玉米×Teosinte远缘杂种后代重要性状的表型及遗传分析
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摘要
玉米由于长期自然和人工选择造成遗传多样性相对于野生近缘种明显降低。现代玉米育种主要是杂种优势利用,多集中在少数骨干自交系的遗传改良,造成组配的杂交种品种遗传基础狭窄。对于我国来说玉米属于外引作物,种植历史仅500余年,可利用的玉米育种材料相对贫乏,很难育成突破性品种。没有种质的不断创新就没有玉米杂种优势水平的提高。因此,种质的创制与拓宽是玉米育种的战略任务,其中开发和利用野生资源是其重要的途径之一。
大色草是玉米近缘野生植物,为一年生草本。植株生长旺盛,具有高分粟性和分枝性,籽粒蛋白质含量高,对多种玉米病害免疫或高抗,是玉米育种可利用的基因源。同时大色草又是典型的短日照植物,在温带不能正常繁殖。本研究以大自草,玉米自交系掖478、综31、3153,以及包括3153x大当草选系在内的76个RILs为材料,进行大色草光周期、蛋白质含量杂种优势、远缘杂交创新玉米种质、大色草分粟性和RIL形态和产量性状的SSR标记关联分析,探索大当草短日照诱导开花的敏感苗龄、最佳的日照时数,通过杂交导入有利性状进行玉米改良与种质资源创新。取得如下结果:
1.19一26d是大色草能否接受诱导的临界苗龄,苗龄(sd一47d)越大诱导效果越好;gh是诱导生殖生长的最佳日照时数;各日照处理的POD活性随苗龄的增大呈现一定的规律性变化,但不能直接反映大当草生长发育状况;a、b、。、d四条POD酶带是大当草处于营养生长阶段的生化标记,e、f、g三条POD酶带的出现表明进入生殖生长阶段,POD酶的相对含量主导着大色草的生长发育方向;采用40一47d苗龄每日gh短日照处理的大色草与掖478、综31花期相遇,通过远缘杂交育成NX3153等自交系。
2.远缘杂种BC:产量性状表现最大值,而且变异系数最小,F:除穗长外,穗粗、穗行数、行粒数都大于Fl,并存在广泛的遗传变异,而穗粗、穗行数、行粒数都比Fl变化幅度大。Fl穗重、穗粒重表现极大的变异系数,F:变异系数大幅度降低,表现最稳定的是穗粗和穗行数。F:抽雄期、开花期、吐丝期、株高、穗位高比Fl、BCI变异幅度大、分离严重,BC:变异系数逐渐变小、分离减缓。Fl、FZ、BCI、BC:花期不遇个体顺序减少。Fl、F:的AIS较长,自身授粉结实困难,BCZ最短,自身可以正常授粉。不同类群玉米与有分粟的F:杂交后代分粟产生的效果不同,属于瑞德类群的65232抑制分粟效果最好。
3.本研究对76个玉米重组自交系的形态、遗传多态性进行研究,并对17个性状与48个SSR位点进行了关联分析。76个重组自交系分为3个主类群5个亚群,通过K检验、回归分析和组合分析分别检测到82、59和40个明显的关联。17个性状单独分析时,Q一SSR与穗部性状之间的关联达27.8%,显著高于其它组合2.3%一6.3%的关联。含有大自草的RILs的变异范围较广,在三个主类群以及五个亚群中都有分布,但分布很不均匀,第一主群的第一亚群中最少,短锥形和短筒形类群中最多。
4.17个性状之间的相关程度存在较大差别。玉米茎秆弯曲、叶宽、玉米雄穗一级分支数和穗上长度与株高的比率4个形态学性状之间、以及与其它31个性状之间不存在显著相关外,其它13个性状彼此都存在相关性。穗位高与穗粗,穗粗与粒行数,粒行数与叶片夹角,穗粗与百粒,重、穗形、开花吐丝间隔,穗形与百粒重、开花吐丝间隔,穗长与粒型都达到极显著正相关;粒行数与行粒数,行粒数与开花吐丝间隔、穗形、穗粗、百粒重,百粒重与粒型、穗长,穗形与穗长,穗粗与粒型都存在极显著负相关。
5.71个性状还存在着星状辐射、三角、四角以及线性等特殊关联。①穗粗与其它9个性状存在关联,表现明显的星状辐射关系,其中5个极显著正相关关系、3个极显著负相关关系、1个显著正相关关系,是所有研究的性状中关联最多、极显著正相关关系最多、极显著正相关关系与极显著负相关关系对立最明显的性状,同时还存在1个间接极显著正相关关系的性状。穗粗除与其他产量因素间存在复杂的关联外,还与穗位、叶片夹角等形态性状的存在关联。其次是穗形,存在7个关联,百粒重存在6个关联;②百粒重、穗形、开花吐丝间隔三者之间以及穗粗、百粒重、穗形三者之间存在极显著正相关三角关联;③穗粗、百粒重、穗形、开花吐丝间隔四者之间的极显著正相关四角关系;④EH一ED一EL反向极显著相关线性关系,表明穗位越高果穗越粗,穗长变短,EH一LA一AB反向显著相关线性关系。
The genetic diversity of maize was significantly reduced compared with wild relative species,due to the effects of long-term natural selection and artificial selection.Maize genetics and breeding is mainly based on the utilization of heterosis and concentrated on the genetic improvement of a small number of main inbred lines,which resulted in the narrow genetic base of the hybrid varieties.Maize originated in the Americas and it has been planted in China only for 500 years,consequently,maize breeding germplasn is relatively scarce,it is difficult to breed breakthrough high-yielding single cross hybrid.Germplasm innovation is needed in raising the level of heterosis.Therefore,development of maize new germplasm is the primary strategic task in maize breeding, in which the development and exploit of wild resources is one of the most important ways.
