梅EST分析与利用及梅、杏合子前生殖隔离的初步证据
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摘要
梅(Prunus mume)是蔷薇科(Rosaceae)李属,原产中国,在东亚国家有着广泛的种植。梅在我国已有7000年以上的应用历史,栽培历史也有3000多年。随着生物技术的快速发展,从基因水平上认识梅的生长发育以及开展梅在核果类植物中的遗传进化研究逐渐得到了重视。近年来分子标记在植物分类和亲缘关系分析等方面得到了广泛的应用。我国拥有丰富的梅种质资源,对梅资源进行分类和亲缘关系的研究,有利于梅种质资源的收集、保存、鉴定、创新和合理利用。本研究以梅品种‘细叶青’为材料,构建了梅花和果实的cDNA文库；结合文库所得的EST序列以及其他核果类植物中的已知功能的部分EST序列,首次通过芯片杂交试验从转录组水平上进行梅和杏果实发育过程中差异基因表达的分析。另外,本研究还对梅、杏之间的隔离机制进行了初步探讨。主要结果如下：
1.以梅品种‘细叶青’不同时期的花和果实为实验材料,应用Creator SMART cDNA Construction Kit,构建了我国果梅第一个花和果实的cDNA文库。结果表明,该文库的库容量约为1.4×106pfu/mL,从扩增文库中随机挑取30个克隆进行PCR鉴定,其重组率达97%,插入片段多分布在1～3kb之间,平均大于1kb,说明本研究成功构建了果梅花和果实的cDNA文库,为进一步开展果梅开花调控和果实发育等相关基因的克隆及其表达与功能分析等研究奠定了重要基础。
2.构建了一个梅花和果实的cDNA文库,共测序10123个克隆,去冗余后得到高质量的EST序列8,656条。经拼接获得组4473unigenes,包括1492个contigs和2981个singletons,并提交到NCBI数据库(IDS:GW868575-GW873047).经BLASTx后,有1294个unigene具有已知或假定功能。
3.利用MISA软件对10,123条梅EST序列进行SSR位点查找,得到含SSR位点的序列935条,SSR位点1,233个,平均每100条EST序列中含有12.18个SSR位点。2核苷酸、3核苷酸重复是最主要的重复类型,分别占35.52%和41.36%。设计了42对EST-SSR引物并进行扩增,有24对引物能扩增出理想的PCR产物,其中17对引物具有较好的扩增多态性。测序后发现14对引物中有73.08%的片段具有相应的SSR位点。利用上述14对EST-SSR引物,在桃、杏和李上进行进一步的转换研究,并对桃、杏、李、梅进行遗传多样性分析。结果发现,100%的引物在杏DNA中都能扩增出目的片段,多态性达87%；89%的引物在桃和李中都能扩增出目的片段,在李中的多态性达65%,而桃中则为46%。说明梅的EST-SSR引物在其它三种核果类树种中有很高的通用性。
4.以两个梅品种(‘细叶青’、‘软条红梅’)和两个杏品种(‘玛瑙’、‘真魁’)为试验材料,利用李属核果类植物的10,641条EST序列作为探针,首次通过芯片杂交试验从转录组水平上进行梅和杏果实发育过程中差异基因表达的分析。结果显示共检测到2倍以上差异表达基因1,418个,其中上调表达基因707个,下调基因711个。GO注释结果表明,差异表达基因中,54%(766)具有功能注释的差异表达基因主要与代谢过程、应激反应、发育、多细胞生物过程、定位、生物调控、信号转导等17个生物过程；Pathway分析结果则表明,梅、杏差异表达基因主要参与糖异生/糖酵解、三羧酸循环、磷酸戊糖代谢、淀粉和蔗糖代谢等多个碳水化合物代谢途径和次生代谢产物的生物合成包括苯丙烷类,黄酮类,类胡萝卜素,激素和木质素的生物合成。随机选择的16个差异表达基因(8个下调表达,8个上调表达)荧光定量验证结果显示,12个基因的表达分析结果与芯片检测结果基本一致,但在不同发育时期的果实中有不同强度的动态表达。
5.利用已获得的梅Unigene与GenBank数据库中已公布的杏、桃EST序列进行同源比对,发现同源序列592条,同源序列的总长度为235,576bp,平均长度和同源性分别为437bp，97.5%；进行Blast比对后发现,所有同源序列中有340个具有相应的功能注释,183个为未知功能蛋白,其余69个为没有相应序列信息的新基因；在所有同源序列中共发现8,818个SNPs,总频率为26bp/SNP.同时,对三个物种的同源序列进行两两比较发现,SNP数量明显少于这三者比较的结果。另外,利用上述所有SNP数据信息对梅、杏、桃3个物种的聚类分析结果说明梅、杏的亲缘关系较近,二者与桃亲缘关系较远。
6.以68个梅品种(花梅36个、果梅32个)为材料,利用SNPs标记对梅品种的遗传多样性和亲缘关系进行了研究。以每个品种的DNA为模板,利用9对引物扩增获得了总长度均为2,916bp的基因组序列,获得多态性SNP位点92个,分布密度为1SNP/32bp,包括：51个转换(55.4%),34个颠换(37%),4个缺失或插入(4.4%),3个其它类型(3.3%)。本研究对花梅和果梅的SNP进行了统计：花梅SNP总数为50,密度为1SNP/58bp;而果梅中总数为72,密度为1SNP/41bp,果梅SNP的出现频率高于花梅,说明果梅品种间的遗传差异较大,也可能是由于这些花梅品种的育种亲本相似或者地理起源相近。根据92个SNPs位点多态性资料,应用PHYLIP程序中的邻位相连法构建了花梅和果梅的系统进化树。结果表明：所有梅品种并没有形成花梅和果梅两大组群,说明果梅与花梅在遗传上具有很高的相似性,是同一树种中某些性状(主要是花器官)存在一定差异的不同品种类型。上述系统进化树将供试68份品种资源分成11个群体,包括5个花梅组群,2个果梅组群,4个花果混合群。至于果梅或花梅小组群的存在,可能由于这些小组群中的品种来源的相似性而具有较近的亲缘关系。本研究结果表明,尽管花梅和果梅的用途以及花性状上的明显差异,但它们在梅的进化来源以及遗传分析方面具有同等的重要性。
