影响甘蓝型油菜每角粒数因素分析
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摘要
为培育高角粒数的油菜品种,研究种子(或胚珠)败育对每角果粒数的影响,选取36份双单倍体(DH)材料,采用自然开放和强制自交、杂交、辅助人工授粉等4种处理方式,分别调查各处理中初花时胚珠数和成熟风干角果中正常成熟籽粒和败育种子的数量与形态变化,探寻种子(或胚珠)败育发生原因。结果发现:材料的初始胚珠数为23. 05～36. 15个,差异极显著。种子成熟后,每角粒数为平均16. 80;胚珠数与每角粒数间的相关系数为0. 77**;各材料每角粒数占初始胚珠数的百分比平均为56. 19%,近44. 00%的胚珠在种子发育过程中会发生败育;胚珠数与每角粒数呈正相关;败育是导致每角粒数减少的主要原因。不同材料胚珠数不同,胚珠数、败育比率均与材料基因型有关。根据胚珠的直径可将败育类型划分为3种:a类型(直径小于0. 1mm)、b类型(直径0. 1～1. 0mm)、c类型(胚败育)。3种败育类型在36个参试材料和4种试验处理中均有发生,前两种类型败育是胚珠(或种子)败育的主要类型,平均发生概率分别为19. 34%和7. 91%,胚败育发生概率较低,仅为1. 54%。胚珠(或种子)败育现象普遍存在于甘蓝型油菜种子发育过程中,原因可能与花粉和柱头发育、花粉与柱头识别、受精以及胚初期发育异常相关;胚败育可能与胚发育停滞有关。
To improve seed setting rate and get more seeds in siliques of rapeseed,abortion of ovules and seeds were investigated in 36 DH(double hyploid) materials under 4 pollinating treatments(open pollination,forced selfing,hybridizing and assisted artificial pollination).The initial number of ovules,normal mature seeds and abortion seeds were investigated to better understand the abortion period and causes.Statistic results showed that the initial ovule numbers of different materials had highly significantly variation,ranging from 23.05 to 36.15 with an average of 29.48.After maturing,the seeds number in single silique was 8.82-26.38,with an average of 16.80.The correlation coefficient between the number of ovules and the number of seeds per siliques was 0.77.Average,56.19% of the initial ovules could develop into mature seeds and nearly 44% ovules aborted during the seed development.The abortion rate was found to be related to genotype as well.The abortion could be divided into 3 types.They were divided by ovules diameter as:a(less than 0.1 mm),b(between 0.1-1.0 mm) and c(embryo abortion).They occurred in all 36 materials under 4 treatments.Type a and b were the main types of ovule(or seed)abortion,with average occurrences of 19.34% and 7.91% respectively.Type c occurred only 1.54%,which might be related to the ceasing of embryonic developmental.These results indicated that the ovules number was related to genotype,and the abortion occurrence and types were also related to genotype.The cause of abortion might be in stigma development,stigma and pollen recognition,and/or in fertilization and initial embryo development.
To improve seed setting rate and get more seeds in siliques of rapeseed,abortion of ovules and seeds were investigated in 36 DH(double hyploid) materials under 4 pollinating treatments(open pollination,forced selfing,hybridizing and assisted artificial pollination).The initial number of ovules,normal mature seeds and abortion seeds were investigated to better understand the abortion period and causes.Statistic results showed that the initial ovule numbers of different materials had highly significantly variation,ranging from 23.05 to 36.15 with an average of 29.48.After maturing,the seeds number in single silique was 8.82-26.38,with an average of 16.80.The correlation coefficient between the number of ovules and the number of seeds per siliques was 0.77.Average,56.19% of the initial ovules could develop into mature seeds and nearly 44% ovules aborted during the seed development.The abortion rate was found to be related to genotype as well.The abortion could be divided into 3 types.They were divided by ovules diameter as:a(less than 0.1 mm),b(between 0.1-1.0 mm) and c(embryo abortion).They occurred in all 36 materials under 4 treatments.Type a and b were the main types of ovule(or seed)abortion,with average occurrences of 19.34% and 7.91% respectively.Type c occurred only 1.54%,which might be related to the ceasing of embryonic developmental.These results indicated that the ovules number was related to genotype,and the abortion occurrence and types were also related to genotype.The cause of abortion might be in stigma development,stigma and pollen recognition,and/or in fertilization and initial embryo development.
