高加索三叶草与白三叶杂交后代遗传特性及育性恢复的研究
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摘要
三叶草属植物具有品质优良、营养丰富、固氮能力强、适口性好、建植迅速等特性。为了培育出结合双亲优良性状的三叶草新品种,以引自新西兰具有强抗逆性的六倍体高加索三叶草为母本,以具有强固氮能力的四倍体大兴安岭野生白三叶为父本进行远缘杂交,通过人工授粉技术成功获得了二者的杂种F1代,但其高度不育。本研究从形态及生物学、细胞遗传学、解剖学及分子标记几个不同水平对高加索三叶草、白三叶及其杂种F1进行鉴定分析,并对杂种育性恢复的途径进行了初步研究,主要结果如下：
1体细胞染色体数目的鉴定结果表明,高加索三叶草与自三叶的杂种F1为五倍体(2n=5x=40),核型公式为2n=5x=40=34m+6sm,是双亲真实的杂交后代。双亲的PMCM I减数分裂行为正常,其中高加索三叶草PMCM I的染色体构型为0.06Ⅰ+23.97Ⅱ,白三叶PMCM I的染色体构型为0.04Ⅰ+15.98Ⅱ。杂种F1的PMCM I染色体构型为9.07Ⅰ+14.33Ⅱ+0.23Ⅲ+0.03Ⅳ,伴随较高频率的单价体和一定频率的多价体,在减数分裂后期Ⅰ,出现滞后染色体及染色体桥。
2高加索三叶草、自三叶及其杂种F]根、茎、叶的解剖结构存在较大差异。初步认定三者的抗旱能力为高加索三叶草>杂种F1>白三叶。
3杂种F1生育期及全年生长日分别为99d和201d,介于双亲之间。发育前、后期生长速度偏向白三叶,中期则显著超过双亲,表现出较强的杂种优势。株型偏向高加索三叶草,除茎长以外,株高、叶长、叶宽、茎粗、根长、根粗、分枝数等形态特征均介于双亲之间,并与双亲有极显著差异。在花的形态方面,杂种F1的小花柄最短而花萼最长,其单株花序数、花序长、花序轴长、小花数、小花长均表现为双亲的中间类型。
4经过远缘杂交得到的杂种F1已经可以结瘤,但固氮性依然缺失。
5对高加索三叶草、白三叶及其杂种F1进行光合生理特性对比分析可知,杂种F1有较高的净光合速率及较低的蒸腾速率,其水分利用效率极显著高于双亲,具备高产潜能。
6MS+2,4-D(0.1mg/L)+6-BA (2mg/L)是杂种F1茎段愈伤组织适宜的诱导培养基；MS+NAA (0.5mg/L)+6-BA (1.0mg/L)+KT(1.0mg/L)为杂种F、茎段愈伤组织适宜的分化培养基,不添加任何激素的1/2MS培养基为适宜的生根培养基。使用蛭石作为炼苗移栽的基质成活率最高,可达93.75%。
7高加索三叶草与白三叶的亲缘关系较远,通过回交法恢复杂种F,育性有一定的困难,本研究中,染色体加倍法是解决其育性问题较合理的途径。
8利用秋水仙素处理杂种F1实体苗的加倍率最高,在2000mg/L秋水仙素结合1.5%的DMSO溶液中,避光处理48h的变异率为6.67%,但嵌合现象较严重；1000mg/L秋水仙素处理杂种F1组培再生苗48h的加倍率为5.88%,但成活率较低,仅为26.47%；处理愈伤组织适宜的秋水仙素溶液浓度为250mg/L,时间为48h,但加倍率较低,仅为4.08%,但获得的植株的倍性纯合,利于稳定遗传。
9利用筛选出的16个ISSR适宜引物对高加索三叶草、自三叶及其杂种F1、F1再生植株进行扩增,共得到170条条带,平均每个引物扩增出10.63条,多态性条带138条,多态性条带百分率达81.18%。亲本高加索三叶草与白三叶遗传差异最大,杂种F1与高加索三叶草遗传距离相对较小,F1组培再生植株与杂种F1间的遗传差异最小,二者遗传相似性为0.9244。
The majority species of Trifolium forage was with the characters of rich nutrition, strong ability of nitrogen fixation, good palatability and planting rapidly. In order to create new forage germplasms of parental superior characters, hexaploid Trifolium ambiguum Bieb. of strong resistance to drought and cold which from New Zealand as the famale parent was hybridized with the tetraploid Trifolium repens L. of strong ability of nitrogen fixation which from Greater Higgnan Mountains, and the hybrid F1which were completely sterile were successfully gotten. In this study, several different levels from the morphology and biology, cytological basis, anatomy and molecular markers of T. ambiguum Bieb., T. repens L. and their hybrid F1were identified by analysis, and fertility restoration of hybrids was preliminarily studied also. The main results were as follows:
1. The identification results of the somatic chromosome number showed that the hybrids F1of T. ambiguum Bieb. and T. repens L. were pentaploid (2n=5x=40), the karyotype formula of them was2n=5x=40=34m+6sm. They were the real hybrid offsprings of T. ambiguum Bieb. and T. repens L.. The PMCM I chromosome pairing configuration of T. ambiguum Bieb., T. repens L. and their hybrids were0.06Ⅰ+23.97Ⅱ,0.04Ⅰ+15.98Ⅱ and9.07Ⅰ+14.33Ⅱ+0.23Ⅲ+0.03Ⅳ respectively. The meiosis of hybrids Fi was irregular. Univalents existed universally and multivalents occurred in a low frequency in metaphase I.Laggards and chromatid bridge appeared in anaphase I
