大白菜抽薹过程中发生叶色转变的分子机制
详细信息    本馆镜像全文|  推荐本文 |  |   获取CNKI官网全文
摘要
大白菜(Brassica rapa L. ssp. pekinensis)的先期抽薹,是在大白菜的营养体还未充分长成之前,在不该抽薹时提前抽薹的现象。抽薹后的大白菜会失去食用价值,大大降低经济效益。因此,研究其抽薹开花相关的分子机制,对白菜育种具有重要意义。
     大白菜在春化后会抽薹开花,在抽薹前表现出一个叶色转变过程,叶片颜色从鲜亮的绿色转为没有光泽的暗灰色后花茎就会伸长并开始现蕾抽薹,叶色转变似乎是大白菜抽薹的一个生理信号,在弱冬性品种上的表现尤为突出,因此推测叶色转变与抽薹开花有关,叶色转变前后基因和蛋白表达必将发生变化。本试验从抽薹开花相关基因表达和蛋白质表达两个方面研究大白菜生殖发育转变过程中的分子机理。以四个冬性不同的大白菜品种为材料,分别采取春化前、春化后、花芽分化、花茎分化、叶色转变、花茎伸长六个时期的植株生长点部位新生幼嫩叶片,来测定叶片激素含量,并进行FT和LFY两个开花相关基因的RT-PCR表达分析和蛋白质双向电泳分析;另外在每个品种的叶色转变期取植株的根、短缩茎、茎端生长点幼嫩叶片、近端叶片、远端叶片五个部位,进行FT和LFY基因的RT-PCR表达分析。
     主要试验结果如下:
     1.通过测定激素含量的变化,得知它与大白菜抽薹开花过程中叶色的转变有一定关系,大白菜抽薹过程中, IAA和GA3含量均在叶色转变期出现峰值,叶色转变后花茎开始伸长,叶色转变是大白菜抽薹的一个生理信号。
     2.FT基因在春化前和春化后也即整个的营养生长阶段不表达或微量表达,在花芽分化期以后表达强度递增,在叶色转变期和花茎伸长期表达强烈,另外还发现FT基因在弱冬性品种中要比在强冬性品种中表达强烈。FT基因在根和短缩茎中不表达,在生长点嫩叶中和近端叶片、远端叶片表达,在弱冬性品种中,FT在近端叶片中强烈表达,在远端叶片中表达极微弱,而在强冬性品种中,FT在近端叶片与远端叶片中表达均较强。
     3.LFY基因在春化前不表达,花芽分化期和叶色转变期和花茎伸长期表达较强,在生长点嫩叶中表达最强烈,在近端叶片和远端叶片中也有较强表达,在短缩茎中有极微弱表达,在根中不表达。
     4.蛋白质表达图谱显示叶色转变后有18种新的特异蛋白质产生,在花茎分化期表达的17种蛋白质在叶色转变期停止表达;有29种蛋白质在叶色转变后表达上调,还有21种蛋白质表达下调。
     5.对17个蛋白点进行质谱鉴定,成功鉴定出其中的16个点,这些蛋白1个与转录调控有关,2个与信号传递和能量传递有关,6个与细胞生长及代谢有关,还有1个是结构蛋白,此外还有1个未知蛋白,5个功能未知的假设性蛋白质。
Premature bolting of Chinese cabbage (Brassica rapa L. ssp. pekinensis) is a phenomenon that bolting at a improper time before its vegetative plants ripen, then the vegetable will lose value,It causes a great loss in production. It has great significance to study the Physiological and Molecular mechanism connected with bolting of Brassica.rapa,especially the genes associated with bolting and flowering in vernalization pathway.