引文
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[10]许汪洁,傅鹰,董红利,等.一个甘蓝型油菜胚珠败育突变体的形态鉴定及生理特性分析[J].中国农业科学,2014,47(10):2062-2068.
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[12]袁豆豆.蜜蜂授粉对新疆油菜增产效果影响[J].新疆农业科技,2016(4):47.
[13]侯燕,侯剑,韩宏仕,等.杂交油菜制种中提高母本结实率的研究Ⅵ.蜜蜂放养对制种结实率和产量的影响[J].贵州农业科学,2008,36(1):22,26.
[14]田祖庆.油菜不结实的原因分析及应对策略[J].作物研究,2015,29(6):665-666.
[15]周斌,胡德厚,胡爱民,等.峡江县油菜结实率低的原因调查与分析[J].安徽农学通报,2009,15(16):123,142.
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[17] Wang X J,Mathieu A,Cournède P H,et al. Variability and regulation of the number of ovules,seeds and pods according to assimilate availability in winter oilseed rape(Brassica napus L.)[J]. Field Crop Res,2011,122(1):60-69.
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[19]胡青,高述民,李凤兰.植物雌性不育的研究进展[J].北京林业大学学报,2004,26(1):87-91.
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[21] Bouttier C,Morgan D G. Ovule development and determination of seed number per pod in oilseed rape(Brassica napus L.)[J]. J Exp Bot,1992,43(5):709-714.
[22]侯燕.杂交油菜制种提高母本结实率的研究Ⅰ不同亲本结实率的差异及其对制种产量的影响[J].种子,2003,22(4):103-104.
[23]陈锋,张洁夫,张维,等.甘蓝型油菜亲本遗传差异与杂种优势表现[J].江西农业学报,2016,28(2):35-38.
[24]赵庆华,黄剑华,颜昌敬.油菜花粉发芽的研究[J].作物学报,1986,12(1):15-20.
[25]李训贞.杂交油菜花粉活力的初步研究[J].作物研究,1989,3(3):20-22.
[26]蒋梁材,刘启鑫.油菜温敏不育系与可育系的花粉活力及育性变化比较[J].西南农业学报,1993,6(3):109-112.
[27]侯国佐.油菜花粉和柱头寿命的研究[J].贵州农业科学,1988,16(5):1-5.
[28]陈利娜,薛辉,牛娟,等.果树雌蕊败育研究进展[J].果树学报,2016,33(9):1156-1163.
[29] Tekleyohans D G,Nakel T,Groβ-Hardt R. Patterning the female gametophyte of flowering plants[J]. Plant Physiol,2017,173(1):122-129.
[30]李春宏,付三雄,陈新军,等.甘蓝型油菜雌性不育突变体FS-M1乳突细胞的细胞学观察[J].植物学报,2012,47(1):36-43.
[31]李春宏,付三雄,戚存扣.甘蓝型油菜授粉功能缺失突变体及其野生型柱头差异蛋白研究[J].中国农业科学,2012,45(22):4728-4737.
[32]韩玉娜,梁冬,申玉贤,等. 3个苜蓿材料花粉活力和柱头可授性比较[J].中国草地学报,2016,38(5):8-12.
[33]姚志敏,刘艳华,戴培刚,等.野生烟草花粉活力与柱头可授性及繁育特性研究[J].西北植物学报,2015,35(3):614-621.
[34]常海龙,张垂明,张伟,等.甘蔗花粉活力和柱头可授性日变化研究[J].广东农业科学,2017,44(4):14-18.
[35]孟佳佳,董树亭,石德杨,等.玉米雌穗分化与籽粒发育及败育的关系[J].作物学报,2013,39(5):912-918.
[36]孙倩倩,武剑,程锋,等.白菜型油菜自交亲和性、角果及种子相关性状的QTL定位[J].中国农业科学,2016,49(13):2449-2458.
[37]胡适宜.植物的受精作用第四讲受精的障碍:不亲和性[J].植物学通报,1984,19(Z1):93-99.
[38]刘后利,付廷栋,杨小牛,等.自交不亲和性的遗传及其在甘兰型油菜杂种优势上的利用[J].华中农学院学报,1981(1):1-12.
[39]安彩泰,陈军.甘蓝型油菜自交不亲和系柱头和花粉粒不亲和互作的扫描电镜研究[J].甘肃农业大学学报,1992,27(2):159-162,180.
