苎麻多胚诱导及单倍体筛选研究
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摘要
植物多胚是一种特殊的生殖现象,其中不仅包含了丰富的遗传信息,还往往能从多胚苗中发现单倍体。目前已有不少多胚植物中的单倍体已经广泛应用于育种工作,成为加快育种进程的珍贵材料。苎麻是雌雄同株的杂合体,其多胚性已经得到证实,然而品种间多胚率差异极大。本试验从促进苎麻多胚种萌发和诱导多胚品系两方面入手,以获得苎麻多胚品系和单倍体植株为目标,同时初步探索苎麻多胚苗在形态学、遗传学、染色体倍性和分子生物学方面的特点。主要方法和结果如下：
(1)多胚苎麻种子萌发的最佳条件是：28～30℃的常规条件下,以湿润滤纸为固体基质。喷洒0.8%的H202可提高苎麻多胚种子萌发率。从国家种质苎麻圃筛选出沅江肉麻等15个多胚率高于5%o的地方栽培品种,通过杂交和自交获得了8份多胚率高于5%o的材料。
(2)本研究对多胚籽苗从形态上进行分类,根据有多胚苗的大小、结构完整性以及有无结合点将苎麻多胚苗划分为五类：双同型、大小型、大残型、双残型和连体型,多胚苗的类型与其遗传背景有关。
(3)构建了靖西青麻×中苎1号F1代多胚群体——09JZ,采用SRAP分子标记对21对双胚苗、1束三胚苗、1株正常单苗及其亲本共48份材料进行分析,24对引物共扩增出159条多态性条带,每对引物平均扩增出6.6个多态性条带,多态性比率平均为63%。48份材料中成对遗传相似系数为0.36～0.88,在遗传相似系数0.51处可将聚类图中全部材料划分为四类群。多胚苗与亲本没有出现扩增谱带完全一致的情况,排除了该群体多胚来源于无融合生殖的可能。
(4)以中苎2号为母本,中苎1号为父本,经延迟授粉获得1对存活双胚苗,以SSR分子标记对父母本和双胚苗进行分析,结果显示双胚苗为亲本杂交后代。
(5)对大量杂交、自交以及正反交子代的多胚苗频率分析表明,受试材料F1代在多胚性上与母本表型相似。
(6)本研究采用染色体计数和流式细胞法对多胚苎麻材料的倍性进行了检测,通过对胚根分生区细胞显微观察发现2个含有n-2n型多胚苗的品系,通过重点培育这两个品系多胚苗,获得1株单倍体植株,与二倍体植株在大小、叶色、叶龄和长势等方面存在巨大差异。
本研究首次将苎麻正反交子代的多胚性与亲本相比较,发现F,代的多胚性与母本一致,这对于多胚苎麻的选育工作具有指导意义,单倍体植株的检出说明从双胚苗中获得单倍体材料可行,这对加快苎麻育种进程有重要意义。
As a peculiar reproduction phenomenon in plants, Polyembryony not only contains abundant genetic information, but also contains spontaneous haploids. At present, quite a few haploids from polyembryonic seedlings have been widely applied to crops breeding researches, becoming a precious resource in promoting breeding process and constructing genetic map. Ramie is a kind of monoecism heterozygote crop, whose property of polyembryony has been confirmed. However, the variation of polyembryonic ratio was significant among breeds. This study aimed to obtain polyembryonic varieties and haplobiont. Two ways were adopted in this study:facilitating the germination of polyembryonic seeds and inducing the formation of polyembryony. Meanwhile, this article tentatively explored the characteristics of polyembryonic ramie seedlings from prospectives of morphology, hereditism, ploidy and molecular biology. Main approaches and results are introduced and explored as follows:
(1) The optimum germination condition of polyembryonic ramie seeds was defined as: environment temperature controlled to28～30℃, and moist filter paper used as solid substrate. Germination ratio of polyembryonic seeds could be increased by spraying0.8%H2O2liquor. In this study,15cultivars were screened out for their polyembryony percentage above5%o. Meanwhile, by hybridizing and self-crossing samples,8varieties with polyembryony percentages above5%o were cultivated.
(2) According to their sizes, structural integrity and whether they are conjoined or not, polyembryonic ramie seedlings can be categorized into5types:semlable, big-small, big-abnormal, both abnormal and siamesed. Further study demonstrated that the categories of polyembryonic ramie seedlings related to its genetic background.
(3)09JZ, a group of ramie seedlings, has been cultivated. This group was consisted with21pairs of twin ramie seedlings, a bunch of tri-embryony seedlings and one normal seedling which all screened out from the F1progenies of Jingxi qing x Zhongzhu No.1. SRAP (Sequence Related Amplified Polymorphism) markers were applied in analyzing the genomes of this group and their parents. The results showed that a total of159polymorphic bands were detected with an average of6.6polymorphic bands per primer pair, whose percentage of polymorphic bands were63%. The genetic similarities of all48materials ranged from0.360to0.876. UPGMA method cluster analysis showed that these materials classified into four cluster groups with genetic similarity of0.51. No individual of09JZ has fully consistent polymorphic bands with their parents, so the possibility of individual apomixes excluded.
(4) One pair of twin ramie seedlings were obtained by delayedpollination between Zhongzhu No.2and Zhongzhu No.1.The twin seedlings and their parents were detected by SSR markers. The results showed that the twin seedlings were real filial generations.
(5) By analyzing the polyembryonic seedlings ratio of filial generation of hybridization and self-crossing, we concluded that polyembryony percentage of filial generation were as same as that of female parents.
(6) Chromosome counting and FCM (Flow Cytometry) were both applied to detecting the ploidy of polyembryonic seedlings.2varieties which contained n-2n type were screened out. What's more,1haplobiont of Yuanjiang Rou was obtained, which was widely different from diploid plants in size, leaf age and growth vigour fields.
