苹果三倍性杂交后代表型及主要性状遗传倾向研究
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摘要
我国苹果生产中的品种大多数为二倍体品种,但是为数不多的三倍体品种却表现优良。三倍体苹果已经成为苹果育种尤其是多倍体育种工作的重要一部分。目前已经通过杂交获得了一部分多倍体资源,但是对这些资源的研究较少,关于遗传变异的机理报道很少,尤其是通过杂交获得的三倍性苹果的遗传变异的分子机理至今报道甚少。本研究以二倍体苹果‘寒富’、‘富士’和‘金冠’为母本,与‘四倍体嘎拉’苹果为父本进行杂交获得的三倍性杂种后代为试材,在对其生物学性状进行调查研究的基础上,对其遗传变异的分子机理进行了探讨,同时对基因组DNA甲基化水平和甲基化模式变异的表观遗传进行了研究,结果如下：
1.‘寒富’与‘四倍体嘎拉’苹果的三倍性杂种后代的叶片、枝条和花性状变异幅度在26.42%～1.33%之间,花粉平均PE值变异幅度为4.46%。大多数性状平均值小于亲中值。果实性状的变异幅度在5.12%～27.82%之间,除平均单果重低于亲中值以外,果形指数、果实硬度、果实可溶性固形物含量均大于亲中值。这6份杂种后代的特征香气大多数受父母本的共同影响,果实香气倾向于父本遗传。果糖是影响这6份三倍性杂种后代的甜味感的最大因素,而苹果酸则是影响其酸味味感的最大因素。
2.‘寒富’与‘四倍体嘎拉’苹果的三倍性杂种后代经ISSR扩增得到的遗传多样性指数中扩增多态性比率(PPB)为60.00%、观察等位基因数(Na)为1.6000、有效等位基因数(Ne)为1.4204、Nei's基因多样性指数(功为0.2368、Shannon信息指数(Ⅰ)为0.3464,均小于AFLP扩增的遗传多样性指数中的相应数值66.54%、1.6654、1.421、0.2405和0.3570。根据ISSR和AFLP得到的UPGMA聚类图显示这7份三倍性杂种后代倾向于母本‘寒富’。在7份三倍性杂种后代中大部分位点来源于父母本,但也存在一定的缺失位点和特有位点。
3.‘富士’与‘四倍体嘎拉’苹果的三倍性杂种后代ISSR遗传多样性指数中多态性比例为77.42%。观察等位基因数为1.7742,有效等位基因数为1.4004,Nei's基因多样性指数为0.2355,Shannon信息指数为0.3572；‘金冠’与‘四倍体嘎拉’苹果的三倍性杂种后代的ISSR遗传多样性指数中多态性比例为36.55%。观察等位基因数为1.9655,有效等位基因数为1.5322,Nei's基因多样性指数为0.3092,Shannon信息指数为0.4641。在这80份三倍性杂种后代中,子代植株的ISSR条带与母本条带相同占多数,且存在一定的条带增加和缺失；具有母本遗传倾向。
4.‘寒富’与‘四倍体嘎拉’苹果的三倍性杂种后代的总甲基化率(含全甲基化和半甲基化)在14.74%～20.00%之间；全甲基化率在5.41%-10.38%之间,半甲基化率在5.85%～14.59%之间；与亲本相比总甲基化率和全甲基化率均呈高于母本,低于父本的趋势,半甲基化率则均高于亲本。‘富士’与‘四倍体嘎拉’苹果的三倍性杂种后代总甲基化率在14.65%～27.96%之间；全甲基化率在8.25%～17.40%之间；半甲基化率在4.19%-10.82%之间；与母本相比,总甲基化率和半甲基化率大多呈增加趋势,全甲基化率呈下降趋势；与父本相比,均呈下降趋势。‘金冠’与‘四倍体嘎拉’苹果的三倍性杂种后代的总甲基化率在10.74%～28.86%之间,全甲基化率在7.87%～24.22%之间,半甲基化率在2.10%-11.84%之间。与母本相比,总甲基化和全甲基化呈上升趋势,半甲基化呈下降趋势；与父本相比,均呈下降趋势。
5.在这三个杂交组合中DNA甲基化共分5种类型：单态性(A)、去甲基化(B)、过或超甲基化(C)、次甲基化(D)和中间型(E)。在苹果三倍性杂种的基因组加倍和重组过程中,发生了大量过或超甲基化和去甲基化变异,少量发生了次甲基化现象,极少部分的甲基化状态介于双亲之间。通过MSAP分析,共获得10条差异片段,有8条能够在苹果基因组中检测到,且同源性较高,这8条片段分别位于第1条、9条、10条、12条和16条染色体的叠连群上,但是具体功能没有描述。
At present in the apple production of China, diploid varieties play a major part, but a few triploid varieties have good performance. The triploid apple breeding has become an important part in fruit polyploid breeding. But there was very little research on that had obtained polyploidy resources by hybridization, and also very few reports of hybrid mutation mechanism. Especially, the molecular mechanism of genetic variation of triploid apple obtained by crossing has rarely been reported. In this study, molecular mechanism of genetic variation and epigenetic variation in genome including the status and patterns of DNA methylation based on the investigation of the botanical characters of triploid hybrid progenies of 'Hanfu'×'4xGala','Fuji'×'4xGala','Golden Delicious'×'4xGala' were studied, and the results are as follows:
1. The coefficient of variation of leaves, branches and flowers of triploid hybrids of 'Hanfu'×'4xGala' were among26.42%～1.33%and that of pollen PE was4.46%. The averages of most of the characters were lower than that of the mid-parents. Coefficient of varation of fruit characters were among5.12%～27.82%, and the same trend with the botanical characters. Characteristic aromas of6hybrids were effect by both parent, and the genetic trend of fruit aromas was paternal inheritance. The major effect factor on the sweetness of6triploid hybrids was fructose, and that of sour flavor was malic acid.