7.对梅、杏之间潜在的生殖隔离因素包括开花物候期和地理分布进行了调查分析。结果显示,地理隔离和开花物候隔离是梅、杏保持两个物种的界限的关键合子前隔离因素。同时,我们也提供了杏和梅之间存在合子前隔离的分子证据。说明杏和梅是遗传相近的两个独立的物种,自然杂交的机率较低。这些比较分析表明合子前隔离在杏和梅的进化过程中发挥着重要的作用。另外,杏梅是这两个物种经杂交产生可育后代。本研究所获得的结果和分子证据为进一步研究杏和梅物种形成及在整个李属植物的演化提供了重要的信息和理论依据。
Mei (Prunus mume) originated in China and belongs to the sub-family Prunus within the Rosaceae family and is widely cultivated in East Asian countries. Mei has been used for7,000years and the cultivation history for over3,000. With the rapid development of biotechnology, it has reported that understanding of the growth and development of Prunus mume from the level of gene were gradually received some attention, as well as the genetic evolution of the stone fruit plants. In recent years, molecular markers have been widely used in plant taxonomy and phylogenetic analysis. And there are rich germplasm resource in China, the studies of classification and phylogenetic relationship of Prunus germplasm resources had conducive to the collection, preservation, identification innovation and rational utilization of Prunus mume germplasm resources. In this study, the cDNA library of the mei flower and fruit was constructed. Combined with part of EST sequences from other stone fruit plants, the differentially expressed genes involved in fruit development between Prunus mume and Prunus armeniaca were analyzed on the level of the transcriptome by using microarray experiments. In addition, the isolation mechanism between Prunus mume and Prunus armeniaca was also discussed in this study. The main results are as follows:
1. The first cDNA library of Prunus mume derived from the RNA isolated from the flower and fruit at different developmental stages was constructed with Creator SMART cDNA Construction Kit. Analysis indicated that the cDNA library had a high titer of1.4×106pfu/mL, with a recombinant efficiency about97%. The insert sizes of the library were1-3kb based on a PCR analysis of30randomly chosen clones, suggesting that the cDNA library of Prunus mume flower and fruit has been successfully constructed and could serve well for the cloning and functional analysis of important genes involved in the development of flower and fruit.
2. In this study, we constructed a cDNA library of Prunus mume flower and fruit, sequenced10,123clones of the library, and obtained8,656expressed sequence tag (EST) sequences with high quality. The ESTs were assembled into4,473unigenes composed of1,492contigs and2,981singletons and that have been deposited in NCBI (accession IDs: GW868575-GW873047), among which1,294unique ESTs were with known or putative functions.
