四川雅安地区凤仙花属植物种质资源及其花粉形态学研究
|
摘要
凤仙花属(Impatiens L.)植物全世界有900余种,我国已知有220余种。凤仙花属植物种类繁多,花形奇特,变化丰富,该属植物的多样性为选育花卉优良品种提供了丰富的基因资源。部分野生种类本身就具有很高的观赏价值,亟待引种栽培和推广。目前,该属的开发与利用很少,对于种质资源保存方面的研究则几乎是空白,因此意义重大。
鉴于此,本研究于2009年6月至2010年12月对雅安周公山、海子山、碧峰峡、喇叭河森林公园、二郎山东坡、太湖寺、龙苍沟森林公园的凤仙花属植物进行实地调查及植物鉴定。调查内容包括立地条件、种群概况,个体植株大小,叶、花、蒴果、种子等方面。同时,在收集种质资源的基础上进行凤仙花属植物孢粉学研究。在扫描电镜下观察测量供试花粉粒的8个定量指标(极轴长P、长赤道轴长E1、短赤道轴长E2、沟长L、沟宽W、萌发沟数目、网眼大小、脊宽)和9个定性指标(极面观形状、外壁纹饰类型、网脊长短、网脊曲直、网脊形态、网眼均匀度、网眼形状、极面和沟间区网眼比较、网眼内颗粒状突起)。取定量、定性指标作运算单位,利用国际通用的SPSS分析软件进行主成分分析,然后作聚类分析。
经调查研究,初步得到以下结果：
1雅安地区凤仙花属植物资源
通过对雅安地区自然分布的凤仙花属植物的资源调查及鉴定,确定所调查的16个种分别是峨眉凤仙、湖北凤仙、匙叶凤仙、菱叶凤仙、荞麦地凤仙、天全凤仙、红雉凤仙、短喙凤仙、扭萼凤仙、波缘凤仙、齿萼凤仙、条纹凤仙、华丽凤仙、耳叶凤仙、太子凤仙、紫萼凤仙；另外,通过观察测定各项数据指标补充和细化了《中国植物志》中对这16种凤仙花属植物的形态特征描述,并详细记录了其生境特点。
但在调查中发现峨眉凤仙、湖北凤仙、扭萼凤仙、紫萼凤仙等的居群很小,甚至只有少数几株,更新很弱,急需对其采取相应的保护措施。另外,如天全凤仙、菱叶凤仙等在雅安地区分布非常广泛、萌蘖性很强的种,当地群众肆意将其采割了做饲料,对其破坏极大,为了保护野生资源,相关部门应当采取措施禁止这些行为的发生。希望通过此次调查能够引起有关部门的重视,加强对野生凤仙花属植物种植资源的保护。
2凤仙花属花粉形态学研究
利用扫描电子显微镜,对收集的16种凤仙花属植物的花粉形态进行系统研究。凤仙花属花粉为等极单粒花粉,花粉粒形状极面观为圆角三角形、椭圆形、矩圆形、哑铃形、赤道面观为扁圆形,具角萌发沟,3沟或4沟,属N3P4C3或N4P4C3型,中等大小,极轴长P:11.5～19.6μm,长赤道轴长E1:25.7～36.2μm,短赤道轴长E2：16.5～22.1μm。从花粉外壁纹饰来看,凤仙花属均为网状纹饰。凤仙花属种间花粉差异主要表现在大小、形状及网状纹饰中网脊宽度、曲直度、网眼大小及网眼内衍生物等方面。凤仙花属花粉特征,特别是花粉网状纹饰中网眼内有无颗粒状突起及颗粒状突起的形态特征,在属内常具种水平上的可见变异,可作为该属分类的重要证据。
有学者通过孢粉学分析表明凤仙花属花粉萌发孔演化趋势为3沟→4沟,具3沟花粉的峨嵋凤仙花等种类为凤仙花属的原始类群,具4沟花粉的菱叶凤仙等种类为该中国凤仙花属为该属的进化类群；从孢粉学分析的聚类树系图可以看出,凤仙花属花粉形态分类与目前已有的外部性状分类能够得出许多相同的结论,如3沟花粉类型与具有4枚侧生萼片的种类关系密切,网眼形状与花色具有一定的相关性,网脊形态、网眼大小与距形态、翼瓣上的长细丝具有一定的相关性。另一方面,在研究中发现花粉形态分类结果与传统形态分类结果有差异,说明花粉性状与其外部形态分类依据存在一定的相关性,但并无直接关联。这些差异与研究方法等方面存在的问题有关,如主观因素的存在,定量数据的测量误差和定性指标的多态性编码以及采用不同的聚类方法等,这些都还需要细致深入的研究。
There are 900 various Impatiens genus in the world, among which approximately 220 species distribute in China. The diversity of Impatiens provides abundant gene resources for breeding herb flower species. But up to now this precious germplasm resources are still left in wild without sufficient researches on the breeding or horticultural culture, especially on the preservation of germplasm, so study on the germplasm resources of Impatiens is of great significance.
