甜瓜属Cucumis×hytivus Chen and Kirkbride种间杂交的分子验证和细胞器基因的遗传分析
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摘要
甜瓜属Cucumis X hytivus Chen and Kirkbride (2n=38)是陈劲枫通过远缘杂交和胚拯救并经体细胞无性系变异染色体数目加倍而育成的异源四倍体新种,该物种的合成对改善黄瓜种质资源,将近缘野生种的有用性状转移到栽培种黄瓜具有重要意义。
为了在分子水平验证Cucumis X hytivus Chen and Kirkbride起源于栽培种黄瓜‘北京截头’与甜瓜属野生种之间的种间远缘杂交并且明确经过5代繁育后是否仍然保持杂种特性,本研究以异源四倍体新种经过5代连续自交的后代S5和栽培种黄瓜‘北京截头’(Cucumis sativus L.'Beijingjietou',2n=14)及母本甜瓜属野生种(Cucumis hystrix Chakr..2n=24)为材料,以6-磷酸葡萄糖酸脱氢酶基因(6-phosphogluconate dehydrogenase gene,6PGDH)为对象进行研究,结果如下：
1.黄瓜6-磷酸葡萄糖酸脱氢酶基因(6-phosphogluconate dehydrogenase gene,6PGDH)的克隆及分析
根据6PGDH保守氨基酸序列设计简并引物,应用RT-PCR技术从黄瓜叶片中获得了640bp的特异片段,以该序列在EST数据库进行同源检索筛选,显示甜瓜EST序列AM715537.2与之高度同源,据此设计引物经RT-PCR扩增、分子克隆和序列拼接,获得了黄瓜6-磷酸葡萄糖酸脱氢酶基因全长序列,命名为Cs6PGDH。序列分析表明该基因全长1,829bp,其中开放读码框(ORF)长1,488bp编码495个氨基酸组成的多肽,编码区内无内含子存在,5’,3’端非翻译区长度各为70bp和271bp。Blast同源性分析表明该基因编码的氨基酸序列与拟南芥、大豆、水稻、玉米、菠菜等物种6PGDH的氨基酸序列有74%以上的同源性,该基因在GenBank登录号为FJ610345。运用半定量RT-PCR技术对6PGDH基因的转录水平进行分析,结果表明该基因在叶、根、茎中均有表达,高温胁迫下的表达量高于常温对照,说明6PGDH基因与热胁迫相关。
2.利用6PGDH基因进行杂交种的分子验证
根据上述黄瓜6PGDH基因序列(GenBank登录号：FJ610345)设计引物,扩增了异源四倍体新种S5、甜瓜属野生种和‘北京截头’的6PGDH基因片段,测序结果表明3个种的6PGDH同源一致性高达99.73%,片断长度分别为1,718bp,1718bp和1,720bp,均包含1,488bp编码495个氨基酸的完整开放阅读框和68-70bp的5’非编码区,162bp的3’非翻译区末端。利用DNAMAN软件检测到异源四倍体新种与野生种和栽培黄瓜‘北京截头’的氨基酸多态性差异位点有4个,其中新种有3个氨基酸位点来自野生种,另1个来从‘北京截头’；碱基多态性差异位点有24个,其中新种有11个位点碱基来自‘北京截头’,另有10个来自野生种,有2个位点同时含有双亲碱基,只有1个位点碱基发生转换突变而与双亲都不相同。
氨基酸和碱基多态性差异位点的遗传分布符合孟德尔遗传学定律,在分子水平证实了异源四倍体新种起源于甜瓜属野生种和栽培黄瓜‘北京截头’的种间杂交,并且在经过5代繁育后仍然保持杂交种特性。
为研究甜瓜属hystrix×sativis种间杂交中细胞器(线粒体、叶绿体)基因的遗传规律,对甜瓜属人工合成的异源四倍体新种(Cucumis×hytivus Chen and Kirkbride)的自交后代S5及其杂交母本甜瓜属野生种(Cucumis hystrix Chakr.)和杂交父本栽培种黄瓜‘北京截头’(Cucumis sativus L.)线粒体基因组中apocytochrome b(cob),NADH dehydrogenase subunit1(NAD1),NADH dehydrogenase subunit7(NAD7)基因序列,叶绿体基因组中matK-trnK和rbcL-accD区域的DNA序列进行了测序与比较分析,结果如下：
3.线粒体基因的遗传分析
在线粒体基因组中,3个种cob基因的909bp长度片断序列中只存在1个多态性碱基位点,新种该位点碱基类型与父本‘北京截头’相同而不同于母本野生种；在长度为943bp的NAD1内含子中有7个多态性碱基位点,其中有6个位点的碱基类型新种与父本‘北京截头’相同而不同于母本野生种,另1个位点的碱基类型与双亲都不相同；在长度为880bp的NAD7内含子序列中存在17个多态性碱基位点,其中有14个位点新种的碱基类型与父本‘北京截头’相同而不同于母本野生种,另3个位点碱基类型与双亲都不相同,这可能主要是由于DNA随机突变造成的。上述结果表明甜瓜属hystrix×sativus种间杂交中线粒体基因主要表现为父系遗传。
4.叶绿体基因的遗传分析
在叶绿体基因组中,3个种在长度为2.036bp Matk-trnK区域中存在着18个多态性碱基位点,其中有16个多态性位点的碱基类型新种与母本野生种相同,只有2个位点的碱基类型与父本‘北京截头’相同；在长度为945bp的rbcL-accD区域中存在着19个多态性位点,其中有18个多态性位点的碱基类型新种与母本野生种相同,只有1个多态性位点碱基类型与父本‘北京截头’相同。这表明甜瓜属hystrix×sativus种间杂交叶绿体基因主要表现为母系遗传。
Cucumis X hytivus Chen and Kirkbride (2n=38) of genus Cucumis was created by Jin-feng Chen via interspecific crossing and embryo rescue, chromosome numbers were doubled through somaclonal variation. This new species was very useful in broadening the germplasm base of cultivated cucumber and transferring some valuable traits from the wild cucumis species to cultivated cucumber.