     Chinese cabbage will bolting and flowering after vernalization , the leaf color changes before the bolting, stems elongate and the plant start to bud and bolt after the leaf color changes from bright green to dark grey without being shiny or lustrous .the changes of the leaf color seems to be a physiological signal of the bolting for Chinese cabbage ,the phenomenon is particularly acute for the weak winterness cultivars, so we speculate that the changes of leaf color is relevant with the bolting and flowering process. the genes and protein expression varies before and after leaf color changes. In present study, we used four cultivars of Chinese cabbage that were different in winter to study the molecular mechanism of reproductive development by testing the related genes and proteins expression. We took samples of fresh young leaves in six development stages for hormone levels Determination and 2-D electrophoresis,which are before and after vernalization, floral bud differentiation period,stem differentiation period,the period of leaf color transition and the stem elongation period. Then we took samples at root,dwarf stem, fresh leaves at growing point, proximal leaves, distal leaves in the period of leaf color transition for FT and LFY RT-PCR all together with the samples mentioned above. Main results were as follows:
     1. It turns out that the hormone content is associated with the leaf color changes in the bolting process, different winterness varieties get content peaks of IAA and GA3 in the period of leaf color transition respectively in bolting process. The stems begin to elongate after this stage.The changes of leaf color is a physiological signal in cabbage bolting.
     2 . FT gene expresses stronger in weak winterness cultivars than in strong winterness cultivars; It doesn’t express or express very weak in the whole vegetative growth stage, it expresses stronger and stronger after the bud differentiation stage, and expresses very strongly in the leaf color transition period and stems elongation period .FT gene doesn’t express in roots and dwarf stems, it expresses in the fresh leaves at growing point, proximal leaves and distal leaves. FT expresses strongly in proximal leaves and very weak in distal leaves in weak winterness cultivars, however, in strong winterness cultivars,FT expresses both strongly in proximal leaves and distal leaves .
     3. LFY gene doesn’t express before vernalization,expresses strongly in the bud differentiation period,the leaf color transition period and stems elongation period,most strongly in the fresh leaves at growing point.it expresses strongly in proximal leaves and distal leaves,it doesn’t express in roots ,and express very weak in dwarf stems.
     4. We found 18 new proteins espresses after leaf color changes,and 17 proteins don’t espresses anymore after this transition, There are also 21 proteins are up-regulation of experssion and 29 are down-regulation of experssion according to analysis and comparation.
     5. 17 protein spots were identified using MS.and 16 of them are successfully identified. 1 of them is related with transcriptional regulation, 2 of them are related with signal or energy transmission,6 proteins participate in cell growth and metabolism,and the last one is structural protein.
引文
奥岩松,李式军,陈广富,等.1996.种子春化与光周期处理对大白菜花芽分化和抽薹的影响.东北农业大学学报.27(3) :250~254
    奥岩松,李式军.1997.大白菜发育过程中可溶性蛋白质的变化.中国蔬菜.2:19~21
    奥岩松.1996.晚抽薹大白菜的种子春化特性与光周期反应,园艺学进展,中国农业出版社,北京. 27(3):250~254
    曹家树.1996.中国白菜起源、演化和分类研究进展.园艺学年评,科学出版社. 1~21
    曹仪植.2002.植物分了生物学.北京:高等教育出版社.
    陈书霞,王晓武,方智远,程智慧,孙培田. 2003.芥蓝x甘蓝的F2群体抽薹期性状QTLs的RAPD标记.园艺学报. 30(4): 421~426
    陈晓,陈彦惠,任永哲.2005.植物开花转换的分子生物学研究.分子植物育种. 3(4):557~565
    程斐,李式军,奥岩松,等. 1999,大白菜抽薹性状的遗传规律研究.南京农业大学学报.22(1):26-28
    程斐,李式军,奥岩松.1998.大白菜抽薹期的遗传力研究.园艺学进展. 2: 524-526
    程斐,张蜀宁,孙朝晖等.1999.春大白菜品种选育的形态与生理指标.园艺学报. 26(2): 120~122
    傅永福,孟繁静.1997.植物成花决定.植物生理学通讯. 33 (2): 81-87
    高伟.2004.玉米光周期反应与类LFY基因克隆的研究.河南农业大学.
    郭春晓,田素波,郑成淑,等.2009.光周期途径植物开花决定关键基因FT.基因组学与应用生物学,28(3):613-618
    胡巍,侯喜林,史公军.2004.植物春化特性及春化作用机理.植物学通报. 21(1):26~36
    惠麦侠,巩振辉,张鲁刚,柯桂兰,张明科.2003.大白菜花芽形态分化的研究.华北农学报. 18(1): 75~78
    惠麦侠.2002.大白菜耐抽薹性及其鉴定方法的研究:〔硕士学位论文〕.陕西:西北农林科技大学.