For that reason, this study investigated and identified the resources of the Impatiens at Zhougong mountain Forest Park, Haizi mountain, Bifeng canyon Ecloogical Tourist Area, Laba river Natural Scenic Spot, Er'lang mountain Forest Park, Taihu Temple, Longcang ravine Forest Park in June 2009 to December 2010. The investigation included the site conditions, general situation of the species group, the size of the plant and the condition of the leaf, the flower, the capsule, the seed. At the same time, the study researched the pollen morphology of the Impatiens which was based on the resources of it. Measured the eight quantitative indexes (polar axis, long equatorial diameter, short equatorial diameter, aperture length, aperture width, aperturate number, lumina size and, muri width) and qualitative indexes (the shape in polar view, the type of sexine ornamentation, muri length, merits degrees of muri, the shape of muri, the degree of uniformity of lumen, the shape of lumen, the difference of the lumen between the polar view and the aperture, the irregular capita in lumen). Use the quantitative and qualitative indexes as the operational unit to do main component analysis, and then made the clustering analysis by SPSS.
The main results are summarized as follows:
1 Resources of Impatiens L. (Balsaminaceae) in Ya'an
By investigating the resources of Impatiens L. natural distributing in Ya'an, sixteen species were found. They were I. omeiana Hook.f.,I. pritzelii Hook.f.,I. spathulata Y.X. Xiong,I. rhombifolia Y.Q. Lu et Y.L.Chen,I. lemeei Levl, I.tienchuanensis Y.L.Chen,I. oxyanthera Hook.f.,I. rostellata Franch.,I. tortisepala Hook.f., I. undulate Y.L.Chen et Y.Q.Lu, I. dicentra Franch. ex Hook.f.,I. vittata Franch., I. faberi Hook.f.,I. delavayi Franch.,I. alpicola Y.L. Chen et Y.Q.Lu,I. platychlaena Hook.f. On the other hand, complemented and detailed the morphological characterstics of the 16 tested species by measured the various data index.
In the investigation, it was found that the groups of I. omeiana Hook.f.,I. pritzelii Hook.f.,I. tortisepala Hook.f.,I. platychlaena Hook.f. were very small, even some relevant protection was badly in need for a few plants that updated weakly. But the dispersedspecies such as I.tienchuanensis Y.L.Chen and I. rhombifolia Y.Q. Lu et Y.L.Chen were picked as feed by the local people, even extensive population couldn't bear such destroy. So we hoped this investigation could cause the attention of concerned branch to strengthen the protecting of the resource of Impatiens.
2 Pollen morphology of Impatiens L. (Balsaminaceae) from Ya'an
Pollen morphology of the sixteen species of Impatiens occurring in Ya'an was examined based on SEM. The pollen of Impatiens was etc extremely kernel pollen and belonged to the type of N3P4C3 or N4P4C3. The outline in polar view was rounded triangular, ellipse, oblong, dumb-bell, and the outline in equatorial view was all oblateness. The aperture was angulaperturate and the number was 3 or 4. The length of polar axis was 11.5～19.6μm, the length of the long equatorial diameter was 25.7～36.2μm, the length of the short equatorial diameter was 16.5～22.1μm. The sexine ornamentation of the Impatiens was reticulate. On the other hand, some characteristic showed difference, existing obvious heterogeneity and diversity, which mainly displayed in the pollen size, shape and the width of muri, the lumina size, the capita absence on the inner locule wall and so on. These differences were a good reason to this genera of the taxonomy. The pollen characteristics especially the capita could be used as the important proof of the classification of the Impatiens.
