花椰菜和大白菜新种质创制
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摘要
花椰菜(Brassica oleracea var. botrytis)为十字花科(Cruciferae)芸薹属甘蓝种的一个变种,生产中常遭受多种病、虫害的威胁,如黑腐病、黑胫病、根肿病和蚜虫等,严重影响其产量和品质。在花椰菜常规种中,上述病、虫害的抗性资源匮乏或者单一,而在十字花科野生植物中存在丰富的异源高抗基因资源。应用体细胞杂交技术,可以克服常规种间杂交不亲和性,获得种间杂种,转移来自不同物种的异源抗性基因,丰富育种材料抗性基因的遗传多样性,创制新的抗病、虫种质。
本研究第一部分是利用体细胞杂交技术向芸薹属甘蓝类蔬菜转移野生抗病、虫等优良性状。
紫罗兰(M. incana)为十字花科紫罗兰属的一个野生种,具有很高的ω-亚麻酸含量以及优良的蚜虫抗性等。本实验建立了花椰菜和紫罗兰原生质体的PEG融合及培养技术,首次成功获得了紫罗兰与花椰菜的属间体细胞杂种再生植株,并对再生株的形态学,分子标记特征,染色体数目,细胞DNA含量等指标进行了系统的分析,为异源优异种质的利用建立了体细胞杂交技术基础。
黑芥(B. nigra)为芸薹属三个基本种之一,多为野生状态存在。黑芥中存在对黑腐病、黑胫病、根肿病等病害高抗、多抗性的基因型。本实验室通过体细胞杂交技术,获得了众多花椰菜与黑芥体细胞杂种。本研究对利用非对称体细胞杂交技术获得并成功移入温室的79个花椰菜和黑芥的杂种进行全生育期的生物学特性调查,内容包括植株形态学观察、细胞DNA含量测定、染色体计数、基因组组成、育性调查等。通过胚挽救技术获得了部分杂种的回交后代植株并对其部分生物学特性也进行了调查,同时对杂种一代及其回交后代进行了黑腐病抗性鉴定。
本研究第二部分是利用远缘有性杂交手段向白菜中转移野生种黑芥的抗病性状。大白菜(Brassica campestris ssp. pekinensis)是我国重要的蔬菜作物之一。在种植过程中,常遭受黑腐病、黑胫病和根肿病等病菌的侵染,致使产量下降。通过远缘有性杂交手段把黑芥的抗病性转移到白菜中,对白菜的育种将具有重要意义。本实验以品质优良的感病大白菜为母本,黑芥为父本进行杂交,并对获得的F1代植株进行了形态学,细胞学和分子标记三种手段分析鉴定。通过胚珠和子房培养等胚挽救技术成功获得了回交后代,初步的黑腐病菌人工接种鉴定结果表明,部分回交二代杂种植株中仍具备良好的黑腐病抗性。
主要实验结果如下：
1.成功建立了紫罗兰和花椰菜属间体细胞杂交体系,获得了118株再生植株,并对它们进行了形态学调查、RAPD和SRAP分子标记鉴定,同工酶谱分析,染色体计数和流式细胞术核DNA含量检测,表明二株为真杂种,且均为混倍体,染色体数变化范围为26到48条。对不同鉴定方法的比较显示,形态学鉴定直观、简便,有较高的判断率,但SRAP分子标记鉴定更准确可靠。获得的紫罗兰和花椰菜属间体细胞杂种植株在移栽入地后,生长缓慢,没有达到成株期。
2.对成功移栽入温室的79株花椰菜和黑芥的体细胞杂种进行了全生育期性状调查分析,内容包括细胞DNA含量测定、染色体计数、植株形态学观察和育性调查等。杂种形态变异广泛,多数呈中间类型,少数呈偏亲性状；杂种细胞DNA含量不一致,呈现大于、等于和小于双亲细胞DNA含量之和的三种类型；DNA含量大于双亲之和的杂种中,有较高比例的材料叶型出现畸形和(或)植株生长势弱,育性差。显示出杂种植株的生长势和育性与细胞的DNA含量具有一定的相关性。通过染色体计数,发现所检测杂种多为混倍体,即同一植株的细胞有多种染色体数目构成。在79个杂种中,筛选出13个目标材料,生长势较强,有较好或一定程度的育性,表明采用体细胞杂交技术手段来改良甘蓝类蔬菜的遗传多样性具有可行性。
3.根据结实与否选取了可育和不育杂种各二株,对其生物学特性进行了对比分析。不育杂种的花粉母细胞(PMCs)减数分裂行为异常的比例远高于可育杂种,出现了大量的染色体桥,染色体滞后,染色体不均等分裂等异常行为；对PMCs终变期观察表明：不育杂种中出现了大量单价体；可育杂种中,二价体数目明显提高,单价体数目较少。基因组原位杂交(GISH)结果显示：可育杂种的体细胞染色体数为供、受体染色体数之和,为对称杂种；不育杂种细胞中含有受体花椰菜的全部基因组以及附加的3-12条供体完整染色体,为低度非对称杂种。且附加的黑芥染色体在减Ⅰ期均出现染色体滞后,染色体不均等分裂等异常行为。对可育和不育杂种的基因组SRAP分子标记分析表明：杂种DNA均有重组发生,且可育杂种的平均新带比例略大于不育杂种。