3. In this study, from10,123Prunus mume ESTs screened using MIS A software, and a total of1233putative simple sequence repeats (SSRs) in the P. mume unigene dataset were mined from935ESTs with frequency of12.18SSR loci distributed over100ESTs. Di-and tri nucleotide repeat EST-SSRs were dominant, accounting for35.52%, and41.36%, respectively. Furthermore, we randomly tested42pairs of PCR primers flanking potential SSRs, with24primer pairs showing amplifications while17were polymorphic, where14pairs were identified and73.08%of the PCR amplification bands as true-to-type SSR loci. We further used the14EST-SSR primer pairs to test the transferability on other Prunus species, such as peach, apricot and plum. The result showed that100%and89%of the primer pairs produced target PCR bands in the apricot and two other species, respectively. A high level of marker polymorphism was observed in the apricot species (89.6%), lower in the plum (65%) and peach (46%), and the clustering analysis of the three species.
4. Differentially expressed transcripts along the entire fruit life cycle base on two P. mume cultivar'Xiyeqing','Ruantiaohongemi'and two P. armeniaca'Manao','Zhenkui' were first identified, by using microarrays spotted with10,641ESTs collected from P. mume and other Prunus EST sequences. A total of1,418ESTs were selected after quality control of microarray spots and analyzed for differential gene expression patterns during fruit development of P. mume and P. armeniaca. Among them,707up-regulated and711down-regulated differentially expressed genes showing more than2.0-fold differences in expression level were annotated by GO based on biological processes, molecular functions and cellular components. And54%(766) genes were found involved in17biological processes such as metabolic process, cellular process, response to stimulus, developmental process, multi-cellular organism process, localization, biological regulation, reproduction, signaling. The KEGG results also showed that those genes encoding enzymes were involved in several pathways primarily focused on Carbohydrate Metabolism including Glycolysis/Gluconeogenesis, Citrate cycle (TCA cycle), Galactose metabolism, Starch and sucrose metabolism, and Biosynthesis of Other Secondary Metabolites containing Phenylpropanoid, Flavonoid, Carotenoid, Hormone and lignin biosynthesis. qRT-PCR expression patterns of12(75%) of these genes were in agreement with their microarray analysis results, and it should be noted that all the selected genes were detected with different expression during the stages of fruit development.
5. In this research, multiple alignments were performed between the unigenes of Prunus mume and both EST database of P. armeniaca and P. Persica published in GenBank,592homologous sequences were found with a total length of235576bp, and the average length and homology were437bp and97.5%, respectively. The Blast results also showed that340of them had the corresponding functional annotation,183were unknown proteins, and the remaining69had new gene sequence information. The amount and frequency of nucleotides were further analyzed, where8,818SNPs were found having a total frequency of26bp per SNP. The amount of SNPs compared in pairs was significantly less than the number among the homologous sequences. In addition, the cluster analysis result by using the obtained SNP information showed that the relationship between Prunus mume and P. armeniaca was closer, and they were distantly related with P. persica,
6. Genetic relationships among68mei cultivars(Prunus mume Sieb. et Zucc.), including36flowering mei and32fruiting mei cultivars, were assessed using single nucleotide polymorphism (SNP) markers. The sequence alignments of nine group genomic sequences consisting of a total length of2,916bp, amplified from68DNA samples using nine pairs of PCR primers, yielded92SNPs with a distribution frequency of one SNP per32bp. Among these SNPs,51(55.4%) were transitions,34(37%) were transversions,4(4.4%) were InDels and3(3.3%) were others. Of the92SNP loci in these68mei cultivars, there are several different existing-states of them among different cultivars:13(14.1%) were shown to be two kinds of homozygotes in different cultivars;22(23.9%) were recognised as heterozygotes in some cultivars and two kinds of homozygotes in other cultivars;50(54.3%) were heterozygotes and only one kind of homozygote in other cultivars. Cluster analysis of the68mei cultivars was carried out using the92SNPs, and a consensus cladogram was constructed. All the mei cultivars could be classified into11groups, instead of two groups consisting of only flowering mei or fruiting mei, respectively. This suggested that flowering and fruiting mei trees are genetically similar differently in their floral and fruit characteristics.