Sporopollen analysis showed that the 3-aperturate types evolved to the 4-aperturate types, the 3-aperturate types such as I. omeiana Hook.f. was the original group, and the 4-aperturate types such as I. rhombifolia Y.Q. Lu et Y.L.Chen was the evolutionary group. Learned from the dendrogram of Q cluster analysis for Impatiens.L, a lot of the same conclusions could be found between the pollen typoiogical classification and external character classification. In the investigation, pollen data supported that the 3-aperturate ones existed certain relevance with the 4-lateral sepals ones, and also the muri shape, the lumina size existed certain relevance with the the spur shape, and the threadlet growing on the Wing. But there were some variant between the pollen typoiogical classification and external character classification, indicating that pollen morphology exist certain relevance with the genetic traits of Impatiens but not the direct relevance. These differences were related to the problems of the study methods, such as subjective factor, measuring error, the polymorphism of coding and the different cluster methods, so it needed to be studied much more carefully and deeply.
鉴于此,本研究于2009年6月至2010年12月对雅安周公山、海子山、碧峰峡、喇叭河森林公园、二郎山东坡、太湖寺、龙苍沟森林公园的凤仙花属植物进行实地调查及植物鉴定。调查内容包括立地条件、种群概况,个体植株大小,叶、花、蒴果、种子等方面。同时,在收集种质资源的基础上进行凤仙花属植物孢粉学研究。在扫描电镜下观察测量供试花粉粒的8个定量指标(极轴长P、长赤道轴长E1、短赤道轴长E2、沟长L、沟宽W、萌发沟数目、网眼大小、脊宽)和9个定性指标(极面观形状、外壁纹饰类型、网脊长短、网脊曲直、网脊形态、网眼均匀度、网眼形状、极面和沟间区网眼比较、网眼内颗粒状突起)。取定量、定性指标作运算单位,利用国际通用的SPSS分析软件进行主成分分析,然后作聚类分析。
经调查研究,初步得到以下结果：
1雅安地区凤仙花属植物资源
通过对雅安地区自然分布的凤仙花属植物的资源调查及鉴定,确定所调查的16个种分别是峨眉凤仙、湖北凤仙、匙叶凤仙、菱叶凤仙、荞麦地凤仙、天全凤仙、红雉凤仙、短喙凤仙、扭萼凤仙、波缘凤仙、齿萼凤仙、条纹凤仙、华丽凤仙、耳叶凤仙、太子凤仙、紫萼凤仙；另外,通过观察测定各项数据指标补充和细化了《中国植物志》中对这16种凤仙花属植物的形态特征描述,并详细记录了其生境特点。
但在调查中发现峨眉凤仙、湖北凤仙、扭萼凤仙、紫萼凤仙等的居群很小,甚至只有少数几株,更新很弱,急需对其采取相应的保护措施。另外,如天全凤仙、菱叶凤仙等在雅安地区分布非常广泛、萌蘖性很强的种,当地群众肆意将其采割了做饲料,对其破坏极大,为了保护野生资源,相关部门应当采取措施禁止这些行为的发生。希望通过此次调查能够引起有关部门的重视,加强对野生凤仙花属植物种植资源的保护。
2凤仙花属花粉形态学研究
利用扫描电子显微镜,对收集的16种凤仙花属植物的花粉形态进行系统研究。凤仙花属花粉为等极单粒花粉,花粉粒形状极面观为圆角三角形、椭圆形、矩圆形、哑铃形、赤道面观为扁圆形,具角萌发沟,3沟或4沟,属N3P4C3或N4P4C3型,中等大小,极轴长P:11.5～19.6μm,长赤道轴长E1:25.7～36.2μm,短赤道轴长E2：16.5～22.1μm。从花粉外壁纹饰来看,凤仙花属均为网状纹饰。凤仙花属种间花粉差异主要表现在大小、形状及网状纹饰中网脊宽度、曲直度、网眼大小及网眼内衍生物等方面。凤仙花属花粉特征,特别是花粉网状纹饰中网眼内有无颗粒状突起及颗粒状突起的形态特征,在属内常具种水平上的可见变异,可作为该属分类的重要证据。
有学者通过孢粉学分析表明凤仙花属花粉萌发孔演化趋势为3沟→4沟,具3沟花粉的峨嵋凤仙花等种类为凤仙花属的原始类群,具4沟花粉的菱叶凤仙等种类为该中国凤仙花属为该属的进化类群；从孢粉学分析的聚类树系图可以看出,凤仙花属花粉形态分类与目前已有的外部性状分类能够得出许多相同的结论,如3沟花粉类型与具有4枚侧生萼片的种类关系密切,网眼形状与花色具有一定的相关性,网脊形态、网眼大小与距形态、翼瓣上的长细丝具有一定的相关性。另一方面,在研究中发现花粉形态分类结果与传统形态分类结果有差异,说明花粉性状与其外部形态分类依据存在一定的相关性,但并无直接关联。这些差异与研究方法等方面存在的问题有关,如主观因素的存在,定量数据的测量误差和定性指标的多态性编码以及采用不同的聚类方法等,这些都还需要细致深入的研究。
There are 900 various Impatiens genus in the world, among which approximately 220 species distribute in China. The diversity of Impatiens provides abundant gene resources for breeding herb flower species. But up to now this precious germplasm resources are still left in wild without sufficient researches on the breeding or horticultural culture, especially on the preservation of germplasm, so study on the germplasm resources of Impatiens is of great significance.
