甘蓝型油菜隐性细胞核雄性不育的基因定位
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摘要
油菜杂种优势利用是提高产量、缓解产量与品质矛盾、增强抗(耐)逆性的有效途径,而高效授粉系统的发掘与利用则是杂交种子生产的关键一环。隐性细胞核雄性不育因其具有不育性彻底且稳定、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,但由于得不到100%的不育系,在杂交制种时需人工拔除不育系株行中50%的可育株,限制了其广泛应用。陈凤祥等(1993)提出了隐性核不育的隐性上位互作模式,为隐性核不育的“三系”配套提供了理论依据,为核不育的广泛应用奠定了基础。
目前国内生产上应用的隐性核不育有双隐性核不育(S45A和117A等)和隐性上位互作核不育(9012A等)。双隐性核不育的育性受两对重叠隐性基因(ms_1和ms_2)控制,在两对基因均为隐性纯合时表现不育,只要有一个位点具有显性基因时即可恢复育性。隐性上位互作核不育的育性受两对重叠隐性不育基因(ms_3和ms_4)和一对隐性上位抑制基因(rfrf)互作控制,隐性上位基因纯合时对隐性不育基因起抑制作用,即三对基因都为隐性纯合时表现可育,与纯合不育系杂交能够产生全不育的群体,可用作保持系,从而解决了核不育制种中拔除50%可育株的难题。陈凤祥等(1998)还证明9012A中两对重叠隐性基因与S45A和117A中的两对重叠隐性基因非等位,且隐性上位基因与不同的重叠隐性不育基因之间的互作可能是非专一性的。为了进一步探索其不育机理,使隐性核不育得到更加广泛的应用,有必要对其核不育基因进行深入研究。
本研究以甘蓝型油菜双隐性核不育两型系117AB、S45AB和隐性上位互作核不育两型系9012AB为材料,采用AFLP技术寻找各个不育系中与可育基因位点紧密连锁的分子标记,构建目标基因区域的连锁图谱,并利用芸苔属作物与拟南芥的同源性和共线性开展目的基因的图位克隆工作,取得的实验结果和主要结论如下:
1.将双隐性核不育两型系117AB中的可育株(117B)与S45AB中的可育株(S45B)进行人工杂交,在F_1代套袋自交,收获10个F_1的自交后代,在F_2代对这些株系进行育性调查,其结果显示:有2个株系出现15:1的育性分离,8个株系出现3:1的育性分离,表明在这两个近等基因系中,117B和S45B的可育基因是不等位的。
2.在甘蓝型油菜双隐性核不育两型系117AB中构建了含152个单株的近等基因系群体,应用BSA法构建了可育基因池和不育基因池,在两个基因池之间一共筛选了512对AFLP引物,找到了6个与目标基因紧密连锁的分子标记,其中有3个标记(K1、K4和K5)位于可育基因位点的同一侧,与可育基因位点的遗传图距分别为4.0 cM、8.0 cM和8.0 cM,一个标记(Y1)位于可育基因位点的另一侧,与可育基因位点的遗传图距为8.0 cM,其它两个标记(K2和K3)在所用的群体中与目标基因共分离。对标记K1、K2、K3和Y1进行回收、克隆、测序,并根
The utilization of heterosis in rapeseed is an efficient pathway to increase yield, to solve the problem between yield and quality, and to improve resistance/tolerance. The study and utilization of high efficient pollination control system is the key step in hybrid seed production. The genic male sterility (GMS) systems had great potential for hybrid seed production in virtue of the characteristics of complete and stable male sterility, no negative cytoplasmic effects on plant growth, and ease of transference of the male sterility gene(s) to diverse genetic backgrounds. However, the production of F1 hybrid seed requires the removal of 50% of segregating fertile plants in the sterile female line which limits its utilization for production of hybrid seed. In 1993, Chen et al. reported a recessive genic male sterility (RGMS) line with an epistatic inhibitor gene, a 100% sterile population can be obtained via temporary maintainer, which made three-line systems available in hybrid seed production, and set up condition for the widely utilization of RGMS lines.Now, the RGMS lines used in China included several RGMS lines (eg. 117A and S45A) and several RGMS lines with an epistatic inhibitor gene (eg. 9012A). The 117AB and S45AB are two RGMS lines in which the sterility is controlled by two duplicate recessive genes named ms1 and ms2, located at two separate loci. One or two of the dominant genes existed at any loci can restore their fertility. The Brassica napus oilseed rape line 9012AB is a recessive epistatic GMS two-type line in which the sterility is controlled by two pairs of recessive duplicate sterile genes (ms3 and ms4) interacting with one pair of a recessive epistatic inhibitor gene (rf). Homozygosity at the rf locus (rfrf) inhibits the expression of the recessive male sterility trait in homozygous ms3 ms3 ms4 ms4 plants. In order to make the RGMS system a wider application, it is necessary to initiate further study on them. In this research, RGMS two-type lines 117AB, S45AB and 9012AB were used to identify AFLP markers linked to the Ms genes of RGMS lines117A and 9012A, construct genetic linkage map surrounding the Ms loci, and initiate comparative mapping between Arabidopsis and Brassica napus surrounding the Ms locus towards map-based cloning of the Ms gene of 9012A. Main results of the present study are as following:1. Crosses were made between the fertile plants of 117AB and S45AB, then selfed the F1 plants, and harvested seeds from 10 F1 individuals. Fertility of the 10 F1-derived F2 populations showed that the dominant Ms loci in 117A and S45A were non-allelic.2. Sterile bulk (BS) and fertile bulk (BF) DNA samples prepared from 9 male sterile and 9 male fertile plants of the two-type line 117AB were subjected to AFLP analysis. From the survey of 512 AFLP primer combinations, six AFLP fragments (Y1, K1, K2, K3, K4 and K5) were identified as being tightly linked to the Ms locus. The genetic distances between the markers and the Ms locus were all less than 8 cM, among which