Teosinte. is one of the available gene sources in maize breeding.lt is an annual herb,one of the maize wild relative species,with the characteristics of high tillering and branching, high protein content in grain, immune or high resistant on a variety of maize diseases. Meanwhile,Teosinte is a typical short-day plant and can not reproduce normally in the temperate regions. In order to study the short-day sensitive seedling age , the optimal day-length for reproductive induction of Teosinte,make it possible to transfer novel genes into elite inbred lines to create new maize germplasms by inducing the flowering time of Teosinte. synchronized with elite maize cultivars,we investigated the photoperiod and proteins content heterosis of Teosinte., studied the Teosinte,'s tillering,maize germplasm innovation through distant hybridization.Also we did the association analysis of the RIL agronomic traits,yield traits and SSR markers,with Teosinte.,maize inbred line Y478,Zong31,3153,and 76 RILs comprise of 6 selected inbred lines from 3153xTeosinte. as the materials.
The main results are as follows:
1. The results show that the sensitive productive induction stage in Teosinte. is between 19d and 26d in seedling age. The induction rate becomes higher with the Teosinte seedlings become older (from 5d to 47d). The optimal day-length for reproductive induction is 9 hours POD activity shows certain correlation with the seedling age, but has no implication on the growth conditions. Four types of POD isoenzymes, i.e. a, b, c and d, are good characteristics for vegetative growth, and the POD isoenzymes of e, f, g feature reproductive growth. Furthermore, the composition of these POD isoenzymes determines whether Zea. luxurians. enters into reproductive growth. Through 9 hours's short-day treatment in the seedling age from 40 to 47d, the flowering time of Teosinte. meet well with Ye 478 and Zong 31,therefore,several inbred lines ,such as ,NX3153 have been bred successfully.
2. The investigation of tiller, tasseling stage, ASI, plant height and ear height from distant hybridization progeny F_i displayed that ,there is no difference in the tillers,plant traits and ear characteristics of reciprocal crosses,but the tiller restraint of different hybridization ways is distinct in progenies, which suggested that evolution level of different maize groups is different.Through transfering beneficial traits to the general maize from Teosinte., a number of excellent maize introgression lines were obtained,which provide evidence for the identification of Teosinte elite gene and molecular markers location.
