抗根肿病桔红心大白菜雄性不育系转育方法研究
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摘要
根肿病（Plasmodiophora brassicae）严重威胁着大白菜等十字花科作物的生产，导致大白菜等十字花科蔬菜产量和质量急剧下降。桔红心大白菜〔Brassica campestrisL. ssp. pekinensis（Lour.）Olsson〕色泽鲜艳，营养丰富，保健价值高，本文以桔红心大白菜和普通心大白菜抗根肿病甲型雄性不育“两用系”为转育材料，进行了抗根肿病桔红心雄性不育系的转育方法研究，并对桔红心大白菜子叶颜色的遗传特征和作为形态标记性状的使用价值进行了探究。主要研究结果如下：
1.通过莲座期对不同品种桔红心材料的叶片抱合方式、叶片平展还是褶皱、有无叶毛等特征的调查，得出124和125与CR9112A的叶片特征比较一致。收获时对桔红心材料经济性状的调查结果显示，123和125的球心色最理想。总起来看，只有125在各方面的性状表现最好，与待转育亲本CR9112A有较多相似性状，所以选125为桔红心亲本，CR9112A为抗源和不育源。对桔红心自交系125以及其与纯合抗性和杂合抗性植株杂交后代的抗性鉴定结果说明，桔红心自交系对根肿病没有抗性，抗性基因型为rr；利用甲型和乙型“两用系”不育株对桔红心自交系育性基因型的测定结果说明，其育性基因型为msms。
2.以普通心抗根肿病大白菜甲型雄性不育“两用系”和桔红心大白菜自交系（LH）为材料，宏观上验证了抗性、育性和桔红心球心色三者的遗传关系，认为三者在遗传上遵循孟德尔的自由组合规律。根据这一规律和核不育“复等位基因”遗传假说，提出了抗根肿病桔红心大白菜100%雄性不育系的转育模式。在转育过程中，F1代回交实现桔红心球心色的纯化，BC1F1代重组纯化抗性，并得到了纯抗的甲型雄性不育“两用系”可育株和临时保持系，甲型雄性不育“两用系”可育株自交后代兄妹交，通过育性调查得到新甲型雄性不育“两用系”，甲型雄性不育“两用系”可育株自交后代不育株与临时保持系杂交得到100%核雄性不育系。转育时先进行抗性鉴定，再进行球心色鉴定和经济性状调查，最后进行育性鉴定。
3.以桔红心大白菜自交系（LH）、普通心大白菜自交系（9112）做亲本对桔红心大白菜子叶颜色的遗传特征和在环境中的稳定性进行了研究，研究结果表明：桔红心和普通心大白菜正反交后代（F1和F1′）子叶颜色均为黄色，F2和BC1F1子叶颜色分离比分别为3:1和1:1（黄色子叶:桔红子叶），这说明桔红心子叶颜色由1对等位核基因控制，与细胞质无关，桔红色对黄色表现为隐性；子叶颜色、花色和球心色三者的遗传表现为一因多效，即桔红子叶植株球心色和花色也为桔红色。在遮光或者弱光处理结束后，光照2h桔红心子叶的颜色可以稳定表现，在低温遮光条件下表现最为显著，催芽时间不同（24h或48h）不影响桔红心子叶颜色的表现。因此，子叶颜色可以作为桔红心大白菜选育中的一个理想的形态标记加以应用。
4.以桔红心的子叶颜色作为形态标记，BC1F1代的桔红子叶幼苗经抗性鉴定得到抗病桔红心植株，经过杂交重组，在BC1F2代得到含有甲型雄性不育“两用系”可育株基因型的抗病桔红心群体。该群体自交后通过抗性鉴定和育性调查筛选到纯合抗病的桔红心甲型雄性不育“两用系”，自交后代进行兄妹交，可育株同时进行测交，通过测交结果得到了纯合抗病的桔红心甲型雄性不育“两用系”。子叶法筛选桔红心植株时期较早，可以在抗性鉴定前完成，因此只有桔红心植株进行抗性鉴定，即抗性鉴定植株减少。不仅如此，经济性状调查时不包括普通心植株，调查群体也减少。
5.转育抗根肿病桔红心不育系以普通心抗病甲型雄性不育“两用系”与桔红心自交系杂交，F1兄妹交抗性和育性同步纯化，第4代得到雄性不育系，比回交模式提前一代。第二代筛选抗根肿病桔红心植株时，球心色法在结球期调查经济性状调查时确定桔红心的，花色法开花时球心色调查和育性调查同时进行，子叶法球心色、抗性、经济性状、育性逐一筛选。无论是BC1F1代还是F2代，子叶法的抗性鉴定群体和经济性状调查群体都是最小，降低了转育过程中的工作强度，所以，三种方法中，子叶法适用范围广，最经济，是筛选桔红心抗病品种和雄性不育系的理想方法。
Orange head Chinese cabbages are known for bright color, rich nutrition and high health value.The clubroot disease in crucifers, caused by Plasmodiophora brassicae, has brought great damagein cruciferous vegetable production, and resulted in Chinese cabbage suffer most seriously forlonger growing period. In the paper utilized AB lines type I of common head Chinese cabbage withhomozygous resistance gene and orange head inbred lines, the genetic male sterile lines of orangeleaf head Chinese cabbage with anti-clubroot were bred. And cotyledon color of orange headChinese cabbages was studied. The main results were described as follows:
1. In order to accelerate the process of transformation, Blade traits and economic traits