For that reason, this study investigated and identified the resources of the Impatiens at Zhougong mountain Forest Park, Haizi mountain, Bifeng canyon Ecloogical Tourist Area, Laba river Natural Scenic Spot, Er'lang mountain Forest Park, Taihu Temple, Longcang ravine Forest Park in June 2009 to December 2010. The investigation included the site conditions, general situation of the species group, the size of the plant and the condition of the leaf, the flower, the capsule, the seed. At the same time, the study researched the pollen morphology of the Impatiens which was based on the resources of it. Measured the eight quantitative indexes (polar axis, long equatorial diameter, short equatorial diameter, aperture length, aperture width, aperturate number, lumina size and, muri width) and qualitative indexes (the shape in polar view, the type of sexine ornamentation, muri length, merits degrees of muri, the shape of muri, the degree of uniformity of lumen, the shape of lumen, the difference of the lumen between the polar view and the aperture, the irregular capita in lumen). Use the quantitative and qualitative indexes as the operational unit to do main component analysis, and then made the clustering analysis by SPSS.
The main results are summarized as follows:
1 Resources of Impatiens L. (Balsaminaceae) in Ya'an
By investigating the resources of Impatiens L. natural distributing in Ya'an, sixteen species were found. They were I. omeiana Hook.f.,I. pritzelii Hook.f.,I. spathulata Y.X. Xiong,I. rhombifolia Y.Q. Lu et Y.L.Chen,I. lemeei Levl, I.tienchuanensis Y.L.Chen,I. oxyanthera Hook.f.,I. rostellata Franch.,I. tortisepala Hook.f., I. undulate Y.L.Chen et Y.Q.Lu, I. dicentra Franch. ex Hook.f.,I. vittata Franch., I. faberi Hook.f.,I. delavayi Franch.,I. alpicola Y.L. Chen et Y.Q.Lu,I. platychlaena Hook.f. On the other hand, complemented and detailed the morphological characterstics of the 16 tested species by measured the various data index.
In the investigation, it was found that the groups of I. omeiana Hook.f.,I. pritzelii Hook.f.,I. tortisepala Hook.f.,I. platychlaena Hook.f. were very small, even some relevant protection was badly in need for a few plants that updated weakly. But the dispersedspecies such as I.tienchuanensis Y.L.Chen and I. rhombifolia Y.Q. Lu et Y.L.Chen were picked as feed by the local people, even extensive population couldn't bear such destroy. So we hoped this investigation could cause the attention of concerned branch to strengthen the protecting of the resource of Impatiens.
2 Pollen morphology of Impatiens L. (Balsaminaceae) from Ya'an
Pollen morphology of the sixteen species of Impatiens occurring in Ya'an was examined based on SEM. The pollen of Impatiens was etc extremely kernel pollen and belonged to the type of N3P4C3 or N4P4C3. The outline in polar view was rounded triangular, ellipse, oblong, dumb-bell, and the outline in equatorial view was all oblateness. The aperture was angulaperturate and the number was 3 or 4. The length of polar axis was 11.5～19.6μm, the length of the long equatorial diameter was 25.7～36.2μm, the length of the short equatorial diameter was 16.5～22.1μm. The sexine ornamentation of the Impatiens was reticulate. On the other hand, some characteristic showed difference, existing obvious heterogeneity and diversity, which mainly displayed in the pollen size, shape and the width of muri, the lumina size, the capita absence on the inner locule wall and so on. These differences were a good reason to this genera of the taxonomy. The pollen characteristics especially the capita could be used as the important proof of the classification of the Impatiens.