    蒋欣梅,于锡宏.2004.低温处理青花菜萌动种子对花芽分化的促进作用.植物生理与生物学学报. 30(4):421~427
    靳文海.多维液相色谱-质谱联用技术在蛋白质组学中的应用[D].中国科学院研究生院(大连化学物理研究所)博士学位论文,2005:6-9
    李季伦,朱彤霞,李永胜等.1980.玉米赤霉烯酮的研究.北京农业大学学报. 6(1): 13~28
    李春刚.2008.大白菜抽薹过程中叶色转变机理初探.东北农业大学硕士研究生论文.
    李利斌,刘立锋,李化银等. 2010,两个大白菜FT基因的生物信息学解析.山东农业科学.3: 1~4, 7
    李林海,刘燕,徐景先.2006.拟南芥等植物的春化分子机理研究进展.西北植物学报. 26(1): 0207-0210
    李梅兰,侯雷平.2005.植物春化作用研究进展.山西农业科学.33(1): 43~45
    李梅兰.2001.DNA甲基化与白菜的生长转变.浙江大学
    李曙轩,李式军.1964.白菜的花芽分化与叶球形成(核糖变化).中国农业科学. 6: 36~40
    李曙轩,寿诚学.1957.春化及光照对于白菜及芥菜发育的影响.植物学报. 6(1): 7~28
    李秀珍,等.1992.冬小麦春化过程中可溶性蛋白质的变化与形态发生的关系,植物学报. 29(2): 492~498
    李子银,陈受宜.1999.水稻抗病基因同源序列的克隆、定位及表达.科学通报.44(7):727~733
    凌俊.1993.春化对油菜叶片硝酸还原酶活性的影响.植物生理学通论. 11: 56~58
    刘春香,韩玉珠,栗长兰等.2001.大白菜的春化特性及未熟抽薹研究进.吉林农业大学学报. 23(3): 61~64
    刘宜生.1996.中国大白菜,中国农业出版社,北京
    刘旭新,Stephen Jackson,李春阳.PVX介导拟南芥Flowering locus T诱导烟草开花研究核农学报.2009 ,23 (1) :65~69
    刘月学,等.2006.普通枇杷和栎叶枇杷APETALA1同源基因的克隆和序列分析.福建农林大学学报:自然科学版. 35(2): 173~176
    卢刚,曹家树,陈杭.1999.芸苔属植物分子标记技术和基因组研究进展.园艺学报. 26(6):384~390
    彭振英.小麦渐渗系耐盐抗旱的蛋白质组学比较研究[D].山东大学博士学位论文,2008:26-9
    阙月美,梁振兴,韩玉珍等.1990.冬小麦和棉花开花结实过程中玉米赤霉烯酮含量的变化.北京农业大学学报. 16(2): 153~155
    任江萍,潘登奎,尹均.1999.小麦春化过程中蛋白质变化研究[J].山西农业大学学报. 19: 298~301
    杉山直仪.1981.蔬菜的发育生理和栽培技术.赖俊铭译.北京:中国农业出版社
    史庆馨,鹿英杰,徐文龙,等.2000.环境条件对大白菜自交系开花期影响初步研究.北方园艺. 6: 6~8
    苏学军,杨衔美,李化银,张焕家.1999.夏播白菜先期抽薹原因及预防.北方园艺,5: 5~6
    孙日飞,张淑江,司家钢等.1999.春化和赤霉素对大白菜抽薹开花的影响.中国蔬菜. 3: 14~17
    孙志宾.盐芥Th TRXh功能分析和盐芥不同生态型的比较蛋白质组学初步研究[D].山东师范大学博士学位论文,2006:10-13
    汪炳良.2004.萝卜春化作用及其与DNA甲基化水平的关系.[博士学位论文].杭州:浙江大学
    王利琳,庞基良,胡江琴,等.2002.温度对植物成花的影响.植物学通报. 19(2): 176~183
    王淑芬,徐文玲,何启伟等.2003.春化深度对萝卜抽薹的影响及抽薹过程GA3和IAA含量的变化.山东农业科学. 6: 23~27
    王淑芬.2002.赤霉素对耐抽薹萝卜抽薹开花的影响.山东农业科学. 6: 14~16
    闻凤英,刘晓晖,王玉龙,等.2001.青麻叶大白菜耐抽薹性研究初报.中国蔬菜.2:30~31
    武峻新.2005.蔬菜耐未熟抽蔓性的研究及利用.北方园艺. 6: 4~6
    夏广清,何启伟,王翠花.2006.赤霉素处理对大白菜开花的诱导效应.中国蔬菜. 5: 19~20
    夏英武,吴殿星,舒庆尧.1996.人工诱发的植物叶绿素突变体的遗传及应用.核农学通报. 17(1): 41~44