of different orange head materials were surveyed in rosette period and in heading period. Blade traits of No.124and125were consistent with CR9112A, and head color of No.123and125were best. So No.123was selected as orange head parent, CR9112A as resistant source
and sterile source. On the other hand, identification results of fertility and resistance of No.125showed that its fertile genotype was msms, and resistant genotype was rr, which was flu on clubroot.
2. On the basis of proving resistance and fertility genotypic identification of orange headChinese cabbage, the model of transfer the genetic male sterile line of anti-clubroot orange headChinese cabbage were proposed. In the light of the model, the new male sterile lines weretransformed obtained by crossing, selfing, sib-mating and testing. During the course oftransformation, resistance was first identified, second head color, third characters, last fertility. It wassolved how to gain successful genetic male sterile lines when three traits need to be transfer.
3. The orange head Chinese cabbage self-line (LH) and the common head Chinese cabbage(9112) self-line were parents, and by cross, reciprocal cross, selfing and back-cross, F1, F1′, F2andBC1F1were gotten. The cotyledon and head of F1and F1′were both yellow, the separation ratio ofF2and BC1F1cotyledon color were3:1and1:1(yellow to orange)respectively, which showed thatcotyledon color of orange leaf head was controlled by a pair of recessive alleles. And the inheritanceof cotyledon color, flower color and leaf head color belongs to pleiotropism. Under the dark or weaklight treatment, seed leaves of orange leaf head Chinese cabbage showed orange after they wereexposed in the sun for about two hours. Under the low temperature (0-3℃)and dark conditions,orange seed leaves of orange leaf head Chinese cabbage were observed more clearly, which were not affected by time of pregermination(24h or48h). So Cotyledon color of orange leaf head Chinesecabbage can be used as an ideal morphological character to apply to the breeding of orange headChinese cabbage.
4. Orange head seedlings, which were gotten by orange as morphological marker inBC1F1, went through resistant identification. After resistant orange head seedlings of BC1F1crossed, there were fertile plants of AB lines type I with orange head and homozygosisresistant genes in BC1F2. The fertile plants went on in self. Passing resistant and fertileidentification of BC1F3, AB lines type I with orange head and homozygosis resistance werecome when the sterile plants in BC1F3sib-mated the fertile plants that should be tested.Cotyledon method screening orange head were finished before resistant identification.Therefore plants of resistant test were less, so did that of character traits.