two fragments, designated as K2 and K3, co-segregated with the target gene in the tested population. The markers detected could be valuable in marker-assisted selection in the utilization of RGMS in B. napus. By the way, we sequenced the AFLP fragments of Yl, Kl, K2, and K3, and compared the sequences with the Arabidopsis data, then primarily mapped the Ms locus to the Arabidopsis chromosome I . And fragment K2 was successfully converted into a SCAR marker.3. Sterile bulk (BS) and fertile bulk (BF) DNA samples prepared from 10 male sterile and 10 male fertile plants of the homozygous two-type line 9012AB were subjected to AFLP analysis. A total of 256 primer combinations were used and seven markers tightly linked to one fertile gene (Ms) in the RGMS line were identified. Among them, six fragments co-segratated with the target gene in the tested population, and the other one had a genetic distance of 5.4 cM. Five of them were successfully converted into dominant SCAR markers, which would facilitate the selection and breeding of temporary maintainers.4. A NIL population including 1506 individuals of 9012AB was developed for further study of the 5 SCAR markers, and a fine linkage map surrounding the Ms locus was constructed. Among them, two markers (SEP5 and SEP8) were located at one side of the target locus with a genetic distance of 0.2 cM and 0.1 cM respectively; and three markers (SEP 10, SEP7 and SEP4) were located at the other side with a genetic distance of 1.4 cM, 0.7 cM and 0.4 cM respectively.5. Isolated flanking sequences of the AFLP markers by PCR walking were submitted to NCBI for BLAST analysis. Information about Arabidopsis of each marker was collected and submitted to the Arabidopsis genome database (TAIL), and a linkage map in Arabidopsis chromosome V were made by 5 out of the 7 AFLP markers in the same order as in B. napus. Then, we could map the Ms locus in Arabidopsis between EP4 and EP8, which spanned a physical distance of about 2.1 Mb.
目前国内生产上应用的隐性核不育有双隐性核不育(S45A和117A等)和隐性上位互作核不育(9012A等)。双隐性核不育的育性受两对重叠隐性基因(ms_1和ms_2)控制,在两对基因均为隐性纯合时表现不育,只要有一个位点具有显性基因时即可恢复育性。隐性上位互作核不育的育性受两对重叠隐性不育基因(ms_3和ms_4)和一对隐性上位抑制基因(rfrf)互作控制,隐性上位基因纯合时对隐性不育基因起抑制作用,即三对基因都为隐性纯合时表现可育,与纯合不育系杂交能够产生全不育的群体,可用作保持系,从而解决了核不育制种中拔除50%可育株的难题。陈凤祥等(1998)还证明9012A中两对重叠隐性基因与S45A和117A中的两对重叠隐性基因非等位,且隐性上位基因与不同的重叠隐性不育基因之间的互作可能是非专一性的。为了进一步探索其不育机理,使隐性核不育得到更加广泛的应用,有必要对其核不育基因进行深入研究。
本研究以甘蓝型油菜双隐性核不育两型系117AB、S45AB和隐性上位互作核不育两型系9012AB为材料,采用AFLP技术寻找各个不育系中与可育基因位点紧密连锁的分子标记,构建目标基因区域的连锁图谱,并利用芸苔属作物与拟南芥的同源性和共线性开展目的基因的图位克隆工作,取得的实验结果和主要结论如下:
1.将双隐性核不育两型系117AB中的可育株(117B)与S45AB中的可育株(S45B)进行人工杂交,在F_1代套袋自交,收获10个F_1的自交后代,在F_2代对这些株系进行育性调查,其结果显示:有2个株系出现15:1的育性分离,8个株系出现3:1的育性分离,表明在这两个近等基因系中,117B和S45B的可育基因是不等位的。
2.在甘蓝型油菜双隐性核不育两型系117AB中构建了含152个单株的近等基因系群体,应用BSA法构建了可育基因池和不育基因池,在两个基因池之间一共筛选了512对AFLP引物,找到了6个与目标基因紧密连锁的分子标记,其中有3个标记(K1、K4和K5)位于可育基因位点的同一侧,与可育基因位点的遗传图距分别为4.0 cM、8.0 cM和8.0 cM,一个标记(Y1)位于可育基因位点的另一侧,与可育基因位点的遗传图距为8.0 cM,其它两个标记(K2和K3)在所用的群体中与目标基因共分离。对标记K1、K2、K3和Y1进行回收、克隆、测序,并根
The utilization of heterosis in rapeseed is an efficient pathway to increase yield, to solve the problem between yield and quality, and to improve resistance/tolerance. The study and utilization of high efficient pollination control system is the key step in hybrid seed production. The genic male sterility (GMS) systems had great potential for hybrid seed production in virtue of the characteristics of complete and stable male sterility, no negative cytoplasmic effects on plant growth, and ease of transference of the male sterility gene(s) to diverse genetic backgrounds. However, the production of F1 hybrid seed requires the removal of 50% of segregating fertile plants in the sterile female line which limits its utilization for production of hybrid seed. In 1993, Chen et al. reported a recessive genic male sterility (RGMS) line with an epistatic inhibitor gene, a 100% sterile population can be obtained via temporary maintainer, which