2. Genetic variation of7triploid hybrids o f'Hanfu'×'4xGala' was investigated based on ISSR and AFLP. The percentage of polymorphic bands (PPB) of ISSR was60.00%, observed number of alleles (Na) was1.6000, effective number of alleles (Ne) was1.4204, Nei's gene diversity (H) was0.2368and Shannon's Information index (I) was0.3464. And these were lower than that of AFLP. The UPMGA cluster showed that most of7triploid hybrids were tend to the female parent'Hanfu'based on ISSR and AFLP. The analysis indicated that ISSR and AFLP both can use to analyze the genetic diversity of apple triploid hybrids, and these triploid hybrids were maternal inheritance.
3. Genetic variation of29triploid hybrids of 'Fuji'×'4xGala' was investigated based on ISSR. The percentage of polymorphic bands (PPB) was77.42%, observed number of alleles (Na) was1.7742, effective number of alleles (Ne) was1.4004, Nei's gene diversity (H) was0.2355and Shannon's Information index (I) was0.3574. Genetic variation of61triploid hybrids of 'Golden Delicious'×'4xGala' were investigated based on ISSR. The percentage of polymorphic bands (PPB) was36.55%, observed number of alleles (Na) was1.9655, effective number of alleles (Ne) was1.5322, Nei's gene diversity (H) was0.3092 and Shannon's Information index (I) was0.4641. In the80triploid hybrid progenies, the bands of hybids were most come from female parent, and there was a certain increase and deletion in the strips, and these triploid hybrids were maternal inheritance.
4. The total methylation ratios of7triploid hybrids of 'Hanfu'×'4xGala' were among14.74%to20.00%, the full methylated ratios were among5.41%to10.38%, and the hemi-methylated ratios of that were among5.85%to14.59%. Compared with 'Hanfu', the total methylation ratios, the full-methylatied ratios and hemi-methylated ratios were mostly tended to increase. But all of that were tended to decrease compared with'4xGala'.There was no obvious trend with that of their parents. The total methylation ratios of29triploid hybrids of 'Fuji'×'4xGala' were among14.65%to27.96%, the full methylated ratios were among8.25%to17.40%, and the hemi-methylated ratios of that were among4.19%to10.82%. Compared with 'Fuji', the total methylation ratios and hemi-methylatied ratios were mostly tended to increase, but full methylated ratios was tended decrease. But all of that were tended to decrease compared with '4xGala'. The total methylation ratios of61triploid hybrids of 'Golden Delicious'×'4xGala' were among10.74%to28.86%, the full methylated ratios were among7.84%to24.22%, and the hemi-methylated ratios of that were among2.10%to11.84%. Compared with 'Golden Delicious', the total methylation ratios and the full-methylatied ratios were mostly tended to increase, but hemi-methylated ratios was tended decrease. But all of that were tended to decrease compared with '4xGala'.