3. In this study,we characterized morphological and genetic diversity for 76 recombinant inbred lines ,and analysed the association of 17 morphological characters or values and 48 SSR markers. The 76 recombinant inbred lines were divided into three major groups and five sub-groups. 82,59 and 40 significant correlation were revealed through the only K-test, only logistic regression and both analysis,respectively. The association between the Q-SSR and the ear characters was 27.8%, significantly higher than the other combiantion( 2.3% -6.3%) when 17 traits were analysed independently.The RILs with teosinte had wide variation range, and were distributed in the three major groups and five subgroups, but the distribution is very uneven, the distribution in the first sub-group of the first main group was least and conversely in the short conical and cylindrical groups.
4. There is a big difference in the correlation of 17 traits. The four morphological traits, corn stalk bending, leaf width, maize tassel branch number and the ratio of length above ear to the plant height,displayed no significant correlation,likewise, with the other 13 traits.But other 13 traits were correlated with each other. Ear height and ear diameter, ear diameter and kernel row number, kernel row number and leaf angle, ear diameter and hundred kernel weight, ear shape, anthesis silking internal have reached a very significant positive correlation,likewise,ear shape and hundred kernel weight, anthesis silking intervaEar Length and kernel type were also significant positive correlation. Kernel row number and kernel number per row,kernel number per row and anthesis silking interval, ear shape, ear diameter, hundred kernel weight displayed a very significant negative correlation. Hundred kernel weight and kernel type,ear length were negatively related.Ear shape and ear length,ear diameter and kernel type also reached very significantly negative correlation.
5. Star emission,triangle, square and linear and other special association were also existed in 17 traits.The results are as below,①ear diameter showed an obvious star emissions association with the other 9 traits, 5 of which were a significant positive correlation,3 of which dispalyed a highly significant negative correlation,I of which showed a highly significant positive correlation,and was a trait that had most correlation,most highly significant positive correlation,the contrast of highly significant positive correlation and highly significant negative correlation of which was most obvious.Meanwhile,there was a trait that had a indirect highly significant positive correlation with ear diameter.Ear diameter was correlated with ear height and leaf angle besides complex correlation with other yield factor.②Hundred kernel weight,ear shape, anthesis silking internal showed a highly significant positive triangle correlation each other,the same as ear diameter, hundred kernel weight,ear shape.③Ear diameter, hundred kernel weight,ear shape, anthesis silking internal had a highly significant positive square correlation each other.(EH-ED-EL showed a reversely highly significant linear correlation,which indicated the higher the ear height,the thicker the ear. EH—LA—AB also had a reversely significantly linear correlation.
大色草是玉米近缘野生植物,为一年生草本。植株生长旺盛,具有高分粟性和分枝性,籽粒蛋白质含量高,对多种玉米病害免疫或高抗,是玉米育种可利用的基因源。同时大色草又是典型的短日照植物,在温带不能正常繁殖。本研究以大自草,玉米自交系掖478、综31、3153,以及包括3153x大当草选系在内的76个RILs为材料,进行大色草光周期、蛋白质含量杂种优势、远缘杂交创新玉米种质、大色草分粟性和RIL形态和产量性状的SSR标记关联分析,探索大当草短日照诱导开花的敏感苗龄、最佳的日照时数,通过杂交导入有利性状进行玉米改良与种质资源创新。取得如下结果:
1.19一26d是大色草能否接受诱导的临界苗龄,苗龄(sd一47d)越大诱导效果越好;gh是诱导生殖生长的最佳日照时数;各日照处理的POD活性随苗龄的增大呈现一定的规律性变化,但不能直接反映大当草生长发育状况;a、b、。、d四条POD酶带是大当草处于营养生长阶段的生化标记,e、f、g三条POD酶带的出现表明进入生殖生长阶段,POD酶的相对含量主导着大色草的生长发育方向;采用40一47d苗龄每日gh短日照处理的大色草与掖478、综31花期相遇,通过远缘杂交育成NX3153等自交系。
2.远缘杂种BC:产量性状表现最大值,而且变异系数最小,F:除穗长外,穗粗、穗行数、行粒数都大于Fl,并存在广泛的遗传变异,而穗粗、穗行数、行粒数都比Fl变化幅度大。Fl穗重、穗粒重表现极大的变异系数,F:变异系数大幅度降低,表现最稳定的是穗粗和穗行数。F:抽雄期、开花期、吐丝期、株高、穗位高比Fl、BCI变异幅度大、分离严重,BC:变异系数逐渐变小、分离减缓。Fl、FZ、BCI、BC:花期不遇个体顺序减少。Fl、F:的AIS较长,自身授粉结实困难,BCZ最短,自身可以正常授粉。不同类群玉米与有分粟的F:杂交后代分粟产生的效果不同,属于瑞德类群的65232抑制分粟效果最好。
3.本研究对76个玉米重组自交系的形态、遗传多态性进行研究,并对17个性状与48个SSR位点进行了关联分析。76个重组自交系分为3个主类群5个亚群,通过K检验、回归分析和组合分析分别检测到82、59和40个明显的关联。17个性状单独分析时,Q一SSR与穗部性状之间的关联达27.8%,显著高于其它组合2.3%一6.3%的关联。含有大自草的RILs的变异范围较广,在三个主类群以及五个亚群中都有分布,但分布很不均匀,第一主群的第一亚群中最少,短锥形和短筒形类群中最多。
4.17个性状之间的相关程度存在较大差别。玉米茎秆弯曲、叶宽、玉米雄穗一级分支数和穗上长度与株高的比率4个形态学性状之间、以及与其它31个性状之间不存在显著相关外,其它13个性状彼此都存在相关性。穗位高与穗粗,穗粗与粒行数,粒行数与叶片夹角,穗粗与百粒,重、穗形、开花吐丝间隔,穗形与百粒重、开花吐丝间隔,穗长与粒型都达到极显著正相关;粒行数与行粒数,行粒数与开花吐丝间隔、穗形、穗粗、百粒重,百粒重与粒型、穗长,穗形与穗长,穗粗与粒型都存在极显著负相关。
5.71个性状还存在着星状辐射、三角、四角以及线性等特殊关联。①穗粗与其它9个性状存在关联,表现明显的星状辐射关系,其中5个极显著正相关关系、3个极显著负相关关系、1个显著正相关关系,是所有研究的性状中关联最多、极显著正相关关系最多、极显著正相关关系与极显著负相关关系对立最明显的性状,同时还存在1个间接极显著正相关关系的性状。穗粗除与其他产量因素间存在复杂的关联外,还与穗位、叶片夹角等形态性状的存在关联。其次是穗形,存在7个关联,百粒重存在6个关联;②百粒重、穗形、开花吐丝间隔三者之间以及穗粗、百粒重、穗形三者之间存在极显著正相关三角关联;③穗粗、百粒重、穗形、开花吐丝间隔四者之间的极显著正相关四角关系;④EH一ED一EL反向极显著相关线性关系,表明穗位越高果穗越粗,穗长变短,EH一LA一AB反向显著相关线性关系。
The genetic diversity of maize was significantly reduced compared with wild relative species,due to the effects of long-term natural selection and artificial selection.Maize genetics and breeding is mainly based on the utilization of heterosis and concentrated on the genetic improvement of a small number of main inbred lines,which resulted in the narrow genetic base of the hybrid varieties.Maize originated in the Americas and it has been planted in China only for 500 years,consequently,maize breeding germplasn is relatively scarce,it is difficult to breed breakthrough high-yielding single cross hybrid.Germplasm innovation is needed in raising the level of heterosis.Therefore,development of maize new germplasm is the primary strategic task in maize breeding, in which the development and exploit of wild resources is one of the most important ways.
Teosinte. is one of the available gene sources in maize breeding.lt is an annual herb,one of the maize wild relative species,with the characteristics of high tillering and branching, high protein content in grain, immune or high resistant on a variety of maize diseases. Meanwhile,Teosinte is a typical short-day plant and can not reproduce normally in the temperate regions. In order to study the short-day sensitive seedling age , the optimal day-length for reproductive induction of Teosinte,make it possible to transfer novel genes into elite inbred lines to create new maize germplasms by inducing the flowering time of Teosinte. synchronized with elite maize cultivars,we investigated the photoperiod and proteins content heterosis of Teosinte., studied the Teosinte,'s tillering,maize germplasm innovation through distant hybridization.Also we did the association analysis of the RIL agronomic traits,yield traits and SSR markers,with Teosinte.,maize inbred line Y478,Zong31,3153,and 76 RILs comprise of 6 selected inbred lines from 3153xTeosinte. as the materials.