    熊延,刘翠敏,王淑芳,王宁宁,王勇.2002.叶绿体基因组与叶绿体基因表达的调节.植物生理学通讯. 38(3): 264~269
    徐继,张玉竹,谭克辉.1994.冬小麦春化过程中6一磷酸葡萄糖脱氢酶活性变化的研究.植物学集刊. 7: 344~348
    徐泽安.2002.白菜和芜菁成花过程中多胺水平变化的研究.湖南农业科学. 5: 9~10
    杨传平,等.2002.高等植物成花基因的研究.遗传HEREDITAS. 24(3): 379~384
    尹均,任江萍,潘登奎等.2002.小麦春化发育相关蛋白质功酶的研究.麦类作物学报. 22(1): 33~38
    俞利荣.曾嵘.夏其昌.蛋白质组研究技术及其进展.生命的化学,1998,18(6):4-9.
    于惠敏.2002.植物的叶绿体基因组.植物生理学通讯. 37:483~488
    余阳俊,陈广,刘琪.1996.大白菜晚抽薹性及其与开花叶片数的关系.北京农业科学.14(3): 32~34
    余阳俊,张凤兰,赵岫云等.2004.大白菜晚抽薹性快速评价方法.中国蔬菜. 6: 16~18
    余阳俊,张凤兰,赵岫云等.2005.大白菜及种内杂种小白菜×大白菜、芜菁×大白菜的晚抽薹遗传效应研究[J].华北农学报,20(3):17-21
    原玉香. 2008.白菜类作物抽薹开花的分子遗传分析.中国农业科学院蔬菜花卉研究所博士论文.
    张德双,徐小谢,徐家炳.2003.春化天数对大白菜、小白菜现蕾和开花德影响.华北农学报. 18(1): 75~78
    张德双,徐小谢,徐家炳.2003.春化天数对大白菜、小白菜现蓄和开花的影响.园艺学报. 30(6): 731~733
    张恩和,黄鹏.1998.春化处理对当归苗生理活性的影响.甘肃农业大学学报. 3: 240~243
    张凤兰,徐家炳,赵灿云等.1996.苗期遮光对易抽薹大白菜抽薹开的影响.中国蔬菜. 22(3): 22~25
    张华锋,孟庆忠,张立军.2000.环境因子在植物成花诱导中的作用及其机理.生物学通报. 35(7): 16~18
    张鲁刚,孔小平,惠麦霞等.2007.大白菜新品种‘冠春’的春化特性及其遗传表现.西北农林科技大学学报(自然科学版). 2(35): 93~96
    张素芝,左建儒.2006.拟南芥开花时间调控的研究进展[J].生物化学与生物物理进展. 33(4): 301~309
    张伟丽,钮心恪,王曾贵.1993.结球白菜组培苗不经低温条件在试管内抽薹开花.中国蔬菜. 5: 62
    张渭,万文鹏,宫晓农等.1996.赤霉素处理对大白菜阶段发育的影响.种子科学. 4: 14~15
    赵香梅,孙守如,张晓伟等.2005.大白菜春化与抽薹特性的研究进展.中国蔬菜. 1:33~35
    种康,雍伟东,谭克辉.1999.高等植物春化作用研究进展.植物学通报. 16(5): 481~487
    周长久,王鸣等.1996现代蔬菜育种学.科学技术文献出版社. 90~91
    朱广廉,杨中汉.1982.SDS一聚丙烯酰胺凝胶电泳法测定蛋白质分子量,植物生理学通讯. 2: 43~47
    朱娜.龙眼LFY同源基因表达及启动子克隆.福建农林大学硕士学位论文.2010
    Alban A,David SO,Bjorkesten L,Andersson C,Sloge E,Lewis S,Currie I.A novel experimental design for comparative two-dimensional gel analysis:two-dimensional difference gel electrophoresis incorporating a pooled internal standard.Proteomics,2003,3:36–44
    Anderson NI,Anderson NG.Proteome and Proteomics:new technologies,new concepts,and new words[J].Electrophoresis,1998,19:1 853-1 861
    Bae,M.s.,et al.,Analysis of the Arabi-dopsis nuclear proteome and its response to cold stress.Plant J.,2003.36:p.652-663.
