CN114369147 A CN 114369147A
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Abstract
本发明公开了BFNE基因在番茄株型改良和生物产量提高中的应用。本发明公开的BFNE基因的CDS序列如序列表中的序列2所示，基因组序列如序列表中的序列3所示，其编码的氨基酸序列如序列表中的序列1所示。本发明还公开了BFNE蛋白质或其相关生物材料在如下1)‑4)任一种中的应用：1)调控番茄株型；2)调控番茄产量；3)培养转基因番茄或番茄育种；4)鉴定或区分野生番茄和栽培番茄。
Description
技术领域
本发明属于植物基因工程领域，具体涉及BFNE基因在番茄株型改良和生物产量提高中的应用。
背景技术
农业是我国的基础产业，粮食生产更是重中之重。粮食是人类生存的基本条件，随着人口的不断增加，耕地的逐渐减少，粮食问题、农业发展问题不断突出。利用高新技术育种农业科技攻关，大力培养优良品种，提高单位面积作物的产量，是当前增加产量最直接最有效的途径。地球的耕地是有限的。随着世界人口的不断增长，人均耕地越来越少了。近50年来，世界人均耕地的数量竟减少了一半以上。全国的人均耕地面积都不乐观，18亿亩红线非常重要。相比内地，新疆的耕地面积算大的，但是耕地面积不一定就代表产量，受限于各种各样的原因，如种植技术、土地盐碱度大、作物品种等原因。
番茄(Solanum lycopersicum)是重要的蔬菜和经济作物，在世界范围内广受欢迎。我国是鲜食番茄第一大生产国，也是加工番茄的第三大生产国，在世界番茄市场中具有举足轻重的地位。由于气候原因，新疆一年番茄只能种一季。番茄株型和果实数量均为影响番茄产量的重要因子，解析株型和产量的遗传基础无疑为分子育种提供重要基因资源。
发明内容
本发明首先提供了一种蛋白质，本发明提供的蛋白质的名称为BFNE，其为a1)或a2)或a3)或a4)：
a1)氨基酸序列是序列1所示的蛋白质；
a2)在序列1所示的蛋白质的N端或/和C端连接标签得到的融合蛋白质；
a3)将序列1所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的与植物耐逆性相关的蛋白质；
a4)将序列1所示的氨基酸序列具有90％同一性、来源于番茄且与番茄株型或产量相关的蛋白质。
上述a2)所述的蛋白质中，所述标签是指利用DNA体外重组技术，与目的蛋白一起融合表达的一种多肽或者蛋白，以便于目的蛋白的表达、检测、示踪和/或纯化。所述标签可为Flag标签、His标签、MBP标签、HA标签、myc标签、GST标签和/或SUMO标签等。
上述a3)所述的蛋白质中，所述一个或几个氨基酸残基的取代和/或缺失和/或添加为不超过10个氨基酸残基的取代和/或缺失和/或添加。
上述a4)所述的蛋白质中，“同一性”包括与本发明的序列1所示的氨基酸序列具有90％或更高，或91％或更高，或92％或更高，或93％或更高，或94％或更高，或95％或更高，或96％或更高，或97％或更高，或98％或更高，或99％或更高同源性的氨基酸序列。
上述a1)或a2)或a3)或a4)所述的蛋白质可人工合成，也可先合成其编码基因，再进行生物表达得到。
本发明又提供了与BFNE蛋白质相关的生物材料，该生物材料为下述A1)至A8)中的任一种：
A1)编码BFNE蛋白质的核酸分子；
A2)含有A1)所述核酸分子的表达盒；
A3)含有A1)所述核酸分子的重组载体；
A4)含有A2)所述表达盒的重组载体；
A5)含有A1)所述核酸分子的重组微生物；
A6)含有A2)所述表达盒的重组微生物；
A7)含有A3)所述重组载体的重组微生物；
A8)含有A4)所述重组载体的重组微生物。
上述生物材料中，A1)所述核酸分子为如下1)或2)或3)所示的基因：
1)其编码序列是序列3所示的基因组DNA分子或序列2所示的cDNA分子；
2)与1)限定的核苷酸序列具有75％或75％以上同一性，且编码BFNE蛋白质的cDNA分子或基因组DNA分子；
3)在严格条件下与1)或2)限定的核苷酸序列杂交，且编码BFNE蛋白质的cDNA分子或基因组DNA分子。
其中，所述核酸分子可以是DNA，如cDNA、基因组DNA或重组DNA；所述核酸分子也可以是RNA，如mRNA或hnRNA等。
本领域普通技术人员可以很容易地采用已知的方法，例如定向进化和点突变的方法，对本发明的编码BFNE蛋白质的核苷酸序列进行突变。那些经过人工修饰的，具有编码BFNE蛋白质的核苷酸序列75％或者更高同一性的核苷酸，只要编码BFNE蛋白质且具有相同功能，均是衍生于本发明的核苷酸序列并且等同于本发明的序列。
这里使用的术语“同一性”指与天然核酸序列的序列相似性。“同一性”包括与本发明的编码序列1所示的氨基酸序列组成的蛋白质的核苷酸序列具有75％或更高，或85％或更高，或90％或更高，或95％或更高同一性的核苷酸序列。同一性可以用肉眼或计算机软件进行评价。使用计算机软件，两个或多个序列之间的同一性可以用百分比(％)表示，其可以用来评价相关序列之间的同一性。
上述75％或75％以上同一性，可为80％、85％、90％或95％以上的同一性。
上述生物材料中，所述严格条件是在2×SSC，0.1％SDS的溶液中，在68℃下杂交并洗膜2次，每次5min，又于0.5×SSC，0.1％SDS的溶液中，在68℃下杂交并洗膜2次，每次15min；或，0.1×SSPE(或0.1×SSC)、0.1％SDS的溶液中，65℃条件下杂交并洗膜。
上述生物材料中，A2)所述的含有编码BFNE蛋白质的核酸分子的表达盒(BFNE基因表达盒)是指能够在宿主细胞中表达BFNE蛋白质的DNA，该DNA不但可包括启动BFNE转录的启动子，还可包括终止BFNE转录的终止子。进一步，所述表达盒还可包括增强子序列。可用于本发明的启动子包括但不限于：组成型启动子；组织、器官和发育特异的启动子及诱导型启动子。合适的转录终止子包括但不限于：农杆菌胭脂碱合成酶终止子(NOS终止子)、花椰菜花叶病毒CaMV 35S终止子、tml终止子、豌豆rbcS E9终止子和胭脂氨酸和章鱼氨酸合酶终止子。