[40]武军艳,孙万仓.利用花粉-柱头的相互作用鉴定5个十字花科油用植物的自交亲和性[J].西北农业学报,2006,15(5):61-64.
[41]赵志刚.人工合成的甘蓝型油菜自交亲和性分析[J].江苏农业科学,2013,41(8):95-98.
[42]李登科,薛晓芳,王永康,等.枣胚胎发育及胚败育动态观察[J].西北农业学报,2016,25(9):1379-1385.
[43]贺佳玉,李云,姜金仲,等.植物胚败育机理及其离体培养挽救技术之研究进展[J].中国农学通报,2008,24(1):141-146.
[44]潘学军,李顺雨,张文娥,等.红宝石无核葡萄胚发育及败育的观察[J].种子,2011,30(11):66-67.
[45] Khoshandam L,Baneh H D,Marandi R J,et al. Effect of BA and ovule developmental stages on embryo rescue in perlette grape(Vitis vinifera L.)cultivar[J]. Europ Online J Nat Soc Sci,2017,6(1):1-9.
[2]王健胜,梁亚红,侯桂玲,等.油菜育种亲本主要性状相关性及主成分分析[J].中国农学通报,2015,31(27):147-152.
[3] Zhang L W,Yang G S,Liu P W,et al. Genetic and correlation analysis of silique-traits in Brassica napus L. by quantitative trait locus mapping[J]. Theor Appl Genet,2011,122(1):21-31.
[4]黄华磊,周燕,李艳华,等.长江上游区十七个油菜品种产量性状的相关性分析[J].南方农业,2015,9(34):46-47,50.
[5]关周博,董育红,李少钦,等.甘蓝型油菜产量性状的遗传分析[J].作物研究,2016,30(3):266-270,315.
[6]陆时万,徐祥生,沈敏健,等.植物学(第二版)[M].北京:高等教育出版社,1991.
[7] Bouttier C,Morgan D G. Ovule development and determination of seed number per pod in oilseed rape(Brassica napus L.)[J]. J Exp Bot,1992,43(5):709-714.
[8] Yang Y H,Wang Y,Zhan J P,et al. Genetic and cytological analyses of the natural variation of seed number per pod in rapeseed(Brassica napus L.)[J]. Front Plant Sci,2017,8:1890.
[9]刘瑞娜,杨太明,陈鹏,等.安徽省油菜花期连阴雨灾害损失评估指标[J].中国农业气象,2016,37(4):471-478.
[10]许汪洁,傅鹰,董红利,等.一个甘蓝型油菜胚珠败育突变体的形态鉴定及生理特性分析[J].中国农业科学,2014,47(10):2062-2068.
[11]王风敏,薛艳,习光清,等.汉中市气候条件下直播油菜单株产量与重要农艺性状相关性分析[J].现代农业科技,2016(10):14,19.
[12]袁豆豆.蜜蜂授粉对新疆油菜增产效果影响[J].新疆农业科技,2016(4):47.
[13]侯燕,侯剑,韩宏仕,等.杂交油菜制种中提高母本结实率的研究Ⅵ.蜜蜂放养对制种结实率和产量的影响[J].贵州农业科学,2008,36(1):22,26.
[14]田祖庆.油菜不结实的原因分析及应对策略[J].作物研究,2015,29(6):665-666.
[15]周斌,胡德厚,胡爱民,等.峡江县油菜结实率低的原因调查与分析[J].安徽农学通报,2009,15(16):123,142.
[16] Wang X J,Mathieu A,Cournède P H,et al. Stochastic models in floral biology and its application to the study of oilseed rape(Brassica napus L.)fertility[C]//Third International Symposium on Plant Growth Modeling,Simulation, Visualization and Applications, Beijing,China,2009. New York,USA:IEEE,2009.
[17] Wang X J,Mathieu A,Cournède P H,et al. Variability and regulation of the number of ovules,seeds and pods according to assimilate availability in winter oilseed rape(Brassica napus L.)[J]. Field Crop Res,2011,122(1):60-69.
[18] Charlesworth D. Why do plants produce so many more ovules than seeds?[J]. Nature,1989,338(6210):21-22.
[19]胡青,高述民,李凤兰.植物雌性不育的研究进展[J].北京林业大学学报,2004,26(1):87-91.