4.对10个株系的15株BC1杂种进行了生物学特性调查,内容包括细胞DNA含量测定、染色体计数、植株形态学观察和花粉活力调查等。杂种形态多呈中间类型,有三棵植株形态明显趋向于受体花椰菜。核DNA含量和染色体数目均不一致,显示出混倍特性,但相比于体细胞杂种一代,其变化范围明显变窄,且有接近于受体花椰菜的趋势。以花椰菜为父本,共有4个BC1植株获得自交和回交种子,一颗植株通过胚挽救技术获得回交二代胚苗。
5.采用从北京,湖南等全国六个省份收集并经过鉴定确定的8个黑腐病菌种以及从英国引进的分别属于RaceⅠ和RaceⅣ的二个生理小种,共10个黑腐菌种对融合亲本花椰菜和黑芥进行苗期抗病性鉴定。筛选出了一个受体(花椰菜)的病情指数为70.4,为“高感(HS)”级别,同时供体(黑芥)的病情指数为8.8,达到“高抗(HR)”级别的黑腐菌种CH5。用筛选的黑腐菌种CH5对体细胞杂种一代及其回交后代植株进行抗病性鉴定,结果显示部分回交二代杂种植株仍具有良好的黑腐病抗性,病情指数为15.7,达到“抗病(R)”级别。
6.对获得的10株白菜×黑芥的F1代植株,经形态学,细胞学和SRAP等方法鉴定出6株为真杂种。其中有一株杂种(H1)体细胞含有26条染色体,GISH结果显示其基因组组成为ABB型,育性相对较高,获得了175粒回交一代种子；其它5株杂种50%以上的体细胞含有18条染色体,GISH结果显示其基因组组成为AB型,花粉完全败育,通过胚挽救技术也未获得回交后代。由此推测,染色体数目的增多和同源染色体的配对有利于提高杂种的育性。
7.白菜-黑芥杂种与白菜的BC1代植株形态学上趋向于母本大白菜,表现出明显的杂种优势。细胞学观察发现不同BC1代杂种细胞染色体数目略有不同,但多为28条,GISH结果显示其基因组组成为AAB型,花粉活力明显提高,得到大量的回交二代种子；BC2代杂种细胞染色体数目不同,从22-28条不等。对回交一代和回交二代杂种植株的黑腐病抗性鉴定结果显示：部分回交二代植株仍然保留良好的黑腐病抗性。
Cauliflower, a variety of Brassica in Cruciferae, is popular with many people as a kind of vegetable. However, cauliflower production always suffers from diseases such as black rot, blackleg, club root and so on, affecting the yield and quality of cauliflower seriously.
Somatic hybridization can overcome the sexual incompatibility barriers to obtain the hybrid between different species, and transfer the alien resistant genes from other species. It can enrich the diversity of pathogen resistant genes of the breeding material, and realize the innovation of crop germplasm.
One object of this study was to transfer multi-resistant genes from wild species to cauliflower by using somatic hybridization technique.
Matthiola incana belongs to Cruciferae family, which possesses desirable characteristics such as the rich a-linolenic acid and the high resistance to aphids. This study produced somatic hybrids between B. oleracea and M. incana for the first time by using the symmetrical somatic hybridization technique, which laid a good foundation for the utilizing of the heterogenous excellent germ plasm. The regenerated plants were investigated on morphology, RAPD or SRAP molecular marker, isozyme pattern, chromosome number and DNA content.