In order to verify in molecular level that Cucumis X hytivus Chen and Kirkbride was originated from interspecific crossing between cultivated cucumber'Beijingjietou'and the wild Cucumis species and whether it still maintained hybridization trait after five generations breeding, in this study we selected cucumber cultivar'Beijingjietou'(C.sativus L.), the wild Cucumis species (C. hystrix Chakr.) and the S5progenies of C. hytivus Chen and Kirkbride.) as materials to investigate, the research results were as follow.
1. The cloning and the analysis of6-phosphogluconate dehydrogenase gene (6PGDH) from cucumber
Degenerated primers were designed according on conserved amino acids region of6-phosphogluconate dehydrogenase gene (6PGDH),a640bp in length cDNA fragment was amplified from Cucumis sativus'Beijingjietou'leaves through RT-PCR method, using fragment sequence as a querying probe, one highly homologous melon EST sequence AM715537.2was obtained from EST database. The full-length of cucumber6-phosphogluconate dehydrogenase gene(designated as Cs6PGDH) was obtained through RT-PCR method with primers designed basing on EST sequence and sequence assembling. Sequence analysis showed that Cs6PGDH (GenBank accession number was FJ610345) was1,829bp, which encompassed a1,488bp open reading frame (ORF) encoding495amino acid residues, no intron existed in ORF region and non-translation of5'and3'-terminal region were70bp,271bp, respectively. The results of Blast homologous analysis demonstrated that deduced amino acid sequence of Cs6PGDH had above74%identity with Arabidopsis thaliana, Glycine max, Oryza saliva, Zea mays, Spinacia oleracea et al. The expression of6PGDH was analyzed by semi-QRT-PCR indicating that the gene had expression in leaf, root and stem, the amount of expression was higher in heat stress than normal temperature, the result showed that expression of6PGDH was related to heat stress.
2. Confirmation of the interspecific hybridization trait by nuclear gene6PGDH
Primers were designed based on the6PGDH gene of cucumber (GenBank accession number was FJ610345), fragments from allotetraploid, the wild Cucumis species and cucumber'Beijingjietou'were obtained in PCR reaction, respectively and sequences alignment were analyzed by DNAMAN software. Four polymorphic amino acid sites were detected among three species, the allotetraploid possessed three amino acids of polymor-phic sites from the wild Cucumis species, and one amino acid from cucumber 'Beijingjietou'; twenty-four polymorphic nucleotide sites were also detected in three species, the allotetraploid possessed eleven bases type of polymorphic nucleotide sites from cucumber'Beijingjietou', ten bases type of polymorphic nucleotide sites from the wild Cucumis, two additive nucleotide sites possessed both different nucleotides sites from cultivar'Beijingjietou'and the wild Cucumis species simultaneously, while one base type of nucleotide sites differ from both species.
