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作者赵会彦（2019）在《白菜型油菜和甘蓝型油菜粒色调控机制差异比较分析》一文中研究指出：油菜是世界上重要的油料作物之一,黄籽油菜与普通黑籽油菜相比,具有油质清澈、油脂和蛋白质含量高、皮壳率低、饼粕饲用价值高等优点,因此受到育种家的重视。目前,世界上广泛种植的芸薹属油料作物栽培种中,白菜型油菜、芥菜型油菜和埃塞俄比亚芥中均存在天然的黄籽资源,但种植面积最大的甘蓝型油菜中却缺乏天然的黄籽资源,且甘蓝型油菜粒色成因复杂,调控甘蓝型油菜粒色主效基因仍不明确,限制了黄籽这一优良性状在油菜产业中的应用。白菜型油菜作为甘蓝型油菜的二倍体亲本之一,与其在遗传进化上密切相关。相对于甘蓝型油菜,其粒色遗传规律较简单,且种植广泛,遗传资源丰富。因此,本研究拟通过对拟南芥、白菜、甘蓝和甘蓝型油菜中各类黄酮途径基因的家族成员的比较鉴定分析,粒色关键基因在不同遗传背景的黄、黑(褐)籽白菜型油菜和甘蓝型油菜材料中的表达特征分析,白菜型油菜和甘蓝型油菜不同发育时期的种子中转录组及代谢组的差异比较分析等方面,明确白菜型油菜和甘蓝型油菜粒色调控的差异机制,为甘蓝型黄籽油菜育种奠定基础。具体研究结果如下:1.十字花科物种类黄酮途径基因成员的全基因组鉴定本研究对拟南芥、芸薹属作物及十字花科其它已测序物种中TT同源基因进行分析,结果在17种十字花科植物中,针对21个类黄酮途径基因共检测到649个基因拷贝,且这些基因拷贝在所有物种中均能被检测到,但不同物种间的类黄酮途径基因之间存在差异,包括染色体间的重组和部分基因家族成员拷贝数的增加或缺失等现象。对每个类黄酮途径基因在17个物种中的同源基因进行系统进化分析发现,17个十字花科植物中的类黄酮途径基因也可大致分为分别与芸薹属(Brassica)和拟南芥属(Arabidopsis)进化关系较近的两类,表明芸薹属作物与拟南芥相比,其类黄酮途径存在一定的差异。通过白菜、甘蓝和甘蓝型油菜中类黄酮途径基因的同源比对发现,甘蓝型油菜中A和C亚基因组的基因分别与白菜和甘蓝中的同源基因更相似,而白菜和甘蓝两者之间的差异则较大;与两个二倍体亲本相比,一些重要的转录因子或结构基因在甘蓝型油菜染色体中的位置发生了变化。2.不同来源白菜型油菜和甘蓝型油菜中粒色相关基因的表达模式对比分析大量研究已证实类黄酮途径基因在种子粒色形成过程中的重要作用,此外,在本课题组前期对甘蓝型油菜粒色主效基因的定位中,筛选得到若干重要候选基因,为探究白菜型油菜和甘蓝型油菜中类黄酮途径基因调控模式的差异,本研究利用不同来源的白菜型油菜和甘蓝型油菜黄、黑籽材料对各类黄酮途径基因和甘蓝型油菜粒色候选基因在不同粒色、不同发育时期种子中的表达进行系统分析。首先对甘蓝型油菜和白菜型油菜中类黄酮途径基因的整体差异分析发现,白菜型油菜黄籽材料中部分类黄酮途径结构基因和调控基因的表达水平极显著低于对应的黑籽材料,但在甘蓝型油菜黄籽材料中,尽管其表达量显著低于黑籽材料,但仍有一定量的表达。白菜中筛选得到的若干粒色主效基因在甘蓝型油菜黄、黑籽材料间均表现为显著差异,而甘蓝型油菜中的大部分粒色主效候选基因在白菜型油菜黄、黑籽材料中表达差异不显著,表明甘蓝型油菜粒色主效基因与白菜型油菜粒色主效基因有一定的差异,且在甘蓝型油菜中,其粒色主效基因对白菜型油菜粒色主效基因的同源基因应具有调控作用。对粒色相关基因的不同成员的表达分析显示,来自于An和Cn亚基因组的同源基因均有一定量的表达,且表达模式也有相似之处,推测C基因组类黄酮途径基因的导入,对A基因组类黄酮途径基因的表达会产生影响。3.不同来源白菜型油菜和甘蓝型油菜种子中代谢组分差异对比分析本研究利用不同来源的白菜型油菜和甘蓝型油菜黄、黑籽品种,通过对不同发育时期的种子进行LC/MS及色素含量测定分析,确定其中主要类黄酮代谢产物的种类和含量的差异,分析白菜型油菜和甘蓝型油菜粒色形成物质之间的差异。在甘蓝型油菜各品种间,检测到更多的单独存在于一个或两个品种中的物质,表明甘蓝型油菜各品种间类黄酮途径的差异大于白菜型油菜各品种之间的差异,由此推测类黄酮途径在甘蓝型油菜品种间的分化更大,而白菜型油菜中的类黄酮代谢途径在品种间更稳定。借鉴前人研究结果,共鉴定得到类黄酮类或含芥子酰基类物质35种,包括9种表儿茶素单体(Epicatechin)或多聚体,8种槲皮素(Quercetin)及其衍生物,3种异鼠李素(Isorhamnetin)及其衍生物,6种山奈酚(Kaempferol)及其衍生物,9种芥酸及含芥子酰基的代谢物。对这些类黄酮类物质和部分未知物进行定量分析并归类,表儿茶素及由表儿茶素缩合形成的不同聚合态的原花青素多聚体,是白菜型油菜和甘蓝型油菜黄、黑籽油菜间差异最显著的一类化合物,而其它类黄酮途径中间产物在白菜型油菜和甘蓝型油菜中的变化动态各异,可能是由于鉴定出的类黄酮类物质种类较少,也可能由于各品种间遗传背景各异、粒色调控基因不同导致的。4.不同发育时期白菜型和甘蓝型油菜种子转录组和粒色主效基因差异对比分析为了探究白菜型油菜和甘蓝型油菜粒色主效基因对类黄酮途径调控方式的差异,本研究利用一个白菜型油菜分离群体,对白菜型油菜粒色主效基因进行进一步的精细定位,对白菜型油菜和甘蓝型油菜中的粒色主效基因进行了进一步的归纳整理,通过转录组测序,比较了白菜型油菜和甘蓝型油菜黄、黑籽材料间种子中基因整体转录水平的差异,对二者粒色主效基因对类黄酮途径的调控机制进行初步预测。通过对白菜型油菜粒色基因的精细定位,结合转录组测序的结果,筛选到11个与黄籽沙逊粒色形成相关的候选基因。通过对白菜型油菜和甘蓝型油菜粒色主效基因的归纳分析发现,甘蓝型油菜中的粒色主效基因多位于A9染色体末端,而白菜中的粒色主效基因位点多定位于A9染色体中段。