枇杷小种子植株多样性分析和优异种质筛选及种子退化机理研究
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摘要
四川农业大学枇杷品种选育课题组独创性地通过小种子(枇杷中度退化种子)无菌萌发再生创制了一批小种子植株,为开拓和创新枇杷种质资源提供了宝贵的材料。本研究第一部分内容从细胞和形态学等层次对小种子植株进行遗传多样性分析和育种价值评价,并通过各种优良性状和特异性状的比较,特别依照枇杷品种选育目标筛选具有较高研究和利用价值的优异种质；第二部分内容进行了花粉母细胞减数分裂、花粉和胚珠的细胞形态学观察,同时首次从雌蕊构造角度入手,探讨雌蕊结构和繁殖特征与种子退化的关系,对全面弄清枇杷退化种子的来源和了解枇杷生殖特性、胚胎发育及科学利用退化种子创新枇杷种质资源具有重要的理论和实践意义。主要研究结果如下：
1小种子植株叶遗传多样性分析和重要类群倍性鉴定
小种子植株叶遗传多样性指数D=0.785,各特征变异系数在9.84%～40.09%之间,具有丰富的多样性。UPGMA法将79份材料分为了6大类群,第Ⅰ类和第Ⅵ类表现最为特殊,明显区别于母本“大五星”,为枇杷特异种质选育的重要类群。其中Ⅰ类表型为叶片大型、叶柄粗长、叶色浓绿、质地厚实、坚硬,具有多倍体特征。利用本实验室优化的适于枇杷染色体计数根尖压片方法,对第Ⅰ类小种子植株进行染色体倍性鉴定,确定1号,115号,137号和249号为三倍体(2n=3x=51),为无籽枇杷育种提供了宝贵的材料。三倍体植株形态学上表现为叶大、花大、果大、钟形,果皮橙黄色,无籽。
2小种子植株花期、果实种子性状多样性与特异优良材料筛选
小种子植株花期多样性丰富,并结合育种和生产实际,筛选到212号早花型单株和6060号晚花型单株,为培育枇杷早熟晚熟品种提供了宝贵的材料；小种子植株在果实种子性状上多样性丰富,变异系数平均为27.92%,多样性指数平均为0.7482；并综合枇杷育种目标,初步筛选出了大果型单株203号,少核型单株C1号和白肉型单株4-1-5号,为培育大果、少核和优质白肉品种提供了宝贵的材料。
3枇杷小种子植株抗寒性调查及抗寒株系初步筛选
小种子植株平均冻害指数为25.34,对冻害的反应表现出多样性。种胚受冻率及二年生秋稍落叶率变幅分别达65.4%、90%。落叶率和种胚受冻率不具有相关性(R=0.2911)；种胚受冻率和花期早晚呈现出显著负相关(R2=0.9137),种胚受冻率随花期的延后逐渐降低；并以种胚受冻率为主要评价指标,结合落叶情况,初步筛选出抗寒株系6-3.10和C2,为抗寒基因的定位及抗寒育种提供了宝贵的材料。
4枇杷花粉母细胞减数分裂及花粉和胚珠细胞形态学观察
枇杷(早钟6号为材料)减数分裂Ⅱ期异常的占3.9%,表现为不分裂或不同步分裂,产生了异常二分体(0.3%),四分体(0.5%)和六分体(0.1%),未减数配子的频率为0.28%；花粉细胞有21.6%的形态结构异常,且均出现在单核期,包括空腔、无细胞核以及畸变小孢子等。枇杷(大五星为材料)胚珠石蜡切片结构观察发现,有13.3%存在异常现象,包括小胚囊、雌性生殖器退化、胚囊退化和胚珠退化等4类。减数分裂异常,花粉异常畸变可导致枇杷花粉败育,从而引起授粉受精不良,同时异常的胚珠结构不能提供受精及胚胎发育的场所,最终导致种子退化。
5枇杷柱头和花柱的形态分析及其与花粉萌发和花粉管生长的关系
枇杷(以“大五星”、“早钟6号”和“龙泉1号”为材料)每花内5个花柱长短差异显著。枇桤(以“大五星”为材料)每花内长、中和短型花柱的柱头乳突细胞、柱头可受性、分泌大量黏液的持续时间,以及花粉萌发数量和花粉管生长均存在着明显的差异,长的花柱比短的花柱更有利于接受花粉及随后的花粉萌发；并且不同长度的花柱其花粉原位萌发和花粉管生长情况也有较明显的差异,表现为短型花柱柱头花粉萌发量、中部和基部的花粉管数量均依次低于中型和长型花柱。由此推论枇杷柱头、花柱的形态繁殖特性与种子高退化率及不同退化程度间具有相关性。
6枇杷的雌蕊结构及其与种子生产和退化的关系
通过徒手切片和去柱头人工控制授粉等措施,首次查明了枇杷的5个柱头和5个心室在形态结构和繁殖特性上均是完美的一一对应关系。枇杷雌蕊结构为子房被隔膜分为5个心室,5个花柱离生,基部分离,并且每个花柱分别对应一个心室；通过去掉4个柱头,只留一个柱头进行人工授粉后,只在一个心室中有种子产生,其它4个心室的胚珠均未受精而萎蔫。其中有76.7%的果实符合模式Ⅰ即同一心室中同时产生两颗正常的种子,有23.3%的果实符合模式Ⅱ,即同一心室中有一颗正常的种子,另外一颗产生了不同程度的退化。在只有一个柱头时,种子的败育率为11.7%。根据以上结果首次讨论了枇杷雌蕊结构与种子败育之间的关系,推论同一心室中的败育种子有一定几率是由刺激产生的单倍体。对去4柱头授粉得到的42粒退化种子进行无菌培养和染色体倍性鉴定。结果得到一份单倍体材料(n=x=17),证实了推论。同时,还获得一份非整倍体材料(n=x-3=14),为非整倍体的获得提供了全新的途径和契机。
The loquat breeding group of Sichuan Agricultural University had developed a protocol for culture of mini-seeds (middle-degraded seeds) and production of plants called "mini-seed derived plant", providing the valuable material for germplasm resource development and innovation. In the present study, a series of experiments were carried out to investigate the genetic diversity of mini-seed derived plants, excellent germplasm selection and seed degeneration mechanism in loquat. In part I, the genetic diversity and breeding value of mini-seed plants were studied