泰沂山区苹果属植物系统学研究
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摘要
苹果属植物具有重要的经济意义,已经成为植物学、园艺学以及其他相关领域的研究热点。但本属植物难于鉴定,尤其栽培种没有模式植物,在一些研究文献中经常出现名不副实的现象,严重影响本属植物的深入研究和利用。因此,急需建立科学规范的分类体系。泰沂山区是我国苹果属植物资源较为丰富的地区之一,除特有的湖北海棠种下茶山海棠、泰山海棠2个类型外,其他种均为栽培类型,变异丰富,杂交现象严重,存在大量的种间过渡类型,因此同名异物、同物异名现象严重。20世纪50年代至70年代末,研究人员对该地区的苹果属植物资源进行过较大规模的收集保存及优良砧木选育工作,为《山东果树志》苹果砧木资源的编写工作奠定了坚实基础。现有关于泰沂山区苹果属植物资源的介绍多局限于简单资源归类,性状描述不够详细,使该区苹果属资源开发利用受到限制。本研究针对泰沂山区苹果属植物资源较为丰富的特点,进行了较大范围收集保存工作,在借鉴国内现有苹果属植物分类体系的基础上,引进数量分类方法,建立简化的中国苹果属植物数量化分类体系,对泰沂山区苹果属植物进行鉴定,明确其归属。结合应用不同来源的DNA序列及cpSSR分子标记技术,从分子系统学角度阐述泰沂山区苹果属植物资源系统学关系,主要研究结果如下。
1.运用数量分类方法对27种中国苹果属植物进行表型系统分类,通过R型聚类明确了中国苹果属植物种间鉴定的49个独立性状,通过Q型聚类将中国苹果属植物分为2大组,即花楸苹果组和真苹果组,真苹果组中又将苹果系与山荆子系明确区分。主成分分析表明,新疆野苹果与所有栽培种聚在一起,花楸苹果组与真苹果组的野生种聚在一起。
运用该数量分类体系对泰沂山区102份资源进行了有效鉴定。鉴定结果表明,泰沂山区苹果属植物主要存在西府海棠、楸子、沙果、苹果、湖北海棠、三叶海棠6个种,每个种均存在2个种下类型。西府海棠2个代表类型分别为难咽和林檎,楸子的代表类型为烟台沙果和茶果子,三叶海棠分黄三叶和红三叶,湖北海棠主要代表类型为平邑甜茶和泰山海棠,沙果主要分黄色品种群和红色品种群。难咽与林檎的主要区分性状为花蕾颜色、果梗长度、果实直径；楸子两大类型区分的关键性状是果实直径、花冠大小、叶片长度；平邑甜茶与泰山海棠区分的主要性状是果实直径、花梗着毛、花蕾颜色、花瓣颜色、花冠大小、叶片宽度；三叶海棠两个类型区分的主要性状是果面盖色、花冠大小、叶片大小；沙果黄色品种群与红色品种群的区别性状主要为果面盖色、花冠大小、叶片长度。
2.泰沂山区苹果属植物ITS序列(含5.8S)全长为593-665bp,各样品种间的碱基组成有一定差异,4种核苷酸量平均为A=17.95%,G=32.36%,T=17.59%,C=32.26%。ITS序列的G+C含量较高,都在50%以上,平均含量为64.46%。当空位(gap)作缺失处理时,ITS全序列的长度为629位点,其中301个变异位点,234个为系统发育的信息位点,变异位点和信息位点分别占47.85%和37.20%。ITS序列在苹果属植物内进化速率较快,提供的变异位点较多,在种间的系统发育重建研究中可以提供较丰富的信息,NJ树与MP树分支结构基本一致,主要有4大分支,野生种山荆子与湖北海棠2个种下类型泰山海棠、平邑甜茶处于同一分支,位于系统树较低部位,第二分支包括少部分沙果和西府海棠,第三分支主要为新疆野苹果和沙果,第四分支包括苹果、沙果、楸子、西府海棠,位于系统树的最顶端,各分支支持率均在60%以上。聚类结果也表明野生种与栽培种可以明确分开,同时也证明栽培种亲缘关系复杂,并且形成途径复杂。沙果、西府海棠、楸子中可能均存在一些进化类型。
泰沂山区苹果属植物trnL-trnF序列,不同试材之间长度差异较大,最长为1064bp,最短为945bp,平均长度为996bp。G+C含量较低,变化范围在31.11%～32.89%之间,材料间几乎无差异。所测样品碱基组成A=33.52%,G=16.02%,T=34.35%,C=16.11%。当空位(gap)作缺失处理时,trnL-trnF全序列的长度为1004位点,其中29个变异位点,12个为系统发育的信息位点,变异位点、信息位点分别占2.89%和1.20%,序列相当保守,无法为泰沂山系苹果属植物的系统学关系提供足够证据。
以trnL-trnF序列与ITS序列构建的系统发育树存在较大差异,但都证明泰沂山区苹果属植物栽培种形成过程及遗传背景复杂,很多资源虽为同一个种但其杂交亲本可能来自不同种,甚至多次杂交。
3.12对供试cpSSR引物中仅有4对适合泰沂山区苹果属植物遗传多样性分析,这些引物分别是ccmp2、ccmp4、ccmp5、cicp9,这4对引物均检测出泰沂山区苹果属植物遗传多样性。4对引物从36份资源中共扩增出177个DNA位点,多态性比率为76.27%；UPGMA聚类结果显示,cpSSR标记能将36份泰沂山区苹果属植物资源完全分开,在距离系数1.82处将其分为2大类,山荆子与湖北海棠的2个种下类型泰山海棠、平邑甜茶处于同一分支,西府海棠、楸子、沙果、苹果位于另一分支,呈交互聚类状,与核基因组ITS序列构建的系统树结果基本一致,同样支持栽培种与野生种分开的观点,也表明栽培种之间亲缘关系复杂。
本研究结果初步表明,数量分类方法可以有效解决中国苹果属植物分类鉴定问题,栽培种和野生种可以独立分开,栽培种的祖先应为新疆野苹果,栽培种形成过程复杂,遗传背景复杂,亲本来源多样。泰沂山区苹果属植物主要包括5个种,10个类型。基于核基因组ITS构建的系统树与基于cpSSR标记构建的系统树结果基本一致,在一定程度上可以支持表型分类结果。基于叶绿体基因组trnL-trnF序列构建的系统树无法有效解决泰沂山区苹果属植物系统学关系。
