百合属（Lilium）植物亲缘关系及通江百合（L.sargentiae）遗传多样性研究
|
摘要
百合属由Linnaeus于1753年建立,是一个北温带的属,北半球广泛分布。自1836年Endlicher最早对百合属属下类群进行划分开始,百合属的分类争议从未停止过,著名学者Baker、Wilson等都曾对百合属下类群划分提出过自己的观点,他们的观点因存在争议并未被广泛认可,直至1949年Comber依据15个形态特征建立了百合属的分类系统,此系统一度被大多数学者所接受并延用至今。分子系统学兴起后,研究者曾对日本、朝鲜等区域的百合属植物进行过系统发育研究,他们的部分研究结果与Comber的系统不完全相同,其中围绕毛百合、湖北百合的系统位置,卷瓣组的设立等问题的争议较多。中国是百合属的分布中心之一,百合属植物在中国分布区内发生着强烈的分化,而针对中国范围内百合属植物的较为全面的分子系统学研究几乎从未开展过,已有的研究只是针对部分区域范围内的百合资源遗传多样性或部分野生种与栽培种的亲缘关系等进行过探索。利用新兴的技术较为全面的研究我国百合属原生种的亲缘关系对于解决目前百合属下类群划分的一些热点问题是有借鉴意义的,也是十分必要的。而了解我国百合原生种的资源特点对于资源合理利用、发展花卉产业也是十分迫切的。
本文主要以收集到的百合属23种2变种为试材,采用SRAP和AFLP两种分子标记技术研究了野生百合种质间的亲缘关系,探讨了有关百合属下类群划分的几个有争议的问题。同时,针对重庆范围内通江百合资源相对较为丰富的特点,采用SRAP标记技术分析了10个居群的遗传多样性和居群遗传结构,提出了种质资源利用与保护建议。主要的研究结果如下
1百合属植物遗传多样性丰富,种间差异明显,遗传基础广。
采用SRAP和AFLP两种分子标记技术分析了我国百合23个原生种,2变种共25份种质的遗传多样性。19对SRAP引物对23份供试材料的基因组DNA进行扩增,共得到清晰条带236条,多态性条带233条,多态性条带比率为98.72%,遗传相似系数变化范围为0.3750～0.7679。8对AFLP引物组合对百合属植物25份材料共扩增出条带395条,其中多态性条带368条,平均多态性条带比率为93.12%,8对引物的多态性条带比率都在90%以上,百合属内相似系数的变化范围为0.4417～0.8126。两种标记的结果都显示我国百合资源遗传多样性丰富,种间差异明显,基因资源丰富。
2分子系统学研究结果与形态分类结果部分吻合。
基于SRAP和AFLP两种分子标记技术共得到2份系统发育与亲缘关系树状图,与百合属形态分类观点部分吻合。SRAP和AFLP分析得到的轮叶组、百合组聚类结果与形态分类观点大致相符,SRAP聚类结果显示轮叶组种质虽然聚在了一起,但并不是自成一支。卷瓣组分成了几个支系。毛百合与卷丹聚在一起,未成独立一支,也没同钟花组其他种质聚在一起。AFLP聚类结果与我国传统分类观点大致相符,供试25份种质被分成了4组,轮叶组和百合组种质聚类结果与传统形态分类结果大部分吻合,卷瓣组,钟花组聚类结果与传统分类观点不尽相同。两种标记的聚类分析说明虽然依据形态性状可以将百合属植物分开,可以进行属下类群划分,但是我们研究结果同时也证实,单纯依据形态特征的分类是有争议的,因为有些种虽然形态特征相似但是其亲缘关系却很远,比如湖北百合与南川百合等。
3卷瓣组的组成成分复杂,组内分化强烈。
卷瓣组以花冠不为钟型,花被片反卷,雄蕊向四周张开而与其它组明显分开。本研究分析了百合属卷瓣组11种2变种13份种质,各种质间除具上述共性之外,在花色,花瓣基部流苏状突起的有无、叶型、节间长度,植株高矮等形态特征上差别明显。SRAP聚类结果表明除湖北百合外,余下10种分别于Ⅲ、Ⅳ类群下聚成3支。AFLP聚类结果也显示除湖北百合外,余下的10种2变种在Ⅱ、Ⅲ类下聚成不同支系,两种标记的聚类结果都显示卷瓣组的种质未能聚成独立的一类,而是分成了几个亚类群,说明卷瓣组组内分化强烈,是个非常复杂的组,可能为多元起源。
4湖北百合与百合组亲缘关系较近。
Comber认为湖北百合与南川百合外部形态特征相似,将二者都划在了卷瓣组下,本研究两种分子标记的聚类结果均显示,湖北百合未能与卷瓣组种质聚在一起,而是与百合组的种质聚成一支,说明湖北百合与百合组(喇叭花组)亲缘关系较近,尤其是与百合组下的岷江百合亚组亲缘关系较近,而与卷瓣组亲缘关系较远。
5卷瓣组与钟花组存在基因交流。
《中国植物志》中卷瓣组和钟花组各自为独立的组,两组种质在花型上差异明显,但本研究显示,无论是SRAP标记还是AFLP标记都未能将两组完全分开,卷瓣组的部分种质,尤其是卷丹常和钟花组种质聚在一起,说明两组之间存在基因交流和渗透。
6青岛百合与东北百合亲缘关系较近。
青岛百合与东北百合均为轮生叶序,花具斑点,在本研究基于SRAP和AFLP标记分析的聚类图中,两者都聚在了一起,说明两者亲缘关系较近,同时也表明传统分类中把二者划归入同一组的正确性。
7毛百合系统位置复杂。
Comber把毛百合列在了单种组毛百合组内,我国传统分类观点认为毛百合与渥丹形态特征相似,所以把二者归入钟花组,表明毛百合的系统位置是有争议的。本研究显示基于SRAP的聚类结果,毛百合没有和钟花组的玫红百合、渥丹百合聚在一起,基于AFLP的聚类结果显示,毛百合和钟花组的其它种质聚在一起,进一步验证了毛百合的系统位置的不确定性,需要进一步的研究、探索。
8通江百合遗传多样性丰富,居群内变异大于居群间变异。
15对SRAP引物对通江百合重庆分布区内10个居群的100个样品的基因组DNA扩增结果表明,通江百合在百合属中群体水平和物种水平上的遗传多样性均较丰富,居群水平的遗传多样性高于其他的单子叶植物。异花传粉、无性繁殖系数低可能是通江百合遗传多样性高于其他单子叶植物的主要原因。通江百合居群内变异大于居群间变异。通江百合居群发生了较大程度的遗传变异,居群间遗传分化系数Gst为0.3672,高于单子叶植物平均分化系数(Gst=0.2310)和多年生草本植物的平均分化系数(Gst=0.2330)。异交,基因流受阻、地理阻隔作用和较小的种群规模是是导致通江百合产生遗传分化的主要原因。
The genus Lilium, established by Linnaeus in 1753, was widely distributed in the Northern Hemisphere. The disputes have never stopped since Endlicher earliest proposed the classification (taxonomy) of the infrageneric in 1836. Baker and Wilson, famous scholar, put forward views on the classification, which was not universally acknowledged. In 1949, Comber established the classification of the genus Lilium according to the morphological characteristics, which was accepted by most of the scholars up to now. But some researchers'late results according to systematic development were not identical with Comber's, the main difference was the systematics position of L. dauricum, L. henryi and Sect. Sinomartagon. The great differentiation of Lilium occurred in China, where was one of the distribution center. Comprehensive molecular systematics of Lilium distributed in China has not been reported. It is necessary to study the phylogenetic relationship to solve the infrageneric of the genus Lilium, also urgent to utilize resources reasonable and develop flower culture industry.
23 lily wild species and 2 varietyies were chosen for the present study, two molecular markers, sequence-related amplified polymorphism (SRAP) and amplified fragment length polymorphism (AFLP), were used to probe the infrageneric phylogenetic relationship and the classification of Lilium. The population genetic diversity and genetic structure of L. sargentiae were analyzed in view of rich resources in Chongqing, and resources utilization and protection were also proposed. The main results were as follows.
