第十章-果实发育.ppt

单击此处编辑母版标题样式,单击此处编辑母版文本样式,第二级,第三级,第四级,第五级,*,第,10,章 果实发育,Seed development:,种子由受精后的胚珠,发育而来,Fruit development:,How fruits develop?,From carpeles and ovary,果实的发育与类型,一、果实的发育：,子房发育成果实。,根据子房在果实发育中的作用分：真果和假果,二、果实的结构,1,、真果：仅由子房发育而成的果实。如桃的果实。,（一）果实的结构组成,外果皮,果皮 中果皮,果实 内果皮,胎座,胚珠 种子,2.,假果的结构,除了子房以外，花托、花被甚至是整个花序参与发育而成的果实。如梨、苹果等。,果实上残留的花萼与果柄分离即为假果,!,根据成熟果实果皮的干燥程度可分为,:,干果,果实可分为,:,干果,肉质果,肉质果,第一节 拟南芥果实发育,拟南芥长角果结构,Stigma,柱头,Style,花柱,Ovary,子房,Valve margin,瓣边缘,Replum,胎座框,Gynophore,雌蕊柄,Nectaries,蜜腺,Abscission zone,脱落层,拟南芥长角果横切图,Valve margin,瓣边缘,Replum,胎座框,Septum,隔膜,Gynophore,雌蕊柄,Nectaries,蜜腺,Abscission zone,脱落层,拟南芥长角果从授粉到裂果的发育过程,花原基起始,花原基隆起并与,SAM,分离,花萼原基形成,花萼近远轴形成,花瓣与雄蕊原基产生,顶端细胞不分裂，导致凹陷形成,两侧发育，形成中空的管状结构,Stage 6,：萼片包围花芽及雌蕊形成,拟南芥长角果从授粉到裂果的发育过程,Stage 7,：中间花药突起及中空状雌蕊形成,Mr,：中间脊,O:,胎座,Stage 8 is marked by the formation of locules in the,anthers.,心皮具有,5-6,层细胞,Stage 8,：花药中小室形成及管状雌蕊增大,Stage 9 is marked by the petal primordia becoming stalked,at the base.,O,：胎座,S,：隔膜,Stage 9,：花瓣原基产生及隔膜形成,Stage 10 begins when the petal primordia reach the length,of the short stamens(Smyth et al.,1990).,Stage 10,：花瓣原基达到较短的花药的高度,The stigmatic papillae cover the entire surface of the stigma,during stage 11(Figure 6A).,funiculus,珠柄,inner integument,内珠被,outer integument,外珠被,n,珠心,Stage 11,：柱头乳突细胞形成，内外珠被开始形成,Stage 12,：花瓣达到较长花药高度，雌蕊成熟，花柱形成,Stage 13,：花开放，传粉、授精,At stage 14,the anthers extend above the top of the stigma,Stage 14,：花药达到了柱头的高度、授精完成,Stage 15,：雌蕊进一步生长,Stage 16,：花瓣、萼片、雄蕊凋谢,Stage 16,：角果发育,木质化层形成，,enb,层木质化,果实成熟及开裂机理,影响角果发育的相关基因,影响心皮特性的调控,控制果实形状,决定种子散发的瓣边缘特性基因,决定瓣发育的基因,决定果脊发育的基因,心皮的特征机构包括：胚珠，柱头组织，花柱组织，瓣细胞，隔膜原基。,AGAMOUS,与心皮发育,AP2,基因突变导致花瓣心皮化,ap2-ag,双突变体仍然保持多心皮专一组织,Spt-ap2-pi-ag,突变体外轮心皮有胚珠状突起,CRC,基因的进一步突变消除了胚珠类似结构,影响心皮数量的基因：,stm-clv-wus,stm,杂合体可以部分抑制由于,clv,变异导致的心皮数量增加,过量表达,WUS,和,AG,均可以导致心皮数量增加,果实形状的控制,ERECTA(ER),编码富含亮氨酸重复序列的受体蛋白激酶，,ER,在茎顶端分生组织中强烈表达，作为细胞外信号调节细胞分裂和扩张，控制果实形状。,决定种子散发的基因调控,SHP,，,IND,，,ALC,，,RPL,，,FUL,SHP,对木质化层和分离层的形成具有重要作用,ALC,影响分离层分化,IND,专一化的决定木质化层的形成,SHP,IND,ALC,FUL,RPL,表达模式与调控关系,FUL,对其他基因的调控用,第二节 番茄果实发育,fw2.2,:A Quantitative Trait Locus Key to the Evolution of Tomato Fruit Size,Anne Frary,T.Clint Nesbitt,.,Steven D.Tanksley,Science 2000 289:85-88,Cultivated tomato can be 1000 times larger than the wild tomatoes.,野生型,栽培种,控制数量性状的基因能通过分子标记定位在遗传图谱上,怎么最终确定这些基因呢？,在番茄遗传图谱中定位了,28,个控制果重的,QTL,，其中主效位点,fw2.2,定位在第二连锁群上,QTL,定位：,利用番茄品种,L.esculentum,和,NIL,构建的分离群体将,fw2.2,进行精细定位,fw2.2,候选基因所在的区段在,XO31,到,XO33,之间。经过序列分析发现四个候选,cDNAs(70,27,38,and 44),和四个柯斯载体,(cos62,84,69,and 50,）在这个重组区域内,.,Arrows represent the cDNAs and the vertical lines are positions of RFLP or CAPs markers.,序列分析发现,cos50,包含有,cDNA44 and ORFX,发现,ORFX,是,QTL,的最有可能的关键候选基因,.,ORFX,和数据库中其它序列比对发现和动物中控制细胞分裂的,RAX,同源,.,no cos50,R1 progeny with cos50(containing the small-fruit allele)Transgenic tomato,Phenotypic effect of the,fw2.2,transgene in the cultivar Mogeor,having cos50,Non-transgenic tomato,fw2.2,:A Quantitative Trait Locus Key to the Evolution of Tomato Fruit Size,Science 2000 289:85-88.,Fruit size is controlled by two processes:cell cycle,（细胞周期）,and organ number determination,（器官数量的决定）,.,The key gene in the