对甘蓝型油菜和白菜型油菜转录组测序的结果进行比较分析发现,在甘蓝型油菜和白菜型油菜中,最显著富集的下调代谢途径与本研究一致,为苯丙烷途径和类黄酮代谢途径,但显著富集的上调代谢通路在两个物种中不同,表明甘蓝型油菜和白菜型油菜中粒色主效基因的功能差异。通过比较粒色主效基因较明确的甘蓝型油菜和白菜型油菜中粒色基因的表达,发现在甘蓝型油菜中起主效调控作用的基因,其在白菜型油菜中的直系同源基因对白菜型油菜粒色的形成,无明显调控作用,表明这些粒色主效基因是在甘蓝型油菜物种形成之后新产生的粒色调控基因,其可能通过直接调控一些已发现的重要粒色调控基因(如TT1,TT8,TTG1等)或通过与其相互作用,来控制类黄酮代谢途径,进一步证实类黄酮途径在芸苔属作物中不断发生进化。综上,TT基因的表达模式分析、代谢组及转录组分析的结果均进一步证明类黄酮代谢途径与油菜粒色形成的重要关系,原花色素是影响白菜型油菜和甘蓝型油菜粒色差异的重要色素物质。与白菜型油菜黄籽材料相比,甘蓝型油菜粒色主效基因沉默后,其类黄酮途径仍未被完全抑制,可能与甘蓝型油菜粒色性状主基因+微效多基因的遗传模式相关;另外,Cn亚基因组的导入后,不但使An亚基因组中TT基因的物理位置发生变化,新导入的TT基因也参与了类黄酮代谢途径的调控,这些都可能是甘蓝型油菜粒色调控机制比白菜型油菜复杂的原因。甘蓝型油菜中新发现的粒色主效基因,它们的同源基因在白菜型油菜中并不参与类黄酮途径的调控,而在甘蓝型油菜中,其可能通过直接或间接调控已知的重要TT基因的表达,来控制甘蓝型油菜种皮色素的合成
Abstract
you cai shi shi jie shang chong yao de you liao zuo wu zhi yi ,huang zi you cai yu pu tong hei zi you cai xiang bi ,ju you you zhi qing che 、you zhi he dan bai zhi han liang gao 、pi ke lv di 、bing po si yong jia zhi gao deng you dian ,yin ci shou dao yo chong jia de chong shi 。mu qian ,shi jie shang an fan chong zhi de yun tai shu you liao zuo wu zai pei chong zhong ,bai cai xing you cai 、gai cai xing you cai he ai sai e bi ya gai zhong jun cun zai tian ran de huang zi zi yuan ,dan chong zhi mian ji zui da de gan lan xing you cai zhong que que fa tian ran de huang zi zi yuan ,ju gan lan xing you cai li se cheng yin fu za ,diao kong gan lan xing you cai li se zhu xiao ji yin reng bu ming que ,xian zhi le huang zi zhe yi you liang xing zhuang zai you cai chan ye zhong de ying yong 。bai cai xing you cai zuo wei gan lan xing you cai de er bei ti qin ben zhi yi ,yu ji zai wei chuan jin hua shang mi qie xiang guan 。xiang dui yu gan lan xing you cai ,ji li se wei chuan gui lv jiao jian chan ,ju chong zhi an fan ,wei chuan zi yuan feng fu 。yin ci ,ben yan jiu ni tong guo dui ni na gai 、bai cai 、gan lan he gan lan xing you cai zhong ge lei huang tong tu jing ji yin de jia zu cheng yuan de bi jiao jian ding fen xi ,li se guan jian ji yin zai bu tong wei chuan bei jing de huang 、hei (he )zi bai cai xing you cai he gan lan xing you cai cai liao zhong de biao da te zheng fen xi ,bai cai xing you cai he gan lan xing you cai bu tong fa yo shi ji de chong zi zhong zhuai lu zu ji dai xie zu de cha yi bi jiao fen xi deng fang mian ,ming que bai cai xing you cai he gan lan xing you cai li se diao kong de cha yi ji zhi ,wei gan lan xing huang zi you cai yo chong dian ding ji chu 。