The distribution of these polymorphic sites was congruence well with Mendel's genetics law of inheritance. Therefore it was proven at molecular level that this synthetic allotetraploid species was the hybrid originated from the wild Cucumis species and cucumber'Beijingjietou' interspecific crossing and still maintained hybridization trait after five generations breeding.
To investigate the inheritance of organellar(mitochondrial and chloroplast) gene in Genus Cucumis, the Nad7, Nad1and cob gene fragments of mitochondrial DNA and the Matk-trnK and rbcL-accD gene region fragments of chloroplast DNA were amplified and sequenced from the S5progenies of the allotetraploid(Cucumis hytivus Chen and Kirkbride.), the wild Cucumis species(C. hystrix Chakr.) the maternal parents and cultivated cucumber(C. sativus L.'Beijingjietou') the paternal parents, respectively.
3. The inheritance of mitochondrial(mt) gene
Polymorphic Loci in Nad7, Nad1and cob mt gene were detected among three species by DNA sequencing and comparison, the results showed that only one polymorphic locus existed in909bp length sequence of cob gene, the base type of polymorphic locus in allotetraploid was identical to cucumber'Beijingjietou'the paternal parents but different from the wild Cucumis species the maternal parents; seven polymorphic loci existed in943bp length intron sequence of NAD1gene, among them six base types of polymorphic loci in allotetraploid were identical to cucumber 'Beijingjietou' but different from the wild Cucumis species, one base type of polymorphic locus was different from both parents; seventeen polymorphic loci were detected in880bp length intron sequence of NAD7gene, among them fourteen base types of allotetraploid were identical to cucumber 'Beijingjietou' but different from the wild Cucumis species, three base types of polymorphic loci were different from both parents.
This result revealed that mt genes was dominantly paternal transmission in hystrixxsativus interspecific crossing.
4. The inheritance of chloroplast(cp) gene
Polymorphic Loci in Matk-trnK and rbcL-accD gene region of cp genomes were also detected among three species by DNA sequencing and comparison, the results showed that eighteen polymorphic loci existed in2,036bp length sequence of Matk-trnK region, among them the base types of sixteen loci in allotetraploid were identical to wild Cucumis species the maternal parents, only two loci's base type in allotetraploid were identical to the cucumber'Beijingjietou' the paternal parents; nineteen polymorphic loci existed in945bp length sequence of rbcL-accD region, among them the base types of eighteen Loci in allotetraploid were identical to wild Cucumis species only one base type of polymorphic locus was identical to the cucumber'Beijingjietou'.
This study revealed that cp genes were dominantly maternal transmission in hystrixxsativus interspecific crossing.