1. The genus Lilium was rich in genetic diversity, the difference was significant on the level of species, genetic foundation was extensive
23 lily wild species,2 varieties were chosen to probe the infrageneric phylogenetic relationship based on SRAP and AFLP. DNA extracted from 23 lily wild species was PCR amplified, and 19 SRAP primer combinations were used. A total of 236 clear DNA bands were amplified,233 of which were polymorphic, the proportion was 98.72% with average 12.3 polymorphic loci per SRAP primer combination. The genetic similarities ranged from 0.3750 to 0.7679. DNA extracted from 25 lily species was PCR amplified, using 8 AFLP primer combinations, a total of 395 clear DNA bands were amplified,368 of which were polymorphic, the average proportion was 93.12%, the similarity index ranged from 0.4417 to 0.8126. The two molecular markers showed that the genetic diversity was rich, the difference was significant on the level of species, and germplasm was rich in genus Lilium in China.
2. Molecular systematics and morphological classification were partially identical
Two phylogenetic trees were constructed based on two molecular markers. The results were partially consistent with morphology classification. The clustering of Sect. Martagon and Sect. Leucolirion was basically identical with morphology classification based on SRAP and AFLP. SRAP dendrogram showed that the germplasms of Sect. Martagon clustered together, Sect. Sinomartagon comprised a few clades, L. dauricum and L. lancifolium clustered together, separated with the germplasms of Sect. Lophophorum. AFLP dendrogram showed that the clustering results were mostly consistent with traditional taxonomy, four major clades including 25 species were delimited, the clustering results of Sect. Martagon and Sect. Leucolirion were mostly consistent with traditional taxonomy, whereas the clustering results of Sect. Sinomartagon and Sect. Lophophorum were different with traditional taxonomy. It demonstrated that it's difficult to classify Lilium only based on the morphological characteristics, because phylogenetic relationship was distant despite that morphological characteristics were similar between some species such as L. henryi and L. rosthornii.
3. The germplasms of Sect. Sinomartagon were complex, the differentiation was great in the section
Sect. Sinomartagon was characterized by no bell-shaped corolla, revolute tepals, open stamen. Using 11 wild species and 2 varieties of Sect. Sinomartagon as material, the morphological characteristics was analyzed, flower colour, fringed outgrowth on the petal base, leaf type, internode length and plant height were apparently different. SRAP dendrogram showed that 3 clades including 10 species were delimited under group III and IV except L. henryi. AFLP dendrogram also showed that 10 wild species and 2 varieties were divided into different clades under group II and III except L. henryi. Two molecular markers showed that Sect. Sinomartagon germplasms were not clustered into one group, but divided into different clades, it demonstrated that the differentiation occurred in Sect. Sinomartagon, the origin of Sect. Sinomartagon could be multielement.
4. L. henryi and Sect. Leucolirion were the closely related systems
Comber classified L. henryi and L. rosthornii into Sect. Sinomartagon according to the similarity of the morphological characteristics. In the present study two molecular markers showed that L. henryi was not clustered into Sect. Sinomartagon germplasms, but clustered one clade into Sect. Leucolirion. It suggested that L. henryi was closely related with Sect. Leucolirion, especially with L. regale subsection, and distantly related with Sect. Sinomartagon.
5. Gene exchanges existed between Sect. Lophophorum and Sect. Sinomartagon
Sect. Sinomartagon and Sect. Lophophorum were separate section in "Flora of China" because of the apparent difference of flower shape. The result showed that two sections germlpasms could not be divided separately by two molecular markers dendrogram, it suggested that gene exchanges existed between the two sections.
6. L. tsingtauense and L. Distichum were the closely related systems
Leaves of L. tsingtauense and L. Distichum were whorled, flowers were spotted, both two phylogenetic trees showed that L. tsingtauense and L. Distichum were clustered into one group, it suggested that L. tsingtauense and L. Distichum were the closely related systems, which was identical with morphology classification.
7. Systematics position of L. dauricum was controversial
Although L. dauricum was classified into Sect. Daurolirion which comprised only one species by Comber, L. dauricum and L. concolor were classified into Sect. Lophophorum, because morphological characteristics of L. dauricum and L. concolor were similar in traditional taxonomy in China. The result showed that L. dauricum was not clustered into one group with L. amoenum and L. concolor of Sect. Lophophorumon the basis of SRAP dendrogram, L. dauricum was clustered into other germplasms on the basis of AFLP dendrogram, which proved that systematics position of L. dauricum is uncertain.
8. L. sargentiae was rich in genetic diversity
100 wild lily samples were collected from 10 populations, and 15 SRAP primer combinations were used for determination. The results showed that L. sargentiae was rich in genetic diversity on the level of populations and species of genus Lilium, the genetic diversity within populations exceeded other monocotyledonous plants, low coefficient of cross-pollination and asexual reproduction might be the main reason. High degree of variation of the population of L. sargentiae occurred, the variation within populations exceeded among populations, the restraint of gene flow, geography obstructs and small population were the main reason of genetic differentiation in L. sargentiae.
本文主要以收集到的百合属23种2变种为试材,采用SRAP和AFLP两种分子标记技术研究了野生百合种质间的亲缘关系,探讨了有关百合属下类群划分的几个有争议的问题。同时,针对重庆范围内通江百合资源相对较为丰富的特点,采用SRAP标记技术分析了10个居群的遗传多样性和居群遗传结构,提出了种质资源利用与保护建议。主要的研究结果如下
1百合属植物遗传多样性丰富,种间差异明显,遗传基础广。
采用SRAP和AFLP两种分子标记技术分析了我国百合23个原生种,2变种共25份种质的遗传多样性。19对SRAP引物对23份供试材料的基因组DNA进行扩增,共得到清晰条带236条,多态性条带233条,多态性条带比率为98.72%,遗传相似系数变化范围为0.3750～0.7679。8对AFLP引物组合对百合属植物25份材料共扩增出条带395条,其中多态性条带368条,平均多态性条带比率为93.12%,8对引物的多态性条带比率都在90%以上,百合属内相似系数的变化范围为0.4417～0.8126。两种标记的结果都显示我国百合资源遗传多样性丰富,种间差异明显,基因资源丰富。
2分子系统学研究结果与形态分类结果部分吻合。
基于SRAP和AFLP两种分子标记技术共得到2份系统发育与亲缘关系树状图,与百合属形态分类观点部分吻合。SRAP和AFLP分析得到的轮叶组、百合组聚类结果与形态分类观点大致相符,SRAP聚类结果显示轮叶组种质虽然聚在了一起,但并不是自成一支。卷瓣组分成了几个支系。毛百合与卷丹聚在一起,未成独立一支,也没同钟花组其他种质聚在一起。AFLP聚类结果与我国传统分类观点大致相符,供试25份种质被分成了4组,轮叶组和百合组种质聚类结果与传统形态分类结果大部分吻合,卷瓣组,钟花组聚类结果与传统分类观点不尽相同。两种标记的聚类分析说明虽然依据形态性状可以将百合属植物分开,可以进行属下类群划分,但是我们研究结果同时也证实,单纯依据形态特征的分类是有争议的,因为有些种虽然形态特征相似但是其亲缘关系却很远,比如湖北百合与南川百合等。
3卷瓣组的组成成分复杂,组内分化强烈。
卷瓣组以花冠不为钟型,花被片反卷,雄蕊向四周张开而与其它组明显分开。本研究分析了百合属卷瓣组11种2变种13份种质,各种质间除具上述共性之外,在花色,花瓣基部流苏状突起的有无、叶型、节间长度,植株高矮等形态特征上差别明显。SRAP聚类结果表明除湖北百合外,余下10种分别于Ⅲ、Ⅳ类群下聚成3支。AFLP聚类结果也显示除湖北百合外,余下的10种2变种在Ⅱ、Ⅲ类下聚成不同支系,两种标记的聚类结果都显示卷瓣组的种质未能聚成独立的一类,而是分成了几个亚类群,说明卷瓣组组内分化强烈,是个非常复杂的组,可能为多元起源。
