摘要
甘蓝型油菜(Brassica napus L., 2n=38,AACC)是我国油菜的主要栽培品种,但其作为异源四倍体种起源发生的历史较短,遗传背景也较狭窄;加之我国甘蓝型油菜引种国外,因此其遗传基础单一的问题尤为突出。鉴于甘蓝型油菜在生产中的重要地位,如何利用现代生物技术将近缘种属中的优良性状(如黄籽性状、抗病基因)导入栽培油菜中去,拓宽甘蓝型油菜遗传背景是非常重要的。油菜黄籽性状具有较好的营养和加工品质,已受到油菜育种学家的重视,而甘蓝型油菜自然种质中不存在黄籽资源,目前甘蓝型油菜中的黄籽资源均来自芸薹属种间的基因流动,尚未见芸薹属以外黄籽种质资源转入的报道。
白芥(Sinapis alba L., 2n=24, SS)属于十字花科白芥属,和油菜亲缘关系非常近,且具有很多优良的农艺性状(如黄籽、抗裂角等),对十字花科多种病虫害具有较高的抗性,如抗病毒病、黑胫病、黑斑病和线虫病等。目前国内有关对白芥的利用报道的很少。本研究是在获得甘蓝型油菜和白芥体细胞杂种的基础上,将属间杂种不断与甘蓝型油菜进行连续多代回交及自交,通过基因组原位杂交分析(GISH),追踪外源染色体(白芥)在各个时期的行为。此外,我们首次将白芥的黄籽性状转入甘蓝型油菜,并对其进行了细胞学及分子生物学鉴定,这对于拓宽油菜的遗传背景以及品质改良和育种具有重要的应用价值。以白芥特异的重复序列为探针进行荧光原位杂交(FISH)检测,在黄籽株系D244-18中检测到白芥染色体的易位片段。
根据拟南芥等的类黄酮合成相关基因的保守序列设计27对兼并引物,共13个基因的16对引物在双亲之间扩增出多态性,其中TT3、TT19和TTG1各有2对引物均扩增出多态性,TT1、TT2、TT8、TT10、TT12、TT18、BAN、PAL、PAP和AHA10各有1对引物扩增出多态性,说明白芥的类黄酮合成基因与甘蓝型油菜存在显著差异。用这16对引物在黄籽后代中进行扩增,除TT2-2引物在黄籽后代中扩增出亲本白芥的特异带外,其它引物在黄籽后代中的扩增带型均与亲本扬油6号一致。而引物TT2-2的PCR扩增结果显示后代黄籽材料同时具有亲本白芥和甘蓝型油菜的特异条带,因此可以推断白芥的部分DNA序列通过易位进入了甘蓝型油菜。对TT2-2引物在黄籽后代D244-18中扩增的白芥特异条带1800bp左右进行克隆、测序。测序结果(1672bp)在NCBI Genbank中对测序结果进行比对(BLAST),结果显示:61-468、664 -1709两个区段的碱基与Brassica rapa BAC clone(LOCUS:AC189336,GI:199580018)同源性分别达到97%和99%,全序列一致性为82%;而469-663区段195个碱基序列在Genbank中无仍何匹配结果。根据测序结果设计引物Sa1,在白芥和黄籽后代中扩增出预期产物(1408bp),而亲本扬油6号的扩增产物比白芥和黄籽后代小200bp左右,进一步说明了黄籽后代中的该特异产物来自白芥。此外, Sa1引物在现有市场上通过种间杂交所获的3个黄籽品种(系)进行比较发现没有白芥所特有的条带,表明我们通过和白芥属杂交所获的黄籽材料与其他来源的黄籽材料存在差异。
The genetic basis of Brassica napus L. (2n=38, AACC), one of the most important oilseed crops worldwide, is quite narrow. The species originated in Europe and was introduced into China as an oil crop in the 1950s. In view of its important role in economic manufacture, how to exploit valuable characters like yellow-seed and disease resistant genes from related species into B. napus using modern biotechnology is in face of breeders. Because of better nutrition and processing quality, yellow-seeded trait is becoming an increasingly important objective of rapeseed breeding. None yellow-seed exists in natural B. napus, and even the existed yellow rapeseed is bred via interspecific hybridization of Brassia species, novel yellow-seeds introduced from the related genus has not been reported.
Sinapis alba L., a member of the Brassicaceae, possesses desirable agronomic characteristics such as yellow seed color, tolerance to drought stress, reduced pod shattering, resistance to virus diseases, blackleg disease, black spot and beet cyst nematodes. Somatic hybrids of B. napus and S. alba obtained by electrofusion have been described previously. For creation of high yielding rapeseed lines with improved disease resistance and seed quality, these hybrids were subsequently backcrossed with B. napus and self-pollinated to obtain a BC3F1 generation with valuable agronomic characteristics derived from S. alba. This material contains various interesting phenotypes that may be useful for rapeseed breeding.
The objective of this study was to carry out character observation and trace the chromosome behaviors through genomic in situ hybridization (GISH). On the other hand, we obtained yellow-seed variants in the BC3F1 progenies, which were characterized by both molecular and cytological analysis. Polymerase chain reaction (PCR) and fluorescence in situ hybridization (FISH) techniques were applied to demonstrate the presence of S. alba chromatin and putative S. alba genome introgressions in B. napus and S. alba interspecific progenies. In addition, we used minisatellite core sequence 33.6 as primer also obtained a 300bp band in D244 which was specific to S. alba. FISH using the PCR-derived fragments of DNA hybridized to mitotic metaphase chromosomes of D244, enabled dependable discrimination of S. alba chromatin introgression, showing a significant hybridization signal on one of the 38 chromosomes.
According to flavonoid biosynthetic genes of Arabidopsis thaliana, 27 primers were designed for PCR identification of S. alba, B. napus and intergeneric hybrid progenies, of which 16 primers derived from 13 genes amplified polymorphism between both parents, suggesting that significant deviation of flavonoid biosynthetic genes exists between B. napus and S. alba. Further identification of yellow seed progenies with above 16 primers indicated that TT2-2 yielded a S. alba-specific band (1800bp) in progeny lines, whereas amplified bands of progenies by other primers were consistent with‘Yangyou 6’. This result further confirmed that part of S. alba genes was transferred to B. napus through introgressions. A gene fragment of 1672bp in length results from the 1800bp band of D244-18, which was amplified by primer TT2-2 was sequenced. BLAST analysis indicated segment similarity of two base region 61-468 and 664-1709 to B. napus BAC clone (LOCUS: AC189336, GI: 199580018) was 97% and 99%, respectively. No matched data in genebank was found resemble to the 195 bases of region 469-663. Another primer Sa1, designed according to the sequencing result, obtained a prospective 1408bp fragment in S. alba and yellow seed lines, which was differ from the 1200bp band of‘Yangyou 6’, proving S. alba gene exists in yellow seed progenies. In addition, above S. alba specific gene fragment was absent in other three yellow seed varieties produced through interspecific hybridization, illuminating our yellow seed germplasm was different from some of other yellow seed resources tested.
引文
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