可用现有的表达载体构建含有所述BFNE基因表达盒的重组载体。所述植物表达载体包括双元农杆菌载体和可用于植物微弹轰击的载体等。如pAHC25、pBin438、pCAMBIA1302、pCAMBIA2301、pCAMBIA1301、pCAMBIA1300、pBI121、pCAMBIA1391-Xa或pCAMBIA1391-Xb等。所述植物表达载体还可包含外源基因的3′端非翻译区域，即包含聚腺苷酸信号和任何其它参与mRNA加工或基因表达的DNA片段。所述聚腺苷酸信号可引导聚腺苷酸加入到mRNA前体的3′端，如农杆菌冠瘿瘤诱导(Ti)质粒基因(如胭脂碱合成酶基因Nos)、植物基因(如大豆贮存蛋白基因)3′端转录的非翻译区均具有类似功能。使用本发明的基因构建植物表达载体时，还可使用增强子，包括翻译增强子或转录增强子，这些增强子区域可以是ATG起始密码子或邻接区域起始密码子等，但必需与编码序列的阅读框相同，以保证整个序列的正确翻译。所述翻译控制信号和起始密码子的来源是广泛的，可以是天然的，也可以是合成的。翻译起始区域可以来自转录起始区域或结构基因。为了便于对转基因植物细胞或植物进行鉴定及筛选，可对所用植物表达载体进行加工，如加入可在植物中表达的编码可产生颜色变化的酶或发光化合物的基因(GUS基因、萤光素酶基因等)、抗生素的标记基因(如赋予对卡那霉素和相关抗生素抗性的nptII基因，赋予对除草剂膦丝菌素抗性的bar基因，赋予对抗生素潮霉素抗性的hph基因，和赋予对氨甲喋呤抗性的dhfr基因，赋予对草甘磷抗性的EPSPS基因)或是抗化学试剂标记基因等(如抗除莠剂基因)、提供代谢甘露糖能力的甘露糖-6-磷酸异构酶基因。从转基因植物的安全性考虑，可不加任何选择性标记基因，直接以逆境筛选转化植株。
上述生物材料中，所述载体可为质粒、黏粒、噬菌体或病毒载体。
上述生物材料中，所述微生物可为酵母、细菌、藻或真菌，如农杆菌。
本发明还提供了上述BFNE蛋白质或上述生物材料的新用途。
本发明提供了上述BFNE蛋白质或上述生物材料在如下1)-4)任一种中的应用：
1)调控番茄株型；
2)调控番茄产量；
3)培养转基因番茄或番茄育种；
4)鉴定或区分野生番茄和栽培番茄；
进一步的，所述番茄株型具体为番茄分枝。所述番茄产量具体包括番茄果实数量和果实重量。
更进一步的，所述调控番茄株型为增加番茄分枝，所述调控番茄产量为提高番茄产量。具体体现为：番茄中的BFNE蛋白质的含量和/或活性越高或BFNE基因表达水平越高，番茄的分枝数越多，番茄的果实数量和重量越多。
本发明还提供了一种转基因番茄的培育方法。
本发明提供的转基因番茄的培育方法包括如下步骤：提高受体番茄中BFNE蛋白质的含量和/或活性，得到转基因植物；所述转基因番茄的分枝数和/或产量高于所述受体番茄。
上述方法中，所述提高受体植物中BFNE蛋白质的含量和/或活性的方法为在受体番茄中过表达BFNE蛋白质。
进一步的，所述过表达的方法为将BFNE蛋白质的编码基因导入受体植物中。
更进一步的，所述BFNE蛋白质的编码基因为序列表中的序列2所示。
本发明最后提供了一种鉴定或区分野生番茄和栽培番茄的方法。
本发明提供的鉴定或区分野生番茄和栽培番茄的方法包括如下步骤：检测待测番茄是否含有BFNE蛋白质或其编码基因：若待测番茄含有BFNE蛋白质(序列1)或其编码基因(序列3)，则待测番茄为野生番茄；若待测番茄不含有BFNE蛋白质或其编码基因(含有序列4所示的蛋白质或序列5所示的基因)，则待测番茄为栽培番茄。
进一步的，检测待测番茄是否含有BFNE蛋白质或其编码基因的方法包括如下步骤：提取待测番茄的基因组DNA，采用序列6所示核苷酸序列和序列7所示核苷酸序列进行扩增，得到扩增产物，然后电泳检测扩增产物，若扩增得到大小为777bp的扩增产物条带，则待测番茄品种为野生番茄，若扩增得到大小为534bp的扩增产物条带，则待测番茄品种为栽培番茄。
更进一步的，所述野生番茄的品种可为如下中的任一种：类番茄Solanumlycopersicoides、多毛番茄Solanum habrochaites、潘那利番茄Solanum pennellii、智利番茄Solanum chilense、秘鲁番茄Solanum peruvianum、多腺番茄Solanumcorneliomulleri、小花番茄Solanum neorickii、克梅留斯基Solanum chmielewskii、醋栗Solanum pimpinellifolium、加拉帕戈斯Solanum galapagense。
所述栽培番茄的品种可为如下中的任一种：樱桃番茄Solanum lycopersicumvar.cerasiforme、栽培番茄M82 Solanum lycopersicum、栽培番茄Heinz-1706 Solanumlycopersicum。
本发明首先利用泛基因组挖掘出一个野生番茄基因，并将其命名为BFNE(branchand fruit number enhencer gene)基因，然后本发明将BFNE基因超表达于番茄中，得到转BFNE基因番茄。通过实验证明：转BFNE基因番茄表现为分枝数增加，果实数量与重量均增多。说明BFNE基因可改良番茄株型、提高番茄产量。最后本发明还发现BFNE基因还可用于鉴定或区分野生番茄和栽培番茄，并用于野生番茄和栽培番茄杂交育种与番茄品种改良。
本发明的优点在于：
(1)本发明利用番茄泛基因组数据，进行多基因组比对，直接挖掘克隆一个新基因BFNE。本发明操作省时高效，避免了传统的QTL基因定位方法耗时耗力的缺点。