5. There were five patterns of DNA methylatiion in these three hybridized combinations including monomophism (A), demthylation (B), hyermethylation (C), hypomethylation (D) and intermediate methylation (E).During the process of genome recombination and triploidization in apple triploid hybrids, a great number of hypermethylations and demethlation were occurred, a small amout of occurrence methane phenomenon and a very small part of the methylation status were between parents. In total10differential fragments were obtained by MSAP.8can be detected in the apple genome, and the high homology fragments were located in the contig of8,9,10,12and16chromosomes, but the specific function is not described.
1.‘寒富’与‘四倍体嘎拉’苹果的三倍性杂种后代的叶片、枝条和花性状变异幅度在26.42%～1.33%之间,花粉平均PE值变异幅度为4.46%。大多数性状平均值小于亲中值。果实性状的变异幅度在5.12%～27.82%之间,除平均单果重低于亲中值以外,果形指数、果实硬度、果实可溶性固形物含量均大于亲中值。这6份杂种后代的特征香气大多数受父母本的共同影响,果实香气倾向于父本遗传。果糖是影响这6份三倍性杂种后代的甜味感的最大因素,而苹果酸则是影响其酸味味感的最大因素。
2.‘寒富’与‘四倍体嘎拉’苹果的三倍性杂种后代经ISSR扩增得到的遗传多样性指数中扩增多态性比率(PPB)为60.00%、观察等位基因数(Na)为1.6000、有效等位基因数(Ne)为1.4204、Nei's基因多样性指数(功为0.2368、Shannon信息指数(Ⅰ)为0.3464,均小于AFLP扩增的遗传多样性指数中的相应数值66.54%、1.6654、1.421、0.2405和0.3570。根据ISSR和AFLP得到的UPGMA聚类图显示这7份三倍性杂种后代倾向于母本‘寒富’。在7份三倍性杂种后代中大部分位点来源于父母本,但也存在一定的缺失位点和特有位点。
3.‘富士’与‘四倍体嘎拉’苹果的三倍性杂种后代ISSR遗传多样性指数中多态性比例为77.42%。观察等位基因数为1.7742,有效等位基因数为1.4004,Nei's基因多样性指数为0.2355,Shannon信息指数为0.3572；‘金冠’与‘四倍体嘎拉’苹果的三倍性杂种后代的ISSR遗传多样性指数中多态性比例为36.55%。观察等位基因数为1.9655,有效等位基因数为1.5322,Nei's基因多样性指数为0.3092,Shannon信息指数为0.4641。在这80份三倍性杂种后代中,子代植株的ISSR条带与母本条带相同占多数,且存在一定的条带增加和缺失；具有母本遗传倾向。
4.‘寒富’与‘四倍体嘎拉’苹果的三倍性杂种后代的总甲基化率(含全甲基化和半甲基化)在14.74%～20.00%之间；全甲基化率在5.41%-10.38%之间,半甲基化率在5.85%～14.59%之间；与亲本相比总甲基化率和全甲基化率均呈高于母本,低于父本的趋势,半甲基化率则均高于亲本。‘富士’与‘四倍体嘎拉’苹果的三倍性杂种后代总甲基化率在14.65%～27.96%之间；全甲基化率在8.25%～17.40%之间；半甲基化率在4.19%-10.82%之间；与母本相比,总甲基化率和半甲基化率大多呈增加趋势,全甲基化率呈下降趋势；与父本相比,均呈下降趋势。‘金冠’与‘四倍体嘎拉’苹果的三倍性杂种后代的总甲基化率在10.74%～28.86%之间,全甲基化率在7.87%～24.22%之间,半甲基化率在2.10%-11.84%之间。与母本相比,总甲基化和全甲基化呈上升趋势,半甲基化呈下降趋势；与父本相比,均呈下降趋势。
5.在这三个杂交组合中DNA甲基化共分5种类型：单态性(A)、去甲基化(B)、过或超甲基化(C)、次甲基化(D)和中间型(E)。在苹果三倍性杂种的基因组加倍和重组过程中,发生了大量过或超甲基化和去甲基化变异,少量发生了次甲基化现象,极少部分的甲基化状态介于双亲之间。通过MSAP分析,共获得10条差异片段,有8条能够在苹果基因组中检测到,且同源性较高,这8条片段分别位于第1条、9条、10条、12条和16条染色体的叠连群上,但是具体功能没有描述。
At present in the apple production of China, diploid varieties play a major part, but a few triploid varieties have good performance. The triploid apple breeding has become an important part in fruit polyploid breeding. But there was very little research on that had obtained polyploidy resources by hybridization, and also very few reports of hybrid mutation mechanism. Especially, the molecular mechanism of genetic variation of triploid apple obtained by crossing has rarely been reported. In this study, molecular mechanism of genetic variation and epigenetic variation in genome including the status and patterns of DNA methylation based on the investigation of the botanical characters of triploid hybrid progenies of 'Hanfu'×'4xGala','Fuji'×'4xGala','Golden Delicious'×'4xGala' were studied, and the results are as follows:
1. The coefficient of variation of leaves, branches and flowers of triploid hybrids of 'Hanfu'×'4xGala' were among26.42%～1.33%and that of pollen PE was4.46%. The averages of most of the characters were lower than that of the mid-parents. Coefficient of varation of fruit characters were among5.12%～27.82%, and the same trend with the botanical characters. Characteristic aromas of6hybrids were effect by both parent, and the genetic trend of fruit aromas was paternal inheritance. The major effect factor on the sweetness of6triploid hybrids was fructose, and that of sour flavor was malic acid.
