摘要
油菜是我国主要的油料作物,而杂种优势利用又是提高油菜产量、缓解产量与品质之间矛盾、增强品种抗(耐)逆性的一条有效途径。油菜隐性细胞核雄性不育具有不育性彻底稳定、细胞质来源丰富、恢复源广等优点,在国内已经成为杂种优势利用的一个重要途径,但在杂交制种时需人工拔除母本行中50%的可育株,限制了其广泛应用。隐性细胞核雄性不育材料S45A的不育性受两对重叠隐性基因控制(潘涛等,1988),S45B的BnMs1位点处于杂合状态,而Bnms2位点处于隐性纯合状态。本研究以S45A和育性正常的普通甘蓝型油菜IL362-2-4为材料通过两亲本的杂交、自交、兄妹交及等位性分析构建了在BnMs2位点分离的近等基因系S4516AB,在此基础上对隐性核不育基因BnMs2进行了精细定位,克隆和转基因研究,一方面为雄性不育的选育提供分子标记,另一方面以期加深对雄性不育机理的认识。主要研究结果如下：
1.对S4516B和S45B的杂合位点进行等位性分析,为S4516AB定位群体的确立打下了基础。通过对NIL系S4516AB的育性调查结果表明,兄妹交后代的育性分离比为1：1,可育株自交后代的育性比例符合3：1,说明S4516B为单位点杂合,结合等位性分析的结果证实了两型系S4516AB的基因型,不育株S4516A为Bnms1Bnms1 Bnms2bnms2,可育株S4516B为Bnms1Bnms1 BnMs2Bnms2。
2.利用AFLP结合BSA的分析方法,通过筛选1024对AFLP引物组合找到了与BnMs2连锁的AFLP标记12个,在262株的NIL分离群体中对BnMs2进行了初步定位,将目标基因锁定在3.5cM内,另外8个AFLP标记与BnMs2共分离。
3.对12个AFLP标记回收克隆测序后,用PCR walking分离了8个共分离标记的侧翼序列,延长后的标记序列长度在815-2661bp之间,并成功转化了5个SCAR标记,其中2个为共显性标记。继续在2650株的不育大群体中采用改良的混合样品作图法和侧翼分子标记分析法对BnMs2进行精细定位,将8个在小群体中共分离的标记定位在BnMs2两侧1.0cM区域内,其中最近标记Lm10和Lm11距目标基因分别只有0.038cM和0.075cM。
4.通过图谱定位将与BnMs2连锁的4个标记整合到2个DH群体(Tapidor×Ningyou7和Quantum×No.2127-17)的连锁群上,成功将BnMs2定位到甘蓝型油菜C基因组N16染色体上。并在TN群体的N16连锁群上找到1个共显性的SSR标记Na12A05。
5.延伸后的标记序列与拟南芥进行同源性比较,在拟南芥第1条染色体下方找到一个包含121个拟南芥基因的共线性区域。基于拟南芥基因组信息及网上大量芸苔属EST、GSS、BAC序列信息,开发了1个ACGM标记B14,同时Lm10经1个白菜BAC为桥梁被定位在拟南芥基因Atlg69510和Atlg69520之间。B14和Lm10将BnMs2在拟南芥上的候选基因缩小到12个。由于BnMs2和BnMs1功能相同,都可以恢复S45A和S4516A不育株的育性,而本定位研究筛选出的12个候选基因又包括BnMs1的2个候选基因,因此BnMs2基因也必在BnMs1的2个候选基因中,将这2个BnMs2的候选基因命名为BnG1和BnG2。
6.对2个候选基因BnG1、BnG2分别构建转基因表达载体p2301G1和p2301G2进行互补测验,以S4516AB为外植体通过农杆菌介导的遗传转化均获得了转基因阳性株。在7株BnG1的转化株中,S4516A没有能恢复育性；在3株BnG2的转化株中,也没有发现S4516A育性恢复。
7.根据BnMs1和BnG2基因cDNA5'端的保守序列构建反义表达载体p2301AntiG2,通过转基因对Westar中的BnMs1和BnG2进行反义抑制,获得了20株阳性株,对其中12株在花期进行育性检测没有发现预期的不育表型。
8.构建候选基因BnG2启动子的GUS融合基因表达载体p121P2GUS,对目标基因启动子进行组织特异性表达分析。经对11株阳性株的GUS染色,成功地分离了BnG2启动子,结果显示该基因仅限在花药某一特定时期(花蕾1-3mm)表达,在其它组织器官中均无表达。
Rapeseed is one of the major oil crops in China and the utilization of heterosis in rapeseed is an efficient way to increase yield, to mitigate the contradiction between yield and quality, and to improve resistance/tolerance. In China the recessive genic male sterility system (GMS) has become one of the most important approaches in virtue of the characteristics of complete and stable sterility, rich sources of cytoplasm and affluent restorer resources. However, the production of F1 hybrid seed requires the removal of 50% of the fertile plants in the female line, which limits its wide application. S45AB is a recessive GMS two-type line in Brassica napus and genetic analysis indicated that two duplicate recessive genes controlled the male sterility in S45A (Pan et al.,1988). For S45B, the BnMsl locus is heterozygous and Bnms2 locus is homozygous. In this study, for the fine mapping and cloning of BnMs2, a near isogenic line S4516AB, derived from S45A×IL362-2-4, was constructed by crossing, selfing, allelism testing and repeated full-sib mating. The objectives of this research are:(1) to develop a set of markers tightly linked to BnMs2 for marker assisted selection (MAS) in breeding of RGMS lines; (2) to isolate the BnMs2 gene and analyse its promoter to better our understanding of the mechanism of male sterility. The main results are as follows:
1. The allelism analysis between S4516B and S45B indicated that the heterozygous locus of S4516B was not allelic to that of S45B. After assaying the segregation ratio of sterile plants and fertile plants in the NIL population of 262 individuals, S4516B was proved to be a single locus segregating line. Combining with the result of allelism study, the genotype of S4516A was Bnms1 Bnms1 Bnms2Bnms2 and that of S4516B was Bnms1Bnms1BnMs2Bnms2.