B. nigra, one of the three Brassica species, exhibited high resistance to black rot, black leg and clubroot diseases. In our privious study, a lot of somatic hybrids between B. nigra and cauliflower were obtained. Seventy-nine hybrids were transplanted into greenhouse successfully, the biological characters of which were studied systemicly in the present research. The backcross progeny of partial somatic hybrids were obtained through embryo rescue technique, the reaiatance of which to X. campestris pv. campestris was also assessed.
The other object of this study was to transfer the alien diseases resistant genes from B. nigra to Chinese cabbage. Chinese cabbage is one of the most important vegetable crops in China. In the process of cultivation, the chinese cabbage production always suffers from diseases such as black rot, blackleg, club root and so on, affecting the yield and the quality seriously. In order to transfer the alien diseases resistant genes from other species, crosses between Brassica campestris pekinensis and B. nigra were made and the F1 plants obtained were investigated on morphology, SRAP molecular marker, chromosome number and genome constitution. The backcross progeny of partial F1 hybrids were obtained through embryo rescue technique, the reaiatance of which to X. campestris pv. campestris was also assessed.
All the results were described as follows:
1. Two somatic hybrids between cauliflower and Matthiola incana were produced and investigated on morphology, RAPD molecular, chromosome number, isozyme pattern and nuclear DNA content. The two hybrids were both allopolyploid and had 26-48 chromosomes. Comparing the different identification method, it was thought that the morphological observation was more simple and direct, and with enough veracity in most of the cases, but the SRAP molecular marker was more exact and dependable. Both hybrids showed limit growth and neither of them can grow to mature plants in the greenhouse.
2. Seventy-nine somatic hybrids were transplanted in greenhouse successfully and were investigated on morphology, SRAP molecular marker, chromosome number and genome size. Morphology of the regenerated plants varied a lot and most of the population was intermediate to that of the parents; The hybrid plants had different nuclear DNA content:higher, lower than the parental sum or equal to it. The hybrids with higher DNA content had abnormal leaf, bad growth vigour and poor fertility, which indicated that the growth vigour and fertility had relations with the nuclear DNA content. By chromosome counting, the majority of the detected hybrids were mixoploid. Among 79 hybrids,13 aimed hybrids were picked out, which all had vigorous growth and partial fertility. Therefore, it was feasible to broaden the genetic variability of cabbage though asymmetric somatic hybridization.
3. The compare of the biological characters of the fertile and sterile hybrids were also studied. The PMCs meiosis behaviors were more abnormal in sterile hybrids than in fertile ones. Laggard chromosomes, chromosome bridges and unequal separation happened more frequently. The results of GISH analysis indicated that the fertile hybrids contained the whole genome of B. nigra and cauliflower with regular meiotic divisions while the sterile hybrids had the whole genome of cauliflower and partial intact chromosomes of B. nigra. Moreover, the additional B. nigra chromosomes had a high occurrence of laggards and unequal distribution. Genome study of the two kinds of hybrids showed that the DNA recombination happened in all hybrids.
4. A total of fifteen BC1 hybrids were obtained, the biological characters of which were studied. Morphology of the BC1 hybrids was intermediate to that of the parents and three ones had similar morphology to cauliflower. The nuclear DNA content and the chromosomes number were both inconsistent, which indicated the trait of aneuploid. Comparing with the somatic hybrids, the range of the BC1 hybrids became narrow, which also showed a tendensy to cauliflower. All the fifteen BC1 plants were used to self-pollination and backcrossing with cauliflower, four of which could obtain BC2 and BC1S1 seeds.
5. Eight local Xcc strains and two foreign Xcc strains belonging to raceⅠand raceⅣrespectively, were used to inoculate parent plants. A Xcc strain CH5 were selected depending on severe chlorosis around the inoculation sites on cauliflower giving a disease severity rating of 9 and no symptom development on B. nigra. The selected Xcc strain CH5 was used to inoculate hybrids, and backcross plants. Some BC2 hybrids showing good disease resistance were obtained.
6. Ten F1 plants were obtained between Chinese cabbage and B. nigra, six of which were characterized to be hybrids through morphology, SRAP molecular marker and cytology analysis. One of the hybrids had 26 chromosomes with ABB genomic constitution by GISH analysis showing partial fertility. The other five hybrids had 18 chromosomes in more than 50% PMCs with AB genomic constitution were entirely sterile. An increase in the chromosomes number and homologous chromosomes pairing mignt improve the hybrids fertility.