7. In this study, the divergent flowering phonologies and geographic distribution were reported that they were in substantial reproductive isolation between two important species of Prunus armeniaca and Prunus mume that can be cross-compatible. The investigation results show that geographical isolation and their flowering phenology isolation were important barrier in crucial mode of pre-zygotic isolation to maintain species boundaries of P. armeniaca and P. mume. The molecular evidence for pre-zygotic isolation between P. armeniaca and P. mume are also presented. Ecological and molecular factors provide adequate proof that P. armeniaca and P. mume could be two independent species that are however genetically related. Conclusions:These comparative analyses demonstrate that pre-zygotic isolation plays an important role in evolution between P. armeniaca and P. mume and shows that'Xingmei'is the sterile hybrid generated by hybridization of these two species. The empirical and molecular evidences in this paper can be informative and helpful for the further studies on evolution and speciation of P. armeniaca and P. mume and even for the entire Prunus genus.
1.以梅品种‘细叶青’不同时期的花和果实为实验材料,应用Creator SMART cDNA Construction Kit,构建了我国果梅第一个花和果实的cDNA文库。结果表明,该文库的库容量约为1.4×106pfu/mL,从扩增文库中随机挑取30个克隆进行PCR鉴定,其重组率达97%,插入片段多分布在1～3kb之间,平均大于1kb,说明本研究成功构建了果梅花和果实的cDNA文库,为进一步开展果梅开花调控和果实发育等相关基因的克隆及其表达与功能分析等研究奠定了重要基础。
2.构建了一个梅花和果实的cDNA文库,共测序10123个克隆,去冗余后得到高质量的EST序列8,656条。经拼接获得组4473unigenes,包括1492个contigs和2981个singletons,并提交到NCBI数据库(IDS:GW868575-GW873047).经BLASTx后,有1294个unigene具有已知或假定功能。
3.利用MISA软件对10,123条梅EST序列进行SSR位点查找,得到含SSR位点的序列935条,SSR位点1,233个,平均每100条EST序列中含有12.18个SSR位点。2核苷酸、3核苷酸重复是最主要的重复类型,分别占35.52%和41.36%。设计了42对EST-SSR引物并进行扩增,有24对引物能扩增出理想的PCR产物,其中17对引物具有较好的扩增多态性。测序后发现14对引物中有73.08%的片段具有相应的SSR位点。利用上述14对EST-SSR引物,在桃、杏和李上进行进一步的转换研究,并对桃、杏、李、梅进行遗传多样性分析。结果发现,100%的引物在杏DNA中都能扩增出目的片段,多态性达87%；89%的引物在桃和李中都能扩增出目的片段,在李中的多态性达65%,而桃中则为46%。说明梅的EST-SSR引物在其它三种核果类树种中有很高的通用性。
4.以两个梅品种(‘细叶青’、‘软条红梅’)和两个杏品种(‘玛瑙’、‘真魁’)为试验材料,利用李属核果类植物的10,641条EST序列作为探针,首次通过芯片杂交试验从转录组水平上进行梅和杏果实发育过程中差异基因表达的分析。结果显示共检测到2倍以上差异表达基因1,418个,其中上调表达基因707个,下调基因711个。GO注释结果表明,差异表达基因中,54%(766)具有功能注释的差异表达基因主要与代谢过程、应激反应、发育、多细胞生物过程、定位、生物调控、信号转导等17个生物过程；Pathway分析结果则表明,梅、杏差异表达基因主要参与糖异生/糖酵解、三羧酸循环、磷酸戊糖代谢、淀粉和蔗糖代谢等多个碳水化合物代谢途径和次生代谢产物的生物合成包括苯丙烷类,黄酮类,类胡萝卜素,激素和木质素的生物合成。随机选择的16个差异表达基因(8个下调表达,8个上调表达)荧光定量验证结果显示,12个基因的表达分析结果与芯片检测结果基本一致,但在不同发育时期的果实中有不同强度的动态表达。
5.利用已获得的梅Unigene与GenBank数据库中已公布的杏、桃EST序列进行同源比对,发现同源序列592条,同源序列的总长度为235,576bp,平均长度和同源性分别为437bp，97.5%；进行Blast比对后发现,所有同源序列中有340个具有相应的功能注释,183个为未知功能蛋白,其余69个为没有相应序列信息的新基因；在所有同源序列中共发现8,818个SNPs,总频率为26bp/SNP.同时,对三个物种的同源序列进行两两比较发现,SNP数量明显少于这三者比较的结果。另外,利用上述所有SNP数据信息对梅、杏、桃3个物种的聚类分析结果说明梅、杏的亲缘关系较近,二者与桃亲缘关系较远。
6.以68个梅品种(花梅36个、果梅32个)为材料,利用SNPs标记对梅品种的遗传多样性和亲缘关系进行了研究。以每个品种的DNA为模板,利用9对引物扩增获得了总长度均为2,916bp的基因组序列,获得多态性SNP位点92个,分布密度为1SNP/32bp,包括：51个转换(55.4%),34个颠换(37%),4个缺失或插入(4.4%),3个其它类型(3.3%)。本研究对花梅和果梅的SNP进行了统计：花梅SNP总数为50,密度为1SNP/58bp;而果梅中总数为72,密度为1SNP/41bp,果梅SNP的出现频率高于花梅,说明果梅品种间的遗传差异较大,也可能是由于这些花梅品种的育种亲本相似或者地理起源相近。根据92个SNPs位点多态性资料,应用PHYLIP程序中的邻位相连法构建了花梅和果梅的系统进化树。结果表明：所有梅品种并没有形成花梅和果梅两大组群,说明果梅与花梅在遗传上具有很高的相似性,是同一树种中某些性状(主要是花器官)存在一定差异的不同品种类型。上述系统进化树将供试68份品种资源分成11个群体,包括5个花梅组群,2个果梅组群,4个花果混合群。至于果梅或花梅小组群的存在,可能由于这些小组群中的品种来源的相似性而具有较近的亲缘关系。本研究结果表明,尽管花梅和果梅的用途以及花性状上的明显差异,但它们在梅的进化来源以及遗传分析方面具有同等的重要性。
7.对梅、杏之间潜在的生殖隔离因素包括开花物候期和地理分布进行了调查分析。结果显示,地理隔离和开花物候隔离是梅、杏保持两个物种的界限的关键合子前隔离因素。同时,我们也提供了杏和梅之间存在合子前隔离的分子证据。说明杏和梅是遗传相近的两个独立的物种,自然杂交的机率较低。这些比较分析表明合子前隔离在杏和梅的进化过程中发挥着重要的作用。另外,杏梅是这两个物种经杂交产生可育后代。本研究所获得的结果和分子证据为进一步研究杏和梅物种形成及在整个李属植物的演化提供了重要的信息和理论依据。