ju ti yan jiu jie guo ru xia :1.shi zi hua ke wu chong lei huang tong tu jing ji yin cheng yuan de quan ji yin zu jian ding ben yan jiu dui ni na gai 、yun tai shu zuo wu ji shi zi hua ke ji ta yi ce xu wu chong zhong TTtong yuan ji yin jin hang fen xi ,jie guo zai 17chong shi zi hua ke zhi wu zhong ,zhen dui 21ge lei huang tong tu jing ji yin gong jian ce dao 649ge ji yin kao bei ,ju zhe xie ji yin kao bei zai suo you wu chong zhong jun neng bei jian ce dao ,dan bu tong wu chong jian de lei huang tong tu jing ji yin zhi jian cun zai cha yi ,bao gua ran se ti jian de chong zu he bu fen ji yin jia zu cheng yuan kao bei shu de zeng jia huo que shi deng xian xiang 。dui mei ge lei huang tong tu jing ji yin zai 17ge wu chong zhong de tong yuan ji yin jin hang ji tong jin hua fen xi fa xian ,17ge shi zi hua ke zhi wu zhong de lei huang tong tu jing ji yin ye ke da zhi fen wei fen bie yu yun tai shu (Brassica)he ni na gai shu (Arabidopsis)jin hua guan ji jiao jin de liang lei ,biao ming yun tai shu zuo wu yu ni na gai xiang bi ,ji lei huang tong tu jing cun zai yi ding de cha yi 。tong guo bai cai 、gan lan he gan lan xing you cai zhong lei huang tong tu jing ji yin de tong yuan bi dui fa xian ,gan lan xing you cai zhong Ahe Cya ji yin zu de ji yin fen bie yu bai cai he gan lan zhong de tong yuan ji yin geng xiang shi ,er bai cai he gan lan liang zhe zhi jian de cha yi ze jiao da ;yu liang ge er bei ti qin ben xiang bi ,yi xie chong yao de zhuai lu yin zi huo jie gou ji yin zai gan lan xing you cai ran se ti zhong de wei zhi fa sheng le bian hua 。2.bu tong lai yuan bai cai xing you cai he gan lan xing you cai zhong li se xiang guan ji yin de biao da mo shi dui bi fen xi da liang yan jiu yi zheng shi lei huang tong tu jing ji yin zai chong zi li se xing cheng guo cheng zhong de chong yao zuo yong ,ci wai ,zai ben ke ti zu qian ji dui gan lan xing you cai li se zhu xiao ji yin de ding wei zhong ,shai shua de dao re gan chong yao hou shua ji yin ,wei tan jiu bai cai xing you cai he gan lan xing you cai zhong lei huang tong tu jing ji yin diao kong mo shi de cha yi ,ben yan jiu li yong bu tong lai yuan de bai cai xing you cai he gan lan xing you cai huang 、hei zi cai liao dui ge lei huang tong tu jing ji yin he gan lan xing you cai li se hou shua ji yin zai bu tong li se 、bu tong fa yo shi ji chong zi zhong de biao da jin hang ji tong fen xi 。