    Barkan A, Goldschmidt-Clermon M. 2000. Participation of nuclear, genes inchloroplast gene expression. Biochimie. 82:559~572
    Bernier G, Kinet M, SachsRM.1981.The PhyBiology of Flowering. Boca Raton F L, USA: CRC Press. 69~82
    Bernier G, Kinet M, SachsRM.1981.The Physiology of Flowering. Boca Raton F L, USA: CRC Press. 69~82
    Blackstock WP,Weri WP.Proteomics:quantitative and physical mapping of cellular proteins.Trends Biotechnol,1999,17(3):121–127
    Blazquez M.A, Green R, Nilsson O, Sussman M.R, Weigel D. 1998. Gibberellins promote flowering of Arabidopsis by activating the LEAFY promoter. Plant Cell. 10(5) : 791-800
    Burn J E, Bagnall D J, Metzger J D, etal.1993.DNA methylation, vernalization and the initiation offlowering . Proc Natl AcadSci USA, 90: 287-291
    Caicedo AL, Stinchcombe JR, Olsen KM, Schmitt J, Purugganan MD. 2004. Epistaticinteraction between Arabidopsis FRI and FLC flowering time genes generates a latitudinal cline in a life history trait. Proc Natl Acad Sci U S A. 101: 15670-15675.
    Calonje M, Cubas P, Martinez-Zapater J.M., and Carmona M.J., 2004. Floral meristem identity genes are expressed duringtendril development in grapevine, Plant Physiol., 135:1491~1501
    Caicedo AL, Stinchcombe JR, Olsen KM, Schmitt J, Purugganan MD. 2004. Epistatic interaction between Arabidopsis FRI and FLC flowering time genes generates a latitudinal cline in a life history trait. Proc Natl Acad Sci U S A. 101: 15670-15675.
    Chandler J, Wilson A, Dean C.1996.Arabidopsis mutants showing an alter edresponse to vernalization,P1antJ, 10(4): 637~644
    Clarke J H, Dean C.1994.Mapping FRI, alocusxonte rolling flowering time and vernalization response in Arabidopsis Thaliana .Mo1GenGenet, 242: 81~89
    Curtis I S.2003.The noble radish: past, present and future. Trends in plant science, 7(8):305~307
    Corbesier L, Coupland G. 2006. The quest for florigen: a review of recent progress. J Exp Bot. 57(13): 3395-3403
    Cui,S.X.,et al.,A proteomic analysis of cold stress responses in rice
    seedlings.Proteomics,2005.5:p.3162-3172.
    Devlin P.F, Patel S.R, Whitelam G.C. 1998. Phytochrome E influences internode elongation and flowering time in Arabidopsis. Plant Cell.10(9):1479-1487
    Fields S.Proteomics in Genomeland.Science,2001,291(2):1 221-1 224.