4湖北百合与百合组亲缘关系较近。
Comber认为湖北百合与南川百合外部形态特征相似,将二者都划在了卷瓣组下,本研究两种分子标记的聚类结果均显示,湖北百合未能与卷瓣组种质聚在一起,而是与百合组的种质聚成一支,说明湖北百合与百合组(喇叭花组)亲缘关系较近,尤其是与百合组下的岷江百合亚组亲缘关系较近,而与卷瓣组亲缘关系较远。
5卷瓣组与钟花组存在基因交流。
《中国植物志》中卷瓣组和钟花组各自为独立的组,两组种质在花型上差异明显,但本研究显示,无论是SRAP标记还是AFLP标记都未能将两组完全分开,卷瓣组的部分种质,尤其是卷丹常和钟花组种质聚在一起,说明两组之间存在基因交流和渗透。
6青岛百合与东北百合亲缘关系较近。
青岛百合与东北百合均为轮生叶序,花具斑点,在本研究基于SRAP和AFLP标记分析的聚类图中,两者都聚在了一起,说明两者亲缘关系较近,同时也表明传统分类中把二者划归入同一组的正确性。
7毛百合系统位置复杂。
Comber把毛百合列在了单种组毛百合组内,我国传统分类观点认为毛百合与渥丹形态特征相似,所以把二者归入钟花组,表明毛百合的系统位置是有争议的。本研究显示基于SRAP的聚类结果,毛百合没有和钟花组的玫红百合、渥丹百合聚在一起,基于AFLP的聚类结果显示,毛百合和钟花组的其它种质聚在一起,进一步验证了毛百合的系统位置的不确定性,需要进一步的研究、探索。
8通江百合遗传多样性丰富,居群内变异大于居群间变异。
15对SRAP引物对通江百合重庆分布区内10个居群的100个样品的基因组DNA扩增结果表明,通江百合在百合属中群体水平和物种水平上的遗传多样性均较丰富,居群水平的遗传多样性高于其他的单子叶植物。异花传粉、无性繁殖系数低可能是通江百合遗传多样性高于其他单子叶植物的主要原因。通江百合居群内变异大于居群间变异。通江百合居群发生了较大程度的遗传变异,居群间遗传分化系数Gst为0.3672,高于单子叶植物平均分化系数(Gst=0.2310)和多年生草本植物的平均分化系数(Gst=0.2330)。异交,基因流受阻、地理阻隔作用和较小的种群规模是是导致通江百合产生遗传分化的主要原因。
The genus Lilium, established by Linnaeus in 1753, was widely distributed in the Northern Hemisphere. The disputes have never stopped since Endlicher earliest proposed the classification (taxonomy) of the infrageneric in 1836. Baker and Wilson, famous scholar, put forward views on the classification, which was not universally acknowledged. In 1949, Comber established the classification of the genus Lilium according to the morphological characteristics, which was accepted by most of the scholars up to now. But some researchers'late results according to systematic development were not identical with Comber's, the main difference was the systematics position of L. dauricum, L. henryi and Sect. Sinomartagon. The great differentiation of Lilium occurred in China, where was one of the distribution center. Comprehensive molecular systematics of Lilium distributed in China has not been reported. It is necessary to study the phylogenetic relationship to solve the infrageneric of the genus Lilium, also urgent to utilize resources reasonable and develop flower culture industry.
23 lily wild species and 2 varietyies were chosen for the present study, two molecular markers, sequence-related amplified polymorphism (SRAP) and amplified fragment length polymorphism (AFLP), were used to probe the infrageneric phylogenetic relationship and the classification of Lilium. The population genetic diversity and genetic structure of L. sargentiae were analyzed in view of rich resources in Chongqing, and resources utilization and protection were also proposed. The main results were as follows.
1. The genus Lilium was rich in genetic diversity, the difference was significant on the level of species, genetic foundation was extensive
23 lily wild species,2 varieties were chosen to probe the infrageneric phylogenetic relationship based on SRAP and AFLP. DNA extracted from 23 lily wild species was PCR amplified, and 19 SRAP primer combinations were used. A total of 236 clear DNA bands were amplified,233 of which were polymorphic, the proportion was 98.72% with average 12.3 polymorphic loci per SRAP primer combination. The genetic similarities ranged from 0.3750 to 0.7679. DNA extracted from 25 lily species was PCR amplified, using 8 AFLP primer combinations, a total of 395 clear DNA bands were amplified,368 of which were polymorphic, the average proportion was 93.12%, the similarity index ranged from 0.4417 to 0.8126. The two molecular markers showed that the genetic diversity was rich, the difference was significant on the level of species, and germplasm was rich in genus Lilium in China.
2. Molecular systematics and morphological classification were partially identical
Two phylogenetic trees were constructed based on two molecular markers. The results were partially consistent with morphology classification. The clustering of Sect. Martagon and Sect. Leucolirion was basically identical with morphology classification based on SRAP and AFLP. SRAP dendrogram showed that the germplasms of Sect. Martagon clustered together, Sect. Sinomartagon comprised a few clades, L. dauricum and L. lancifolium clustered together, separated with the germplasms of Sect. Lophophorum. AFLP dendrogram showed that the clustering results were mostly consistent with traditional taxonomy, four major clades including 25 species were delimited, the clustering results of Sect. Martagon and Sect. Leucolirion were mostly consistent with traditional taxonomy, whereas the clustering results of Sect. Sinomartagon and Sect. Lophophorum were different with traditional taxonomy. It demonstrated that it's difficult to classify Lilium only based on the morphological characteristics, because phylogenetic relationship was distant despite that morphological characteristics were similar between some species such as L. henryi and L. rosthornii.
3. The germplasms of Sect. Sinomartagon were complex, the differentiation was great in the section
Sect. Sinomartagon was characterized by no bell-shaped corolla, revolute tepals, open stamen. Using 11 wild species and 2 varieties of Sect. Sinomartagon as material, the morphological characteristics was analyzed, flower colour, fringed outgrowth on the petal base, leaf type, internode length and plant height were apparently different. SRAP dendrogram showed that 3 clades including 10 species were delimited under group III and IV except L. henryi. AFLP dendrogram also showed that 10 wild species and 2 varieties were divided into different clades under group II and III except L. henryi. Two molecular markers showed that Sect. Sinomartagon germplasms were not clustered into one group, but divided into different clades, it demonstrated that the differentiation occurred in Sect. Sinomartagon, the origin of Sect. Sinomartagon could be multielement.
4. L. henryi and Sect. Leucolirion were the closely related systems
Comber classified L. henryi and L. rosthornii into Sect. Sinomartagon according to the similarity of the morphological characteristics. In the present study two molecular markers showed that L. henryi was not clustered into Sect. Sinomartagon germplasms, but clustered one clade into Sect. Leucolirion. It suggested that L. henryi was closely related with Sect. Leucolirion, especially with L. regale subsection, and distantly related with Sect. Sinomartagon.
5. Gene exchanges existed between Sect. Lophophorum and Sect. Sinomartagon
Sect. Sinomartagon and Sect. Lophophorum were separate section in "Flora of China" because of the apparent difference of flower shape. The result showed that two sections germlpasms could not be divided separately by two molecular markers dendrogram, it suggested that gene exchanges existed between the two sections.
6. L. tsingtauense and L. Distichum were the closely related systems
Leaves of L. tsingtauense and L. Distichum were whorled, flowers were spotted, both two phylogenetic trees showed that L. tsingtauense and L. Distichum were clustered into one group, it suggested that L. tsingtauense and L. Distichum were the closely related systems, which was identical with morphology classification.