2. The anatomical structures of roots, stems, leaves among T. ambiguum Bieb., T. repens L. and their hybrids were quite different. Initially determined that the drought tolerance was T. ambiguum Bieb.> hybrid F1> T. repens L.
3. The growth period and full-year growth day of hybrids F1was99d and201d respectively, which were between their parents. The growth rate of hybrids F1similar to T. repens L. in earl)'and later growth period but significantly more than their parents in the medium period, showed strong heterosis. Plant type of F1was similar to T. ambiuuum Bieb.. the morphological features in plant height, leaf length, leaf width, stem diameter, root length, root diameter, number of branches were all in the middle of both parents except stem length and showed significant difference.
4. Root nodule were found on the hybrids F1of T. ambiguum Bieb. and T. repens L..but the ability of nitrogen fixation was still absent.
5. Photosynthetic physiological characteristics of T. ambiguum Bieb., T. repens L. and their hybridsshowed that the hybrids F1with a higher net photosynthetic rate and lower transpiration rate.Their water use efficiency was significantly higher than parents, which suggested a potential of high yield.
6. MS+2,4-D (0.1mg/L)+6-BA (2mg/L) were the appropriate induction media of stems of hybrids F,. MS+NAA (0.5mg/L)+6-BA (1.0mg/L)+KT(1.0mg/L) was the suitable differentiation media for the callus from stems of F], and the1/2MS medium which had no any hormones was the suitable rooting medium, vermiculite was the suitable matrix to harden and transplant the regenerated plants of hybrids F1, with highest survival rate up to93.75%.
7. Backcross have not succeeded in seeding, the chromosome doubling is the reasonable method of resuming fertility to the hybrid.
8. Highest efficiency of chromosome doubling was to treat the seedlings of the hybrids F1, the suitable colchicine concentration and treatment time for them was2000mg/L and48h and the variation rate was6.67%. but with frequent chimerism. The suitable colchicine concentration and treatment time for regenerated plants of hybrids F1was1000mg/L and48h and the variation rate was5.88%, but with lower survival rate which was26.47%. The suitable colchicine concentration and treatment time for callus of hybrids F1was250mg/L and48h and with a lower variation rate which was4.08%. but easily to get the pure decaploid.