Malus Mill.has become the hot topic of botany, horticulture and other related fields of research due to its important economic significance. However, there are some difficulties in the identification of this species, even to some experts in this field. For example, some mistakes in the rename of some species often appeared even in research literatures, thus affecting its research. Therefore, a more scientific, standardized and simple classification system needs to be built.'TaiYi'mountain is one of the areas in China that many Malus species originated, including two Malus hupehensis (var'Pingyitiancha'and'Taishanhaitang') and many cultivated varieties (including some homonyms and synonyms) due to long-term sexual reproduction. From1950s to late1970s in this area, some Malus Mill.species were collected and investigated, mainly for purpose of breeding apple rootstocks. Although Malus species in the'TaiYi'mountain area have received extensive attention, further systematic research should be needed. Some Malus Mill, species in this area were described in the book 'Tree fruit in Shandong", but this description was insufficient and the classification of different species is too simple due to lack of more detailed investigation and systematic research, thus limiting the utilization of the rich Malus resource in this area. In this study, the rich Malus Mill, species in'TaiYi'mountain area were collected, preserved and more importantly classified, base on a more simplified, quantitative classification system for China Malus species which was developed based on the theory of quantitative classification, as well as the former classification system. On this basis, the phylogenetic relationship between different Malus Mill, species in'TaiYi'mountain area was suggested from the molecular perspective, based on DNA sequence and cpSSR marker. The main results are as follows.
1. A quantitative classification system was adopted for phynotypic system classification of27kinds of Malus Mill, species in china.49independent traits of Malus Mill, species were explicated by R cluster, and the Chinese Malus Mill, species were indentified into two large groups by Q cluster, which were Sect.Sorbomalus-ser. and Section Eumalus., even more the wild and cultivated type of Section Eumalus. were also clearly distinct. The principal component analysis indicated that Malus sieversii and all cultivated species were together, while the Sect.Sorbomalus-ser. and the wild type of Section Eumalus were together.
