甘蓝型油菜硼高效近等基因系的构建及硼高效基因BnBE2的定位
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摘要
硼是高等植物生长发育所必需的矿质元素。甘蓝型油菜是需硼较多的作物,对硼缺乏比较敏感。我国油菜主产区土壤普遍缺硼,缺硼导致油菜“花而不实”,大量减产。农业生产上主要依赖增施硼肥解决耕地缺硼问题,然而硼是不可再生的矿质资源。不同甘蓝型油菜品种的硼效率性状存在显著差异,而且受遗传控制,这为硼效率性状的遗传改良提供基础。我们课题组在筛选获得甘蓝型油菜硼效率种质资源的基础上,开展硼高效基因的初步定位工作,研究表明甘蓝型油菜硼效率受多个基因控制。为了精细定位和克隆甘蓝型油菜硼高效基因,本论文建立了营养液培养条件下甘蓝型油菜苗期硼效率的鉴定体系,以甘蓝型油菜苗期硼效率性状为基础,采用随机的分子标记辅助背景选择构建了硼高效近等基因系;并在此基础上开展了硼高效位点BnBE2的定位研究,主要获得以下结果:
1甘蓝型油菜苗期硼效率鉴定体系的建立
在不同硼水平的营养液培养条件下,调查不同基因型甘蓝型油菜苗期的生长势,结果表明在大量和微量元素分别采用优级纯和分析纯化学试剂、营养液母液和工作液均用去离子水配制(＞12 MΩ)时,在0.25μMB水平下不同基因型甘蓝型油菜的生长势表现出明显的差异,硼高效基因型生长较好,而硼低效基因型植株矮小且表现出明显的缺硼症状。该鉴定体系不但适合不同基因型甘蓝型油菜品种苗期硼效率的鉴定,也适合分离群体各单株硼效率差异的研究,其实验条件易控制,重复性好。
2甘蓝型油菜硼高效近等基因系的构建
在营养液培养条件下(0.25μMB),通过连续四代回交,且在每一回交世代的苗期进行硼效率鉴定,采用分子标记对回交世代中硼高效单株进行遗传背景辅助选择,构建了甘蓝型油菜硼高效近等基因系。并证明在回交过程中采用随机分子标记辅助背景选择的有效性和必要性,且适宜在回交早世代进行。
3硼高效基因的定位
3.1回交群体中硼效率性状的遗传分析
采用营养液培养(0.25μM B)对回交分离群体苗期硼效率性状进行调查,根据I_(18)株系的BC_4S_1,BC_5和BC_6三个回交分离群体中高、低效单株的分离比例,表明I_(18)株系的硼效率性状受单个孟德尔因子控制。该株系中目标位点纯合的单株与轮回亲本组成了一对硼高效近等基因系,可供开展硼高效基因的精细定位和克隆、以及硼高效位点生理功能等的研究。
3.2硼高效基因连锁标记的筛选与定位
在I_(18)株系的BC_5回交群体苗期硼效率鉴定的基础上,以极端硼效率表型分组构建了近等基因池,结合AFLP和SRAP分子标记技术,筛选到6个与目标基因连锁的分子标记。利用BC_6、TNDH和BQDH两个双单倍体遗传群体对连锁标记进行定位,结果表明目标基因位于甘蓝型油菜N14连锁群上,该硼高效位点被命名为BnBE2。对这些连锁的分子标记进行测序并与Brassica数据库中的序列进行BLAST分析,发现与连锁标记序列高度同源的克隆均来自甘蓝(Brassica oleracea),从侧面证明了本实验定位结果的可靠性。
3.3连锁的AFLP标记向SSCP标记的转化
将与目标基因连锁的AFLP标记S6M32的序列在Brassica数据库进行比对,发现它与甘蓝的一个BAC克隆BOHZW41高度同源,根据目标片段的侧翼序列设计引物,将该AFLP标记成功地转化为SSCP标记,证明了该转化标记方法的可行性。通过对该SSCP标记的片段测序并采用软件RNA structure 3.2对该SSCP标记的序列进行二级结构预测,发现两个亲本之间因为3个碱基的差异导致了构象的改变,并揭示了产生该SSCP标记的理论依据。
3.4利用比较作图发展连锁标记
甘蓝型油菜的基因组由A、C基因组组成,分别起源于白菜和甘蓝,且甘蓝与白菜基因组间序列具有高度同源性,将不同亲本构建的甘蓝型油菜遗传图谱A4上的3个SSR标记定位在C4上,且与目标基因BnBE2连锁。
4.硼高效基因BnBE2的生理功能
比较I_(18)株系的硼高效回交单株(BC_4S_1,BC_5)以及硼高效亲本青油10号和硼低效亲本Bakow在0.25μMB水平条件下硼吸收累积、生物学产量等性状,推测该株系硼高效QTL位点的硼高效生理基础可能为硼利用高效。
5.连锁标记与甘蓝型油菜硼效率的关联分析
通过分析目标基因BnBE2侧翼的连锁分子标记在不同硼效率甘蓝型油菜品种中的带型,发现该连锁标记的多态性与硼效率表型没有较好的相关性,进一步证明甘蓝型油菜硼效率是一个受多基因位点控制的复杂性状。
Boron(B) is an essential mineral nutrient for the growth and development in higher plants.Rapeseed(Brassica napus) is acutely sensitive to B deficiency.The main production area of rapeseed in China is deficient in soil available B.Boron deficiency causes the symptom of "flowering without seed setting",or even no seed harvest.At present,applying B fertilizer is the main method to settle this problem.However,B isn't a regenerated mineral resource.Considerable genotypic variations in response to B deficiency exist among rapeseed varieties.This raises the possibility of genetic improvement for the trait tolerant to low B stress.Based on the identification of B efficiency for some rapeseed cultivars with different genotype and QTL mapping for B efficiency,the B efficiency was proved to be controlled by quantitative trait loci in Brassica napus.To free mapping and clone these genes,this study was involved in developing a stable and available solution culture method to validate the B efficiency of Brassica napus at the seedling stage,constructing B-efficient near isogenic lines with random markers assisted background selection,mapping a B-efficient locus,named BnBE2.The main results were as follows:
1.Development a system for evaluating B efficiency at the seedling stage