    Finnegan E J, Genger R K, Kovac K, etal.1998.DNA methylation and the promotion of flowering byvernalization . Proc Natl Acad Sci USA. 95: 5824~5829
    Fornara F, Parenicova L, Falasca G., Pelucchi N., Masiero S., CiannameaS., Lopez-Dee Z., Altamura M.M, Colombo L, and Kater M.M., 2004. Functional characterization of OsMADS18 a member of the AP1/SQOA subfamilyof MADS box genes, PlantPhysiol, 135(4): 2207~2219
    Gi Cheol, Jo Jeong Lae, Mun Ji Hye.2003.Effect of triazole chemicals on bolting retardation of Chinese cabbage ( Brassica pekinensis ) in spring cultivation.Journal of the Korean Scociety for Horticultural Science , 44 (4): 434~437
    Guttormsen G ,Moe R. 1985.Effect of plant age temperature on bolting in Chinese cabbage. Scientic Horticulture, 25:217~224
    Gygi S P,Corthals G L,Zhang Y et al.Evaluation of two-dimentional gel electrophoresis-based proteome analysis techonology.PNAS,2000,97:9390-9395
    Kim SY, Park BS, Kwon SJ, Kim J, Lim MH, Park YD, Kim DY, Suh SC, Jin YM, Ahn JH, Lee YH. 2007. Delayed flowering time in Arabidopsis and Brassica rapa by the over-expression of FLOWERING LOCUS C (FLC) homologs isolated from Chinese cabbage (Brassica raga L. ssp. pekinensis). Plant Cell Report.26 (3): 327-336
    Koornneef. M., Alonso-Blanco. C., Peeters. A.J.M. and Soppe. W. 1998. Genetic control offlowering time in Arabidopsis. Ann. Rev. Plant Physiol. Plant Mol. Biol.49:345-370
    Levy YY, Mesnage S, Mylne J.S, Gendall A.R, Dean C.2002.Multiple roles of Arabidopsis VRN1 in vernalization and flowering time control.Science.297(5579):243-246.
    LeeI, Amasino R M.1995. Effect of vernalization, Photo period and light quality on the flowering phenotype of Arabidop-sis plants containing the FRIGID Agene . P1antPhysi-ology, 108: 157~162
    Lim M.H, Kim YS, Chung K.S, Seo Y.H, Lee I, Kim J, Hong C.B, Kim H.J, Park C.M. 2004. A new Arabidopsis gene, FLK, encodes an RNA binding protein with K homology motifs and regulates flowering time via FLOWERING LOCUS C. Plant Cell.16(3):731-740.
    Link A.J.,Eng J.,Schieltz D.M.,et al.Direct analysis of protein complexes using mass spectrometry.Nature Biotechnol,1999,17:676-682
    Li Z, Zhao L, Cui C, Kai G, Zhang L, Sun X, Tang K.2005.Molecular cloning and characterization of anti-bolting related gene (BrpFLC) from Brassica rapa ssp. pekinensis. Plant Sci..168: 407-413
    Mariko Oka, Yasushi Tasaka, Masaki Iwabuchi, etal.2001.Elevated sensitivity to gibberellin byvernalization in the vegetative rosette plants of Eustoma grandif lorum and Arabidopsis thaliana.P1antScience, 160: 1237~1245
    Mero C E ,Honma S.1985.Inheritance of bolting resistance in an interspecific Chinesecabbage ×turnip cross. Hort . Science , 20: 881~882
    Mero E ,Honma S.1984.Inheritane of bolt resistance in an interspecific crossof Brassica spe cies. I. Brassica napus L.×B . ampestris L . ssp.pekinensis . J . Heredity ,75: 407~470
    Michaels S.D, Amasino R.M. 1999. FLOWERING LOCUS C encodes anovel MARS domain protein that acts as a repressor of flowering. Plant Ce11.11(5): 949-956.
    Michaels S.D, Amasino R.M. 2001. Loss of FLOWERING LOCUS C activity eliminates the late-flowering phenotype of FRIGIDA and autonomous pathway mutations but not responsiveness to vernalization. Plant Cell.13(4): 935-941.
    Michaels SD, Bezerra IC and Amasino RM. 2004. FRIGIDA-related genes are required for the winter-annual habit in Arabidopsis. Proc Natl Acad Sci U S A.101: 3281-3285
    Miguel A, Green R, Nilsson O, Michael R, Weigel D. 1998. Gibberellins promote flowering of Arabidopsis by activating the LEAFY promoter. The Plant Cell,.10(5): 791-800.
    Mouradov A, Cremer F, Coupland G.2002.Control of flowering time:interacting pathways as a basis for diversity. Plant Cell, 14(Suppl):S111~S130
    Murai K, Miyamae M, Kato H., Takumi S., and Ogihar Y.,2003.WAP1, awheat APETALA1 homolog,plays a centralrole in the phase transition fro mvegetative to reprodctivegrowth, Plant Cell Physiol., 44(12): 1255~1265
    Oaks A, Wallace W, Stevens D.1972.Synthesis and turnover of nitrate reductase in core roots. Plant Physiol, 50:649
    Oka M, Tasaka Y, Iwabuchi M.2001.Elevated sensivity to gibberellin by vernalization in thevegetabiverosette plants of Eustoma grandif loram and Arabid opsist haliana .Plant Sci, 160: 1237~1245
    Osborn TC, Kole C, Parkin IAP, Sharpe AG, Kuiper M, Lydiate DJ and Trick M. 1997. Comparison of flowering time genes in Brassica rapa, B.napus and Arabidopsis thaliana. Genetics. 146:1123一1129.