2. Genetic variation of7triploid hybrids o f'Hanfu'×'4xGala' was investigated based on ISSR and AFLP. The percentage of polymorphic bands (PPB) of ISSR was60.00%, observed number of alleles (Na) was1.6000, effective number of alleles (Ne) was1.4204, Nei's gene diversity (H) was0.2368and Shannon's Information index (I) was0.3464. And these were lower than that of AFLP. The UPMGA cluster showed that most of7triploid hybrids were tend to the female parent'Hanfu'based on ISSR and AFLP. The analysis indicated that ISSR and AFLP both can use to analyze the genetic diversity of apple triploid hybrids, and these triploid hybrids were maternal inheritance.
3. Genetic variation of29triploid hybrids of 'Fuji'×'4xGala' was investigated based on ISSR. The percentage of polymorphic bands (PPB) was77.42%, observed number of alleles (Na) was1.7742, effective number of alleles (Ne) was1.4004, Nei's gene diversity (H) was0.2355and Shannon's Information index (I) was0.3574. Genetic variation of61triploid hybrids of 'Golden Delicious'×'4xGala' were investigated based on ISSR. The percentage of polymorphic bands (PPB) was36.55%, observed number of alleles (Na) was1.9655, effective number of alleles (Ne) was1.5322, Nei's gene diversity (H) was0.3092 and Shannon's Information index (I) was0.4641. In the80triploid hybrid progenies, the bands of hybids were most come from female parent, and there was a certain increase and deletion in the strips, and these triploid hybrids were maternal inheritance.
4. The total methylation ratios of7triploid hybrids of 'Hanfu'×'4xGala' were among14.74%to20.00%, the full methylated ratios were among5.41%to10.38%, and the hemi-methylated ratios of that were among5.85%to14.59%. Compared with 'Hanfu', the total methylation ratios, the full-methylatied ratios and hemi-methylated ratios were mostly tended to increase. But all of that were tended to decrease compared with'4xGala'.There was no obvious trend with that of their parents. The total methylation ratios of29triploid hybrids of 'Fuji'×'4xGala' were among14.65%to27.96%, the full methylated ratios were among8.25%to17.40%, and the hemi-methylated ratios of that were among4.19%to10.82%. Compared with 'Fuji', the total methylation ratios and hemi-methylatied ratios were mostly tended to increase, but full methylated ratios was tended decrease. But all of that were tended to decrease compared with '4xGala'. The total methylation ratios of61triploid hybrids of 'Golden Delicious'×'4xGala' were among10.74%to28.86%, the full methylated ratios were among7.84%to24.22%, and the hemi-methylated ratios of that were among2.10%to11.84%. Compared with 'Golden Delicious', the total methylation ratios and the full-methylatied ratios were mostly tended to increase, but hemi-methylated ratios was tended decrease. But all of that were tended to decrease compared with '4xGala'.
5. There were five patterns of DNA methylatiion in these three hybridized combinations including monomophism (A), demthylation (B), hyermethylation (C), hypomethylation (D) and intermediate methylation (E).During the process of genome recombination and triploidization in apple triploid hybrids, a great number of hypermethylations and demethlation were occurred, a small amout of occurrence methane phenomenon and a very small part of the methylation status were between parents. In total10differential fragments were obtained by MSAP.8can be detected in the apple genome, and the high homology fragments were located in the contig of8,9,10,12and16chromosomes, but the specific function is not described.
引文
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