为了在分子水平验证Cucumis X hytivus Chen and Kirkbride起源于栽培种黄瓜‘北京截头’与甜瓜属野生种之间的种间远缘杂交并且明确经过5代繁育后是否仍然保持杂种特性,本研究以异源四倍体新种经过5代连续自交的后代S5和栽培种黄瓜‘北京截头’(Cucumis sativus L.'Beijingjietou',2n=14)及母本甜瓜属野生种(Cucumis hystrix Chakr..2n=24)为材料,以6-磷酸葡萄糖酸脱氢酶基因(6-phosphogluconate dehydrogenase gene,6PGDH)为对象进行研究,结果如下：
1.黄瓜6-磷酸葡萄糖酸脱氢酶基因(6-phosphogluconate dehydrogenase gene,6PGDH)的克隆及分析
根据6PGDH保守氨基酸序列设计简并引物,应用RT-PCR技术从黄瓜叶片中获得了640bp的特异片段,以该序列在EST数据库进行同源检索筛选,显示甜瓜EST序列AM715537.2与之高度同源,据此设计引物经RT-PCR扩增、分子克隆和序列拼接,获得了黄瓜6-磷酸葡萄糖酸脱氢酶基因全长序列,命名为Cs6PGDH。序列分析表明该基因全长1,829bp,其中开放读码框(ORF)长1,488bp编码495个氨基酸组成的多肽,编码区内无内含子存在,5’,3’端非翻译区长度各为70bp和271bp。Blast同源性分析表明该基因编码的氨基酸序列与拟南芥、大豆、水稻、玉米、菠菜等物种6PGDH的氨基酸序列有74%以上的同源性,该基因在GenBank登录号为FJ610345。运用半定量RT-PCR技术对6PGDH基因的转录水平进行分析,结果表明该基因在叶、根、茎中均有表达,高温胁迫下的表达量高于常温对照,说明6PGDH基因与热胁迫相关。
2.利用6PGDH基因进行杂交种的分子验证
根据上述黄瓜6PGDH基因序列(GenBank登录号：FJ610345)设计引物,扩增了异源四倍体新种S5、甜瓜属野生种和‘北京截头’的6PGDH基因片段,测序结果表明3个种的6PGDH同源一致性高达99.73%,片断长度分别为1,718bp,1718bp和1,720bp,均包含1,488bp编码495个氨基酸的完整开放阅读框和68-70bp的5’非编码区,162bp的3’非翻译区末端。利用DNAMAN软件检测到异源四倍体新种与野生种和栽培黄瓜‘北京截头’的氨基酸多态性差异位点有4个,其中新种有3个氨基酸位点来自野生种,另1个来从‘北京截头’；碱基多态性差异位点有24个,其中新种有11个位点碱基来自‘北京截头’,另有10个来自野生种,有2个位点同时含有双亲碱基,只有1个位点碱基发生转换突变而与双亲都不相同。
氨基酸和碱基多态性差异位点的遗传分布符合孟德尔遗传学定律,在分子水平证实了异源四倍体新种起源于甜瓜属野生种和栽培黄瓜‘北京截头’的种间杂交,并且在经过5代繁育后仍然保持杂交种特性。
为研究甜瓜属hystrix×sativis种间杂交中细胞器(线粒体、叶绿体)基因的遗传规律,对甜瓜属人工合成的异源四倍体新种(Cucumis×hytivus Chen and Kirkbride)的自交后代S5及其杂交母本甜瓜属野生种(Cucumis hystrix Chakr.)和杂交父本栽培种黄瓜‘北京截头’(Cucumis sativus L.)线粒体基因组中apocytochrome b(cob),NADH dehydrogenase subunit1(NAD1),NADH dehydrogenase subunit7(NAD7)基因序列,叶绿体基因组中matK-trnK和rbcL-accD区域的DNA序列进行了测序与比较分析,结果如下：
3.线粒体基因的遗传分析
在线粒体基因组中,3个种cob基因的909bp长度片断序列中只存在1个多态性碱基位点,新种该位点碱基类型与父本‘北京截头’相同而不同于母本野生种；在长度为943bp的NAD1内含子中有7个多态性碱基位点,其中有6个位点的碱基类型新种与父本‘北京截头’相同而不同于母本野生种,另1个位点的碱基类型与双亲都不相同；在长度为880bp的NAD7内含子序列中存在17个多态性碱基位点,其中有14个位点新种的碱基类型与父本‘北京截头’相同而不同于母本野生种,另3个位点碱基类型与双亲都不相同,这可能主要是由于DNA随机突变造成的。上述结果表明甜瓜属hystrix×sativus种间杂交中线粒体基因主要表现为父系遗传。
4.叶绿体基因的遗传分析
在叶绿体基因组中,3个种在长度为2.036bp Matk-trnK区域中存在着18个多态性碱基位点,其中有16个多态性位点的碱基类型新种与母本野生种相同,只有2个位点的碱基类型与父本‘北京截头’相同；在长度为945bp的rbcL-accD区域中存在着19个多态性位点,其中有18个多态性位点的碱基类型新种与母本野生种相同,只有1个多态性位点碱基类型与父本‘北京截头’相同。这表明甜瓜属hystrix×sativus种间杂交叶绿体基因主要表现为母系遗传。
Cucumis X hytivus Chen and Kirkbride (2n=38) of genus Cucumis was created by Jin-feng Chen via interspecific crossing and embryo rescue, chromosome numbers were doubled through somaclonal variation. This new species was very useful in broadening the germplasm base of cultivated cucumber and transferring some valuable traits from the wild cucumis species to cultivated cucumber.
In order to verify in molecular level that Cucumis X hytivus Chen and Kirkbride was originated from interspecific crossing between cultivated cucumber'Beijingjietou'and the wild Cucumis species and whether it still maintained hybridization trait after five generations breeding, in this study we selected cucumber cultivar'Beijingjietou'(C.sativus L.), the wild Cucumis species (C. hystrix Chakr.) and the S5progenies of C. hytivus Chen and Kirkbride.) as materials to investigate, the research results were as follow.