7. Systematics position of L. dauricum was controversial
Although L. dauricum was classified into Sect. Daurolirion which comprised only one species by Comber, L. dauricum and L. concolor were classified into Sect. Lophophorum, because morphological characteristics of L. dauricum and L. concolor were similar in traditional taxonomy in China. The result showed that L. dauricum was not clustered into one group with L. amoenum and L. concolor of Sect. Lophophorumon the basis of SRAP dendrogram, L. dauricum was clustered into other germplasms on the basis of AFLP dendrogram, which proved that systematics position of L. dauricum is uncertain.
8. L. sargentiae was rich in genetic diversity
100 wild lily samples were collected from 10 populations, and 15 SRAP primer combinations were used for determination. The results showed that L. sargentiae was rich in genetic diversity on the level of populations and species of genus Lilium, the genetic diversity within populations exceeded other monocotyledonous plants, low coefficient of cross-pollination and asexual reproduction might be the main reason. High degree of variation of the population of L. sargentiae occurred, the variation within populations exceeded among populations, the restraint of gene flow, geography obstructs and small population were the main reason of genetic differentiation in L. sargentiae.
引文
陈郦俊,马虹,左开井,唐克轩,唐东芹,曾丽.利用RAPD标记分析百合种质的遗传多样性.上海交通大学学报(农业科学版),2009,27(5)：475-479
陈俊愉,程继珂.中国花卉[M].上海：上海文化出版社.1999
陈之端.分类学中新的增长点----植物分子系统学.植物杂志,2001,(5)：33～34
陈机译,P·玛海莆瓦里.著.被子植物胚胎学引论.北京：科学出版社.1966
傅立国.中国高等植物13卷.山东：青岛出版社.2002,118-133
戴小红,张延龙,牛立新.百合属4种植物的核型研究.西北植物学报,2006,26(1)：50-56
梁松筠.百合科(狭义)植物的分布区对中国植物区系研究的意义.植物分类学报,1995,33(1)：25～27
葛颂.酶电泳资料和系统与进化植物学研究综述.武汉植物研究,1994,12(1)：71～84
国家药典委员会.中国药典[s],北京：化学工业出版社.2005
何林,张洁,郭启高,梁国鲁.东方百合Tiber多倍体诱导及其快繁研究.西南农业大学学报,2006,28(6)：945～949
高丽霞,胡秀,刘念,黄邦海,李正军,李严.中国姜花属基于SRAP分子标记的聚类分析.植物分类学报,2008,46(6)：899-905
黄济明.百合的组织培养和试管内诱发多倍体试验.园艺学报,1983,10(2)：125-127
黄济明,赵晓艺,张国民,倪跃元.玫红百合为亲本育成百合种间杂种.园艺学报,1990,17(2)：153～157
何纯莲,向建南,尹霞,任凤莲.百合中秋水仙碱超临界流体萃取工艺的研究.湖南大学学报(自然科学版),2003,30(6)：37-40
何纯莲,杨小红,黄浩,杨球桢.百合多糖的抗疲劳作用.湖南师范大学学报,2009,6(3)：9-15
何丽萍,李龙根,庞祖云.野生泸定百合种子休眠和萌发习性的研究.种子,2010(29)：46-49
何显静,立标,虞泓.紫斑百合居群生物学研究.云南大学学报(自然科学版.),2003(25)(增刊)：78-83
何显静,李标,周利,虞泓.云南5种百合的形态多样性研究.黑龙江农业科学,2003(6)：26-28
胡文彦,段金廒,钱大玮,王大为.卷丹化学成分研究.中国中药杂志,2007,32(16)：1656-1659
胡敏敏,蔡宝昌,张志杰,张慧芳.百合多糖的药效学研究.中药新药与临床药理,2007,18(2)：107-109
蒋细望,司怀军.百合的组织培养技术综述.湖北农业科学,2004(1)：78-81.
贾月慧,张克中,张文学,张福锁.辐照花粉辅助授粉对百合远缘杂交结实的影响及杂交早期分子鉴定.核农学报,2010,24(1)：25～30
纪海珊.温度对百合划分特性及授粉行为之研究。中华农学会报,2001,2：65～72
柯贤锋,智丽,畦顺照等.野百合AFLP反应体系建立及优化.西南大学学报,2007,29(12)：100-103
龙雅宜,张金政,张兰年.百合——球根花卉之王.金盾出版社,1999.
龙雅宜,张金政.百合属植物资源的保护和利用.植物资源与环境,1998,7(1)：40-44
路安民.种子植物科属地理.北京：科学出版社,1999：542-564.
刘祥军,杨利平,刘政,李瑞军,刘明远.毛百合与山丹幼苗发育的比较形态学.植物研究,1992,12(3)：277～284
刘菊华,余志强,徐碧云,郑思乡,刘志敏.龙牙百合的植株再生与遗传转化.分子植物育种,2003,1(4)：465-474
刘全儒,郭延平,于明,译.植物系统分类学(Singh G):综合理论及方法[M].北京：化学工业出版社,2008
刘光欣,胡凤荣,席梦利等.泸定百合根尖染色体C-带分析.南京林业大学学报(自然科学版),2007,31(6)：91-93
李俊清.植物遗传多样性保护及其分子生物学研究方法.生态学杂志,1994,13(6)：27～33
李新华,弥曼,李汾,任利君,王多宁,杨颖.百合多糖免疫调节作用的实验研究.现代预防医学,2010,37(14)：2708-2709
李恩香,黄玉琴,蒋满英等.江西龙牙百合种质资源遗传多样性研究.园艺学报,2010,37(5)：811-816
罗凤霞,年玉欣,孙晓梅、李智辉,王贤.四种授粉方法对切花百合不同杂交组合结籽量的影 响.园艺学报,2005,32(4)：729-731
罗凤霞,年玉欣,孙晓梅、徐艇,王贤.贮藏温度对不同发育期东方百合花粉生命力的影响.沈阳农业大学学报,2005,36(3)：298～300
陆春霞,韦鹏霄,岑秀芬,梁贵秋.百合多倍体的研究进展.广西农业卡学,2009,40(1)：76～80
陆长旬,黄善武,梁励,赵祥云,张克忠.辐射亚洲百合鳞茎(M1)染色体畸变研究.核农学报,2002,16(3)：148～151
林祖军,孙继霞,迮福慧,崔广琴,辛国胜,刘学庆,于建.电子束在花卉诱变育种上的应用.核农学报,2002,16(6)：351-354
连雪斌.兰州百合多倍体诱导试验报告.甘肃农业科技,1995,6：14～15
贾琳,卢巧玲,虞泓.泸定百合遵义居群与牟定居群的遗传多样性研究.云南大学学报(自然科学版)(增刊),2003(25)：84-90
马克平.论生物多样性的概念.生物多样性,1993,1(1)：20～22
毛钧,张明宇,虞泓.泸定百合普洱居群遗传与变异研究.云南大学学报(自然科学版),2003(25)(增刊)：91-96
穆鼎.观赏百合：生理、栽培、种球生产与育种[M].北京：中国农业出版社,2005
倪武,周明行,王国俊.秋水仙碱拮抗肿瘤坏死因子的肝毒性作用.第二军医大学学报,1998,19(4)：353～355
彭隆金.百合资源与栽培[M].云南民族出版社.2002
曲伟红,周日宝,童巧珍,贺又舜,罗目和,高静.百合的化学成分研究进展.湖南中医药导报,2004,10(3)：75～76
屈云慧,吴丽芳,杨春梅.百合幼胚培养成株体系的建立.云南农林科技,2002(2)：9-13
荣立苹,雷家军,王志刚.东北地区野生百合遗传多样性的RAPD分析.江苏农业学报,2009,25(4)：843～846.
