白菜矮化突变体cd1生理特性及初步定位

白菜矮化突变体cd1生理特性及初步定位DissertationforMaster’sDegreePrimarymappingandphysiologicalcharacterizationofChinesecabbagedwarfmutantcd1Candidate：HuJunYunSupervisor：AssociateProf.WangYugangSpeciality：VegetableScienceResearchField：BreedingandBiotechnologyShenyangAgriculturalUniversityJune，2020第四章讨论4.1cd1是一个新型的白菜矮化基因本试验根据外源施加赤霉素处理发现，cd1突变体对喷施赤霉素敏感。喷施赤霉素后，突变体株高表现为多分蘖并且恢复其株高。这表明出现矮化性状可能与赤霉素的合成途径有关。关于赤霉素调控白菜株高的研究也鲜有报道，所以，cd1作为一个新型的白菜矮化基因，能够帮助我们了解赤霉素对白菜株高的调控机理。但是，考虑到矮化突变体cd1授粉败育以及种子数量有限，目前没有进行充分的重复和处理组合，仍需继续深入研究来确定cd1基因与赤霉素合成代谢途径的关系。4.2白菜矮化突变体cd1的产生原因可能为细胞分裂过程受阻导致通过细胞水平观察发现，造成矮化突变体的原因有三种情况，一种为细胞的伸长，另一种为细胞分裂过程受阻，最后一种为前两者情况同时发生。（杨意宏等2018）。经细胞学研究，在水稻的等位基因矮杆株系上，发现节间长度与细胞数目成正相关，与细胞长度没有显著相关性，这表明节间变短很很可能是由于细胞数目减少变成的。（WangN.,etal.2017）本试验通过对白菜矮化突变体cd1和野生型植株的茎进行石蜡切片观察发现，矮化突变体cd1茎的细胞数目显著少于野生型植株。这表明cd1在细胞分裂过程中受到阻碍。因此cd1表现为矮化性状可能是由于细胞分裂受阻所致。4.3cd1的矮化性状明显，可用于杂交育种本试验矮化突变体cd1表现为极度矮化，是目前在白菜作物上发现为数不多的矮化突变体，这说明cd1对白菜株高的调控作用十分明显，能有效的较低白菜的株高。但是，如何合理利用cd1降低白菜株高，还需我们更进一步了解cd1的调控机制。突变体cd1株高虽然极度矮化，但是与产量相关的性状表现较差，花粉活力检测结果表明，育性极差。很难应用于白菜育种工作中。因此，本试验结论可为后续工作做一个基础。4.4本试验的后续工作本试验是以白菜矮化突变体cd1为材料，利用BSR-Seq分析与验证最终将矮化突变体cd1基因初步定位在引物L06-09与L06-14之间（物理位置21621766-24683923）。后续工作将针对矮化突变体cd1的遗传特性与生理特性，结合BSR-Seq分析结果，对突变体cd1进行精细定位。再对其对突变体cd1和远缘亲本‘YellowSarson’做转录组分析，进一步探究矮化突变体cd1的分子机制并验证候选基因。克隆其全长cDNA，进行序列分析、时空表达分析、亚细胞定位等。第五章结论1.通过对材料突变体cd1与野生型植株的农艺性状进行测量发现：与野生型相对比，植株生长缓慢、下胚轴极短、叶面积变小；叶片颜色加深、有刺毛；根的长度也明显收到抑制，主根变短，侧根数目也明显减少；突变体cd1植株的叶长、叶宽以及株高等数据均极显著低于野生型植株。2、对其生理指标测量发现：突变体cd1叶绿素a、叶绿素b、总叶绿素占比都显著大于野生型，因为其叶绿素占比相对较高，所以突变体cd1光合速率、胞间CO2浓度、气孔导度也高于野生型，可以说明，颜色加深有助于光合作用，由于蒸腾速率低于野生型，说明突变体cd1对外界环境的适应能力较强。但是，我们通过花粉活力测定，我们发现，突变体cd1的花粉活力极低。3、通过对突变体cd1茎秆和叶片进行石蜡切片细胞学观察发现，在叶片上，我们推测叶片表型的不同是由于细胞数目减少和细胞不规则排列造成的。在茎上，我们认为矮化突变体cd1的茎秆组织的薄壁细胞和中柱细胞未能在纵向上正常伸长，细胞数目减少可能是导致茎秆变短的原因。4、根据外源施加赤霉素处理发现，矮化突变体cd1对赤霉素敏感。外源喷施后，突变体的株高可以恢复其高度。这表明矮化性状的出现可能与赤霉素合成途径有关。5、基于对BSR-Seq结果展开的一系列验证与评估，进一步有效完成相应的矮化植株cd1染色体定位。其次凭借大白菜3.0数据库、Primer5展开相应的SSR引物设计，最后有效筛选出具备多态性引物以及与相应突变体cd1密切连锁的标记。最后将相应的矮化突变体cd1基因定位在引物L06-09与L06-14之间（物理位置21621766-24683923）。6、本研究对其候选基因进行差异表达分析，最终确定三个候选基，分别为：BrA06g031210.3C、BrA06g034110.3C、BrA06g033010.3C，这三个候选基因将在后续精细定位中进一步验证。参考文献白丽君,尹淑霞.植物矮化突变体的来源及矮化机理研究进展.生物技术通报,2014,(06):34-39.白丽君，尹淑霞.植物矮化突变体的来源及矮化机理研究进展[J].生物技术通报，2014，6：34-39.陈晶晶,胡玉林,胡会刚,等.植物矮化相关基因的研究进展.广东农业科学,2014,41(15):126-132.程云,王枟刘,杨静,等.种植密度对夏玉米基部节间性状与倒伏的影响［J］.玉米科学,2015,23（5）：112-116.勾玲,黄建军,孙锐,等.玉米不同耐密植品种茎秆穿刺强度的变化特征［J］.农业工程学报,2010,26（11）：156-161.黄海,陈德龙,常莹,等.玉米品种抗倒能力差异及其机制研究［J］.南京农业大学学报,2014,37（4）：22-30.贾继增,丁寿康,李月华,张辉.1992.中国小麦的主要矮秆基因及矮源的研究.中国农业科学,25(l):1~5李合生.现代植物生理学,高等教育出版社,2012.李素燕,小麦株高相关基因的分离克隆研究.