shou xian dui gan lan xing you cai he bai cai xing you cai zhong lei huang tong tu jing ji yin de zheng ti cha yi fen xi fa xian ,bai cai xing you cai huang zi cai liao zhong bu fen lei huang tong tu jing jie gou ji yin he diao kong ji yin de biao da shui ping ji xian zhe di yu dui ying de hei zi cai liao ,dan zai gan lan xing you cai huang zi cai liao zhong ,jin guan ji biao da liang xian zhe di yu hei zi cai liao ,dan reng you yi ding liang de biao da 。bai cai zhong shai shua de dao de re gan li se zhu xiao ji yin zai gan lan xing you cai huang 、hei zi cai liao jian jun biao xian wei xian zhe cha yi ,er gan lan xing you cai zhong de da bu fen li se zhu xiao hou shua ji yin zai bai cai xing you cai huang 、hei zi cai liao zhong biao da cha yi bu xian zhe ,biao ming gan lan xing you cai li se zhu xiao ji yin yu bai cai xing you cai li se zhu xiao ji yin you yi ding de cha yi ,ju zai gan lan xing you cai zhong ,ji li se zhu xiao ji yin dui bai cai xing you cai li se zhu xiao ji yin de tong yuan ji yin ying ju you diao kong zuo yong 。dui li se xiang guan ji yin de bu tong cheng yuan de biao da fen xi xian shi ,lai zi yu Anhe Cnya ji yin zu de tong yuan ji yin jun you yi ding liang de biao da ,ju biao da mo shi ye you xiang shi zhi chu ,tui ce Cji yin zu lei huang tong tu jing ji yin de dao ru ,dui Aji yin zu lei huang tong tu jing ji yin de biao da hui chan sheng ying xiang 。3.bu tong lai yuan bai cai xing you cai he gan lan xing you cai chong zi zhong dai xie zu fen cha yi dui bi fen xi ben yan jiu li yong bu tong lai yuan de bai cai xing you cai he gan lan xing you cai huang 、hei zi pin chong ,tong guo dui bu tong fa yo shi ji de chong zi jin hang LC/MSji se su han liang ce ding fen xi ,que ding ji zhong zhu yao lei huang tong dai xie chan wu de chong lei he han liang de cha yi ,fen xi bai cai xing you cai he gan lan xing you cai li se xing cheng wu zhi zhi jian de cha yi 。zai gan lan xing you cai ge pin chong jian ,jian ce dao geng duo de chan du cun zai yu yi ge huo liang ge pin chong zhong de wu zhi ,biao ming gan lan xing you cai ge pin chong jian lei huang tong tu jing de cha yi da yu bai cai xing you cai ge pin chong zhi jian de cha yi ,you ci tui ce lei huang tong tu jing zai gan lan xing you cai pin chong jian de fen hua geng da ,er bai cai xing you cai zhong de lei huang tong dai xie tu jing zai pin chong jian geng wen ding 。