Sporopollen analysis showed that the 3-aperturate types evolved to the 4-aperturate types, the 3-aperturate types such as I. omeiana Hook.f. was the original group, and the 4-aperturate types such as I. rhombifolia Y.Q. Lu et Y.L.Chen was the evolutionary group. Learned from the dendrogram of Q cluster analysis for Impatiens.L, a lot of the same conclusions could be found between the pollen typoiogical classification and external character classification. In the investigation, pollen data supported that the 3-aperturate ones existed certain relevance with the 4-lateral sepals ones, and also the muri shape, the lumina size existed certain relevance with the the spur shape, and the threadlet growing on the Wing. But there were some variant between the pollen typoiogical classification and external character classification, indicating that pollen morphology exist certain relevance with the genetic traits of Impatiens but not the direct relevance. These differences were related to the problems of the study methods, such as subjective factor, measuring error, the polymorphism of coding and the different cluster methods, so it needed to be studied much more carefully and deeply.
引文
[1]周勃,郑亚萍.中国野生花卉的研究现状.信阳农业高等专科技术学校报,2007.17(2)：111-113
[2]张金政.孙国锋,石雷.中国观赏植物资源现状与展望.园艺学报,2002,29(增刊)：671-678
[3]陈艺林.中国植物志·第四十七卷第二分册(凤仙花科).北京,科学出版社,2001,1-28
[4]王文采.植物分类学的历史回顾与展望[J].生物学通报,2008,43(6)：1-4
[5]Wakabayashi M and Ohba H. Chromosome evolution in Himalayan Impatiens Balsaminaceae [J]. Bot.J.Linn.Soc.,1992,109:247-257.
[6]Akiyama S and Ohba H. Inflorescences of the Himalayan species of Impatiens (Balsaminaceae)[J]. Journal of Japanese Botany,2000,75(4):226-240
[7]Song.Y and Yuan.Y.M..Chromosomal evolution in Balsaminaceae,with cytological observations on 45 species from Southeast Asia[J].Caryologia,2003,56(4):463-481.
[8]Ikeda Hiroshi and Akiyama Shinobu et. A Cytotaxonomic study of Five species of Impatiens (Balsaminaceae) in Java and Borneo, Malesia[J]. Journal of Japanese Botany,2005,80(5):271-277
[9]Perveen A and Qaiser M. Pollen Flora of Pakistan-ⅩⅩⅥ.Balsaminaceae[J].Turkish Journal of Bota-ny,2001,25:35-38
[10]Janssens S and Lens F. Palynological variation in Balsaminoid Ericales. Ⅱ. Balsaminaceae, Tetram eristaceae,Pellicieraceae and general conclusions[J].Annals of Botany,2005,96:1061-1073
[11]Yi Song and Yong-Ming Yuan, Philippe Kupfer.Seedcoat micromorphology of Impatiens (Balsa-minaceae) from China[J].Botanical Journal of the Linnean Society,2005,149(2):195-208
[12]鲁迎青.凤仙花属花粉形态及其分类学意义[J].植物分类学报,1991,29(4)：352-357
[13]蔡秀珍,刘克明等.凤仙花属两种植物花粉的扫描电镜制样研究[J].激光生物学报,2007,16(1),109-111
[14]蔡秀珍,刘克明等.凤仙花属(Impatiens L.)10种植物花粉形态的扫描电镜观察[J].植物研究,2007,27(3)：279-283
[15]蔡秀珍,刘克明等.凤仙花属部分植物的花粉形态[J].园艺学报,2008,35(3)：389-394
[16]J.D.Hooker and T.Thomson. Balsaminaceae. In:Praecursores ad Floram Indicam[J]. Botanical Jour-nal of the Linnean Society,1859,15(4):106-157.
[17]J.D.Hooker. Flora of British India [M]. Kent, L. Reeve,1875,440-483.