[硕士学位论文].北京：中国农业科学,2003.李祖亮.玉米矮化突变体gad5表型分析和基因克隆[硕士学位论文].河南:河南大学,2015.刘秉华，杨丽，丁表珍.1993.小麦显性矮秆基因Rht10与着丝点间遗传距离的测定.科学通报，38(2):1128~1130.刘平,杨慧,孟雪,等.植物矮化研究进展.安徽农业科学,2010,38(27):15442-15446.刘鑫,谢瑞芝,牛兴奎,等.种植密度对东北地区不同年代玉米生产主推品种抗倒伏性能的影响［J］.作物杂志,2012,5（33）：126-130.孙荣锦,杨之刚.1993.小麦矮秆突变系原冬96株高单体分析.核农学报,7(1):57~60王翠红，马建，王帅，等.一个新的水稻D1基因等位突变体的遗传鉴定与基因功能分析[J].作物学报，2016，42(9)：1261-1272.王洪梅,周显昌,周志军,等.赤霉素促进针叶树开花结实技术的研究进展.林业科技,2011,36(3):11-15.王武全，曹本高，员海燕.玉米矮秆突变体的激素敏感性分析[J].西北农林科技大学学报(自然科学版)，2017，45(8)：51-55.王月华，韩烈保，曾会明，等.植物赤霉素矮化突变体研究进展[J].中国生物工程杂志，2006，26(8)：22-27.严建兵,汤华,黄益勤,等.不同发育时期玉米株高QTL的动态分析.科学通报,2003,(18):1959-1964.杨天章,张晓科,刘宏伟,王中华.1993.矮秆小麦XN0004的矮秆基因Rht21的染色体臂定位.西北农林科技大学（自然科学版）,21(4):13~17.姚敏娜,施志国,薛军,等.种植密度对玉米茎秆皮层结构及抗倒伏能力的影响［J］.新疆农业科学,2013,50（11）：2006-2014.于永红，斯华敏.水稻矮化相关基因的研究进展[J].植物遗传资源学报，2005，6(3)：344-347.虞慧芳，曹家树，王永勤.植物矮化突变体的激素调控[J].生命科学，2002，14(2)：85-88.张洪生,盖伟玲,李玲燕,等.种植密度对不同玉米品种抗倒伏性及产量的影响［J］.中国种业,2009,6（17）：34-35.张京,丁寿康.1989.春小麦园柱的矮秆基因定位.作物品种资源,27(1):21~27.张志胜,谢利,萧爱兴,等.秋水仙素处理兰花原球茎对其生长和诱变效应的影响.核农学报,2005,(01):19-23.赵久然,王帅,李明,等.玉米育种行业创新现状与发展趋势［J］.植物遗传资源学报，2018,19（3）：435-446.左圆圆,周杨杨,李小平.植物激素-赤霉素(GA)细胞信号转导机制.淮北师范大学学报(自然科学版),2011,32(2):49-54.AhmadM,SorrellsME.2002.DistributionofmicrosatellitealleleslinkedtoRht8dwarfinggeneinwheat.Euphytica，123:235~240AkiyamaK.,MatsuzakiK.,HayashiH.Plantsesquiterpenesinducehyphalbranchinginarbuscularmycorrhizalfungi.Nature,2005,435(7043):824-7.AlderA.,JamilM.,MarzoratiM.,etal.Thepathfromβ-carotenetocarlactone,astrigolactone-likeplanthormone.Science,2012,335(6074):1348-51.ArberdN.P.,FreelingM.Geneticsofdominantgibberellin-insensitivedwarfisminMaize.Genetics,1989,121(4):827-838.AshikariM，WuJZ，YanoM，etal.Ricegibberellin-insensitivedwarfmutantgeneDwarf1encodesthea-subunitofGTP-bindingprotein[J].ProceedingsoftheNationalAcademyofSciencesofUSA，1999，96：10284-10289.AshikariM，WuJ，YanoM，etal.Ricegibberellin-insensitivedwarfmutantgeneDwarf1encodesthealphasubunitofGTP-bindingprotein[J].ProceedingsoftheNationalAcademyofSciencesoftheUnitedStatesofAmerica，1999，96(18)：102842-10289.BensenR.J.CloningandcharacterizationoftheMaizean1gene.ThePlantCellOline,1995,7(1):75-84.BirnbaumK.A.Geneexpressionmapofthearabidopsisroot.Science,2003,302(5652):1956-1960.BömkeC.,RojasMC.,GongF.,etal.IsolationandcharacterizationofthegibberellinbiosyntheticgeneclusterinSphacelomamanihoticola.Applied&EnvironmentalMicrobiology,2010,74(17):5325-5339.BookerJ.,SiebererT.,WrightW.,etal.MAX1encodesacytochromeP450familymemberthatactsdownstreamofMAX3/4toproduceacarotenoid-derive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