Mei (Prunus mume) originated in China and belongs to the sub-family Prunus within the Rosaceae family and is widely cultivated in East Asian countries. Mei has been used for7,000years and the cultivation history for over3,000. With the rapid development of biotechnology, it has reported that understanding of the growth and development of Prunus mume from the level of gene were gradually received some attention, as well as the genetic evolution of the stone fruit plants. In recent years, molecular markers have been widely used in plant taxonomy and phylogenetic analysis. And there are rich germplasm resource in China, the studies of classification and phylogenetic relationship of Prunus germplasm resources had conducive to the collection, preservation, identification innovation and rational utilization of Prunus mume germplasm resources. In this study, the cDNA library of the mei flower and fruit was constructed. Combined with part of EST sequences from other stone fruit plants, the differentially expressed genes involved in fruit development between Prunus mume and Prunus armeniaca were analyzed on the level of the transcriptome by using microarray experiments. In addition, the isolation mechanism between Prunus mume and Prunus armeniaca was also discussed in this study. The main results are as follows:
1. The first cDNA library of Prunus mume derived from the RNA isolated from the flower and fruit at different developmental stages was constructed with Creator SMART cDNA Construction Kit. Analysis indicated that the cDNA library had a high titer of1.4×106pfu/mL, with a recombinant efficiency about97%. The insert sizes of the library were1-3kb based on a PCR analysis of30randomly chosen clones, suggesting that the cDNA library of Prunus mume flower and fruit has been successfully constructed and could serve well for the cloning and functional analysis of important genes involved in the development of flower and fruit.
2. In this study, we constructed a cDNA library of Prunus mume flower and fruit, sequenced10,123clones of the library, and obtained8,656expressed sequence tag (EST) sequences with high quality. The ESTs were assembled into4,473unigenes composed of1,492contigs and2,981singletons and that have been deposited in NCBI (accession IDs: GW868575-GW873047), among which1,294unique ESTs were with known or putative functions.
3. In this study, from10,123Prunus mume ESTs screened using MIS A software, and a total of1233putative simple sequence repeats (SSRs) in the P. mume unigene dataset were mined from935ESTs with frequency of12.18SSR loci distributed over100ESTs. Di-and tri nucleotide repeat EST-SSRs were dominant, accounting for35.52%, and41.36%, respectively. Furthermore, we randomly tested42pairs of PCR primers flanking potential SSRs, with24primer pairs showing amplifications while17were polymorphic, where14pairs were identified and73.08%of the PCR amplification bands as true-to-type SSR loci. We further used the14EST-SSR primer pairs to test the transferability on other Prunus species, such as peach, apricot and plum. The result showed that100%and89%of the primer pairs produced target PCR bands in the apricot and two other species, respectively. A high level of marker polymorphism was observed in the apricot species (89.6%), lower in the plum (65%) and peach (46%), and the clustering analysis of the three species.