(2)本发明挖掘克隆一个影响番茄分枝和生物产量的多效基因BFNE。BFNE的功能为本发明首次报道，具有极高的首创性。该基因调控番茄的分枝和生物产量，对进一步改良番茄理想品种奠定基础。
附图说明
图1为番茄泛基因组挖掘到的存在/缺失变异PAV多效性基因BFNE示意图。
图2为BFNE基因在野生番茄和栽培番茄各个组织的表达量。
图3为转BFNE基因番茄的鉴定。A为PCR鉴定。M：2000；1500；1000；750；500；250；100。1-5分别为加拉帕戈斯Solanum galapagense、小花番茄Solanum neorickii、栽培番茄M82 Solanum lycopersicum、麦克汤姆小番茄(Micro Tom小番茄)和转BFNE基因番茄植株。B为BFNE基因表达水平鉴定。
图4为转BFNE基因番茄和野生植株的株型对照图。
图5为转BFNE基因番茄和野生植株的生物产量对照图。
图6为转BFNE基因番茄和野生植株的红果果重对比(单位为克)。
图7为PAV-F/PAV-R引物对在野生番茄和栽培番茄中的扩增结果。M：2000；1500；1000；750；500；250；100。1：类番茄Solanum lycopersicoides；2：多毛番茄Solanumhabrochaites；3：潘那利番茄Solanum pennellii；4：智利番茄Solanum chilense；5：秘鲁番茄Solanum peruvianum；6：多腺番茄Solanum corneliomulleri；7：小花番茄Solanumneorickii；8：克梅留斯基Solanum chmielewskii；9：醋栗Solanum pimpinellifolium；10：加拉帕戈斯Solanum galapagense；11：樱桃番茄Solanum lycopersicumvar.cerasiforme；12：栽培番茄M82 Solanum lycopersicum；13：栽培番茄Heinz-1706Solanum lycopersicum。
具体实施方式
以下的实施例便于更好地理解本发明，但并不限定本发明。下述实施例中的试验方法，如无特殊说明，均为常规方法。下述实施例中所用的试验材料，如无特殊说明，均为自常规生化试剂商店购买得到的。以下实施例中的定量试验，均设置三次重复实验，结果取平均值。
下述实施例中的类番茄Solanum lycopersicoides、多毛番茄Solanumhabrochaites、潘那利番茄Solanum pennellii、智利番茄Solanum chilense、秘鲁番茄Solanum peruvianum、多腺番茄Solanum corneliomulleri、小花番茄Solanum neorickii、克梅留斯基Solanum chmielewskii、醋栗Solanum pimpinellifolium、加拉帕戈斯Solanumgalapagense均记载于文献“Spooner DM,Peralta IE,Knapp S.Comparison of AFLPswith Other Markers for Phylogeneti:Inference in Wild Tomatoes[SolanumL.Section Lycopersicon(Mill.)Wettst.][J].Taxon,2005,54(1):43-61.”中，公众可从新疆农业科学院园艺作物研究所获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
下述实施例中的樱桃番茄Solanum lycopersicum var.cerasiforme记载于文献“Ranc N,
S,Santoni S,et al.A clarified position for solanum lycopersicumvar.cerasiforme in the evolutionary history of tomatoes(solanaceae)[J].BMCPlant Biology,2008,8.”中，公众可从新疆农业科学院园艺作物研究所获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
下述实施例中的栽培番茄M82 Solanum lycopersicum记载于文献“Carter J D,Pereira A,Veilleux D.An active ac/ds transposon system for activation taggingin tomato cultivar m82 using clonal propagation.[J].Plant Physiology,2013,162(1):145-156.”中，公众可从新疆农业科学院园艺作物研究所获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
下述实施例中的栽培番茄Heinz-1706 Solanum lycopersicum记载于文献“Aureliano B,Naama M,Tecle I Y,et al.The Sol Genomics Network(solgenomics.net):growing tomatoes using Perl[J].Nucleic Acids Research,2011,39(Database issue):1149-55.”中，公众可从新疆农业科学院园艺作物研究所获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