dased on cytology and morphology research, in order to selecte excellent germplasm. In part II, the seed degeneration mechanism was stuied by pollen mother cell meiosis and the unnormal phenomenon of pollen and ovule in morphocytology; meanwhile, the relationship between pistil structure and seed abortion was clarified for the first time in loquat, aiming at providing data for degenerated seed origin. The main results were as follows.
1Diversity of mini-seed plants based on leaf morphology and chromosome number of group I identified
There was abundant genetic diversity in leaf morphology (D=0.785). The values obtained varied broadly with coefficients of variation ranging from9.8%to40.1%. Analysis of the79accessions divided them into6groups by the UPGMA method. The leaves of group Ⅵ were light green, small, thin and soft with slender petioles; while the leaves of group I were dark green, large, hard and thick with long and thick petioles. These two groups were important excellent germplasm. And1#,115#,137#and249#in group I were triploids (2n=3x=51), which provided precious materials for breeding seedless varieties. The triploid plants had the typical morphological appearances with large leaves, big flowers, big and bell-shaped fruits and orange yellow fruit peel.
2Diversity in florescence, fruit and seed morphology of mini-seed derived plants and excellent germplasm selection
There was abundant genetic diversity in florescence in the mini-seed derived plants and2lines were selected with special traits, early flowering (212#) and late flowering (6060#). There was also abundant diversity in fruit and seed morphology in the mini-seed derived plants (D=0.7482) and the coefficient of variation was27.92%. Considerating the goal of breeding new cultivars in loquat,3lines were selected from the mini-seed derived plants with special traits, big fruit (203#), few seeds (C1#) and white flesh (4-1-5#),respectively.
3Investigation on the hardiness and preliminary screening for cold resistant material from mini-seed derived plants
There were abundant genetic diversity in mini-seed plants in response to the frost, and the range of embryo injury rate and defoliation rate were65.4%and90%, respectively; there was no correlation between the rate of embryo injury and defoliation rate (R=0.2911); a significant negative correlation was found between the rate of embryo injury and blooming time, the rate of embryo injury decreased with the blooming delay; the average frost index of mini-seed derived plants was25.34, and cold-resistant6-3-10and C2were preliminary selected based on the rate of embryo injury as the main index combined with defoliation rate of the biennial autumn shoots.