made three-line systems available in hybrid seed production, and set up condition for the widely utilization of RGMS lines.Now, the RGMS lines used in China included several RGMS lines (eg. 117A and S45A) and several RGMS lines with an epistatic inhibitor gene (eg. 9012A). The 117AB and S45AB are two RGMS lines in which the sterility is controlled by two duplicate recessive genes named ms1 and ms2, located at two separate loci. One or two of the dominant genes existed at any loci can restore their fertility. The Brassica napus oilseed rape line 9012AB is a recessive epistatic GMS two-type line in which the sterility is controlled by two pairs of recessive duplicate sterile genes (ms3 and ms4) interacting with one pair of a recessive epistatic inhibitor gene (rf). Homozygosity at the rf locus (rfrf) inhibits the expression of the recessive male sterility trait in homozygous ms3 ms3 ms4 ms4 plants. In order to make the RGMS system a wider application, it is necessary to initiate further study on them. In this research, RGMS two-type lines 117AB, S45AB and 9012AB were used to identify AFLP markers linked to the Ms genes of RGMS lines117A and 9012A, construct genetic linkage map surrounding the Ms loci, and initiate comparative mapping between Arabidopsis and Brassica napus surrounding the Ms locus towards map-based cloning of the Ms gene of 9012A. Main results of the present study are as following:1. Crosses were made between the fertile plants of 117AB and S45AB, then selfed the F1 plants, and harvested seeds from 10 F1 individuals. Fertility of the 10 F1-derived F2 populations showed that the dominant Ms loci in 117A and S45A were non-allelic.2. Sterile bulk (BS) and fertile bulk (BF) DNA samples prepared from 9 male sterile and 9 male fertile plants of the two-type line 117AB were subjected to AFLP analysis. From the survey of 512 AFLP primer combinations, six AFLP fragments (Y1, K1, K2, K3, K4 and K5) were identified as being tightly linked to the Ms locus. The genetic distances between the markers and the Ms locus were all less than 8 cM, among which
two fragments, designated as K2 and K3, co-segregated with the target gene in the tested population. The markers detected could be valuable in marker-assisted selection in the utilization of RGMS in B. napus. By the way, we sequenced the AFLP fragments of Yl, Kl, K2, and K3, and compared the sequences with the Arabidopsis data, then primarily mapped the Ms locus to the Arabidopsis chromosome I . And fragment K2 was successfully converted into a SCAR marker.3. Sterile bulk (BS) and fertile bulk (BF) DNA samples prepared from 10 male sterile and 10 male fertile plants of the homozygous two-type line 9012AB were subjected to AFLP analysis. A total of 256 primer combinations were used and seven markers tightly linked to one fertile gene (Ms) in the RGMS line were identified. Among them, six fragments co-segratated with the target gene in the tested population, and the other one had a genetic distance of 5.4 cM. Five of them were successfully converted into dominant SCAR markers, which would facilitate the selection and breeding of temporary maintainers.4. A NIL population including 1506 individuals of 9012AB was developed for further study of the 5 SCAR markers, and a fine linkage map surrounding the Ms locus was constructed. Among them, two markers (SEP5 and SEP8) were located at one side of the target locus with a genetic distance of 0.2 cM and 0.1 cM respectively; and three markers (SEP 10, SEP7 and SEP4) were located at the other side with a genetic distance of 1.4 cM, 0.7 cM and 0.4 cM respectively.5. Isolated flanking sequences of the AFLP markers by PCR walking were submitted to NCBI for BLAST analysis. Information about Arabidopsis of each marker was collected and submitted to the Arabidopsis genome database (TAIL), and a linkage map in Arabidopsis chromosome V were made by 5 out of the 7 AFLP markers in the same order as in B. napus. Then, we could map the Ms locus in Arabidopsis between EP4 and EP8, which spanned a physical distance of about 2.1 Mb.
引文
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