first process has been isolated and corresponds to fw2.2.,How about the second process?,fw2.2,a,The locule,（小室）,number in tomato can be varied with the carpel number,Progression of fruit size increase during tomato domestication,a,b,c,A closely related wild species,An intermediate-sized processing variety,A large-sized fresh market variety,Jumbo Red,a b:Increasing in cell division,（细胞分裂速度增加）,.,b c:Increasing in locule number,（小室数目增加）,.,The research group identified a major genetic,fas,determinant of large tomato size that increases the number of female reproductive organs in a tomato flower,and thus the number of compartments in the fruit.,fas,基因通过图位克隆方法被鉴定出来,The mutant allele of the gene,fas,(,fasciated,),was inserted by a 8-kb sequence caused fail of gene expressing,FAS is expressed in young carpel to negatively control the locule number,Regulatory change in YABBY-like transcription factor led to evolution of extreme fruit size during tomato domestication,Nature Genetics 2008,fw2.2,:A Quantitative Trait Locus Key to the Evolution of Tomato Fruit Size,Science 2000,1 Fruit size,2 Fruit maturation,肉质果实成熟常常有呼吸峰出现,伴随乙烯的大量产生，如番茄、香蕉。,Qestion:,What the ethylene works for?,Fruit maturing,氨基酸丙烷羧化酶,(ACC),：乙烯合成的限速酶,Qestion:,How to keep fresh of tomato fruit with the information of ACC by means of genetic manipulation?,Antisense 35S:,ACC,Inhibition of tomato fruit ripening by antisense ACC synthase constructs.,百日鲜,-,华中农大第三节 肉质果实色泽形成,Plant flower color formation,Betanin,Lutein,Carotenoid pigment,Cyanidin,anthocyanidin,Anthocyanidins,矢车菊色素,天竺葵色素,花翠素,花青素与健康,This states that the French have only 30%the rate of heart attacks of Americans,despite eating four times more butter,having higher blood cholesterol and higher blood pressure,and has been attributed to the mododerate consumption of red wine on a daily basis by the French.,花青素具有较高抗氧化作用，可以保护遭受氧化性损伤，减少心血管疾病的发生,.,花青素合成代谢途径,丙二酰,C,O,A,-,香豆酰,C,O,A,Blood Oranges,主要起源,:,意大利西西里，西班牙，中国,摩洛血橙,塔罗科血橙,桑吉耐劳血橙,已有的研究发现，植物果实中的花青素合成多收到,R2R3 MYB,转录因子的调控,根据已有的,R2R3,转录因子设计引物，从血橙中扩增获得一个,cDNA,片段，而在黄色橙中没有扩增到，把这段序列命名为,Ruby,.,With,Ruby,Without,Ruby,Ruby Is a Regulator of Anthocyanin Biosynthesis,转基因验证表明，,Ruby,可以增加花青素合成,与其他因子共表达进一步加强了,Ruby,的调控效果,:(1),35S:Ruby,(2)35S:Ruby 3 35S:Delila,(3)35S:Ruby 3 35S:Mutabilis,(4)35S:Rosea1 3 35S:Delila,and(5)35S:Rosea1 3 35S:Mutabilis.,(2)35S:Ruby,(1)Wild type,35S:Rubya and 35S:Rubyc represent two independent transformants,A cDNA encoding a clade 3f,bHLH protein(Cs MYC2)has been identified in orange(Cultrone et al.,2010).,The Cs MYC2 gene(encoding the,bHLH,partner in the MBW complex)is expressed at detectable levels in both blond and blood oranges.,The expression of,Ruby,limits anthocyanin biosynthesis in oranges!,基因结构不同导致花青素合成有别,转座子表达导致,Ruby,在血橙中高表达,温度敏感,An Independent Blood Orange Accession fromJingxian,China,The blood phenotypes in both Jingxian and the Sicilian group,of blood oranges are therefore of independent origin,showing,that different members of the same family of retrotransposons,may alter the expression of nearby genes through parallel but,distinct mechanisms.,硕果累累！,