9.1SSR molecular maker was used to analyze the genetic relationship among T. ambiguum Bieb.. T. repens L., hybrids F1and regenerated plants of hybrids F1. The results showed that170bands of ISSR markers were amplified with16suitable primers, the average was10.63.138bands were polymorphic, and the percentage of polymorphism bands reached81.18%. The genetic distance between Caucasian clover and white clover was larger; the hybrid F1was closer to female parents than male parents and there was just a slight variation between F1hybrids and regeneration of F1
1体细胞染色体数目的鉴定结果表明,高加索三叶草与自三叶的杂种F1为五倍体(2n=5x=40),核型公式为2n=5x=40=34m+6sm,是双亲真实的杂交后代。双亲的PMCM I减数分裂行为正常,其中高加索三叶草PMCM I的染色体构型为0.06Ⅰ+23.97Ⅱ,白三叶PMCM I的染色体构型为0.04Ⅰ+15.98Ⅱ。杂种F1的PMCM I染色体构型为9.07Ⅰ+14.33Ⅱ+0.23Ⅲ+0.03Ⅳ,伴随较高频率的单价体和一定频率的多价体,在减数分裂后期Ⅰ,出现滞后染色体及染色体桥。
2高加索三叶草、自三叶及其杂种F]根、茎、叶的解剖结构存在较大差异。初步认定三者的抗旱能力为高加索三叶草>杂种F1>白三叶。
3杂种F1生育期及全年生长日分别为99d和201d,介于双亲之间。发育前、后期生长速度偏向白三叶,中期则显著超过双亲,表现出较强的杂种优势。株型偏向高加索三叶草,除茎长以外,株高、叶长、叶宽、茎粗、根长、根粗、分枝数等形态特征均介于双亲之间,并与双亲有极显著差异。在花的形态方面,杂种F1的小花柄最短而花萼最长,其单株花序数、花序长、花序轴长、小花数、小花长均表现为双亲的中间类型。
4经过远缘杂交得到的杂种F1已经可以结瘤,但固氮性依然缺失。
5对高加索三叶草、白三叶及其杂种F1进行光合生理特性对比分析可知,杂种F1有较高的净光合速率及较低的蒸腾速率,其水分利用效率极显著高于双亲,具备高产潜能。
6MS+2,4-D(0.1mg/L)+6-BA (2mg/L)是杂种F1茎段愈伤组织适宜的诱导培养基；MS+NAA (0.5mg/L)+6-BA (1.0mg/L)+KT(1.0mg/L)为杂种F、茎段愈伤组织适宜的分化培养基,不添加任何激素的1/2MS培养基为适宜的生根培养基。使用蛭石作为炼苗移栽的基质成活率最高,可达93.75%。
7高加索三叶草与白三叶的亲缘关系较远,通过回交法恢复杂种F,育性有一定的困难,本研究中,染色体加倍法是解决其育性问题较合理的途径。
8利用秋水仙素处理杂种F1实体苗的加倍率最高,在2000mg/L秋水仙素结合1.5%的DMSO溶液中,避光处理48h的变异率为6.67%,但嵌合现象较严重；1000mg/L秋水仙素处理杂种F1组培再生苗48h的加倍率为5.88%,但成活率较低,仅为26.47%；处理愈伤组织适宜的秋水仙素溶液浓度为250mg/L,时间为48h,但加倍率较低,仅为4.08%,但获得的植株的倍性纯合,利于稳定遗传。
9利用筛选出的16个ISSR适宜引物对高加索三叶草、自三叶及其杂种F1、F1再生植株进行扩增,共得到170条条带,平均每个引物扩增出10.63条,多态性条带138条,多态性条带百分率达81.18%。亲本高加索三叶草与白三叶遗传差异最大,杂种F1与高加索三叶草遗传距离相对较小,F1组培再生植株与杂种F1间的遗传差异最小,二者遗传相似性为0.9244。
The majority species of Trifolium forage was with the characters of rich nutrition, strong ability of nitrogen fixation, good palatability and planting rapidly. In order to create new forage germplasms of parental superior characters, hexaploid Trifolium ambiguum Bieb. of strong resistance to drought and cold which from New Zealand as the famale parent was hybridized with the tetraploid Trifolium repens L. of strong ability of nitrogen fixation which from Greater Higgnan Mountains, and the hybrid F1which were completely sterile were successfully gotten. In this study, several different levels from the morphology and biology, cytological basis, anatomy and molecular markers of T. ambiguum Bieb., T. repens L. and their hybrid F1were identified by analysis, and fertility restoration of hybrids was preliminarily studied also. The main results were as follows:
1. The identification results of the somatic chromosome number showed that the hybrids F1of T. ambiguum Bieb. and T. repens L. were pentaploid (2n=5x=40), the karyotype formula of them was2n=5x=40=34m+6sm. They were the real hybrid offsprings of T. ambiguum Bieb. and T. repens L.. The PMCM I chromosome pairing configuration of T. ambiguum Bieb., T. repens L. and their hybrids were0.06Ⅰ+23.97Ⅱ,0.04Ⅰ+15.98Ⅱ and9.07Ⅰ+14.33Ⅱ+0.23Ⅲ+0.03Ⅳ respectively. The meiosis of hybrids Fi was irregular. Univalents existed universally and multivalents occurred in a low frequency in metaphase I.Laggards and chromatid bridge appeared in anaphase I