[18]J.D.Hook.An epitome of British Indian species of Impatiens[J].Records of the Botanical Survey of India,1910,4:1-58
[19]蔡秀珍.中国凤仙花属Impatiens L一些系统学问题的研究[D].湖南,湖南师范大学.2008
[20]C.Grex-wilson.Impatiens bicornuta and its allies Studies in Balsaminaceae:Ⅶ [J]. Kew Bull, 1988.227(35):61-72
[21]C.Grey-wilson.Impatiens cymbifera and its allies Studies in Balsaminaceae:ⅩⅠ [J]. Kew Bull,1989, 122(44):711-714
[22]V.Bhaskar. Impatiens clavata Bhaskar sp.nov.-a new scapigerous balsam (Balsaminaceae) from Bisle Ghat, Western Ghats, South India[J].Current Science,2006,91(9):1138-1140
[23]C.Grey-wilson.New taxa in African Impatiens(Balsaminaceae)[J].Kew Bull,1978,5(3):641
[24]C.Grey-wilson.Impatiens in Papuasia Studies in Balsaminaceae:I[J] Kew Bull,1979,9(12):661
[25]C.Grey-wilson.Impatiens of Africa[M].Balkema,Rotterdam.1980
[26]Fisher E.Contributions to the Flora of Central Africa V:Two new species of Impatiens (Balsamin-aceae) from Eastern Zaire [J]. Bull. Jard. Bot. Nat. Belg,1997,66:63-71
[27]Cheer M and Fischer E.A tuberous and epiphytic new species of Impatiens Balsaminaceae from southwest Cameroon[J].Kew Bull,1999,54:471-475
[28]Frimodt-mler C and C.Grey-wilson.Two new taxa of Impatiens (Balsaminaceae)from the Udzung-wa Mountains,Tanzania[J].Kew Bull,1999,54:179-184.
[29]Martin Cheek and Laszlo Csiba.A new epiphytic species of Impatiens (Balsaminaceae) from west-ern Cameroon.Kew Bulletin,2002,57:669-674
[30]Fischer E and Dhetchuvi J.P, Ntaganda C.A new species of Impatiens Balsaminaceae from Nyyun-gwe Forest,Rwanda[J].Syst.Geography PI,2003,73:91-101
[31]Fischer E. New taxa of Impatiens (Balsaminaceae) from Madagascar. Ⅱ.A collection from Masoala Peninsula[J].Adansonia,ser.3,2003,25(1):17-31
[32]Fischer E and Rahelivololona M. E.New taxa of Impatiens (Balsaminaceae) from Madagascar. Ⅲ [J].Adansonia, ser.3.2004,26(1):37-52
[33]Fischer E and Rahelivololona M. E. New taxa of Impatiens (Balsaminaceae) from Madagascar. Ⅴ. New species of Impatiens from Masoala Peninsula. Adansonia, ser.3,2007,29 (2):317-332
[34]S.B. Janssens and E.B. Knox,S.Dessein et.Impaties msisimwanensis (Balsaminaceae):description, pollen morphology and phylogenetic position of a new East African species[J].South African Journal of Botany,2009,75:104-109
[35]刘克明,湖南凤仙花属一新种[J].植物分类学报,1999,37(2)：201-203
[36]黄素华,税玉民,陈文红.云南凤仙花属新类群[J].云南植物研究,2003,25(3)：261-280
[37]胡光万,刘克明,刘林愉.湖南的新记录植物(五)[J].植物研究,2003,23(4)：399-401
[38]黄文新,刘克明,蔡秀珍,田建平,胡光万.湖南的新记录植物(六)[J],植物研究,2004,24 (4)：397-399
[39]陈艺林.中国植物志第47卷第二分册[M].北京：科学出版社,2001
[40]陈艺林.国产凤仙花属六新种[J].植物分类学报,1999,37(1)：88-99
[41]陈艺林.国产凤仙花属三新种[J].植物分类学报.2000.38(3)：557-562
[42]陈艺林,浙江凤仙花属的订止--国产凤仙花属植物的研究(二).植物分类学报,1988,8(2)：1-15
[43]于胜祥,陈艺林,覃海宁.广西凤仙花属(凤仙花科)一新变种—瑶山凤仙花.植物分类学报,2007,45(5)：708-712
[44]熊源新.贵州凤仙花科植物的地理分布与区系[J].贵州农学院学报,1990,9(1)：51-57
[45]熊源新.贵州凤仙花属二新种[J].植物分类学报,1996,34(1),98-101
[46]金孝锋,丁炳扬.浙江凤仙花属野生花卉资源及开发.中国野生植物资源,2000,19(4)：27-29
[47]丛义艳.云南高黎贡山凤仙花属(Impatiens L.)植物区系研究[D].湖南：湖南师范大学.2007
[48]Yi-Yan CONG, Ke-Ming LIU et.Impatiens fugongensis (Balsaminaceae),a new species from Yunnan, China,Botanical Studies,2008,49:161-165
[49]王越,温放.“明黄复嫩紫,淡粉换轻红.熠熠还生辉,灼灼又成丛.”--记原产我国的凤仙花(一)[J].中国花卉盆景,2007,4：8-10
[50]王越,温放.“明黄复嫩紫,淡粉换轻红.熠熠还生辉,灼灼又成丛.”--记原产我国的凤仙花(二)[J].中国花卉盆景,2007,5：2-4
[51]新疆八一农学院.植物分类学[M].新疆.农业出版社.1980：5
[52]王开发,王宪曾.孢粉学概论[M].北京.北京大学出版社,1983：4-5
[53]陈学森,郭延奎等.不同制样方法对几种落叶果树花粉形态的影响[J].果树科学,1992,9(4)：198-202.