下述实施例中的Micro Tom小番茄记载于文献“Chetty V J,Ceballos N,GarciaD,et al.Evaluation of four Agrobacterium tumefaciens strains for the genetictransformation of tomato(Solanum lycopersicum L.)cultivar Micro-Tom[J].PlantCell Reports,2013,32(2):239-247.”中，公众可从新疆农业科学院园艺作物研究所获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
下述实施例中的植物表达载体pCAMBIA1300记载于文献“Das S S,Sanan M N.Adirect method for genetically transforming rice seeds modelled with FHVB2,asuppressor of RNAi[J].Plant Cell Tissue&Organ Culture,2015,120(1):277-289.”，公众可从新疆农业科学院园艺作物研究所获得，该生物材料只为重复本发明的相关实验所用，不可作为其它用途使用。
实施例1、BFNE基因的克隆及其在野生番茄和栽培番茄各个组织的表达量分析一、BFNE基因的获得
本发明利用番茄泛基因组数据，进行多基因组比对，直接挖掘克隆得到BFNE基因，具体步骤如下：
1、采用PacBio+Hi-C策略，组装了番茄属11个番茄(9个野生番茄和2个栽培番茄)高质量染色体级的基因组序列。
2、应用四种从头预测软件，包括RepeatScout、LTR-FINDER、MITE-hunter和PILER-DF，形成一个重复序列数据库。通过使用RepeatMasker对最终合并的数据库进行同源搜索，识别重复序列注释。
3、核心基因与非核心基因分析，包括功能富集分析、保守性分析、重复序列区域富集等。
4、基于基因组序列比对进行基因组SVs的检测，可以对SVs特征进行分析，如对SVs类型、长度分布、基因组分布、重复含量进行比较等。
5、最终找到野生番茄特有基因BFNE，结果如图1。
野生番茄中的特有基因BFNE的基因组序列如序列表中序列3所示，CDS序列如序列表中序列2所示，BFNE基因编码的蛋白的氨基酸序列如序列表中序列1所示。
二、BFNE基因在野生番茄和栽培番茄各个组织的表达量分析
采集野生番茄加拉帕戈斯和栽培番茄M82的根、茎、叶、幼苗组织样品，提取RNA反转录，使用LightCycler96实时PCR***进行定量RT-PCR。番茄Actin作为内参，在qRT-PCR实验中，每个处理三个样本(生物重复)。引物序列具体如下：
RTBF4-F：GTGATCATCGCGTTGCTGTT；
RTBF4-R：TCCAAGATTTGGTGTTGCTGC；
Actin-F：GGTGTTATGGTCGGAATGGG；
Actin-R：CAGGGTGTTCTTCAGGAGCAA。
结果如图2所示，结果表明：BFNE基因在野生番茄加拉帕戈斯根、茎、叶、幼苗中的表达量均高于栽培番茄M82，尤其是在叶中。
实施例2、BFNE基因在番茄株型改良和生物产量提高中的应用
一、转BFNE基因番茄的制备
1、重组表达载体的构建
通过酶切和连接，将序列2所示的多效性基因BFNE整合到植物表达载体pCAMBIA1300的XbaI和KpnI酶切位点间，且保持植物表达载体pCAMBIA1300的其它序列不变，得到pCAMBIA1300-BFNE载体。
2、转基因番茄植株的获得
将上述步骤1获得的pCAMBIA1300-BFNE载体导入GV3101农杆菌电转感受态细胞(北京博迈德基因技术有限公司，货号为BC308-01)中，经鉴定得到重组菌pCAMBIA1300-BFNE/GV3101，然后将重组菌pCAMBIA1300-BFNE/GV3101通过叶盘法转化到Micro Tom小番茄中，获得T0代转BFNE基因番茄植株。
3、转基因番茄植株的鉴定
将T0代转BFNE基因番茄植株种植于温室，自交获得T1代转BFNE基因番茄植株，并对其进行鉴定。
1)PCR鉴定
分别提取加拉帕戈斯Solanum galapagense、小花番茄Solanum neorickii、栽培番茄M82 Solanum lycopersicum、麦克汤姆小番茄(Micro Tom小番茄)和T1代转BFNE基因番茄植株的基因组DNA，采用BFNE-F：GCTGCTAAACAACATCCAGAAGAG和BFNE-R：TCCGCAGACGAGACAATGA进行PCR扩增，并电泳检测PCR扩增产物。
结果如图3A所示。从图中可以看出：加拉帕戈斯Solanum galapagense、小花番茄Solanum neorickii和转BFNE基因番茄植株中均扩增得到大小为186bp的条带，而栽培番茄M82 Solanum lycopersicum和麦克汤姆小番茄(Micro Tom小番茄)中均没有扩增出条带。
2)BFNE基因表达量检测
通过RT-PCR分别检测经PCR鉴定为阳性的T1代转BFNE基因番茄植株的根、茎和叶中的BFNE基因表达水平，同时以Micro Tom小番茄作为对照。BFNE基因特异引物序列如下：RTBF1-F：TAGTTGCAGCAATGGGCACT；RTBF1-R：TTCCATTGCCTCGTGAGGTG。内参基因引物序列如下：Actin-F：GGTGTTATGGTCGGAATGGG；Actin-R：CAGGGTGTTCTTCAGGAGCAA。
结果如图3B所示。从图中可以看出：与野生型Micro Tom小番茄相比，T1代转BFNE基因番茄植株茎和叶中的BFNE基因表达水平均显著提高。
二、转BFNE基因番茄的株型与产量分析
将T1代转BFNE基因番茄植株和野生型植株(Micro Tom小番茄)同时种植在温室，于成熟期观察表型，并统计分枝数和果实数量与重量。
结果表明：转BFNE基因番茄分枝(侧枝)增多，而野生型植株无分枝(表1、图4)。而且转BFNE基因番茄的单株总果数量和总果重均高于野生型番茄(表2、图5和图6)。