jie jian qian ren yan jiu jie guo ,gong jian ding de dao lei huang tong lei huo han gai zi xian ji lei wu zhi 35chong ,bao gua 9chong biao er cha su chan ti (Epicatechin)huo duo ju ti ,8chong hu pi su (Quercetin)ji ji yan sheng wu ,3chong yi shu li su (Isorhamnetin)ji ji yan sheng wu ,6chong shan nai fen (Kaempferol)ji ji yan sheng wu ,9chong gai suan ji han gai zi xian ji de dai xie wu 。dui zhe xie lei huang tong lei wu zhi he bu fen wei zhi wu jin hang ding liang fen xi bing gui lei ,biao er cha su ji you biao er cha su su ge xing cheng de bu tong ju ge tai de yuan hua qing su duo ju ti ,shi bai cai xing you cai he gan lan xing you cai huang 、hei zi you cai jian cha yi zui xian zhe de yi lei hua ge wu ,er ji ta lei huang tong tu jing zhong jian chan wu zai bai cai xing you cai he gan lan xing you cai zhong de bian hua dong tai ge yi ,ke neng shi you yu jian ding chu de lei huang tong lei wu zhi chong lei jiao shao ,ye ke neng you yu ge pin chong jian wei chuan bei jing ge yi 、li se diao kong ji yin bu tong dao zhi de 。4.bu tong fa yo shi ji bai cai xing he gan lan xing you cai chong zi zhuai lu zu he li se zhu xiao ji yin cha yi dui bi fen xi wei le tan jiu bai cai xing you cai he gan lan xing you cai li se zhu xiao ji yin dui lei huang tong tu jing diao kong fang shi de cha yi ,ben yan jiu li yong yi ge bai cai xing you cai fen li qun ti ,dui bai cai xing you cai li se zhu xiao ji yin jin hang jin yi bu de jing xi ding wei ,dui bai cai xing you cai he gan lan xing you cai zhong de li se zhu xiao ji yin jin hang le jin yi bu de gui na zheng li ,tong guo zhuai lu zu ce xu ,bi jiao le bai cai xing you cai he gan lan xing you cai huang 、hei zi cai liao jian chong zi zhong ji yin zheng ti zhuai lu shui ping de cha yi ,dui er zhe li se zhu xiao ji yin dui lei huang tong tu jing de diao kong ji zhi jin hang chu bu yu ce 。tong guo dui bai cai xing you cai li se ji yin de jing xi ding wei ,jie ge zhuai lu zu ce xu de jie guo ,shai shua dao 11ge yu huang zi sha xun li se xing cheng xiang guan de hou shua ji yin 。tong guo dui bai cai xing you cai he gan lan xing you cai li se zhu xiao ji yin de gui na fen xi fa xian ,gan lan xing you cai zhong de li se zhu xiao ji yin duo wei yu A9ran se ti mo duan ,er bai cai zhong de li se zhu xiao ji yin wei dian duo ding wei yu A9ran se ti zhong duan 。