[54]黄锦岭,刘宪章.开口箭属一新种及其核型和花粉形态[J].植物分类学报,1996,34(6)：592-596
[55]李树林,曲泽洲等.枣品种资源的花粉学研究[J].河北农业大学学报,1987,10(3)：1-9
[56]刘海涛,关佩聪.黄花芥蓝与自花芥蓝的分类学关系[J].华南农业大学学报,1997,18(2)：13-16.
[57]刘玉莲.桂花品种分类与木犀属种质资源的利用[J].植物资源与环境,1993,2(2)：44-48
[58]袁涛,王莲英.几个牡丹野生种的花粉形态及其演化.分类的探讨[J].北京林业大学学报,1999,21(1)：17-21
[59]陈智忠.牡丹花粉形态研究初报[J].林业科技通讯,1999,5：33-34
[60]孙俊娅,万惠敏等.芍药属牡丹组花粉形态的研究进展[J].河北农业科学,2009,13(4)：18-19
[61]魏乐.牡丹种间花粉粒形态差异性比较[J].青海大学学报(自然科学版),2007,25(6)：52-54
[62]何丽霞,李睿等.中国野生牡丹花粉形态的研究[J].兰州大学学报(自然科学版).2005,41(4)：43-49
[63]袁涛,王莲英.根据花粉形态探讨中国栽培牡丹的起源[J].北京林业大学学报,2002,24(1)：5-1]
[64]康素红,包满珠.梅花品种分类的花粉形态学研究[J].园艺学报,1997,24(2)：170-174
[65]方从兵,盛炳成等.花粉形态特征与梅品种分类研究[J].安徽农业大学学报,2002,29(2)：137-142
[66]赵小兰,姚崇怀.桂花花粉形态种内变异的研究[J].湖北民族学院学报(自然版),1999,17(1)：16-20
[67]蔡丹.四川桂花品种分类地花粉形态学研究[D].雅安,四川农业大学.2006
[68]王文莉.玫瑰花粉形态学及生活力研究[D].山东,山东农业大学.2004
[69]冯立国,邵大伟等.野生玫瑰的花粉形态及其起源与演化的探讨[J].林业科学,2007,43(12)：76-80
[70]李玉舒,张启翔.中国玫瑰品种资源花粉形态的观察与比较[J].北方园艺,2009,8：179-182
[71]高连明,张长芹.杜鹃花族花粉形态及其系统学意义[J].云南植物研究,2002,24(4)：471482
[72]王玉国,李光照等.杜鹃花属植物花粉形态及其分类学意义[J].广西植物,2006,26(3)：113-119
[73]周兰英,王永清等.26种杜鹃花属植物花粉形态及分类学研究[J].林业科学,2008,44(2)：55-63
[74]刘永翔,龙毅等.贵州几种杜鹃花属植物的花粉形态[J].贵州农业科学,2009,37(3)：24-26
[75]周兰英,董平等.杜鹃花属植物孢粉学特性种内变异研究[J].安徽农业科学,2009,37(18)：8441-8444,8448
[76]曾小英,赵庆芳等.百合品种的花粉形态研究[J].西北师范大学学报(自然科学版),2004,40(2)：66-68,113
[77]张丽娜,雷家军等.百合属(Lilium)几种植物花粉形态的观察[J].西北农业学报,2006,15(6)：174-178
[78]苏立娟.百合科(Liliaceae)植物花粉形态及花粉发育的研究[D].北京,首都师范大学.2006
[79]吴祝华,施季森等.百合种质资源花粉形态及亲缘关系研究[J].浙江林学院学报,2007,24(4)：406-412
[80]张秀英,王雁等.桃花种质资源花粉形态的观察与比较[J].北京林业大学学报,1997,19(2)：57-62
[81]陈亮,童启庆.山茶属8种1变种花粉形态比较[J],茶叶科学,1997,17(2)：183-188
[82]傅玉兰.寒菊新品种花粉形态特征的研究[J].北京林业大学学报,1998.20(2)：110-113
[83]马燕,陈俊愉.一些蔷薇属植物的花粉形态研究[J].植物研究,1991,11(3)：69-76
[84]方从兵.我国果树孢粉学研究进展[J].安徽农业大学学报,2002,29(2)：154-157.