表1、转BFNE基因番茄与野生型番茄的分枝数
注：重复一、重复二、重复三分别表示WT和转BFNE基因番茄分别选取的三株重复单株。
表2、转BFNE基因番茄与野生型番茄的果实数量与总重
注：BFNE-1、BFNE-2和BFNE-3均为T1代转BFNE基因番茄植株；WT-1、WT-2和WT-3均为野生番茄(Micro Tom小番茄)；
1、2和3表示T1代转BFNE基因番茄植株和野生番茄(Micro Tom小番茄)分别选取的三株重复单株。
以上结果表明：野生番茄BFNE基因可以使Micro Tom小番茄的分枝增多，果实产量提高。BFNE在改良番茄株型及提高番茄生物产量的分子育种中具有重要应用价值。
实施例3、BFNE基因在鉴定或区分野生番茄和栽培番茄中的应用
由于野生番茄中的BFNE的基因组序列如序列表中序列3所示，其编码的蛋白的氨基酸序列如序列表中序列1所示；栽培番茄中的BFNE的基因组序列如序列表中序列5所示，其编码的蛋白的氨基酸序列如序列表中序列4所示，因此可根据野生番茄和栽培番茄中的BFNE蛋白或其编码基因序列来鉴定或区分野生番茄和栽培番茄。具体方法如下：
实验材料：
1：类番茄Solanum lycopersicoides。
2：多毛番茄Solanum habrochaites。
3：潘那利番茄Solanum pennellii。
4：智利番茄Solanum chilense。
5：秘鲁番茄Solanum peruvianum。
6：多腺番茄Solanum corneliomulleri。
7：小花番茄Solanum neorickii。
8：克梅留斯基Solanum chmielewskii。
9：醋栗Solanum pimpinellifolium。
10：加拉帕戈斯Solanum galapagense。
11：樱桃番茄Solanum lycopersicum var.cerasiforme。
12：栽培番茄M82 Solanum lycopersicum。
13：栽培番茄Heinz-1706 Solanum lycopersicum。
其中，编号1-10均为野生番茄；编号11-13均为栽培番茄。
实验方法：提取待测番茄的基因组DNA，采用PAV-F：CTTGCTTTGCTATCAGACACAC和PAV-R：GCAAGTCAAGTCAGCATTCA进行扩增，得到扩增产物，然后电泳检测扩增产物大小。
实验结果：所有野生番茄中均扩增得到大小为777bp的扩增产物条带，所有栽培番茄中均扩增得到大小为534bp的扩增产物条带(图7)。
因此，在实际应用中，可按照如下方法鉴定待测番茄品种为野生番茄还是栽培番茄：提取待测番茄的基因组DNA，采用PAV-F/PAV-R引物对进行扩增，得到扩增产物，然后电泳检测扩增产物，若扩增得到大小为777bp的扩增产物条带，则待测番茄品种为野生番茄，若扩增得到大小为534bp的扩增产物条带，则待测番茄品种为栽培番茄。
以上所述仅是本发明的优选实施方式，应当指出，对于本技术领域的普通技术人员来说，在不脱离本发明技术原理的前提下，还可以做出若干改进和润饰，这些改进和润饰也应视为本发明的保护范围。
序列表
<110> 新疆农业科学院园艺作物研究所
<120> BFNE基因在番茄株型改良和生物产量提高中的应用
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atcatcgcgt tgctgtttat cttagtgcaa agatttcgtc tttgaatgct gacttgactt 2160
gcttgatggt gtttggaaag aagtacatgg atgaagattt ggacaagagg ggattcaaag 2220
ctcttgtcaa agaggttgac gaattagcag caacaccaaa tcttggagat tttatcccct 2280
tcctcggtgt aattgatctc cagggactca ctcgccggct caaggatctt tcaaaagtgt 2340
ttgatgactt tcttgagaag attatcgatg aacatgtcca gtctggtgac cagaagcaat 2400
ctaaagactt tgttgacacc atgttggaca ttatgcaatc aggagaagca gaatttcaat 2460
ttggccgtag ccatataaaa gctatactct ttgtacgtta ttttgatcat cccatctatg 2520
ttcttctatt attaggtttg tttgtgctaa tttccccttt ctataacagg atatgttagt 2580
tgcagcaatg ggcacttcag caacagcaat agaatggata ttaacagagc ttcttcagca 2640
tcctcatgtg atgaagaaac tccaaaaaga gttggaacaa gtggtaggac ttgaaaaaat 2700
ggtaaaagaa tcagacttgg agaacttgaa ctacttagac atggttgtaa aggagcgctt 2760
gaggctgcat gccgtagcgc cactagcacc tcacgaggca atggaagatt gtgtagtcaa 2820
tagcttccac atacaaaaag gatccggatc ctgattaact tttatgctgt tcaaagggat 2880
ccaaatattt ggcctgaacc tgaaaagttt ttggccgaga ggttttctgg gagcagtata 2940
gacattcgtg gacatgactt ccaactttta ccatttggct cgggtagaag aagttgtcct 3000