A quantitative classification system was applied for the identification of102samples germplasm in'taiyi'mountain. The results showed that Malus resources in'TaiYi'mountain area belong to6species which is Malus micromalus Makino, Malus prunifolia (Wild.) Borth., Malus asiatica Nakai., Malus domestica subsp. chinensis Li Y.N. Malus hupehensis (Pamp.) Rehd. and Malus sieboldii (Reg.) Rend., with each species having2subspecies. Malus micromalus has tow subspecies which was Malus micromalus Mak. and Malus asiatica Nakai. characterized by different flower color, pedicel length and fruit diameter. M.prunifoli (Willd.) Borkh and chaguozi belong to Malus prunifolia, characterized by different fruit diameter, corolla size and leaf length. Pingyitiancha and taishanhaitang are Malus hupehensis, which is different in fruit diameter, hair in flower pedicel, flower color, petal color, corolla size and leaf wide. Malus sieboldii has huangsanye and hongsanye with different leaf size, corolla size and petal length. Malus asiatica posses yellow variety groups and red variety groups, with different fruit color, corolla size, and leaf length.
2. Systematic relationships between different Malus Mill, species in'TaiYi'mountain area were studied using ITS nuclear sequence and chloroplasts trrL-trnF's. It was found that the full length of the ITS sequence (including5.8S) for Malus Mill, species in this area was from593bp to665bp, with nucleotides A, G, T and C averaging17.95%,32.36%,17.59%and32.26%, respectively. Nucleotides G+C of the ITS sequence was much higher than A+T, with each species exceeding50%and the mean content being64.46%. If the gap was considered as missing, the full length of the ITS sequence was629bp, with301variation sites and234sites for systematic development, accounting for47.85%and37.20%, respectively. As ITS sequence of Malus Mill, species is much more evolved with plenty of variation sites, it can provide much information in the reconstruction of systematic development of Malus Mill, species. The branch structure of NJ tree and MP tree were basically the same, there were four main large branches, Malus baccata of the wild species and2subspecies of Malus hupehensis were in the same branch in the lower part of the system tree; the second branch including a little part of Malus asitica and Malus micromalus; the third branch mainly including Malus sieversii and Malus asitica; the forth branch including Malus domestica, Malus prinifolia and Malus asiatica located in the top of the system tree, each branch support more than60%. The clustering results also showed that the wild species and cultivated species can be clearly seprated, but also proved that the genetic relationship of the cultivated species and the formation ways were complex. And there might be some evolutionary type in Malus micromalus, Malus asitica and Malus prunifolia.
The trnL-trnF sequences of Malus in'taiyi'mountain area differ greatly among different species from945bp to1064bp, averaging996bp. The ratio of G+C of diffrent Malus Mill, species in this area is relatively low (each below40%) and there is no significant difference among species, ranging from31.11%to32.89%. The ratio of alkaline base for A, G, T and C is33.52%,16.02%,34.35%and16.11%, respectively. If the gap was considered as missing, the full length of trnL-trnF sequence was1004sites, including29variation sites and12phylogenetic information sites, consisting2.89%and1.20%of the total length, respectively. The trnL-trnF sequence was much conserved and it was not expected to solve the systematic relationships among different Malus Mill, species in'taiyi'mountains area.
Although there was obvious difference between the two phylogenetic trees based on ITS and trnL-trnF sequences, It could be both proved that the formation and genetic background of taiyi'mountains malus was complicated, a lot of germplasms belong to the same species, but their hybrid parents may come from different species, even experienced many hybrids.
3. Four cpSSR primers ccmp2, ccmp4, ccmp5and cicp9were proved to be suitable for genetic diversity analysis of Malus Mill, species in'TaiYi'mountain area, with each primer pairs having good results. Totally177DNA loci was amplified from36samples for these4primer pairs, with a polymorphism ratio of76.27%. UPGMA analysis showed that36Malus Mill,species sampled in this research can be totally distinguished using the method of cpSSR mark. With these36samples, two categories can be obtained at similarity coefficient level of1.82,Malus baccata and2subspecies of Malus hupehensis, Malus hupehensis var. pinyiensis and Malus hupehensis var. taishanesis were in the same branch. Malus micromalus, Malus prunifolia, Malus asitica and Malus domestica were in another branch, showing the interaction cluster-like, they were basically the same with the result of the system tree constructed by ITS sequences of nuclear genome, also supported the view that cultivated species and wild species separated with each other, it also showed that the genetic relationship among cultivars were complex.