4Pollen mother cell meiosis and the unnormal phenomenon of pollen and ovule in morphocytology
About3.9%of pollen mother cells (material cv. Zaozhong6) showed nondivision or nonsynchronous division at meiosis, and gave rise to dyads (0.3%) triads (0.5%) and hexads (0.1%) after anaphase Ⅱ, indicating a frequency of unreduced gametes of0.28%. There were about21.6%unnormal pollen cells including cavity, with no nucleolus and distorted microspores, which all appeared at unicleate stage. The anatomise observation (material cv. Dawuxing) showed that structure of13.3%ovules was abnormal, which included small embryo sac, female genitalia degeneration, embryo sac degeneration and ovule degeneration. Unnormal meiosis and pollen would result in pollen abortion, and fertilization and embryo development could not be satisfied in abnormal ovules, which all resulted in seed degeneration.
5Morphological analysis of styles and stigmas and their relation to pollen germination and pollen-tube growth
The length of styles significantly differed in a flower in loquat (material cv. Dawuxing, Zaozhong6, Longquan1); and there were significant differences in matoid cell characteristics, stigma receptivity, mucus secretion, in situ pollen germination, and pollen tube growth in different lengh styles in a flower. Pollen tube quantity and pollen germination were remarkably different, depending on style length: pollen germination in the stigma of long and middle sized styles was greater than that of short styles, and the number of pollen tubes growing in the short styles was significantly smaller than in the long and middle styles. The results indicated that the long and middle styles were more receptive to pollen germination and pollen tube growth than short styles.
6The pistil structure and its relation to seed set and degeneration
Based on investigating pistil structure through free-hand section and observation on seed set under manual control-pollination, the results, for the first time, demonstrated that the5styles and5lobules had a one-to-one relationship in both the morphylogical and reproductive characteristics in loquat. Obturator originates from the carpellary margins, formation of the septum in the ovary, extends radiating in5lobules; styles5, free at base, each one leading through the placenta to one of the5locules. After hand-pollination with removal of4styles (only one style left), there are seeds only in one lobule per fruit with two types of seed set patterns:76.7%fruits had two normal seeds, which were in accordance with pattern I;23.3%fruits had only one normal seed and one degenerative seed, which were in accordance with pattern II, while all the8ovules in other4lobules were unfertilized and wilted. Seed degradation rate was11.7%when only one style was remained. The results demonstrated that the5styles and5lobules had a one-to-one relationship in reproductive characteristics in loquat. Moreover, the degenerative seed beside the normal seed in the same lobule was possibly resulted from the stimulation by plant growth substances secreted by the normal embryo in the same locule. In the in vitro culture of the seeds, a haploid (n=x=17) and an aneuploid (n-3=14) were obtained.