2. From survey of 1024 AFLP primer combinations,12 AFLP markers tightly linked to the BnMs2 gene were idendified by the method of AFLP and BSA. In the NIL population of 262 plants, BnMs2 was mapped in the interval of 3.5cM and 8 AFLP markers co-segregated with BnMs2.
3. Twelve AFLP markers were cloned and sequenced. By PCR walking we extended all the sequences of the eight co-segregation AFLP markers, the size range of which was 815-2661bp. Five SCAR markers were successfully obtained and two of them were co-dominant markers. Fine mapping was conducted in 2650 sterile plants with the methodology of pooled sampling strategy and flanking marker analysis. The eight AFLP markers and five SCAR markers were all mapped in a 1.0cM region around BnMs2 gene. Among these flanking markers of BnMs2 gene, AFLP markers Lm10 and Lm11 were the most closely linked ones, which were 0.038cM and 0.075cM from BnMs2 gene, respectively.
4. We integrated four BnMs2 linked AFLP or SCAR markers to two doubled-haploid (DH) populations (Tapidor×Ningyou7 and Quantum×No.2127-17), and BnMs2 was mapped on N16. On the N16 linkage map one co-dominant SSR marker Na12A05 was identified by analyzing the selfing progeny of S4516B and the NIL population of 262 plants.
5. BLAST searches were carried out against the Arabidopsis genome with the extended marker sequences and a collinear region comprising 121 Arabidopsis genes was identified. On the basis of Arabidopsis genome information and many Brassica EST, GSS and BAC sequences, one ACGM marker B14 was developed and Lm10 was mapped between two Arabidopsis genes i.e., Atlg69510 and Atlg69520. Thus marker B14 and Lm10 delimited a region only containing 12 Arabidopsis genes in which the homolog of BnMs2 might exist. As both of BnMs2 and BnMs1 can restore the sterility of S45A and S4516A, and the 2 candidate genes of BnMs1 was included in the 12 candidate genes of BnMs2, therefore, we conclude that BnMs2 candidates should be the homologs of the two candidates of BnMs1. The two candidates of BnMs2 were then designated as BnG1 and BnG2, respectively.
6. For the 2 candidate genes of BnG1 and BnG2, plant expression vectors p2301G1 and p2301G2 were constructed, respectively. Using S4516AB as explants, transgenic positive plants were obtained by Agrobacterium-mediated transformation. In the 7 BnG1 transgenic plants, no recovery of fertility was observed. In the 3 BnG2 transgenic plants, no plants showed recovery of fertility, as well.
7. Antisense expression vector p2301AntiG2 was constructed according to the 5' reserved cDNA sequence of BnMsl and BnG2. Using Westar(Brassica napus) as explants,20 positive transgenic plants were recovered,12 of which were assayed as fertile at flowering time. No expected phenotype was found.
8. Gus fusion gene drived by the putative BnG2 promoter was introduced into Westar and 11 positive plants were assayed for GUS activity at different developmental stages. Results indicated that the expression of GUS was restricted specifically to the developing anthers (flower buds of 2-3mm). No GUS expression was observed in vegetative organs of plants.
引文
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