荣立苹,雷家军.东北地区野生百合数量分类研究.植物遗传资源学报,2010,11(1)：99-102
荣丽苹,雷家军,郑洋,高玉福.东北地区野生百合的核型研究.吉林农业大学学报,2009(6)：711-716
孙启时,石猛,付沛云.东北百合属植物的研究.植物研究,1989,9(3)：133-141
孙晓梅,崔文山,刘艳芬.不同授粉方法对新中心和普瑞头两种百合杂交结实影响的研究.辽宁林业科技,2002(2)：6-10
孙晓泽.沈阳地区11个东方百合品种几种同工酶的研究.科技情报开发与经济,2007,17(13)：203～204
四川植物志编辑委员会.四川植物志(第四卷).成都：四川人民出版社,1989
田爱梅,郑日如,王国强,陈发菊,曹家树.中国野生百合资源的研究保护和利用.安徽农业科学,2007,35(31)：9987-9990
王中仁.植物等位酶分析.北京：科学出版社,1996
王瑞波,何平,胡世俊等.南川百合与泸定百合种群格局关联维数的比较研究.重庆林业科技,2008(1)：26-34
王瑞波,张燕平,胡世俊,何平,张春平,邓洪平.两种百合种群空间分布格局对高温干旱气候的响应.林业科学研究,2009,22(2)：249-255
王欣,雷家军,李明旭.渥丹百合不同居群核型研究.中国农学通报,2006,22()：281-284
王欣,雷家军,张丽娜,韩凌.渥丹百合种内居群形态多样性研究.沈阳农业大学学报,2007,38(6)：796-800
王亚斌,杨国锋,安俊学.百合幼胚离体培养的研究[J].沈阳农业大学学报,2002,33(4)：249-251
王红霞,杨宝胜.通江百合形态变异式样研究.河南科技学院学报,2005,25(3)：8-10
王红霞,胡琼华,陈晓兰.通江百合的组织培养.植物生理学通讯,2000,36(2)：132-133
吴祝华,施季森,席梦利,刘光欣.百合种质资源花粉形态及亲缘关系研究.浙江林学院学报,2007,24(4)：406～412
吴祝华,徐进,宋春雷,席梦利,刘光欣,施季森.宜昌百合、泸定百合核型分析.林业科技开发,2007,27(3)：29～31
吴刚,虞泓,梁淑云等.云南川百合DALP遗传变异分析.广东农业科学,2009,(7)：52-55
徐宏发,王静.分子系统学研究进展.生态学杂志,2001,20(3)：41-46
向地英,张延龙,牛立新.秦巴山区及毗邻地区野生百合的形态多样性研究.武汉植物学研究,2005,23(4)：385～388
谢树贵.44个百合品种同工酶遗传多样性研究.西部林业科学,2009,38(4)：27～35
杨利平,丁冰,刘香环,张敩方.东北百合属植物的细胞遗传多样性.东北林业大学学报,1996,24(5)：19～33
杨利平,张敩方,丁冰等.条叶百合×王百合种间杂种的育成.东北林业大学学报,1998,26(3)：37-43
杨红霞,王宝胜.岷江百合居群核型变异研究.河南科技学院学报,2003(3)：40-43
易杨杰,张新全,黄琳凯,凌瑶,马啸,刘伟.野生狗牙根遗传多样性的SRAP分析.遗传,2008,30(1)：94～100
虞泓,黄瑞复,魏蓉城.川百合种内核型多样性研究.云南植物研究,1996,18(增刊)：1～14
虞泓,王红霞,游丹.泸定百合居群染色体形态研究.云南大学学报(自然科学版),2000,22(1)：60-67
于晓英,吴铁明,倪沛,杨梦玲,彭尽晖,于江南.百合幼胚的离体培养和植株再生.湖南农业大学学报,2000,26(4)：286-288
姚青菊,任全尽,熊豫宁,杨军.岷江流域网百合与通江百合资源特征的初步研究.江苏农业科学,2007(5)：245～247
余桂红,马鸿翔,周淼平,黄益洪,陆维忠.百合农杆菌介导的遗传转化受体系统的建立.江西农业科学,2004,16(2)：15～19
杨利平,马宪红,丁冰等.百合花卉种间杂交种的培育.东北林业大学学报,1997,25(1)：29-33
鄢家俊,白使且,张新全.青藏高原东南缘老芒麦自然居群遗传多样性的SRAP和SSR分析.草业学报,2010,19(4)：122-134
杨秀伟,吴云山,崔育新,刘雪辉,肖诗鹰.卷丹中新甾体化合物的分离和鉴定.药学学报,2002,37(11)：863-866
岳玲,雷家军,王欣.辽宁4种(Lilium ssp.)野生百合的核型研究.辽宁农业科学,2006(4).5-8
张炜茹,张启翔.中国百合种质资源调查与引种驯化研究进展.安徽农业科学,2008,36(19)：8065～8067
赵宇,刘庆华等.青岛百合鳞叶扦插繁殖技术研究.山东林业科技,2007(3)：16～18
张克中,贾月慧,张启翔,王洁琼.部分中国野生百合亲缘关系的AFLP分析.东北林业大学学报,2008,36(2)：19～22
张克中,贾月慧,赵祥云,张启翔.部分中国野生百合亲缘关系的RAPD分析.东北林业大学学报,2006,34(6)：43-45
张克中,赵祥云,黄善武,陆长甸,梁励.辐射对百合鳞片生成不定芽及M1植株的影响.北京农学院学报,2002,17(4)：19～25
赵庆芳,马世荣,曾小英,李巧峡,崔艳.百合栽培品种资源的RAPD分析.兰州大学学报(自然科学版),2005,41(2)：30～33
赵祥云,王树栋,刘建斌..鲜切花百合生产原理及实用技术.北京：中国林业出版社.2005
赵祥云,陈新露,方海,张云芳.用RAPD标记评价百合品种间遗传关系.北京农学院学报,1995,(2)：58-63.
左志锐,穆鼎,高俊平,刘春.百合遗传多样性及亲缘关系的RAPD分析.园艺学报,2005,32(3)：468～472
中国科学院中国植物志编委会.中国植物志V o114.北京：科学出版社：1980,118-157
中国科学院昆明植物研究所.云南植物志(第七卷).北京：科学出版社,1997,789～821
中国科学院西北植物研究所.秦岭植物志(第一卷).北京：科学出版社,1976：362-368.-121
邹喻萍,葛颂,王晓东.系统与进化植物学中的分子标记.北京：科学出版社,2001
周厚高.几个百合品种遗传关系的等位酶分析.西南农业学报,1999(4)：92～95
周延清.DNA分子标记技术在植物研究中的应用.北京：化学工业出版社,2005
张西丽,周厚高,周焱,宁云芬.百合品种间的数量分类研究.广西植物,2000(4)：325-328
张兴翠,周昌华,殷家明,张建.药用百合的多倍体诱导及快速繁殖.西南农业大学学报,2003.25(1)：14-17
郑思乡,章海龙,董志渊,雷小云,胡秀,周丽娟.东方百合多倍体诱导及种球繁育的研究.西南农业大学学报,2004,20(3)：260～263
Asano Y. Fertility of a hybrid between distantly related species in Lilium. Cytologia,1984,49:447～ 456
AsanoY.A. numerical taxonomic study of the genus Lilium in Japan, Journal of the Faculty of Agriculture Hokkaido Univ,1986,62:333～341
Baranova MV. A synopsis of the system of the genus Lilium(Liliaceae) (in Russian), Bot Zhur,1988, 73:1319～1329.