1小种子植株叶遗传多样性分析和重要类群倍性鉴定
小种子植株叶遗传多样性指数D=0.785,各特征变异系数在9.84%～40.09%之间,具有丰富的多样性。UPGMA法将79份材料分为了6大类群,第Ⅰ类和第Ⅵ类表现最为特殊,明显区别于母本“大五星”,为枇杷特异种质选育的重要类群。其中Ⅰ类表型为叶片大型、叶柄粗长、叶色浓绿、质地厚实、坚硬,具有多倍体特征。利用本实验室优化的适于枇杷染色体计数根尖压片方法,对第Ⅰ类小种子植株进行染色体倍性鉴定,确定1号,115号,137号和249号为三倍体(2n=3x=51),为无籽枇杷育种提供了宝贵的材料。三倍体植株形态学上表现为叶大、花大、果大、钟形,果皮橙黄色,无籽。
2小种子植株花期、果实种子性状多样性与特异优良材料筛选
小种子植株花期多样性丰富,并结合育种和生产实际,筛选到212号早花型单株和6060号晚花型单株,为培育枇杷早熟晚熟品种提供了宝贵的材料；小种子植株在果实种子性状上多样性丰富,变异系数平均为27.92%,多样性指数平均为0.7482；并综合枇杷育种目标,初步筛选出了大果型单株203号,少核型单株C1号和白肉型单株4-1-5号,为培育大果、少核和优质白肉品种提供了宝贵的材料。
3枇杷小种子植株抗寒性调查及抗寒株系初步筛选
小种子植株平均冻害指数为25.34,对冻害的反应表现出多样性。种胚受冻率及二年生秋稍落叶率变幅分别达65.4%、90%。落叶率和种胚受冻率不具有相关性(R=0.2911)；种胚受冻率和花期早晚呈现出显著负相关(R2=0.9137),种胚受冻率随花期的延后逐渐降低；并以种胚受冻率为主要评价指标,结合落叶情况,初步筛选出抗寒株系6-3.10和C2,为抗寒基因的定位及抗寒育种提供了宝贵的材料。
4枇杷花粉母细胞减数分裂及花粉和胚珠细胞形态学观察
枇杷(早钟6号为材料)减数分裂Ⅱ期异常的占3.9%,表现为不分裂或不同步分裂,产生了异常二分体(0.3%),四分体(0.5%)和六分体(0.1%),未减数配子的频率为0.28%；花粉细胞有21.6%的形态结构异常,且均出现在单核期,包括空腔、无细胞核以及畸变小孢子等。枇杷(大五星为材料)胚珠石蜡切片结构观察发现,有13.3%存在异常现象,包括小胚囊、雌性生殖器退化、胚囊退化和胚珠退化等4类。减数分裂异常,花粉异常畸变可导致枇杷花粉败育,从而引起授粉受精不良,同时异常的胚珠结构不能提供受精及胚胎发育的场所,最终导致种子退化。
5枇杷柱头和花柱的形态分析及其与花粉萌发和花粉管生长的关系
枇杷(以“大五星”、“早钟6号”和“龙泉1号”为材料)每花内5个花柱长短差异显著。枇桤(以“大五星”为材料)每花内长、中和短型花柱的柱头乳突细胞、柱头可受性、分泌大量黏液的持续时间,以及花粉萌发数量和花粉管生长均存在着明显的差异,长的花柱比短的花柱更有利于接受花粉及随后的花粉萌发；并且不同长度的花柱其花粉原位萌发和花粉管生长情况也有较明显的差异,表现为短型花柱柱头花粉萌发量、中部和基部的花粉管数量均依次低于中型和长型花柱。由此推论枇杷柱头、花柱的形态繁殖特性与种子高退化率及不同退化程度间具有相关性。
6枇杷的雌蕊结构及其与种子生产和退化的关系
通过徒手切片和去柱头人工控制授粉等措施,首次查明了枇杷的5个柱头和5个心室在形态结构和繁殖特性上均是完美的一一对应关系。枇杷雌蕊结构为子房被隔膜分为5个心室,5个花柱离生,基部分离,并且每个花柱分别对应一个心室；通过去掉4个柱头,只留一个柱头进行人工授粉后,只在一个心室中有种子产生,其它4个心室的胚珠均未受精而萎蔫。其中有76.7%的果实符合模式Ⅰ即同一心室中同时产生两颗正常的种子,有23.3%的果实符合模式Ⅱ,即同一心室中有一颗正常的种子,另外一颗产生了不同程度的退化。在只有一个柱头时,种子的败育率为11.7%。根据以上结果首次讨论了枇杷雌蕊结构与种子败育之间的关系,推论同一心室中的败育种子有一定几率是由刺激产生的单倍体。对去4柱头授粉得到的42粒退化种子进行无菌培养和染色体倍性鉴定。结果得到一份单倍体材料(n=x=17),证实了推论。同时,还获得一份非整倍体材料(n=x-3=14),为非整倍体的获得提供了全新的途径和契机。
The loquat breeding group of Sichuan Agricultural University had developed a protocol for culture of mini-seeds (middle-degraded seeds) and production of plants called "mini-seed derived plant", providing the valuable material for germplasm resource development and innovation. In the present study, a series of experiments were carried out to investigate the genetic diversity of mini-seed derived plants, excellent germplasm selection and seed degeneration mechanism in loquat. In part I, the genetic diversity and breeding value of mini-seed plants were studied dased on cytology and morphology research, in order to selecte excellent germplasm. In part II, the seed degeneration mechanism was stuied by pollen mother cell meiosis and the unnormal phenomenon of pollen and ovule in morphocytology; meanwhile, the relationship between pistil structure and seed abortion was clarified for the first time in loquat, aiming at providing data for degenerated seed origin. The main results were as follows.