dui gan lan xing you cai he bai cai xing you cai zhuai lu zu ce xu de jie guo jin hang bi jiao fen xi fa xian ,zai gan lan xing you cai he bai cai xing you cai zhong ,zui xian zhe fu ji de xia diao dai xie tu jing yu ben yan jiu yi zhi ,wei ben bing wan tu jing he lei huang tong dai xie tu jing ,dan xian zhe fu ji de shang diao dai xie tong lu zai liang ge wu chong zhong bu tong ,biao ming gan lan xing you cai he bai cai xing you cai zhong li se zhu xiao ji yin de gong neng cha yi 。tong guo bi jiao li se zhu xiao ji yin jiao ming que de gan lan xing you cai he bai cai xing you cai zhong li se ji yin de biao da ,fa xian zai gan lan xing you cai zhong qi zhu xiao diao kong zuo yong de ji yin ,ji zai bai cai xing you cai zhong de zhi ji tong yuan ji yin dui bai cai xing you cai li se de xing cheng ,mo ming xian diao kong zuo yong ,biao ming zhe xie li se zhu xiao ji yin shi zai gan lan xing you cai wu chong xing cheng zhi hou xin chan sheng de li se diao kong ji yin ,ji ke neng tong guo zhi jie diao kong yi xie yi fa xian de chong yao li se diao kong ji yin (ru TT1,TT8,TTG1deng )huo tong guo yu ji xiang hu zuo yong ,lai kong zhi lei huang tong dai xie tu jing ,jin yi bu zheng shi lei huang tong tu jing zai yun tai shu zuo wu zhong bu duan fa sheng jin hua 。zeng shang ,TTji yin de biao da mo shi fen xi 、dai xie zu ji zhuai lu zu fen xi de jie guo jun jin yi bu zheng ming lei huang tong dai xie tu jing yu you cai li se xing cheng de chong yao guan ji ,yuan hua se su shi ying xiang bai cai xing you cai he gan lan xing you cai li se cha yi de chong yao se su wu zhi 。yu bai cai xing you cai huang zi cai liao xiang bi ,gan lan xing you cai li se zhu xiao ji yin chen mo hou ,ji lei huang tong tu jing reng wei bei wan quan yi zhi ,ke neng yu gan lan xing you cai li se xing zhuang zhu ji yin +wei xiao duo ji yin de wei chuan mo shi xiang guan ;ling wai ,Cnya ji yin zu de dao ru hou ,bu dan shi Anya ji yin zu zhong TTji yin de wu li wei zhi fa sheng bian hua ,xin dao ru de TTji yin ye can yu le lei huang tong dai xie tu jing de diao kong ,zhe xie dou ke neng shi gan lan xing you cai li se diao kong ji zhi bi bai cai xing you cai fu za de yuan yin 。gan lan xing you cai zhong xin fa xian de li se zhu xiao ji yin ,ta men de tong yuan ji yin zai bai cai xing you cai zhong bing bu can yu lei huang tong tu jing de diao kong ,er zai gan lan xing you cai zhong ,ji ke neng tong guo zhi jie huo jian jie diao kong yi zhi de chong yao TTji yin de biao da ,lai kong zhi gan lan xing you cai chong pi se su de ge cheng
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