Budak H.Shearman R.C. Parmaksiz I., Gaussoin R.E., Riordan T.P., Dewikat I. Molecular characterization of buffalograss germplasm using sequence-related amplified polymorphism markers, Theoretical and Applied Genetics,2004,108:328-334
Baker JG. The gardeners'chronicle.1871,104:1650
Comber H.F. A new classification of the genus Lilium, London:Royale Horticulture Society, Lily Yearbook.1949,86～105
Cozzolino S.Cafasso D, Pellegrino G. Fine-Scale phylogeographical analysis of Mediterranean Anacamptis palustris(Orchidaceae) populations based on chloroplast minisatellite and microsatellite variation. Molecular Ecology,2003,12:2783～2792
Diwan N, Mcltosh MS, Bauchan G R. Methods of developing a core collection of annual Medicago species.Theoritical and Applied Genetics,1995,90:755～761
Dubouzet J.G., Shinoda K. Phylogenetic analysis of the internal transcribed spacer region of Japanese Lilium species, Theoretical and Applied Genetics,1999,98:954～960
Ferriol M, Pico B, Nuez F. Genetic diversity of a germplasm collection of Cucurbita pepo using SRAP and AFLP markers. Theoretical and Applied Genetics,2003,107:271～282
Ferriol M, Pico B, Nuez F. Morphological and molecular diversity of a collection of Cucurbita maxima landraces. Journal of American Society for Horticultural Science,2004,129(1):60～69
Haruki K., Hosoki T., Nako Y., Ohta K. Possibility of classification in some species of Lilium by PCR-RFLP of ribulose-1,5-bisphosphate carboxylase large subunit (rbcl) gene and ribosomal RNA gene.Journal of the Japanese Society for Horticultural Science,1997,66(1):189～192
Haw SG.The taxonomy of Chinese lilies. In:The lilies of China. Portland:Timber Press.1986,57～ 69
Hamrick J L.Godt M J W. Effects of life history traits on genetic diversity in plant species Philosophical Transactions of the Royal Society B. Biological Sciences,1996,351:1291
Hayashi K. Native lilies in Japan 111, The natural habitats of Lilium alexandrae in Amami O-shima. J Phytogeogr Taxon,1983,31:111～119
Jefferson-Brown M. The lily species. In:The gardener's guide to growing lilies. Portland:Timber Press.1995,60～69
Li G., Quiros C.F. Sequence-related amplified polymorphism (SRAP), a new marker system based on a simple PCR reaction:its application to mapping and gene tagging in Brassica, Theoretical and Applied Genetics.2001,103:455～461
Leslie AC. The international lily register,3rd ed. Royal Horticultural Society, London.1982
Lighty R.W. Evolutionary trends in lilies. Lily Year Book.1968,31:40～44
Li L Y, Chen D X, Qin S Y, Li Q S, Zhong G Y. Studies on genetic relationship of seven species of Curculigo plant from China using SRAP. Journal of Chinese Materia Medica,2008,33:117～120
Lee, W. B., S. Y. Choi, Y. S. Kim. An application of random amplified polymorphic DNA(RAPD) to systematics of some species of Lilium in Korea. Kor. J. Plant Tax,1993,23:43～56 (in Korean)
Lee C.S., Kim S.C., Yeau S.H., and Lee N.S. Major lineages of the genus Lilium (Liliaceae) based on nrDNA sequences, with special emphasis on the Korean species. Journal of Plant Biology,2011, 54(3):159～171
Li En Xiang, Yi Sun, Qiu Ying Xiong. Phylogeography of two East Asian Species in Croomia (Stemonaceae) inferred from chloroplast DNA and ISSR fingerprinting variation. Molecular phylogenetics and Evolution,2008,49:702～714
Mitchell R M. Species DNA research report. NALS Q Bull.1998:52 (1):8-9
Nybom H, Bartish T V. Effects of life history traits and sampling strategies on genetics diversity estimates obtained with RAPD markers in plants. Perspectives in plant Ecology Evolution and Systematics,2000,3:93-114
Nishikawa T., Okazaki K., Arakawa K., Nagamine T. Phylogenetic analysis of Section Sinomartagon in genus Lilium using sequences of the internal transcribed spacer region in nuclear ribosomal DNA, Breeding Science,2001,51:39～46
ROBB SM. The culture of excised tissue from bulb scales of Lilium speciosum. J Exp. Bot,1957, 8:348～352
Sarg T M, B-dormiaty M M. Bishrhr M M. Thin layer chormatographic scanner, spectrophotometric and high-pherphomance liquid chromatographic methods for the determination of colchicines. Analyse.1989,114(5):575～578
Sun S J, Gao W, Lin S Q, Zhu J, Xie B G, Lin Z B. Analysis of genetic diversity in Ganoderma population with a novel molecular marker SRAP. Applied Microbiology and Biotechnology,2006, 72:537～543
Solbrig Q T, (ed) From genus to ecosysytems a reseach agenda for biodiversity, IUBS, Paris 1991
Wilson E H. The lilies of eastern Asia. Dulau and Co., London.1925
Wright S. Evolution and the genetics of populations. Chicargo:University of Chicargo press,1968
Yamagishi M. Detection of section-specific random amplified polymorphic DNA (RAPD) in Lilium, Theoretical and Applied Genetics,1995,91:830～835
陈俊愉,程继珂.中国花卉[M].上海：上海文化出版社.1999
陈之端.分类学中新的增长点----植物分子系统学.植物杂志,2001,(5)：33～34
陈机译,P·玛海莆瓦里.著.被子植物胚胎学引论.北京：科学出版社.1966
傅立国.中国高等植物13卷.山东：青岛出版社.2002,118-133
戴小红,张延龙,牛立新.百合属4种植物的核型研究.西北植物学报,2006,26(1)：50-56
梁松筠.百合科(狭义)植物的分布区对中国植物区系研究的意义.植物分类学报,1995,33(1)：25～27
葛颂.酶电泳资料和系统与进化植物学研究综述.武汉植物研究,1994,12(1)：71～84
国家药典委员会.中国药典[s],北京：化学工业出版社.2005
何林,张洁,郭启高,梁国鲁.东方百合Tiber多倍体诱导及其快繁研究.西南农业大学学报,2006,28(6)：945～949
高丽霞,胡秀,刘念,黄邦海,李正军,李严.中国姜花属基于SRAP分子标记的聚类分析.植物分类学报,2008,46(6)：899-905
黄济明.百合的组织培养和试管内诱发多倍体试验.园艺学报,1983,10(2)：125-127
黄济明,赵晓艺,张国民,倪跃元.玫红百合为亲本育成百合种间杂种.园艺学报,1990,17(2)：153～157
何纯莲,向建南,尹霞,任凤莲.百合中秋水仙碱超临界流体萃取工艺的研究.湖南大学学报(自然科学版),2003,30(6)：37-40
何纯莲,杨小红,黄浩,杨球桢.百合多糖的抗疲劳作用.湖南师范大学学报,2009,6(3)：9-15
何丽萍,李龙根,庞祖云.野生泸定百合种子休眠和萌发习性的研究.种子,2010(29)：46-49
何显静,立标,虞泓.紫斑百合居群生物学研究.云南大学学报(自然科学版.),2003(25)(增刊)：78-83
何显静,李标,周利,虞泓.云南5种百合的形态多样性研究.黑龙江农业科学,2003(6)：26-28
胡文彦,段金廒,钱大玮,王大为.卷丹化学成分研究.中国中药杂志,2007,32(16)：1656-1659
胡敏敏,蔡宝昌,张志杰,张慧芳.百合多糖的药效学研究.中药新药与临床药理,2007,18(2)：107-109
蒋细望,司怀军.百合的组织培养技术综述.湖北农业科学,2004(1)：78-81.
贾月慧,张克中,张文学,张福锁.辐照花粉辅助授粉对百合远缘杂交结实的影响及杂交早期分子鉴定.核农学报,2010,24(1)：25～30
纪海珊.温度对百合划分特性及授粉行为之研究。中华农学会报,2001,2：65～72
柯贤锋,智丽,畦顺照等.野百合AFLP反应体系建立及优化.西南大学学报,2007,29(12)：100-103
龙雅宜,张金政,张兰年.百合——球根花卉之王.金盾出版社,1999.
龙雅宜,张金政.百合属植物资源的保护和利用.植物资源与环境,1998,7(1)：40-44
路安民.种子植物科属地理.北京：科学出版社,1999：542-564.