    Pressman E, Aviram H.1986.Inhibition of flower in Chinese cabbage by applying heat and gro wth retardants to transplants. Plant Growth Regulation , 4:187~194
    Pressman E, Shaked R. 1988.Bolting and flowering of Chinese cabbage as affected by the intensity and source of supplementary light. Scientia Horculturae, 34(4): 177~187
    ReevesP R, 2000.Couplant G Response of plant development to environment: control of flowering by daylength and temperature. Current Opinion in Plant Biology, 3: 37~42
    ReevesP R, Couplant G .2000.Response of plant development to environment: control of flowering by daylength and temperature . Current Opinion in Plant Biology, 3: 37~42
    RooRNNEEF M, Alonso-BLANCOC, PEETERs A J M.1998.Genetic control of flowering inArabidopsis. Ann. Rev. Plant Physiol. Plant Mol. Biol. 49: 345~370
    Shchennikova A.V., Shulga O.A., Immink R., Skryabin K.G.,and Angenent G.C. 2004.Identifi cation and characterization of four chrysanthemum MADS-box genes, belonging
    Shen,S.andY.Jng,Proteomics approach to identify wound-response related proteins from rice leaf sheath,Proteomics,2003.3:p.527-535
    Sheldon C C, Burn J E, Perez P P, etal.1999.The Flfmads box gene: are pressor of flowering in Arabidopsisregulated by vernalization and methylation . Plant Cell, 11:445~458
    Schranz ME, Quijada P, Sung S, Lukens L, Amasino R, Osborn T. 2002. Characterization and effects of the replicated flowering time gene FLC in Brassica rapa. Genetics.162: 1457- 1468.
    Simpson G.G, Deao C. 2002. Arabidopsis, the rosetta stone of flowering time? Science. 296(5566): 285-289.
    Sung S, Amkasimo RM. 2004. Vernalization in Arabidopsis thaliana is mediated by the PHD finger protein VIN3. Nature.427(6970):159-164.
    The Arabidopsis Genome Initiative.Analysis of the genome sequence of the flowering plant Arabidopsis thaliana.Nature,2000,408:796~815.
    Tonge R,Shaw J,Middleton B,Rowlinson R,Rayner S,Young J,Pognan F,Hawkins E,Currie I,DavisonM.Validation and development of fluorescent two-dimensional gel electrophoresis proteomics technology.Proteomics,2001,1:377–396
    Tranquilli G,DubcovsKyJ.2000.Epistatic interaction between vernalization genes Vrn-Aml and Vrn-Amindiploid wheat . Journal of Heredity. 91(4):304~306
    Wilkins M R,Sanchez J C,Williams K L,et al.Progress with proteome projects:why all proteins expressed by a genome should be identified and how to do it.Biotechnology andgenetic Engineering Reviews,1995,13:19-45
    Yui S J, Yoshikawa H, Kuginuki Y.1991 .Breeding of slow bolting Brasica campestris varietywith no low temperature sensitivity. Cruciferae-Newsletter, No.14-1 5,56~57
    Yui S, Yoshikawa H. 1991.Bolting resistant breeding of Chinese cabbage 1 Flower induction of latebolting variety without chilling treatment. Euphytica, 52(3): 171~176
    Yui S. Yoshikawa H. 1992. Breeding of bolting resistance in Chinese cabbage critical day length for flower of late bolting material with no chilling requirement. Journal of the Japanese Society for Horticultural Science.61:3,565-568

© 2004-2018 中国地质图书馆版权所有 京ICP备05064691号 京公网安备11010802017129号

地址:北京市海淀区学院路29号 邮编:100083

电话:办公室:(+86 10)66554848;文献借阅、咨询服务、科技查新:66554700