gcaatgcagt tggcaattgt ccttgtccaa tttacagtgg cacaactggt gcattgcttt 3060
gattgggagc ttccaaatgg tatgcagcct tgtgaaatgg atgttgagga gcactttggg 3120
ttagcaacaa gcagagcaaa tcatttaatg gccattccta cttatagact aaacaagaat 3180
gcttaatctc aaacgatgca atatggtgcg ataatcagtt aaaggaagat caaatgagag 3240
attatataca taataataat ggagttggct gaaatagttg accttaatta aaccatttct 3300
tctcttcttt actggtcatg ctatctctca gactaggctc ctttgtaatt tgtttttctc 3360
tgtagtattt gtggagctaa taagaaatgt attcgtttca caatgtaata caaaatacgt 3420
tgcgaaaata tcataggcta actagcacac aatcataaat aaagcaaaaa gagaagaaaa 3480
t 3481
<210> 4
<211> 306
<212> PRT
<213> Artificial Sequence
<400> 4
Thr Ala Glu Lys Val Leu Lys Thr Tyr Asp His Ile Phe Ala Ser Arg
1 5 10 15
Pro His Asn Glu Ala Ala Tyr Tyr Leu Ala Tyr Gly Gln Arg Asn Met
20 25 30
Ile Ser Ala Lys His Gly Pro Tyr Trp Arg Asn Met Arg Lys Leu Cys
35 40 45
Thr Gln His Leu Phe Ser Asn Gln Lys Ile Asn Ser Phe Gln Ser Met
50 55 60
Arg Arg Gln Gln Val Glu Val Met Ile Lys Ser Leu Lys Asn Gly Thr
65 70 75 80
Cys Asp His Arg Val Ala Val Tyr Leu Ser Ala Lys Ile Ser Ser Leu
85 90 95
Asn Ala Asp Leu Thr Cys Leu Met Val Phe Gly Lys Lys Tyr Met Asp
100 105 110
Glu Asp Leu Asp Lys Arg Gly Phe Lys Ala Leu Val Lys Glu Val Asp
115 120 125
Glu Leu Ala Ala Thr Pro Asn Leu Gly Asp Phe Ile Pro Phe Leu Gly
130 135 140
Val Ile Asp Leu Gln Gly Leu Thr Arg Arg Leu Lys Asp Leu Ser Lys
145 150 155 160
Val Phe Asp Asp Phe Leu Glu Lys Ile Ile Asp Glu His Val Gln Ser
165 170 175
Gly Asp Gln Lys Gln Ser Lys Asp Phe Val Asp Thr Met Leu Asp Ile
180 185 190
Met Gln Ser Gly Glu Ala Glu Phe Gln Phe Gly Arg Ser His Ile Lys
195 200 205
Ala Ile Leu Phe Asp Met Leu Val Ala Ala Met Gly Thr Ser Ala Thr
210 215 220
Ala Ile Glu Trp Ile Leu Thr Glu Leu Leu Gln His Pro His Val Met
225 230 235 240
Lys Lys Leu Gln Lys Glu Leu Glu Gln Val Val Gly Leu Glu Lys Met
245 250 255
Val Lys Glu Ser Asp Leu Glu Asn Leu Asn Tyr Leu Asp Met Val Val
260 265 270
Lys Glu Arg Leu Arg Leu His Ala Val Ala Pro Leu Ala Pro His Glu
275 280 285
Ala Met Glu Asp Cys Val Val Asn Ser Phe His Ile Gln Lys Gly Ser
290 295 300
Gly Ser
305
<210> 5
<211> 3239
<212> DNA
<213> Artificial Sequence
<400> 5
gttttccgaa aacaaaagtt cacattaccc acttttctct ctctaaattt ttcaccaaat 60
tctatcatta tcttttctct ttcacgattt attccattat ttcatattct caagtaactt 120
aaaccttgtc tttttaaaat caaattcctc gatttttcca agggatctca taaatcatgt 180
aaaagttttt gttacctttg ttgaagtgaa ctcagttgat ttgtgattgt ttccattagg 240
gaagaaaatt ttgaaataat cgattgggat atcgaagtga attcaactga tttcgttgaa 300