5. Making use of AB lines type I fertile plants with common head and homozygosisresistance and orange head self-line, new genetic male sterile lines were transformed. Becauseresistance and orange head of F2were homozygous in step by F1sib-mating, male sterile lineswere obtained in the fourth generation. When orange head plants with anti-cluroot were screened inthe second generation, orange heads were picked out at the stage of surveying economic traits byhead color method, and at the blooming stage by flower color method, and at the seedling stage bycotyledon color method. Identification groups of cotyledon color method were the smallest insurveying resistance and economic traits in BC1F1or F2.
1.通过莲座期对不同品种桔红心材料的叶片抱合方式、叶片平展还是褶皱、有无叶毛等特征的调查，得出124和125与CR9112A的叶片特征比较一致。收获时对桔红心材料经济性状的调查结果显示，123和125的球心色最理想。总起来看，只有125在各方面的性状表现最好，与待转育亲本CR9112A有较多相似性状，所以选125为桔红心亲本，CR9112A为抗源和不育源。对桔红心自交系125以及其与纯合抗性和杂合抗性植株杂交后代的抗性鉴定结果说明，桔红心自交系对根肿病没有抗性，抗性基因型为rr；利用甲型和乙型“两用系”不育株对桔红心自交系育性基因型的测定结果说明，其育性基因型为msms。
2.以普通心抗根肿病大白菜甲型雄性不育“两用系”和桔红心大白菜自交系（LH）为材料，宏观上验证了抗性、育性和桔红心球心色三者的遗传关系，认为三者在遗传上遵循孟德尔的自由组合规律。根据这一规律和核不育“复等位基因”遗传假说，提出了抗根肿病桔红心大白菜100%雄性不育系的转育模式。在转育过程中，F1代回交实现桔红心球心色的纯化，BC1F1代重组纯化抗性，并得到了纯抗的甲型雄性不育“两用系”可育株和临时保持系，甲型雄性不育“两用系”可育株自交后代兄妹交，通过育性调查得到新甲型雄性不育“两用系”，甲型雄性不育“两用系”可育株自交后代不育株与临时保持系杂交得到100%核雄性不育系。转育时先进行抗性鉴定，再进行球心色鉴定和经济性状调查，最后进行育性鉴定。
3.以桔红心大白菜自交系（LH）、普通心大白菜自交系（9112）做亲本对桔红心大白菜子叶颜色的遗传特征和在环境中的稳定性进行了研究，研究结果表明：桔红心和普通心大白菜正反交后代（F1和F1′）子叶颜色均为黄色，F2和BC1F1子叶颜色分离比分别为3:1和1:1（黄色子叶:桔红子叶），这说明桔红心子叶颜色由1对等位核基因控制，与细胞质无关，桔红色对黄色表现为隐性；子叶颜色、花色和球心色三者的遗传表现为一因多效，即桔红子叶植株球心色和花色也为桔红色。在遮光或者弱光处理结束后，光照2h桔红心子叶的颜色可以稳定表现，在低温遮光条件下表现最为显著，催芽时间不同（24h或48h）不影响桔红心子叶颜色的表现。因此，子叶颜色可以作为桔红心大白菜选育中的一个理想的形态标记加以应用。
4.以桔红心的子叶颜色作为形态标记，BC1F1代的桔红子叶幼苗经抗性鉴定得到抗病桔红心植株，经过杂交重组，在BC1F2代得到含有甲型雄性不育“两用系”可育株基因型的抗病桔红心群体。该群体自交后通过抗性鉴定和育性调查筛选到纯合抗病的桔红心甲型雄性不育“两用系”，自交后代进行兄妹交，可育株同时进行测交，通过测交结果得到了纯合抗病的桔红心甲型雄性不育“两用系”。子叶法筛选桔红心植株时期较早，可以在抗性鉴定前完成，因此只有桔红心植株进行抗性鉴定，即抗性鉴定植株减少。不仅如此，经济性状调查时不包括普通心植株，调查群体也减少。
5.转育抗根肿病桔红心不育系以普通心抗病甲型雄性不育“两用系”与桔红心自交系杂交，F1兄妹交抗性和育性同步纯化，第4代得到雄性不育系，比回交模式提前一代。第二代筛选抗根肿病桔红心植株时，球心色法在结球期调查经济性状调查时确定桔红心的，花色法开花时球心色调查和育性调查同时进行，子叶法球心色、抗性、经济性状、育性逐一筛选。无论是BC1F1代还是F2代，子叶法的抗性鉴定群体和经济性状调查群体都是最小，降低了转育过程中的工作强度，所以，三种方法中，子叶法适用范围广，最经济，是筛选桔红心抗病品种和雄性不育系的理想方法。
Orange head Chinese cabbages are known for bright color, rich nutrition and high health value.The clubroot disease in crucifers, caused by Plasmodiophora brassicae, has brought great damagein cruciferous vegetable production, and resulted in Chinese cabbage suffer most seriously forlonger growing period. In the paper utilized AB lines type I of common head Chinese cabbage withhomozygous resistance gene and orange head inbred lines, the genetic male sterile lines of orangeleaf head Chinese cabbage with anti-clubroot were bred. And cotyledon color of orange headChinese cabbages was studied. The main results were described as follows:
1. In order to accelerate the process of transformation, Blade traits and economic traits
of different orange head materials were surveyed in rosette period and in heading period. Blade traits of No.124and125were consistent with CR9112A, and head color of No.123and125were best. So No.123was selected as orange head parent, CR9112A as resistant source