4. Differentially expressed transcripts along the entire fruit life cycle base on two P. mume cultivar'Xiyeqing','Ruantiaohongemi'and two P. armeniaca'Manao','Zhenkui' were first identified, by using microarrays spotted with10,641ESTs collected from P. mume and other Prunus EST sequences. A total of1,418ESTs were selected after quality control of microarray spots and analyzed for differential gene expression patterns during fruit development of P. mume and P. armeniaca. Among them,707up-regulated and711down-regulated differentially expressed genes showing more than2.0-fold differences in expression level were annotated by GO based on biological processes, molecular functions and cellular components. And54%(766) genes were found involved in17biological processes such as metabolic process, cellular process, response to stimulus, developmental process, multi-cellular organism process, localization, biological regulation, reproduction, signaling. The KEGG results also showed that those genes encoding enzymes were involved in several pathways primarily focused on Carbohydrate Metabolism including Glycolysis/Gluconeogenesis, Citrate cycle (TCA cycle), Galactose metabolism, Starch and sucrose metabolism, and Biosynthesis of Other Secondary Metabolites containing Phenylpropanoid, Flavonoid, Carotenoid, Hormone and lignin biosynthesis. qRT-PCR expression patterns of12(75%) of these genes were in agreement with their microarray analysis results, and it should be noted that all the selected genes were detected with different expression during the stages of fruit development.
5. In this research, multiple alignments were performed between the unigenes of Prunus mume and both EST database of P. armeniaca and P. Persica published in GenBank,592homologous sequences were found with a total length of235576bp, and the average length and homology were437bp and97.5%, respectively. The Blast results also showed that340of them had the corresponding functional annotation,183were unknown proteins, and the remaining69had new gene sequence information. The amount and frequency of nucleotides were further analyzed, where8,818SNPs were found having a total frequency of26bp per SNP. The amount of SNPs compared in pairs was significantly less than the number among the homologous sequences. In addition, the cluster analysis result by using the obtained SNP information showed that the relationship between Prunus mume and P. armeniaca was closer, and they were distantly related with P. persica,
6. Genetic relationships among68mei cultivars(Prunus mume Sieb. et Zucc.), including36flowering mei and32fruiting mei cultivars, were assessed using single nucleotide polymorphism (SNP) markers. The sequence alignments of nine group genomic sequences consisting of a total length of2,916bp, amplified from68DNA samples using nine pairs of PCR primers, yielded92SNPs with a distribution frequency of one SNP per32bp. Among these SNPs,51(55.4%) were transitions,34(37%) were transversions,4(4.4%) were InDels and3(3.3%) were others. Of the92SNP loci in these68mei cultivars, there are several different existing-states of them among different cultivars:13(14.1%) were shown to be two kinds of homozygotes in different cultivars;22(23.9%) were recognised as heterozygotes in some cultivars and two kinds of homozygotes in other cultivars;50(54.3%) were heterozygotes and only one kind of homozygote in other cultivars. Cluster analysis of the68mei cultivars was carried out using the92SNPs, and a consensus cladogram was constructed. All the mei cultivars could be classified into11groups, instead of two groups consisting of only flowering mei or fruiting mei, respectively. This suggested that flowering and fruiting mei trees are genetically similar differently in their floral and fruit characteristics.
7. In this study, the divergent flowering phonologies and geographic distribution were reported that they were in substantial reproductive isolation between two important species of Prunus armeniaca and Prunus mume that can be cross-compatible. The investigation results show that geographical isolation and their flowering phenology isolation were important barrier in crucial mode of pre-zygotic isolation to maintain species boundaries of P. armeniaca and P. mume. The molecular evidence for pre-zygotic isolation between P. armeniaca and P. mume are also presented. Ecological and molecular factors provide adequate proof that P. armeniaca and P. mume could be two independent species that are however genetically related. Conclusions:These comparative analyses demonstrate that pre-zygotic isolation plays an important role in evolution between P. armeniaca and P. mume and shows that'Xingmei'is the sterile hybrid generated by hybridization of these two species. The empirical and molecular evidences in this paper can be informative and helpful for the further studies on evolution and speciation of P. armeniaca and P. mume and even for the entire Prunus genus.
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