ctatgagatt gtcggtgaaa tacaaaaaca atcaggtatt cattaacggt tgattcttta 360
atttttttgg aattgatacg aaaatggtag ttgaaaaaag tgatttagaa atggatctag 420
atcgtatttt gtactgttgg accttcacat gaagatcttt ataagttact atccaaatat 480
tggatttaaa ggttagacag tcactaaggt attcgatttg ggagtttcag gtcaatattt 540
catccaaggc actatgttgg acaacatcaa taattatagt tatcttttca atgaattgaa 600
gtatttatgt atctttttcg catgcagaag gacccaaagg agatgaaagt acagaatcac 660
cagccatatt ttaaagtaaa aaatatacaa atttttaatt tgtaatatgt ttgggataca 720
aatagctagt gaatgattga aatattttaa ttttttttta ggtcgtatac ttggaataga 780
aaataatttt gaaatctatt tagtaggtgt tttataaatt agacattgaa tataatatat 840
ttttaaaata cataacagat aagaaataca attttttttt aaatacaaat tttgattgct 900
accatgattg taagtttaca tgtgtttgat gactaaaata caaattttat cttacatttt 960
ttataaatat acaaacagat gttgtatcat ttctgaatga aaatgaattt acacaagtat 1020
atggatacaa atatgcttct taaataacta caaatccatt tggtatacat attagaatca 1080
atacaaagta gtaaagaaat tcaaatgtca ttatataaac cctaatatga aaatcataat 1140
gaacacaaat acacatatca tacatcttga aatgaataca atcaaaagta aattttagga 1200
ataacaaata taaaaatcat attagctctc tatagctaca gttttgctat ttgcactcca 1260
tataatgcat aactatcttg acagtactag aattagtaat acttgtacaa ctttctaaac 1320
caacttgcag ccagtcaaat aattcaagta gtttctaaaa agccttatag caacttattc 1380
ccaaggcttg ctttgctatc agacacacaa cttgaaatat ggctcattta gttgtcttct 1440
gacttttttc tgaaagcacc atatcttcca ttaccttgat aatcaatact tctcatccac 1500
atttttcaaa aataatatct aacattacca aacacagcca ttatatgatt gagatctcta 1560
gacccattca attagagatc ttttagaagc taataaattt agtaagtatt aggcacagcg 1620
gagaaggtcc tcaagacata tgatcatatc tttgctagta gacctcacaa tgaggcagct 1680
tattatttag catatggaca gaggaatatg atatccgcaa agcatggacc atactggcgc 1740
aacatgcgca aattgtgcac tcagcacctt ttcagtaacc aaaagatcaa ttcatttcaa 1800
tctatgagaa ggcaacaagt tgaggttatg atcaaatcac ttaaaaatgg aacttgtgat 1860
catcgtgttg ctgtttatct tagtgcaaag atttcgtctt tgaatgctga cttgacttgc 1920
ttgatggtgt ttggaaagaa gtacatggat gaagatttgg acaagagggg attcaaagct 1980
cttgtcaaag aggttgacga attagcagca acaccaaatc ttggagattt tatccccttc 2040
ctcggtgtaa ttgatctcca gggactcact cgccggctca aggatctttc aaaagtgttt 2100
gatgactttc ttgagaagat tatcgatgaa catgtccagt ctggtgacca gaagcaatct 2160
aaagactttg ttgacaccat gttggacatt atgcaatcag gagaagcaga atttcaattt 2220
ggccgtagcc atataaaagc tatactcttt gtacgttatt ttgatcatcc catctatgtt 2280
cttctattat taggtttgtt tgtgctaatt tcccgtttct ataacaggat atgttagttg 2340
cagcaatggg cacttcagca acagcaatag aatggatatt aacagagttt cttcggcatc 2400