2. The anatomical structures of roots, stems, leaves among T. ambiguum Bieb., T. repens L. and their hybrids were quite different. Initially determined that the drought tolerance was T. ambiguum Bieb.> hybrid F1> T. repens L.
3. The growth period and full-year growth day of hybrids F1was99d and201d respectively, which were between their parents. The growth rate of hybrids F1similar to T. repens L. in earl)'and later growth period but significantly more than their parents in the medium period, showed strong heterosis. Plant type of F1was similar to T. ambiuuum Bieb.. the morphological features in plant height, leaf length, leaf width, stem diameter, root length, root diameter, number of branches were all in the middle of both parents except stem length and showed significant difference.
4. Root nodule were found on the hybrids F1of T. ambiguum Bieb. and T. repens L..but the ability of nitrogen fixation was still absent.
5. Photosynthetic physiological characteristics of T. ambiguum Bieb., T. repens L. and their hybridsshowed that the hybrids F1with a higher net photosynthetic rate and lower transpiration rate.Their water use efficiency was significantly higher than parents, which suggested a potential of high yield.
6. MS+2,4-D (0.1mg/L)+6-BA (2mg/L) were the appropriate induction media of stems of hybrids F,. MS+NAA (0.5mg/L)+6-BA (1.0mg/L)+KT(1.0mg/L) was the suitable differentiation media for the callus from stems of F], and the1/2MS medium which had no any hormones was the suitable rooting medium, vermiculite was the suitable matrix to harden and transplant the regenerated plants of hybrids F1, with highest survival rate up to93.75%.
7. Backcross have not succeeded in seeding, the chromosome doubling is the reasonable method of resuming fertility to the hybrid.
8. Highest efficiency of chromosome doubling was to treat the seedlings of the hybrids F1, the suitable colchicine concentration and treatment time for them was2000mg/L and48h and the variation rate was6.67%. but with frequent chimerism. The suitable colchicine concentration and treatment time for regenerated plants of hybrids F1was1000mg/L and48h and the variation rate was5.88%, but with lower survival rate which was26.47%. The suitable colchicine concentration and treatment time for callus of hybrids F1was250mg/L and48h and with a lower variation rate which was4.08%. but easily to get the pure decaploid.
9.1SSR molecular maker was used to analyze the genetic relationship among T. ambiguum Bieb.. T. repens L., hybrids F1and regenerated plants of hybrids F1. The results showed that170bands of ISSR markers were amplified with16suitable primers, the average was10.63.138bands were polymorphic, and the percentage of polymorphism bands reached81.18%. The genetic distance between Caucasian clover and white clover was larger; the hybrid F1was closer to female parents than male parents and there was just a slight variation between F1hybrids and regeneration of F1
引文
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