7. The BC1 hybrids were more homophylic to the female parent on morphology, showing visible heterosis. Most plants had 28 chromosomes with with AAB genomic constitution by GISH analysis. The male fertility increased obviously and many BC2 seeds were obtained. The number of BC2 hybrids chromosomes was not consistent, having 22-36 chromosomes. Some BC2 hybrids showing good Xcc resistance were obtained.
本研究第一部分是利用体细胞杂交技术向芸薹属甘蓝类蔬菜转移野生抗病、虫等优良性状。
紫罗兰(M. incana)为十字花科紫罗兰属的一个野生种,具有很高的ω-亚麻酸含量以及优良的蚜虫抗性等。本实验建立了花椰菜和紫罗兰原生质体的PEG融合及培养技术,首次成功获得了紫罗兰与花椰菜的属间体细胞杂种再生植株,并对再生株的形态学,分子标记特征,染色体数目,细胞DNA含量等指标进行了系统的分析,为异源优异种质的利用建立了体细胞杂交技术基础。
黑芥(B. nigra)为芸薹属三个基本种之一,多为野生状态存在。黑芥中存在对黑腐病、黑胫病、根肿病等病害高抗、多抗性的基因型。本实验室通过体细胞杂交技术,获得了众多花椰菜与黑芥体细胞杂种。本研究对利用非对称体细胞杂交技术获得并成功移入温室的79个花椰菜和黑芥的杂种进行全生育期的生物学特性调查,内容包括植株形态学观察、细胞DNA含量测定、染色体计数、基因组组成、育性调查等。通过胚挽救技术获得了部分杂种的回交后代植株并对其部分生物学特性也进行了调查,同时对杂种一代及其回交后代进行了黑腐病抗性鉴定。
本研究第二部分是利用远缘有性杂交手段向白菜中转移野生种黑芥的抗病性状。大白菜(Brassica campestris ssp. pekinensis)是我国重要的蔬菜作物之一。在种植过程中,常遭受黑腐病、黑胫病和根肿病等病菌的侵染,致使产量下降。通过远缘有性杂交手段把黑芥的抗病性转移到白菜中,对白菜的育种将具有重要意义。本实验以品质优良的感病大白菜为母本,黑芥为父本进行杂交,并对获得的F1代植株进行了形态学,细胞学和分子标记三种手段分析鉴定。通过胚珠和子房培养等胚挽救技术成功获得了回交后代,初步的黑腐病菌人工接种鉴定结果表明,部分回交二代杂种植株中仍具备良好的黑腐病抗性。
主要实验结果如下：
1.成功建立了紫罗兰和花椰菜属间体细胞杂交体系,获得了118株再生植株,并对它们进行了形态学调查、RAPD和SRAP分子标记鉴定,同工酶谱分析,染色体计数和流式细胞术核DNA含量检测,表明二株为真杂种,且均为混倍体,染色体数变化范围为26到48条。对不同鉴定方法的比较显示,形态学鉴定直观、简便,有较高的判断率,但SRAP分子标记鉴定更准确可靠。获得的紫罗兰和花椰菜属间体细胞杂种植株在移栽入地后,生长缓慢,没有达到成株期。
2.对成功移栽入温室的79株花椰菜和黑芥的体细胞杂种进行了全生育期性状调查分析,内容包括细胞DNA含量测定、染色体计数、植株形态学观察和育性调查等。杂种形态变异广泛,多数呈中间类型,少数呈偏亲性状；杂种细胞DNA含量不一致,呈现大于、等于和小于双亲细胞DNA含量之和的三种类型；DNA含量大于双亲之和的杂种中,有较高比例的材料叶型出现畸形和(或)植株生长势弱,育性差。显示出杂种植株的生长势和育性与细胞的DNA含量具有一定的相关性。通过染色体计数,发现所检测杂种多为混倍体,即同一植株的细胞有多种染色体数目构成。在79个杂种中,筛选出13个目标材料,生长势较强,有较好或一定程度的育性,表明采用体细胞杂交技术手段来改良甘蓝类蔬菜的遗传多样性具有可行性。
3.根据结实与否选取了可育和不育杂种各二株,对其生物学特性进行了对比分析。不育杂种的花粉母细胞(PMCs)减数分裂行为异常的比例远高于可育杂种,出现了大量的染色体桥,染色体滞后,染色体不均等分裂等异常行为；对PMCs终变期观察表明：不育杂种中出现了大量单价体；可育杂种中,二价体数目明显提高,单价体数目较少。基因组原位杂交(GISH)结果显示：可育杂种的体细胞染色体数为供、受体染色体数之和,为对称杂种；不育杂种细胞中含有受体花椰菜的全部基因组以及附加的3-12条供体完整染色体,为低度非对称杂种。且附加的黑芥染色体在减Ⅰ期均出现染色体滞后,染色体不均等分裂等异常行为。对可育和不育杂种的基因组SRAP分子标记分析表明：杂种DNA均有重组发生,且可育杂种的平均新带比例略大于不育杂种。
4.对10个株系的15株BC1杂种进行了生物学特性调查,内容包括细胞DNA含量测定、染色体计数、植株形态学观察和花粉活力调查等。杂种形态多呈中间类型,有三棵植株形态明显趋向于受体花椰菜。核DNA含量和染色体数目均不一致,显示出混倍特性,但相比于体细胞杂种一代,其变化范围明显变窄,且有接近于受体花椰菜的趋势。以花椰菜为父本,共有4个BC1植株获得自交和回交种子,一颗植株通过胚挽救技术获得回交二代胚苗。