The results of this study preliminary showed that the quantitative classification system can effectively solve the Chinese Malus Mill,classification and identification problem, cultivated species and wild species can be separated, the ancestor of cultivated species should be Malus seversii, the formation and genetic background of cultivated species were complex, and their parents were various. Malus Mill, of taiyi"mountain mainly including5species,10types. The results of the system tree construted by ITS sequences of nuclear genome and the system tree by cpSSR were basically same, could support the phenotypic classification results. The system tree constructed by trhL-trnF sequence can not slove the systematic relationship of tai yi'mountain.
1.运用数量分类方法对27种中国苹果属植物进行表型系统分类,通过R型聚类明确了中国苹果属植物种间鉴定的49个独立性状,通过Q型聚类将中国苹果属植物分为2大组,即花楸苹果组和真苹果组,真苹果组中又将苹果系与山荆子系明确区分。主成分分析表明,新疆野苹果与所有栽培种聚在一起,花楸苹果组与真苹果组的野生种聚在一起。
运用该数量分类体系对泰沂山区102份资源进行了有效鉴定。鉴定结果表明,泰沂山区苹果属植物主要存在西府海棠、楸子、沙果、苹果、湖北海棠、三叶海棠6个种,每个种均存在2个种下类型。西府海棠2个代表类型分别为难咽和林檎,楸子的代表类型为烟台沙果和茶果子,三叶海棠分黄三叶和红三叶,湖北海棠主要代表类型为平邑甜茶和泰山海棠,沙果主要分黄色品种群和红色品种群。难咽与林檎的主要区分性状为花蕾颜色、果梗长度、果实直径；楸子两大类型区分的关键性状是果实直径、花冠大小、叶片长度；平邑甜茶与泰山海棠区分的主要性状是果实直径、花梗着毛、花蕾颜色、花瓣颜色、花冠大小、叶片宽度；三叶海棠两个类型区分的主要性状是果面盖色、花冠大小、叶片大小；沙果黄色品种群与红色品种群的区别性状主要为果面盖色、花冠大小、叶片长度。
2.泰沂山区苹果属植物ITS序列(含5.8S)全长为593-665bp,各样品种间的碱基组成有一定差异,4种核苷酸量平均为A=17.95%,G=32.36%,T=17.59%,C=32.26%。ITS序列的G+C含量较高,都在50%以上,平均含量为64.46%。当空位(gap)作缺失处理时,ITS全序列的长度为629位点,其中301个变异位点,234个为系统发育的信息位点,变异位点和信息位点分别占47.85%和37.20%。ITS序列在苹果属植物内进化速率较快,提供的变异位点较多,在种间的系统发育重建研究中可以提供较丰富的信息,NJ树与MP树分支结构基本一致,主要有4大分支,野生种山荆子与湖北海棠2个种下类型泰山海棠、平邑甜茶处于同一分支,位于系统树较低部位,第二分支包括少部分沙果和西府海棠,第三分支主要为新疆野苹果和沙果,第四分支包括苹果、沙果、楸子、西府海棠,位于系统树的最顶端,各分支支持率均在60%以上。聚类结果也表明野生种与栽培种可以明确分开,同时也证明栽培种亲缘关系复杂,并且形成途径复杂。沙果、西府海棠、楸子中可能均存在一些进化类型。
泰沂山区苹果属植物trnL-trnF序列,不同试材之间长度差异较大,最长为1064bp,最短为945bp,平均长度为996bp。G+C含量较低,变化范围在31.11%～32.89%之间,材料间几乎无差异。所测样品碱基组成A=33.52%,G=16.02%,T=34.35%,C=16.11%。当空位(gap)作缺失处理时,trnL-trnF全序列的长度为1004位点,其中29个变异位点,12个为系统发育的信息位点,变异位点、信息位点分别占2.89%和1.20%,序列相当保守,无法为泰沂山系苹果属植物的系统学关系提供足够证据。
以trnL-trnF序列与ITS序列构建的系统发育树存在较大差异,但都证明泰沂山区苹果属植物栽培种形成过程及遗传背景复杂,很多资源虽为同一个种但其杂交亲本可能来自不同种,甚至多次杂交。
3.12对供试cpSSR引物中仅有4对适合泰沂山区苹果属植物遗传多样性分析,这些引物分别是ccmp2、ccmp4、ccmp5、cicp9,这4对引物均检测出泰沂山区苹果属植物遗传多样性。4对引物从36份资源中共扩增出177个DNA位点,多态性比率为76.27%；UPGMA聚类结果显示,cpSSR标记能将36份泰沂山区苹果属植物资源完全分开,在距离系数1.82处将其分为2大类,山荆子与湖北海棠的2个种下类型泰山海棠、平邑甜茶处于同一分支,西府海棠、楸子、沙果、苹果位于另一分支,呈交互聚类状,与核基因组ITS序列构建的系统树结果基本一致,同样支持栽培种与野生种分开的观点,也表明栽培种之间亲缘关系复杂。
本研究结果初步表明,数量分类方法可以有效解决中国苹果属植物分类鉴定问题,栽培种和野生种可以独立分开,栽培种的祖先应为新疆野苹果,栽培种形成过程复杂,遗传背景复杂,亲本来源多样。泰沂山区苹果属植物主要包括5个种,10个类型。基于核基因组ITS构建的系统树与基于cpSSR标记构建的系统树结果基本一致,在一定程度上可以支持表型分类结果。基于叶绿体基因组trnL-trnF序列构建的系统树无法有效解决泰沂山区苹果属植物系统学关系。
Malus Mill.has become the hot topic of botany, horticulture and other related fields of research due to its important economic significance. However, there are some difficulties in the identification of this species, even to some experts in this field. For example, some mistakes in the rename of some species often appeared even in research literatures, thus affecting its research. Therefore, a more scientific, standardized and simple classification system needs to be built.'TaiYi'mountain is one of the areas in China