1Diversity of mini-seed plants based on leaf morphology and chromosome number of group I identified
There was abundant genetic diversity in leaf morphology (D=0.785). The values obtained varied broadly with coefficients of variation ranging from9.8%to40.1%. Analysis of the79accessions divided them into6groups by the UPGMA method. The leaves of group Ⅵ were light green, small, thin and soft with slender petioles; while the leaves of group I were dark green, large, hard and thick with long and thick petioles. These two groups were important excellent germplasm. And1#,115#,137#and249#in group I were triploids (2n=3x=51), which provided precious materials for breeding seedless varieties. The triploid plants had the typical morphological appearances with large leaves, big flowers, big and bell-shaped fruits and orange yellow fruit peel.
2Diversity in florescence, fruit and seed morphology of mini-seed derived plants and excellent germplasm selection
There was abundant genetic diversity in florescence in the mini-seed derived plants and2lines were selected with special traits, early flowering (212#) and late flowering (6060#). There was also abundant diversity in fruit and seed morphology in the mini-seed derived plants (D=0.7482) and the coefficient of variation was27.92%. Considerating the goal of breeding new cultivars in loquat,3lines were selected from the mini-seed derived plants with special traits, big fruit (203#), few seeds (C1#) and white flesh (4-1-5#),respectively.
3Investigation on the hardiness and preliminary screening for cold resistant material from mini-seed derived plants
There were abundant genetic diversity in mini-seed plants in response to the frost, and the range of embryo injury rate and defoliation rate were65.4%and90%, respectively; there was no correlation between the rate of embryo injury and defoliation rate (R=0.2911); a significant negative correlation was found between the rate of embryo injury and blooming time, the rate of embryo injury decreased with the blooming delay; the average frost index of mini-seed derived plants was25.34, and cold-resistant6-3-10and C2were preliminary selected based on the rate of embryo injury as the main index combined with defoliation rate of the biennial autumn shoots.
4Pollen mother cell meiosis and the unnormal phenomenon of pollen and ovule in morphocytology
About3.9%of pollen mother cells (material cv. Zaozhong6) showed nondivision or nonsynchronous division at meiosis, and gave rise to dyads (0.3%) triads (0.5%) and hexads (0.1%) after anaphase Ⅱ, indicating a frequency of unreduced gametes of0.28%. There were about21.6%unnormal pollen cells including cavity, with no nucleolus and distorted microspores, which all appeared at unicleate stage. The anatomise observation (material cv. Dawuxing) showed that structure of13.3%ovules was abnormal, which included small embryo sac, female genitalia degeneration, embryo sac degeneration and ovule degeneration. Unnormal meiosis and pollen would result in pollen abortion, and fertilization and embryo development could not be satisfied in abnormal ovules, which all resulted in seed degeneration.
5Morphological analysis of styles and stigmas and their relation to pollen germination and pollen-tube growth
The length of styles significantly differed in a flower in loquat (material cv. Dawuxing, Zaozhong6, Longquan1); and there were significant differences in matoid cell characteristics, stigma receptivity, mucus secretion, in situ pollen germination, and pollen tube growth in different lengh styles in a flower. Pollen tube quantity and pollen germination were remarkably different, depending on style length: pollen germination in the stigma of long and middle sized styles was greater than that of short styles, and the number of pollen tubes growing in the short styles was significantly smaller than in the long and middle styles. The results indicated that the long and middle styles were more receptive to pollen germination and pollen tube growth than short styles.
6The pistil structure and its relation to seed set and degeneration
Based on investigating pistil structure through free-hand section and observation on seed set under manual control-pollination, the results, for the first time, demonstrated that the5styles and5lobules had a one-to-one relationship in both the morphylogical and reproductive characteristics in loquat. Obturator originates from the carpellary margins, formation of the septum in the ovary, extends radiating in5lobules; styles5, free at base, each one leading through the placenta to one of the5locules. After hand-pollination with removal of4styles (only one style left), there are seeds only in one lobule per fruit with two types of seed set patterns:76.7%fruits had two normal seeds, which were in accordance with pattern I;23.3%fruits had only one normal seed and one degenerative seed, which were in accordance with pattern II, while all the8ovules in other4lobules were unfertilized and wilted. Seed degradation rate was11.7%when only one style was remained. The results demonstrated that the5styles and5lobules had a one-to-one relationship in reproductive characteristics in loquat. Moreover, the degenerative seed beside the normal seed in the same lobule was possibly resulted from the stimulation by plant growth substances secreted by the normal embryo in the same locule. In the in vitro culture of the seeds, a haploid (n=x=17) and an aneuploid (n-3=14) were obtained.
引文
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