刘祥军,杨利平,刘政,李瑞军,刘明远.毛百合与山丹幼苗发育的比较形态学.植物研究,1992,12(3)：277～284
刘菊华,余志强,徐碧云,郑思乡,刘志敏.龙牙百合的植株再生与遗传转化.分子植物育种,2003,1(4)：465-474
刘全儒,郭延平,于明,译.植物系统分类学(Singh G):综合理论及方法[M].北京：化学工业出版社,2008
刘光欣,胡凤荣,席梦利等.泸定百合根尖染色体C-带分析.南京林业大学学报(自然科学版),2007,31(6)：91-93
李俊清.植物遗传多样性保护及其分子生物学研究方法.生态学杂志,1994,13(6)：27～33
李新华,弥曼,李汾,任利君,王多宁,杨颖.百合多糖免疫调节作用的实验研究.现代预防医学,2010,37(14)：2708-2709
李恩香,黄玉琴,蒋满英等.江西龙牙百合种质资源遗传多样性研究.园艺学报,2010,37(5)：811-816
罗凤霞,年玉欣,孙晓梅、李智辉,王贤.四种授粉方法对切花百合不同杂交组合结籽量的影 响.园艺学报,2005,32(4)：729-731
罗凤霞,年玉欣,孙晓梅、徐艇,王贤.贮藏温度对不同发育期东方百合花粉生命力的影响.沈阳农业大学学报,2005,36(3)：298～300
陆春霞,韦鹏霄,岑秀芬,梁贵秋.百合多倍体的研究进展.广西农业卡学,2009,40(1)：76～80
陆长旬,黄善武,梁励,赵祥云,张克忠.辐射亚洲百合鳞茎(M1)染色体畸变研究.核农学报,2002,16(3)：148～151
林祖军,孙继霞,迮福慧,崔广琴,辛国胜,刘学庆,于建.电子束在花卉诱变育种上的应用.核农学报,2002,16(6)：351-354
连雪斌.兰州百合多倍体诱导试验报告.甘肃农业科技,1995,6：14～15
贾琳,卢巧玲,虞泓.泸定百合遵义居群与牟定居群的遗传多样性研究.云南大学学报(自然科学版)(增刊),2003(25)：84-90
马克平.论生物多样性的概念.生物多样性,1993,1(1)：20～22
毛钧,张明宇,虞泓.泸定百合普洱居群遗传与变异研究.云南大学学报(自然科学版),2003(25)(增刊)：91-96
穆鼎.观赏百合：生理、栽培、种球生产与育种[M].北京：中国农业出版社,2005
倪武,周明行,王国俊.秋水仙碱拮抗肿瘤坏死因子的肝毒性作用.第二军医大学学报,1998,19(4)：353～355
彭隆金.百合资源与栽培[M].云南民族出版社.2002
曲伟红,周日宝,童巧珍,贺又舜,罗目和,高静.百合的化学成分研究进展.湖南中医药导报,2004,10(3)：75～76
屈云慧,吴丽芳,杨春梅.百合幼胚培养成株体系的建立.云南农林科技,2002(2)：9-13
荣立苹,雷家军,王志刚.东北地区野生百合遗传多样性的RAPD分析.江苏农业学报,2009,25(4)：843～846.
荣立苹,雷家军.东北地区野生百合数量分类研究.植物遗传资源学报,2010,11(1)：99-102
荣丽苹,雷家军,郑洋,高玉福.东北地区野生百合的核型研究.吉林农业大学学报,2009(6)：711-716
孙启时,石猛,付沛云.东北百合属植物的研究.植物研究,1989,9(3)：133-141
孙晓梅,崔文山,刘艳芬.不同授粉方法对新中心和普瑞头两种百合杂交结实影响的研究.辽宁林业科技,2002(2)：6-10
孙晓泽.沈阳地区11个东方百合品种几种同工酶的研究.科技情报开发与经济,2007,17(13)：203～204
四川植物志编辑委员会.四川植物志(第四卷).成都：四川人民出版社,1989
田爱梅,郑日如,王国强,陈发菊,曹家树.中国野生百合资源的研究保护和利用.安徽农业科学,2007,35(31)：9987-9990
王中仁.植物等位酶分析.北京：科学出版社,1996
王瑞波,何平,胡世俊等.南川百合与泸定百合种群格局关联维数的比较研究.重庆林业科技,2008(1)：26-34
王瑞波,张燕平,胡世俊,何平,张春平,邓洪平.两种百合种群空间分布格局对高温干旱气候的响应.林业科学研究,2009,22(2)：249-255
王欣,雷家军,李明旭.渥丹百合不同居群核型研究.中国农学通报,2006,22()：281-284
王欣,雷家军,张丽娜,韩凌.渥丹百合种内居群形态多样性研究.沈阳农业大学学报,2007,38(6)：796-800
王亚斌,杨国锋,安俊学.百合幼胚离体培养的研究[J].沈阳农业大学学报,2002,33(4)：249-251
王红霞,杨宝胜.通江百合形态变异式样研究.河南科技学院学报,2005,25(3)：8-10
王红霞,胡琼华,陈晓兰.通江百合的组织培养.植物生理学通讯,2000,36(2)：132-133
吴祝华,施季森,席梦利,刘光欣.百合种质资源花粉形态及亲缘关系研究.浙江林学院学报,2007,24(4)：406～412
吴祝华,徐进,宋春雷,席梦利,刘光欣,施季森.宜昌百合、泸定百合核型分析.林业科技开发,2007,27(3)：29～31
吴刚,虞泓,梁淑云等.云南川百合DALP遗传变异分析.广东农业科学,2009,(7)：52-55
徐宏发,王静.分子系统学研究进展.生态学杂志,2001,20(3)：41-46
向地英,张延龙,牛立新.秦巴山区及毗邻地区野生百合的形态多样性研究.武汉植物学研究,2005,23(4)：385～388
谢树贵.44个百合品种同工酶遗传多样性研究.西部林业科学,2009,38(4)：27～35
杨利平,丁冰,刘香环,张敩方.东北百合属植物的细胞遗传多样性.东北林业大学学报,1996,24(5)：19～33
杨利平,张敩方,丁冰等.条叶百合×王百合种间杂种的育成.东北林业大学学报,1998,26(3)：37-43
杨红霞,王宝胜.岷江百合居群核型变异研究.河南科技学院学报,2003(3)：40-43
易杨杰,张新全,黄琳凯,凌瑶,马啸,刘伟.野生狗牙根遗传多样性的SRAP分析.遗传,2008,30(1)：94～100
虞泓,黄瑞复,魏蓉城.川百合种内核型多样性研究.云南植物研究,1996,18(增刊)：1～14
虞泓,王红霞,游丹.泸定百合居群染色体形态研究.云南大学学报(自然科学版),2000,22(1)：60-67
于晓英,吴铁明,倪沛,杨梦玲,彭尽晖,于江南.百合幼胚的离体培养和植株再生.湖南农业大学学报,2000,26(4)：286-288
姚青菊,任全尽,熊豫宁,杨军.岷江流域网百合与通江百合资源特征的初步研究.江苏农业科学,2007(5)：245～247
余桂红,马鸿翔,周淼平,黄益洪,陆维忠.百合农杆菌介导的遗传转化受体系统的建立.江西农业科学,2004,16(2)：15～19
杨利平,马宪红,丁冰等.百合花卉种间杂交种的培育.东北林业大学学报,1997,25(1)：29-33
鄢家俊,白使且,张新全.青藏高原东南缘老芒麦自然居群遗传多样性的SRAP和SSR分析.草业学报,2010,19(4)：122-134
杨秀伟,吴云山,崔育新,刘雪辉,肖诗鹰.卷丹中新甾体化合物的分离和鉴定.药学学报,2002,37(11)：863-866
岳玲,雷家军,王欣.辽宁4种(Lilium ssp.)野生百合的核型研究.辽宁农业科学,2006(4).5-8
张炜茹,张启翔.中国百合种质资源调查与引种驯化研究进展.安徽农业科学,2008,36(19)：8065～8067
赵宇,刘庆华等.青岛百合鳞叶扦插繁殖技术研究.山东林业科技,2007(3)：16～18
张克中,贾月慧,张启翔,王洁琼.部分中国野生百合亲缘关系的AFLP分析.东北林业大学学报,2008,36(2)：19～22
张克中,贾月慧,赵祥云,张启翔.部分中国野生百合亲缘关系的RAPD分析.东北林业大学学报,2006,34(6)：43-45
张克中,赵祥云,黄善武,陆长甸,梁励.辐射对百合鳞片生成不定芽及M1植株的影响.北京农学院学报,2002,17(4)：19～25
赵庆芳,马世荣,曾小英,李巧峡,崔艳.百合栽培品种资源的RAPD分析.兰州大学学报(自然科学版),2005,41(2)：30～33
赵祥云,王树栋,刘建斌..鲜切花百合生产原理及实用技术.北京：中国林业出版社.2005
赵祥云,陈新露,方海,张云芳.用RAPD标记评价百合品种间遗传关系.北京农学院学报,1995,(2)：58-63.