5.采用从北京,湖南等全国六个省份收集并经过鉴定确定的8个黑腐病菌种以及从英国引进的分别属于RaceⅠ和RaceⅣ的二个生理小种,共10个黑腐菌种对融合亲本花椰菜和黑芥进行苗期抗病性鉴定。筛选出了一个受体(花椰菜)的病情指数为70.4,为“高感(HS)”级别,同时供体(黑芥)的病情指数为8.8,达到“高抗(HR)”级别的黑腐菌种CH5。用筛选的黑腐菌种CH5对体细胞杂种一代及其回交后代植株进行抗病性鉴定,结果显示部分回交二代杂种植株仍具有良好的黑腐病抗性,病情指数为15.7,达到“抗病(R)”级别。
6.对获得的10株白菜×黑芥的F1代植株,经形态学,细胞学和SRAP等方法鉴定出6株为真杂种。其中有一株杂种(H1)体细胞含有26条染色体,GISH结果显示其基因组组成为ABB型,育性相对较高,获得了175粒回交一代种子；其它5株杂种50%以上的体细胞含有18条染色体,GISH结果显示其基因组组成为AB型,花粉完全败育,通过胚挽救技术也未获得回交后代。由此推测,染色体数目的增多和同源染色体的配对有利于提高杂种的育性。
7.白菜-黑芥杂种与白菜的BC1代植株形态学上趋向于母本大白菜,表现出明显的杂种优势。细胞学观察发现不同BC1代杂种细胞染色体数目略有不同,但多为28条,GISH结果显示其基因组组成为AAB型,花粉活力明显提高,得到大量的回交二代种子；BC2代杂种细胞染色体数目不同,从22-28条不等。对回交一代和回交二代杂种植株的黑腐病抗性鉴定结果显示：部分回交二代植株仍然保留良好的黑腐病抗性。
Cauliflower, a variety of Brassica in Cruciferae, is popular with many people as a kind of vegetable. However, cauliflower production always suffers from diseases such as black rot, blackleg, club root and so on, affecting the yield and quality of cauliflower seriously.
Somatic hybridization can overcome the sexual incompatibility barriers to obtain the hybrid between different species, and transfer the alien resistant genes from other species. It can enrich the diversity of pathogen resistant genes of the breeding material, and realize the innovation of crop germplasm.
One object of this study was to transfer multi-resistant genes from wild species to cauliflower by using somatic hybridization technique.
Matthiola incana belongs to Cruciferae family, which possesses desirable characteristics such as the rich a-linolenic acid and the high resistance to aphids. This study produced somatic hybrids between B. oleracea and M. incana for the first time by using the symmetrical somatic hybridization technique, which laid a good foundation for the utilizing of the heterogenous excellent germ plasm. The regenerated plants were investigated on morphology, RAPD or SRAP molecular marker, isozyme pattern, chromosome number and DNA content.
B. nigra, one of the three Brassica species, exhibited high resistance to black rot, black leg and clubroot diseases. In our privious study, a lot of somatic hybrids between B. nigra and cauliflower were obtained. Seventy-nine hybrids were transplanted into greenhouse successfully, the biological characters of which were studied systemicly in the present research. The backcross progeny of partial somatic hybrids were obtained through embryo rescue technique, the reaiatance of which to X. campestris pv. campestris was also assessed.