This study innovatively reveals the fact that polyembryoney property of filial generation is the same as that of the female, which has guiding significance in ramie's polyembryonic breeding. The feasibility of obtaining haplobiont from twin seedlings can greatly promote the development of ramie breeding research.
(1)多胚苎麻种子萌发的最佳条件是：28～30℃的常规条件下,以湿润滤纸为固体基质。喷洒0.8%的H202可提高苎麻多胚种子萌发率。从国家种质苎麻圃筛选出沅江肉麻等15个多胚率高于5%o的地方栽培品种,通过杂交和自交获得了8份多胚率高于5%o的材料。
(2)本研究对多胚籽苗从形态上进行分类,根据有多胚苗的大小、结构完整性以及有无结合点将苎麻多胚苗划分为五类：双同型、大小型、大残型、双残型和连体型,多胚苗的类型与其遗传背景有关。
(3)构建了靖西青麻×中苎1号F1代多胚群体——09JZ,采用SRAP分子标记对21对双胚苗、1束三胚苗、1株正常单苗及其亲本共48份材料进行分析,24对引物共扩增出159条多态性条带,每对引物平均扩增出6.6个多态性条带,多态性比率平均为63%。48份材料中成对遗传相似系数为0.36～0.88,在遗传相似系数0.51处可将聚类图中全部材料划分为四类群。多胚苗与亲本没有出现扩增谱带完全一致的情况,排除了该群体多胚来源于无融合生殖的可能。
(4)以中苎2号为母本,中苎1号为父本,经延迟授粉获得1对存活双胚苗,以SSR分子标记对父母本和双胚苗进行分析,结果显示双胚苗为亲本杂交后代。
(5)对大量杂交、自交以及正反交子代的多胚苗频率分析表明,受试材料F1代在多胚性上与母本表型相似。
(6)本研究采用染色体计数和流式细胞法对多胚苎麻材料的倍性进行了检测,通过对胚根分生区细胞显微观察发现2个含有n-2n型多胚苗的品系,通过重点培育这两个品系多胚苗,获得1株单倍体植株,与二倍体植株在大小、叶色、叶龄和长势等方面存在巨大差异。
本研究首次将苎麻正反交子代的多胚性与亲本相比较,发现F,代的多胚性与母本一致,这对于多胚苎麻的选育工作具有指导意义,单倍体植株的检出说明从双胚苗中获得单倍体材料可行,这对加快苎麻育种进程有重要意义。
As a peculiar reproduction phenomenon in plants, Polyembryony not only contains abundant genetic information, but also contains spontaneous haploids. At present, quite a few haploids from polyembryonic seedlings have been widely applied to crops breeding researches, becoming a precious resource in promoting breeding process and constructing genetic map. Ramie is a kind of monoecism heterozygote crop, whose property of polyembryony has been confirmed. However, the variation of polyembryonic ratio was significant among breeds. This study aimed to obtain polyembryonic varieties and haplobiont. Two ways were adopted in this study:facilitating the germination of polyembryonic seeds and inducing the formation of polyembryony. Meanwhile, this article tentatively explored the characteristics of polyembryonic ramie seedlings from prospectives of morphology, hereditism, ploidy and molecular biology. Main approaches and results are introduced and explored as follows:
(1) The optimum germination condition of polyembryonic ramie seeds was defined as: environment temperature controlled to28～30℃, and moist filter paper used as solid substrate. Germination ratio of polyembryonic seeds could be increased by spraying0.8%H2O2liquor. In this study,15cultivars were screened out for their polyembryony percentage above5%o. Meanwhile, by hybridizing and self-crossing samples,8varieties with polyembryony percentages above5%o were cultivated.
(2) According to their sizes, structural integrity and whether they are conjoined or not, polyembryonic ramie seedlings can be categorized into5types:semlable, big-small, big-abnormal, both abnormal and siamesed. Further study demonstrated that the categories of polyembryonic ramie seedlings related to its genetic background.
(3)09JZ, a group of ramie seedlings, has been cultivated. This group was consisted with21pairs of twin ramie seedlings, a bunch of tri-embryony seedlings and one normal seedling which all screened out from the F1progenies of Jingxi qing x Zhongzhu No.1. SRAP (Sequence Related Amplified Polymorphism) markers were applied in analyzing the genomes of this group and their parents. The results showed that a total of159polymorphic bands were detected with an average of6.6polymorphic bands per primer pair, whose percentage of polymorphic bands were63%. The genetic similarities of all48materials ranged from0.360to0.876. UPGMA method cluster analysis showed that these materials classified into four cluster groups with genetic similarity of0.51. No individual of09JZ has fully consistent polymorphic bands with their parents, so the possibility of individual apomixes excluded.
(4) One pair of twin ramie seedlings were obtained by delayedpollination between Zhongzhu No.2and Zhongzhu No.1.The twin seedlings and their parents were detected by SSR markers. The results showed that the twin seedlings were real filial generations.
(5) By analyzing the polyembryonic seedlings ratio of filial generation of hybridization and self-crossing, we concluded that polyembryony percentage of filial generation were as same as that of female parents.
(6) Chromosome counting and FCM (Flow Cytometry) were both applied to detecting the ploidy of polyembryonic seedlings.2varieties which contained n-2n type were screened out. What's more,1haplobiont of Yuanjiang Rou was obtained, which was widely different from diploid plants in size, leaf age and growth vigour fields.
This study innovatively reveals the fact that polyembryoney property of filial generation is the same as that of the female, which has guiding significance in ramie's polyembryonic breeding. The feasibility of obtaining haplobiont from twin seedlings can greatly promote the development of ramie breeding research.
引文
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