The main results are as follows:
1. The results show that the sensitive productive induction stage in Teosinte. is between 19d and 26d in seedling age. The induction rate becomes higher with the Teosinte seedlings become older (from 5d to 47d). The optimal day-length for reproductive induction is 9 hours POD activity shows certain correlation with the seedling age, but has no implication on the growth conditions. Four types of POD isoenzymes, i.e. a, b, c and d, are good characteristics for vegetative growth, and the POD isoenzymes of e, f, g feature reproductive growth. Furthermore, the composition of these POD isoenzymes determines whether Zea. luxurians. enters into reproductive growth. Through 9 hours's short-day treatment in the seedling age from 40 to 47d, the flowering time of Teosinte. meet well with Ye 478 and Zong 31,therefore,several inbred lines ,such as ,NX3153 have been bred successfully.
2. The investigation of tiller, tasseling stage, ASI, plant height and ear height from distant hybridization progeny F_i displayed that ,there is no difference in the tillers,plant traits and ear characteristics of reciprocal crosses,but the tiller restraint of different hybridization ways is distinct in progenies, which suggested that evolution level of different maize groups is different.Through transfering beneficial traits to the general maize from Teosinte., a number of excellent maize introgression lines were obtained,which provide evidence for the identification of Teosinte elite gene and molecular markers location.
3. In this study,we characterized morphological and genetic diversity for 76 recombinant inbred lines ,and analysed the association of 17 morphological characters or values and 48 SSR markers. The 76 recombinant inbred lines were divided into three major groups and five sub-groups. 82,59 and 40 significant correlation were revealed through the only K-test, only logistic regression and both analysis,respectively. The association between the Q-SSR and the ear characters was 27.8%, significantly higher than the other combiantion( 2.3% -6.3%) when 17 traits were analysed independently.The RILs with teosinte had wide variation range, and were distributed in the three major groups and five subgroups, but the distribution is very uneven, the distribution in the first sub-group of the first main group was least and conversely in the short conical and cylindrical groups.
4. There is a big difference in the correlation of 17 traits. The four morphological traits, corn stalk bending, leaf width, maize tassel branch number and the ratio of length above ear to the plant height,displayed no significant correlation,likewise, with the other 13 traits.But other 13 traits were correlated with each other. Ear height and ear diameter, ear diameter and kernel row number, kernel row number and leaf angle, ear diameter and hundred kernel weight, ear shape, anthesis silking internal have reached a very significant positive correlation,likewise,ear shape and hundred kernel weight, anthesis silking intervaEar Length and kernel type were also significant positive correlation. Kernel row number and kernel number per row,kernel number per row and anthesis silking interval, ear shape, ear diameter, hundred kernel weight displayed a very significant negative correlation. Hundred kernel weight and kernel type,ear length were negatively related.Ear shape and ear length,ear diameter and kernel type also reached very significantly negative correlation.
5. Star emission,triangle, square and linear and other special association were also existed in 17 traits.The results are as below,①ear diameter showed an obvious star emissions association with the other 9 traits, 5 of which were a significant positive correlation,3 of which dispalyed a highly significant negative correlation,I of which showed a highly significant positive correlation,and was a trait that had most correlation,most highly significant positive correlation,the contrast of highly significant positive correlation and highly significant negative correlation of which was most obvious.Meanwhile,there was a trait that had a indirect highly significant positive correlation with ear diameter.Ear diameter was correlated with ear height and leaf angle besides complex correlation with other yield factor.②Hundred kernel weight,ear shape, anthesis silking internal showed a highly significant positive triangle correlation each other,the same as ear diameter, hundred kernel weight,ear shape.③Ear diameter, hundred kernel weight,ear shape, anthesis silking internal had a highly significant positive square correlation each other.(EH-ED-EL showed a reversely highly significant linear correlation,which indicated the higher the ear height,the thicker the ear. EH—LA—AB also had a reversely significantly linear correlation.
引文
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