The other object of this study was to transfer the alien diseases resistant genes from B. nigra to Chinese cabbage. Chinese cabbage is one of the most important vegetable crops in China. In the process of cultivation, the chinese cabbage production always suffers from diseases such as black rot, blackleg, club root and so on, affecting the yield and the quality seriously. In order to transfer the alien diseases resistant genes from other species, crosses between Brassica campestris pekinensis and B. nigra were made and the F1 plants obtained were investigated on morphology, SRAP molecular marker, chromosome number and genome constitution. The backcross progeny of partial F1 hybrids were obtained through embryo rescue technique, the reaiatance of which to X. campestris pv. campestris was also assessed.
All the results were described as follows:
1. Two somatic hybrids between cauliflower and Matthiola incana were produced and investigated on morphology, RAPD molecular, chromosome number, isozyme pattern and nuclear DNA content. The two hybrids were both allopolyploid and had 26-48 chromosomes. Comparing the different identification method, it was thought that the morphological observation was more simple and direct, and with enough veracity in most of the cases, but the SRAP molecular marker was more exact and dependable. Both hybrids showed limit growth and neither of them can grow to mature plants in the greenhouse.
2. Seventy-nine somatic hybrids were transplanted in greenhouse successfully and were investigated on morphology, SRAP molecular marker, chromosome number and genome size. Morphology of the regenerated plants varied a lot and most of the population was intermediate to that of the parents; The hybrid plants had different nuclear DNA content:higher, lower than the parental sum or equal to it. The hybrids with higher DNA content had abnormal leaf, bad growth vigour and poor fertility, which indicated that the growth vigour and fertility had relations with the nuclear DNA content. By chromosome counting, the majority of the detected hybrids were mixoploid. Among 79 hybrids,13 aimed hybrids were picked out, which all had vigorous growth and partial fertility. Therefore, it was feasible to broaden the genetic variability of cabbage though asymmetric somatic hybridization.
3. The compare of the biological characters of the fertile and sterile hybrids were also studied. The PMCs meiosis behaviors were more abnormal in sterile hybrids than in fertile ones. Laggard chromosomes, chromosome bridges and unequal separation happened more frequently. The results of GISH analysis indicated that the fertile hybrids contained the whole genome of B. nigra and cauliflower with regular meiotic divisions while the sterile hybrids had the whole genome of cauliflower and partial intact chromosomes of B. nigra. Moreover, the additional B. nigra chromosomes had a high occurrence of laggards and unequal distribution. Genome study of the two kinds of hybrids showed that the DNA recombination happened in all hybrids.
4. A total of fifteen BC1 hybrids were obtained, the biological characters of which were studied. Morphology of the BC1 hybrids was intermediate to that of the parents and three ones had similar morphology to cauliflower. The nuclear DNA content and the chromosomes number were both inconsistent, which indicated the trait of aneuploid. Comparing with the somatic hybrids, the range of the BC1 hybrids became narrow, which also showed a tendensy to cauliflower. All the fifteen BC1 plants were used to self-pollination and backcrossing with cauliflower, four of which could obtain BC2 and BC1S1 seeds.
5. Eight local Xcc strains and two foreign Xcc strains belonging to raceⅠand raceⅣrespectively, were used to inoculate parent plants. A Xcc strain CH5 were selected depending on severe chlorosis around the inoculation sites on cauliflower giving a disease severity rating of 9 and no symptom development on B. nigra. The selected Xcc strain CH5 was used to inoculate hybrids, and backcross plants. Some BC2 hybrids showing good disease resistance were obtained.
6. Ten F1 plants were obtained between Chinese cabbage and B. nigra, six of which were characterized to be hybrids through morphology, SRAP molecular marker and cytology analysis. One of the hybrids had 26 chromosomes with ABB genomic constitution by GISH analysis showing partial fertility. The other five hybrids had 18 chromosomes in more than 50% PMCs with AB genomic constitution were entirely sterile. An increase in the chromosomes number and homologous chromosomes pairing mignt improve the hybrids fertility.
7. The BC1 hybrids were more homophylic to the female parent on morphology, showing visible heterosis. Most plants had 28 chromosomes with with AAB genomic constitution by GISH analysis. The male fertility increased obviously and many BC2 seeds were obtained. The number of BC2 hybrids chromosomes was not consistent, having 22-36 chromosomes. Some BC2 hybrids showing good Xcc resistance were obtained.
引文
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