左志锐,穆鼎,高俊平,刘春.百合遗传多样性及亲缘关系的RAPD分析.园艺学报,2005,32(3)：468～472
中国科学院中国植物志编委会.中国植物志V o114.北京：科学出版社：1980,118-157
中国科学院昆明植物研究所.云南植物志(第七卷).北京：科学出版社,1997,789～821
中国科学院西北植物研究所.秦岭植物志(第一卷).北京：科学出版社,1976：362-368.-121
邹喻萍,葛颂,王晓东.系统与进化植物学中的分子标记.北京：科学出版社,2001
周厚高.几个百合品种遗传关系的等位酶分析.西南农业学报,1999(4)：92～95
周延清.DNA分子标记技术在植物研究中的应用.北京：化学工业出版社,2005
张西丽,周厚高,周焱,宁云芬.百合品种间的数量分类研究.广西植物,2000(4)：325-328
张兴翠,周昌华,殷家明,张建.药用百合的多倍体诱导及快速繁殖.西南农业大学学报,2003.25(1)：14-17
郑思乡,章海龙,董志渊,雷小云,胡秀,周丽娟.东方百合多倍体诱导及种球繁育的研究.西南农业大学学报,2004,20(3)：260～263
Asano Y. Fertility of a hybrid between distantly related species in Lilium. Cytologia,1984,49:447～ 456
AsanoY.A. numerical taxonomic study of the genus Lilium in Japan, Journal of the Faculty of Agriculture Hokkaido Univ,1986,62:333～341
Baranova MV. A synopsis of the system of the genus Lilium(Liliaceae) (in Russian), Bot Zhur,1988, 73:1319～1329.
Budak H.Shearman R.C. Parmaksiz I., Gaussoin R.E., Riordan T.P., Dewikat I. Molecular characterization of buffalograss germplasm using sequence-related amplified polymorphism markers, Theoretical and Applied Genetics,2004,108:328-334
Baker JG. The gardeners'chronicle.1871,104:1650
Comber H.F. A new classification of the genus Lilium, London:Royale Horticulture Society, Lily Yearbook.1949,86～105
Cozzolino S.Cafasso D, Pellegrino G. Fine-Scale phylogeographical analysis of Mediterranean Anacamptis palustris(Orchidaceae) populations based on chloroplast minisatellite and microsatellite variation. Molecular Ecology,2003,12:2783～2792
Diwan N, Mcltosh MS, Bauchan G R. Methods of developing a core collection of annual Medicago species.Theoritical and Applied Genetics,1995,90:755～761
Dubouzet J.G., Shinoda K. Phylogenetic analysis of the internal transcribed spacer region of Japanese Lilium species, Theoretical and Applied Genetics,1999,98:954～960
Ferriol M, Pico B, Nuez F. Genetic diversity of a germplasm collection of Cucurbita pepo using SRAP and AFLP markers. Theoretical and Applied Genetics,2003,107:271～282
Ferriol M, Pico B, Nuez F. Morphological and molecular diversity of a collection of Cucurbita maxima landraces. Journal of American Society for Horticultural Science,2004,129(1):60～69
Haruki K., Hosoki T., Nako Y., Ohta K. Possibility of classification in some species of Lilium by PCR-RFLP of ribulose-1,5-bisphosphate carboxylase large subunit (rbcl) gene and ribosomal RNA gene.Journal of the Japanese Society for Horticultural Science,1997,66(1):189～192
Haw SG.The taxonomy of Chinese lilies. In:The lilies of China. Portland:Timber Press.1986,57～ 69
Hamrick J L.Godt M J W. Effects of life history traits on genetic diversity in plant species Philosophical Transactions of the Royal Society B. Biological Sciences,1996,351:1291
Hayashi K. Native lilies in Japan 111, The natural habitats of Lilium alexandrae in Amami O-shima. J Phytogeogr Taxon,1983,31:111～119
Jefferson-Brown M. The lily species. In:The gardener's guide to growing lilies. Portland:Timber Press.1995,60～69
Li G., Quiros C.F. Sequence-related amplified polymorphism (SRAP), a new marker system based on a simple PCR reaction:its application to mapping and gene tagging in Brassica, Theoretical and Applied Genetics.2001,103:455～461
Leslie AC. The international lily register,3rd ed. Royal Horticultural Society, London.1982
Lighty R.W. Evolutionary trends in lilies. Lily Year Book.1968,31:40～44
Li L Y, Chen D X, Qin S Y, Li Q S, Zhong G Y. Studies on genetic relationship of seven species of Curculigo plant from China using SRAP. Journal of Chinese Materia Medica,2008,33:117～120
Lee, W. B., S. Y. Choi, Y. S. Kim. An application of random amplified polymorphic DNA(RAPD) to systematics of some species of Lilium in Korea. Kor. J. Plant Tax,1993,23:43～56 (in Korean)
Lee C.S., Kim S.C., Yeau S.H., and Lee N.S. Major lineages of the genus Lilium (Liliaceae) based on nrDNA sequences, with special emphasis on the Korean species. Journal of Plant Biology,2011, 54(3):159～171
Li En Xiang, Yi Sun, Qiu Ying Xiong. Phylogeography of two East Asian Species in Croomia (Stemonaceae) inferred from chloroplast DNA and ISSR fingerprinting variation. Molecular phylogenetics and Evolution,2008,49:702～714
Mitchell R M. Species DNA research report. NALS Q Bull.1998:52 (1):8-9
Nybom H, Bartish T V. Effects of life history traits and sampling strategies on genetics diversity estimates obtained with RAPD markers in plants. Perspectives in plant Ecology Evolution and Systematics,2000,3:93-114
Nishikawa T., Okazaki K., Arakawa K., Nagamine T. Phylogenetic analysis of Section Sinomartagon in genus Lilium using sequences of the internal transcribed spacer region in nuclear ribosomal DNA, Breeding Science,2001,51:39～46
ROBB SM. The culture of excised tissue from bulb scales of Lilium speciosum. J Exp. Bot,1957, 8:348～352
Sarg T M, B-dormiaty M M. Bishrhr M M. Thin layer chormatographic scanner, spectrophotometric and high-pherphomance liquid chromatographic methods for the determination of colchicines. Analyse.1989,114(5):575～578
Sun S J, Gao W, Lin S Q, Zhu J, Xie B G, Lin Z B. Analysis of genetic diversity in Ganoderma population with a novel molecular marker SRAP. Applied Microbiology and Biotechnology,2006, 72:537～543
Solbrig Q T, (ed) From genus to ecosysytems a reseach agenda for biodiversity, IUBS, Paris 1991
Wilson E H. The lilies of eastern Asia. Dulau and Co., London.1925
Wright S. Evolution and the genetics of populations. Chicargo:University of Chicargo press,1968
Yamagishi M. Detection of section-specific random amplified polymorphic DNA (RAPD) in Lilium, Theoretical and Applied Genetics,1995,91:830～835