The growth of several genotypes was investigated under different B levels with solution culture.The results showed that significant difference in the phenotype growth among the different genotypes was found at the B level of 0.25μM.Thus,the B concentration is defined as a critical B stress level at which B-efficient cultivars had normal growth without B-deficient symptom while B-inefficient cultivars showed typical signs of B deficiency.The system is suitable for B efficiency evaluation not only for different rapeseed cukivars but also for a genetic segregating population.Under the optimum experiment conditions the method has good repeatability.
2.Constructing B efficient near isogenic lines
Based on the identification for B efficiency of each BC generation at the critical B level(0.25μM B),B efficient NIL were developed with random marker assisted background selection.It was proved that random molecular marker assisted background selection is effective and necessary in an early backcrossing generation to accelerate the construction of NILs.
3.Mapping B-efficient gene,BnBE2
3.1 Genetic analysis for B efficiency in BC populations
According to the result of phenotypic investigation in three BC populations(BC_4S_1, BC_5,BC_6) under the low B level at the seedling stage,the ratio of B-efficient individuals to B-inefficient individuals was 1:1 in the BC_5 and BC_6 population and 3:1 in the BC_4S_1 population,respectively.These results indicated that the B efficiency in the BC line was considered as Mendel factor,controlled by a single locus.Thus the homozygous of the B-efficient BC plants and the recurrent parent would be considered to be NILs for B efficiency,which should be useful for further free mapping and cloning the B-efficient gene.
3.2 Identification of linked markers and mapping the B efficient gene Based on the B efficiency identification of BC_5 population,DNA bulks were constructed from ten extreme B-efficient and ten extreme B-inefficient plants, respectively.Six markers were identified linked to BnBE2 using bulked segregant analysis(BSA) in combination with molecular markers AFLP and SRAP techniques.The six markers were all mapped in a region of C4.All of the most homologous sequences were from the Brassica oleracea genome based on the analysis of sequence BLAST against the Brassica database,which was an indication of the correctness of the mapping.
3.3 Conversion of the linked AFLP marker into an SSCP marker
The sequence of the linked marker $6M32 was performed a BLAST against the Brassica database and found to be high homologous to a BAC clone BOHZW41 of Brassica oleracea(CC).Primer was designed based on the flanking sequence and the PCR product was resolved on an 8%nondenaturing polyacrylamide gel and an SSCP marker was successfully detected.The AFLP marker was converted into an SSCP marker successfully,which proved the feasibility of this method in converting markers.The secondary structure was predicted for the sequences of the two parents using the software of RNA structure 3.2.The sequence difference,including 3 base pairs difference between the two parents,revealed the reason producing the difference of conformation.
3.4 Development of linked markers by comparative mapping
The genome of Brassica napus(AACC) includes A and C genome,which originated from Brassica rapa and Brassica oleracea.A genome and C genome share orthologous regions in the genus Brassica.Therefore three SSR markers within A4 on other Brassica napus genetic map were mapped on the target region of BriBE2 on C4.