and sterile source. On the other hand, identification results of fertility and resistance of No.125showed that its fertile genotype was msms, and resistant genotype was rr, which was flu on clubroot.
2. On the basis of proving resistance and fertility genotypic identification of orange headChinese cabbage, the model of transfer the genetic male sterile line of anti-clubroot orange headChinese cabbage were proposed. In the light of the model, the new male sterile lines weretransformed obtained by crossing, selfing, sib-mating and testing. During the course oftransformation, resistance was first identified, second head color, third characters, last fertility. It wassolved how to gain successful genetic male sterile lines when three traits need to be transfer.
3. The orange head Chinese cabbage self-line (LH) and the common head Chinese cabbage(9112) self-line were parents, and by cross, reciprocal cross, selfing and back-cross, F1, F1′, F2andBC1F1were gotten. The cotyledon and head of F1and F1′were both yellow, the separation ratio ofF2and BC1F1cotyledon color were3:1and1:1(yellow to orange)respectively, which showed thatcotyledon color of orange leaf head was controlled by a pair of recessive alleles. And the inheritanceof cotyledon color, flower color and leaf head color belongs to pleiotropism. Under the dark or weaklight treatment, seed leaves of orange leaf head Chinese cabbage showed orange after they wereexposed in the sun for about two hours. Under the low temperature (0-3℃)and dark conditions,orange seed leaves of orange leaf head Chinese cabbage were observed more clearly, which were not affected by time of pregermination(24h or48h). So Cotyledon color of orange leaf head Chinesecabbage can be used as an ideal morphological character to apply to the breeding of orange headChinese cabbage.
4. Orange head seedlings, which were gotten by orange as morphological marker inBC1F1, went through resistant identification. After resistant orange head seedlings of BC1F1crossed, there were fertile plants of AB lines type I with orange head and homozygosisresistant genes in BC1F2. The fertile plants went on in self. Passing resistant and fertileidentification of BC1F3, AB lines type I with orange head and homozygosis resistance werecome when the sterile plants in BC1F3sib-mated the fertile plants that should be tested.Cotyledon method screening orange head were finished before resistant identification.Therefore plants of resistant test were less, so did that of character traits.
5. Making use of AB lines type I fertile plants with common head and homozygosisresistance and orange head self-line, new genetic male sterile lines were transformed. Becauseresistance and orange head of F2were homozygous in step by F1sib-mating, male sterile lineswere obtained in the fourth generation. When orange head plants with anti-cluroot were screened inthe second generation, orange heads were picked out at the stage of surveying economic traits byhead color method, and at the blooming stage by flower color method, and at the seedling stage bycotyledon color method. Identification groups of cotyledon color method were the smallest insurveying resistance and economic traits in BC1F1or F2.
引文
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