that many Malus species originated, including two Malus hupehensis (var'Pingyitiancha'and'Taishanhaitang') and many cultivated varieties (including some homonyms and synonyms) due to long-term sexual reproduction. From1950s to late1970s in this area, some Malus Mill.species were collected and investigated, mainly for purpose of breeding apple rootstocks. Although Malus species in the'TaiYi'mountain area have received extensive attention, further systematic research should be needed. Some Malus Mill, species in this area were described in the book 'Tree fruit in Shandong", but this description was insufficient and the classification of different species is too simple due to lack of more detailed investigation and systematic research, thus limiting the utilization of the rich Malus resource in this area. In this study, the rich Malus Mill, species in'TaiYi'mountain area were collected, preserved and more importantly classified, base on a more simplified, quantitative classification system for China Malus species which was developed based on the theory of quantitative classification, as well as the former classification system. On this basis, the phylogenetic relationship between different Malus Mill, species in'TaiYi'mountain area was suggested from the molecular perspective, based on DNA sequence and cpSSR marker. The main results are as follows.
1. A quantitative classification system was adopted for phynotypic system classification of27kinds of Malus Mill, species in china.49independent traits of Malus Mill, species were explicated by R cluster, and the Chinese Malus Mill, species were indentified into two large groups by Q cluster, which were Sect.Sorbomalus-ser. and Section Eumalus., even more the wild and cultivated type of Section Eumalus. were also clearly distinct. The principal component analysis indicated that Malus sieversii and all cultivated species were together, while the Sect.Sorbomalus-ser. and the wild type of Section Eumalus were together.
A quantitative classification system was applied for the identification of102samples germplasm in'taiyi'mountain. The results showed that Malus resources in'TaiYi'mountain area belong to6species which is Malus micromalus Makino, Malus prunifolia (Wild.) Borth., Malus asiatica Nakai., Malus domestica subsp. chinensis Li Y.N. Malus hupehensis (Pamp.) Rehd. and Malus sieboldii (Reg.) Rend., with each species having2subspecies. Malus micromalus has tow subspecies which was Malus micromalus Mak. and Malus asiatica Nakai. characterized by different flower color, pedicel length and fruit diameter. M.prunifoli (Willd.) Borkh and chaguozi belong to Malus prunifolia, characterized by different fruit diameter, corolla size and leaf length. Pingyitiancha and taishanhaitang are Malus hupehensis, which is different in fruit diameter, hair in flower pedicel, flower color, petal color, corolla size and leaf wide. Malus sieboldii has huangsanye and hongsanye with different leaf size, corolla size and petal length. Malus asiatica posses yellow variety groups and red variety groups, with different fruit color, corolla size, and leaf length.