4.Physiological mechanism for the B efficient locus,BnBE2
The result obtained from study on uptake,accumulation and utilization of B showed that the B-efficient NIL has lower B content and higher B utilization efficiency under the low B condition as compared with the B-inefficient recurrent parent,and the B efficiency is attributed to the higher B utilization efficiency or less demand for B,and this locus is different from BE1 and thus named as BnBE2.
5.Association analysis of molecular markers flanked to BnBE2 for several lines of Brassica napus with different B efficiency
An analysis of the flanking markers linked to BnBE2 in different B efficiency rapeseed cultivars was illustrated and the result showed that there is poor correlation between the flanking markers polymorphism bands and B efficiency,which indicates that the B efficiency is a complex trait controlled by multiple loci in Brassica napus.
1甘蓝型油菜苗期硼效率鉴定体系的建立
在不同硼水平的营养液培养条件下,调查不同基因型甘蓝型油菜苗期的生长势,结果表明在大量和微量元素分别采用优级纯和分析纯化学试剂、营养液母液和工作液均用去离子水配制(＞12 MΩ)时,在0.25μMB水平下不同基因型甘蓝型油菜的生长势表现出明显的差异,硼高效基因型生长较好,而硼低效基因型植株矮小且表现出明显的缺硼症状。该鉴定体系不但适合不同基因型甘蓝型油菜品种苗期硼效率的鉴定,也适合分离群体各单株硼效率差异的研究,其实验条件易控制,重复性好。
2甘蓝型油菜硼高效近等基因系的构建
在营养液培养条件下(0.25μMB),通过连续四代回交,且在每一回交世代的苗期进行硼效率鉴定,采用分子标记对回交世代中硼高效单株进行遗传背景辅助选择,构建了甘蓝型油菜硼高效近等基因系。并证明在回交过程中采用随机分子标记辅助背景选择的有效性和必要性,且适宜在回交早世代进行。
3硼高效基因的定位
3.1回交群体中硼效率性状的遗传分析
采用营养液培养(0.25μM B)对回交分离群体苗期硼效率性状进行调查,根据I_(18)株系的BC_4S_1,BC_5和BC_6三个回交分离群体中高、低效单株的分离比例,表明I_(18)株系的硼效率性状受单个孟德尔因子控制。该株系中目标位点纯合的单株与轮回亲本组成了一对硼高效近等基因系,可供开展硼高效基因的精细定位和克隆、以及硼高效位点生理功能等的研究。
3.2硼高效基因连锁标记的筛选与定位
在I_(18)株系的BC_5回交群体苗期硼效率鉴定的基础上,以极端硼效率表型分组构建了近等基因池,结合AFLP和SRAP分子标记技术,筛选到6个与目标基因连锁的分子标记。利用BC_6、TNDH和BQDH两个双单倍体遗传群体对连锁标记进行定位,结果表明目标基因位于甘蓝型油菜N14连锁群上,该硼高效位点被命名为BnBE2。对这些连锁的分子标记进行测序并与Brassica数据库中的序列进行BLAST分析,发现与连锁标记序列高度同源的克隆均来自甘蓝(Brassica oleracea),从侧面证明了本实验定位结果的可靠性。
3.3连锁的AFLP标记向SSCP标记的转化
将与目标基因连锁的AFLP标记S6M32的序列在Brassica数据库进行比对,发现它与甘蓝的一个BAC克隆BOHZW41高度同源,根据目标片段的侧翼序列设计引物,将该AFLP标记成功地转化为SSCP标记,证明了该转化标记方法的可行性。通过对该SSCP标记的片段测序并采用软件RNA structure 3.2对该SSCP标记的序列进行二级结构预测,发现两个亲本之间因为3个碱基的差异导致了构象的改变,并揭示了产生该SSCP标记的理论依据。
3.4利用比较作图发展连锁标记
甘蓝型油菜的基因组由A、C基因组组成,分别起源于白菜和甘蓝,且甘蓝与白菜基因组间序列具有高度同源性,将不同亲本构建的甘蓝型油菜遗传图谱A4上的3个SSR标记定位在C4上,且与目标基因BnBE2连锁。
4.硼高效基因BnBE2的生理功能
比较I_(18)株系的硼高效回交单株(BC_4S_1,BC_5)以及硼高效亲本青油10号和硼低效亲本Bakow在0.25μMB水平条件下硼吸收累积、生物学产量等性状,推测该株系硼高效QTL位点的硼高效生理基础可能为硼利用高效。
5.连锁标记与甘蓝型油菜硼效率的关联分析
通过分析目标基因BnBE2侧翼的连锁分子标记在不同硼效率甘蓝型油菜品种中的带型,发现该连锁标记的多态性与硼效率表型没有较好的相关性,进一步证明甘蓝型油菜硼效率是一个受多基因位点控制的复杂性状。
Boron(B) is an essential mineral nutrient for the growth and development in higher plants.Rapeseed(Brassica napus) is acutely sensitive to B deficiency.The main production area of rapeseed in China is deficient in soil available B.Boron deficiency causes the symptom of "flowering without seed setting",or even no seed harvest.At present,applying B fertilizer is the main method to settle this problem.However,B isn't a regenerated mineral resource.Considerable genotypic variations in response to B deficiency exist among rapeseed varieties.This raises the possibility of genetic improvement for the trait tolerant to low B stress.Based on the identification of B efficiency for some rapeseed cultivars with different genotype and QTL mapping for B efficiency,the B efficiency was proved to be controlled by quantitative trait loci in Brassica napus.To free mapping and clone these genes,this study was involved in developing a stable and available solution culture method to validate the B efficiency of Brassica napus at the seedling stage,constructing B-efficient near isogenic lines with random markers assisted background selection,mapping a B-efficient locus,named BnBE2.The main results were as follows:
1.Development a system for evaluating B efficiency at the seedling stage
The growth of several genotypes was investigated under different B levels with solution culture.The results showed that significant difference in the phenotype growth among the different genotypes was found at the B level of 0.25μM.Thus,the B concentration is defined as a critical B stress level at which B-efficient cultivars had normal growth without B-deficient symptom while B-inefficient cultivars showed typical signs of B deficiency.The system is suitable for B efficiency evaluation not only for different rapeseed cukivars but also for a genetic segregating population.Under the optimum experiment conditions the method has good repeatability.