[20]李永鹏,程焱,蔡光勤,等.油菜每角果粒数差异的细胞学基础和分子机理[J].中国科学:生命科学,2014,44(8):822-831.
[21] Bouttier C,Morgan D G. Ovule development and determination of seed number per pod in oilseed rape(Brassica napus L.)[J]. J Exp Bot,1992,43(5):709-714.
[22]侯燕.杂交油菜制种提高母本结实率的研究Ⅰ不同亲本结实率的差异及其对制种产量的影响[J].种子,2003,22(4):103-104.
[23]陈锋,张洁夫,张维,等.甘蓝型油菜亲本遗传差异与杂种优势表现[J].江西农业学报,2016,28(2):35-38.
[24]赵庆华,黄剑华,颜昌敬.油菜花粉发芽的研究[J].作物学报,1986,12(1):15-20.
[25]李训贞.杂交油菜花粉活力的初步研究[J].作物研究,1989,3(3):20-22.
[26]蒋梁材,刘启鑫.油菜温敏不育系与可育系的花粉活力及育性变化比较[J].西南农业学报,1993,6(3):109-112.
[27]侯国佐.油菜花粉和柱头寿命的研究[J].贵州农业科学,1988,16(5):1-5.
[28]陈利娜,薛辉,牛娟,等.果树雌蕊败育研究进展[J].果树学报,2016,33(9):1156-1163.
[29] Tekleyohans D G,Nakel T,Groβ-Hardt R. Patterning the female gametophyte of flowering plants[J]. Plant Physiol,2017,173(1):122-129.
[30]李春宏,付三雄,陈新军,等.甘蓝型油菜雌性不育突变体FS-M1乳突细胞的细胞学观察[J].植物学报,2012,47(1):36-43.
[31]李春宏,付三雄,戚存扣.甘蓝型油菜授粉功能缺失突变体及其野生型柱头差异蛋白研究[J].中国农业科学,2012,45(22):4728-4737.
[32]韩玉娜,梁冬,申玉贤,等. 3个苜蓿材料花粉活力和柱头可授性比较[J].中国草地学报,2016,38(5):8-12.
[33]姚志敏,刘艳华,戴培刚,等.野生烟草花粉活力与柱头可授性及繁育特性研究[J].西北植物学报,2015,35(3):614-621.
[34]常海龙,张垂明,张伟,等.甘蔗花粉活力和柱头可授性日变化研究[J].广东农业科学,2017,44(4):14-18.
[35]孟佳佳,董树亭,石德杨,等.玉米雌穗分化与籽粒发育及败育的关系[J].作物学报,2013,39(5):912-918.
[36]孙倩倩,武剑,程锋,等.白菜型油菜自交亲和性、角果及种子相关性状的QTL定位[J].中国农业科学,2016,49(13):2449-2458.
[37]胡适宜.植物的受精作用第四讲受精的障碍:不亲和性[J].植物学通报,1984,19(Z1):93-99.
[38]刘后利,付廷栋,杨小牛,等.自交不亲和性的遗传及其在甘兰型油菜杂种优势上的利用[J].华中农学院学报,1981(1):1-12.
[39]安彩泰,陈军.甘蓝型油菜自交不亲和系柱头和花粉粒不亲和互作的扫描电镜研究[J].甘肃农业大学学报,1992,27(2):159-162,180.
[40]武军艳,孙万仓.利用花粉-柱头的相互作用鉴定5个十字花科油用植物的自交亲和性[J].西北农业学报,2006,15(5):61-64.
[41]赵志刚.人工合成的甘蓝型油菜自交亲和性分析[J].江苏农业科学,2013,41(8):95-98.
[42]李登科,薛晓芳,王永康,等.枣胚胎发育及胚败育动态观察[J].西北农业学报,2016,25(9):1379-1385.
[43]贺佳玉,李云,姜金仲,等.植物胚败育机理及其离体培养挽救技术之研究进展[J].中国农学通报,2008,24(1):141-146.
[44]潘学军,李顺雨,张文娥,等.红宝石无核葡萄胚发育及败育的观察[J].种子,2011,30(11):66-67.
[45] Khoshandam L,Baneh H D,Marandi R J,et al. Effect of BA and ovule developmental stages on embryo rescue in perlette grape(Vitis vinifera L.)cultivar[J]. Europ Online J Nat Soc Sci,2017,6(1):1-9.