1. The cloning and the analysis of6-phosphogluconate dehydrogenase gene (6PGDH) from cucumber
Degenerated primers were designed according on conserved amino acids region of6-phosphogluconate dehydrogenase gene (6PGDH),a640bp in length cDNA fragment was amplified from Cucumis sativus'Beijingjietou'leaves through RT-PCR method, using fragment sequence as a querying probe, one highly homologous melon EST sequence AM715537.2was obtained from EST database. The full-length of cucumber6-phosphogluconate dehydrogenase gene(designated as Cs6PGDH) was obtained through RT-PCR method with primers designed basing on EST sequence and sequence assembling. Sequence analysis showed that Cs6PGDH (GenBank accession number was FJ610345) was1,829bp, which encompassed a1,488bp open reading frame (ORF) encoding495amino acid residues, no intron existed in ORF region and non-translation of5'and3'-terminal region were70bp,271bp, respectively. The results of Blast homologous analysis demonstrated that deduced amino acid sequence of Cs6PGDH had above74%identity with Arabidopsis thaliana, Glycine max, Oryza saliva, Zea mays, Spinacia oleracea et al. The expression of6PGDH was analyzed by semi-QRT-PCR indicating that the gene had expression in leaf, root and stem, the amount of expression was higher in heat stress than normal temperature, the result showed that expression of6PGDH was related to heat stress.
2. Confirmation of the interspecific hybridization trait by nuclear gene6PGDH
Primers were designed based on the6PGDH gene of cucumber (GenBank accession number was FJ610345), fragments from allotetraploid, the wild Cucumis species and cucumber'Beijingjietou'were obtained in PCR reaction, respectively and sequences alignment were analyzed by DNAMAN software. Four polymorphic amino acid sites were detected among three species, the allotetraploid possessed three amino acids of polymor-phic sites from the wild Cucumis species, and one amino acid from cucumber 'Beijingjietou'; twenty-four polymorphic nucleotide sites were also detected in three species, the allotetraploid possessed eleven bases type of polymorphic nucleotide sites from cucumber'Beijingjietou', ten bases type of polymorphic nucleotide sites from the wild Cucumis, two additive nucleotide sites possessed both different nucleotides sites from cultivar'Beijingjietou'and the wild Cucumis species simultaneously, while one base type of nucleotide sites differ from both species.
The distribution of these polymorphic sites was congruence well with Mendel's genetics law of inheritance. Therefore it was proven at molecular level that this synthetic allotetraploid species was the hybrid originated from the wild Cucumis species and cucumber'Beijingjietou' interspecific crossing and still maintained hybridization trait after five generations breeding.
To investigate the inheritance of organellar(mitochondrial and chloroplast) gene in Genus Cucumis, the Nad7, Nad1and cob gene fragments of mitochondrial DNA and the Matk-trnK and rbcL-accD gene region fragments of chloroplast DNA were amplified and sequenced from the S5progenies of the allotetraploid(Cucumis hytivus Chen and Kirkbride.), the wild Cucumis species(C. hystrix Chakr.) the maternal parents and cultivated cucumber(C. sativus L.'Beijingjietou') the paternal parents, respectively.
3. The inheritance of mitochondrial(mt) gene
Polymorphic Loci in Nad7, Nad1and cob mt gene were detected among three species by DNA sequencing and comparison, the results showed that only one polymorphic locus existed in909bp length sequence of cob gene, the base type of polymorphic locus in allotetraploid was identical to cucumber'Beijingjietou'the paternal parents but different from the wild Cucumis species the maternal parents; seven polymorphic loci existed in943bp length intron sequence of NAD1gene, among them six base types of polymorphic loci in allotetraploid were identical to cucumber 'Beijingjietou' but different from the wild Cucumis species, one base type of polymorphic locus was different from both parents; seventeen polymorphic loci were detected in880bp length intron sequence of NAD7gene, among them fourteen base types of allotetraploid were identical to cucumber 'Beijingjietou' but different from the wild Cucumis species, three base types of polymorphic loci were different from both parents.
This result revealed that mt genes was dominantly paternal transmission in hystrixxsativus interspecific crossing.
4. The inheritance of chloroplast(cp) gene
Polymorphic Loci in Matk-trnK and rbcL-accD gene region of cp genomes were also detected among three species by DNA sequencing and comparison, the results showed that eighteen polymorphic loci existed in2,036bp length sequence of Matk-trnK region, among them the base types of sixteen loci in allotetraploid were identical to wild Cucumis species the maternal parents, only two loci's base type in allotetraploid were identical to the cucumber'Beijingjietou' the paternal parents; nineteen polymorphic loci existed in945bp length sequence of rbcL-accD region, among them the base types of eighteen Loci in allotetraploid were identical to wild Cucumis species only one base type of polymorphic locus was identical to the cucumber'Beijingjietou'.
This study revealed that cp genes were dominantly maternal transmission in hystrixxsativus interspecific crossing.
引文
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