2.Constructing B efficient near isogenic lines
Based on the identification for B efficiency of each BC generation at the critical B level(0.25μM B),B efficient NIL were developed with random marker assisted background selection.It was proved that random molecular marker assisted background selection is effective and necessary in an early backcrossing generation to accelerate the construction of NILs.
3.Mapping B-efficient gene,BnBE2
3.1 Genetic analysis for B efficiency in BC populations
According to the result of phenotypic investigation in three BC populations(BC_4S_1, BC_5,BC_6) under the low B level at the seedling stage,the ratio of B-efficient individuals to B-inefficient individuals was 1:1 in the BC_5 and BC_6 population and 3:1 in the BC_4S_1 population,respectively.These results indicated that the B efficiency in the BC line was considered as Mendel factor,controlled by a single locus.Thus the homozygous of the B-efficient BC plants and the recurrent parent would be considered to be NILs for B efficiency,which should be useful for further free mapping and cloning the B-efficient gene.
3.2 Identification of linked markers and mapping the B efficient gene Based on the B efficiency identification of BC_5 population,DNA bulks were constructed from ten extreme B-efficient and ten extreme B-inefficient plants, respectively.Six markers were identified linked to BnBE2 using bulked segregant analysis(BSA) in combination with molecular markers AFLP and SRAP techniques.The six markers were all mapped in a region of C4.All of the most homologous sequences were from the Brassica oleracea genome based on the analysis of sequence BLAST against the Brassica database,which was an indication of the correctness of the mapping.
3.3 Conversion of the linked AFLP marker into an SSCP marker
The sequence of the linked marker $6M32 was performed a BLAST against the Brassica database and found to be high homologous to a BAC clone BOHZW41 of Brassica oleracea(CC).Primer was designed based on the flanking sequence and the PCR product was resolved on an 8%nondenaturing polyacrylamide gel and an SSCP marker was successfully detected.The AFLP marker was converted into an SSCP marker successfully,which proved the feasibility of this method in converting markers.The secondary structure was predicted for the sequences of the two parents using the software of RNA structure 3.2.The sequence difference,including 3 base pairs difference between the two parents,revealed the reason producing the difference of conformation.
3.4 Development of linked markers by comparative mapping
The genome of Brassica napus(AACC) includes A and C genome,which originated from Brassica rapa and Brassica oleracea.A genome and C genome share orthologous regions in the genus Brassica.Therefore three SSR markers within A4 on other Brassica napus genetic map were mapped on the target region of BriBE2 on C4.
4.Physiological mechanism for the B efficient locus,BnBE2
The result obtained from study on uptake,accumulation and utilization of B showed that the B-efficient NIL has lower B content and higher B utilization efficiency under the low B condition as compared with the B-inefficient recurrent parent,and the B efficiency is attributed to the higher B utilization efficiency or less demand for B,and this locus is different from BE1 and thus named as BnBE2.
5.Association analysis of molecular markers flanked to BnBE2 for several lines of Brassica napus with different B efficiency
An analysis of the flanking markers linked to BnBE2 in different B efficiency rapeseed cultivars was illustrated and the result showed that there is poor correlation between the flanking markers polymorphism bands and B efficiency,which indicates that the B efficiency is a complex trait controlled by multiple loci in Brassica napus.
引文
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