ctcatgtgat gaagaaactc caaaaagagt tggaacaagt ggtaggactt gaaaaaatgg 2460
taaaagaatc agacttggag aacttgaact acttagacat ggttgtaaag gagcgcttga 2520
ggctgcatgc cgtagcgcca ctagcacctc acgaggcaat ggaagattgt gtagtcaata 2580
gcttccacat acaaaaagga tccggatcct gattaacttt tatgctgttc aaagggatcc 2640
aaatatttgg cctgaacctg aaaagttttt ggccgagagg ttttctggga gcagtataga 2700
cattcgtgga catgacttcc aacttttacc atttggctcg ggtagaagaa gttgtcctgc 2760
aatgcagttg gcaattgtcc ttgtccaatt tacagtggca caactggtgc attgctttga 2820
ttgggagctt ctaaatggta tgcagccttg tgaaatggat gttgaggagc actttgggtt 2880
agcaacaagc agagcaaatc atttaatagc cattcctact tatagactaa acaagaatgc 2940
ttaatctcaa acgatgcaat atggtgcgat aatcaattaa aggaagatca aatgagagat 3000
tatatacata ataataatgg agttggctga aatagttgac cttaattaaa ccatttcttc 3060
tcttctttac tggtcatgct atctctcaga ctaggctcct ttgtaatttg tttttctctg 3120
tagtatttgt ggagctaata agaaatgtat tcgtttcaca atgtaataca aaatacgttg 3180
cgaaaatatc ataggctaac tagcacacaa tcataaataa agcaaaaaga gaagaaaat 3239
<210> 6
<211> 22
<212> DNA
<213> Artificial Sequence
<400> 6
cttgctttgc tatcagacac ac 22
<210> 7
<211> 20
<212> DNA
<213> Artificial Sequence
<400> 7
gcaagtcaag tcagcattca 20
Claims (10)
1.一种蛋白质，为a1)或a2)或a3)或a4)：
a1)氨基酸序列是序列1所示的蛋白质；
a2)在序列1所示的蛋白质的N端或/和C端连接标签得到的融合蛋白质；
a3)将序列1所示的氨基酸序列经过一个或几个氨基酸残基的取代和/或缺失和/或添加得到的与植物耐逆性相关的蛋白质；
a4)将序列1所示的氨基酸序列具有90％同一性、来源于番茄且与番茄株型或产量相关的蛋白质。
2.与权利要求1所述的蛋白质相关的生物材料，为下述A1)至A8)中的任一种：
A1)编码权利要求1所述的蛋白质的核酸分子；
A2)含有A1)所述核酸分子的表达盒；
A3)含有A1)所述核酸分子的重组载体；
A4)含有A2)所述表达盒的重组载体；
A5)含有A1)所述核酸分子的重组微生物；
A6)含有A2)所述表达盒的重组微生物；
A7)含有A3)所述重组载体的重组微生物；
A8)含有A4)所述重组载体的重组微生物。
3.根据权利要求2所述的相关生物材料，其特征在于：A1)所述核酸分子为如下1)或2)或3)所示的基因：
1)其编码序列是序列3所示的基因组DNA分子或序列2所示的cDNA分子；
2)与1)限定的核苷酸序列具有75％或75％以上同一性，且编码权利要求1所述的蛋白质的cDNA分子或基因组DNA分子；
3)在严格条件下与1)或2)限定的核苷酸序列杂交，且编码权利要求1所述的蛋白质的cDNA分子或基因组DNA分子。
4.权利要求1所述的蛋白质或权利要求2或3所述的生物材料在如下1)-4)任一种中的应用：
1)调控番茄株型；
2)调控番茄产量；
3)培养转基因番茄或番茄育种；
4)鉴定或区分野生番茄和栽培番茄。
5.根据权利要求4所述的应用，其特征在于：所述番茄株型为番茄分枝。
6.一种转基因番茄的培育方法，包括如下步骤：提高受体番茄中权利要求1所述蛋白质的含量和/或活性，得到转基因植物；所述转基因番茄的分枝数和/或产量高于所述受体番茄。
7.根据权利要求6所述的方法，其特征在于：所述提高受体植物中权利要求1所述蛋白质的含量和/或活性的方法为在受体番茄中过表达权利要求1所述蛋白质。
8.根据权利要求7所述的方法，其特征在于：所述过表达的方法为将权利要求1所述蛋白质的编码基因导入受体植物中。
9.一种鉴定或区分野生番茄和栽培番茄的方法，包括如下步骤：检测待测番茄是否含有权利要求1所述蛋白质或其编码基因：若待测番茄含有权利要求1所述蛋白质或其编码基因，则待测番茄为野生番茄；若待测番茄不含有权利要求1所述蛋白质或其编码基因，则待测番茄为栽培番茄。
10.根据权利要求9所述的方法，其特征在于：检测待测番茄是否含有权利要求1所述蛋白质或其编码基因的方法包括如下步骤：提取待测番茄的基因组DNA，采用序列6所示核苷酸序列和序列7所示核苷酸序列进行扩增，得到扩增产物，然后电泳检测扩增产物，若扩增得到大小为777bp的扩增产物条带，则待测番茄品种为野生番茄，若扩增得到大小为534bp的扩增产物条带，则待测番茄品种为栽培番茄。
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