2. Systematic relationships between different Malus Mill, species in'TaiYi'mountain area were studied using ITS nuclear sequence and chloroplasts trrL-trnF's. It was found that the full length of the ITS sequence (including5.8S) for Malus Mill, species in this area was from593bp to665bp, with nucleotides A, G, T and C averaging17.95%,32.36%,17.59%and32.26%, respectively. Nucleotides G+C of the ITS sequence was much higher than A+T, with each species exceeding50%and the mean content being64.46%. If the gap was considered as missing, the full length of the ITS sequence was629bp, with301variation sites and234sites for systematic development, accounting for47.85%and37.20%, respectively. As ITS sequence of Malus Mill, species is much more evolved with plenty of variation sites, it can provide much information in the reconstruction of systematic development of Malus Mill, species. The branch structure of NJ tree and MP tree were basically the same, there were four main large branches, Malus baccata of the wild species and2subspecies of Malus hupehensis were in the same branch in the lower part of the system tree; the second branch including a little part of Malus asitica and Malus micromalus; the third branch mainly including Malus sieversii and Malus asitica; the forth branch including Malus domestica, Malus prinifolia and Malus asiatica located in the top of the system tree, each branch support more than60%. The clustering results also showed that the wild species and cultivated species can be clearly seprated, but also proved that the genetic relationship of the cultivated species and the formation ways were complex. And there might be some evolutionary type in Malus micromalus, Malus asitica and Malus prunifolia.
The trnL-trnF sequences of Malus in'taiyi'mountain area differ greatly among different species from945bp to1064bp, averaging996bp. The ratio of G+C of diffrent Malus Mill, species in this area is relatively low (each below40%) and there is no significant difference among species, ranging from31.11%to32.89%. The ratio of alkaline base for A, G, T and C is33.52%,16.02%,34.35%and16.11%, respectively. If the gap was considered as missing, the full length of trnL-trnF sequence was1004sites, including29variation sites and12phylogenetic information sites, consisting2.89%and1.20%of the total length, respectively. The trnL-trnF sequence was much conserved and it was not expected to solve the systematic relationships among different Malus Mill, species in'taiyi'mountains area.
Although there was obvious difference between the two phylogenetic trees based on ITS and trnL-trnF sequences, It could be both proved that the formation and genetic background of taiyi'mountains malus was complicated, a lot of germplasms belong to the same species, but their hybrid parents may come from different species, even experienced many hybrids.
3. Four cpSSR primers ccmp2, ccmp4, ccmp5and cicp9were proved to be suitable for genetic diversity analysis of Malus Mill, species in'TaiYi'mountain area, with each primer pairs having good results. Totally177DNA loci was amplified from36samples for these4primer pairs, with a polymorphism ratio of76.27%. UPGMA analysis showed that36Malus Mill,species sampled in this research can be totally distinguished using the method of cpSSR mark. With these36samples, two categories can be obtained at similarity coefficient level of1.82,Malus baccata and2subspecies of Malus hupehensis, Malus hupehensis var. pinyiensis and Malus hupehensis var. taishanesis were in the same branch. Malus micromalus, Malus prunifolia, Malus asitica and Malus domestica were in another branch, showing the interaction cluster-like, they were basically the same with the result of the system tree constructed by ITS sequences of nuclear genome, also supported the view that cultivated species and wild species separated with each other, it also showed that the genetic relationship among cultivars were complex.
The results of this study preliminary showed that the quantitative classification system can effectively solve the Chinese Malus Mill,classification and identification problem, cultivated species and wild species can be separated, the ancestor of cultivated species should be Malus seversii, the formation and genetic background of cultivated species were complex, and their parents were various. Malus Mill, of taiyi"mountain mainly including5species,10types. The results of the system tree construted by ITS sequences of nuclear genome and the system tree by cpSSR were basically same, could support the phenotypic classification results. The system tree constructed by trhL-trnF sequence can not slove the systematic relationship of tai yi'mountain.
引文
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