[85]Huynh K L. Cinq especes D Impatiens au pollen insolte[J].Pollen et Spores,1966,8:455-460
[86]王荷生.植物区系地理[M].北京.科学出版社.1992：32-40
[87]李净.北京绣线菊属种质资源调查及其园林应用研究[D].北京,北京林业大学.2007
[88]埃尔特曼G.孢粉学手册[M].中国科学院植物研究所译.北京.科学出版社.1978：1-45.
[89]Punt W., Hoen P.P., Blackmore S., Nilsson S., Le Thomas A. Glossary of pollen and spore terminology. Review of Palaeobotany and Palynology,2007,143:1-81
[90]陈俊愉主编.中国花卉品种分类学[M].北京.中国林业出版社.2001：101-123
[91]曹宏主.雅安地区自然地理志.雅安.2000：263-268
[92]中共雅安地委政策研究室,雅安地区地方志办公室合编.雅安地区概况.雅安.四川省雅安地区印刷厂.1986：119-240
[93]四川省天全县人民政府.天全年鉴.雅安.2004：5-7
[94]于胜祥.广西凤仙花属植物分类学修订—兼论国产凤仙花属的系统发育[D].北京,中国科学院植物研究所.2008
[2]张金政.孙国锋,石雷.中国观赏植物资源现状与展望.园艺学报,2002,29(增刊)：671-678
[3]陈艺林.中国植物志·第四十七卷第二分册(凤仙花科).北京,科学出版社,2001,1-28
[4]王文采.植物分类学的历史回顾与展望[J].生物学通报,2008,43(6)：1-4
[5]Wakabayashi M and Ohba H. Chromosome evolution in Himalayan Impatiens Balsaminaceae [J]. Bot.J.Linn.Soc.,1992,109:247-257.
[6]Akiyama S and Ohba H. Inflorescences of the Himalayan species of Impatiens (Balsaminaceae)[J]. Journal of Japanese Botany,2000,75(4):226-240
[7]Song.Y and Yuan.Y.M..Chromosomal evolution in Balsaminaceae,with cytological observations on 45 species from Southeast Asia[J].Caryologia,2003,56(4):463-481.
[8]Ikeda Hiroshi and Akiyama Shinobu et. A Cytotaxonomic study of Five species of Impatiens (Balsaminaceae) in Java and Borneo, Malesia[J]. Journal of Japanese Botany,2005,80(5):271-277
[9]Perveen A and Qaiser M. Pollen Flora of Pakistan-ⅩⅩⅥ.Balsaminaceae[J].Turkish Journal of Bota-ny,2001,25:35-38
[10]Janssens S and Lens F. Palynological variation in Balsaminoid Ericales. Ⅱ. Balsaminaceae, Tetram eristaceae,Pellicieraceae and general conclusions[J].Annals of Botany,2005,96:1061-1073
[11]Yi Song and Yong-Ming Yuan, Philippe Kupfer.Seedcoat micromorphology of Impatiens (Balsa-minaceae) from China[J].Botanical Journal of the Linnean Society,2005,149(2):195-208
[12]鲁迎青.凤仙花属花粉形态及其分类学意义[J].植物分类学报,1991,29(4)：352-357
[13]蔡秀珍,刘克明等.凤仙花属两种植物花粉的扫描电镜制样研究[J].激光生物学报,2007,16(1),109-111
[14]蔡秀珍,刘克明等.凤仙花属(Impatiens L.)10种植物花粉形态的扫描电镜观察[J].植物研究,2007,27(3)：279-283
[15]蔡秀珍,刘克明等.凤仙花属部分植物的花粉形态[J].园艺学报,2008,35(3)：389-394
[16]J.D.Hooker and T.Thomson. Balsaminaceae. In:Praecursores ad Floram Indicam[J]. Botanical Jour-nal of the Linnean Society,1859,15(4):106-157.
[17]J.D.Hooker. Flora of British India [M]. Kent, L. Reeve,1875,440-483.
[18]J.D.Hook.An epitome of British Indian species of Impatiens[J].Records of the Botanical Survey of India,1910,4:1-58
[19]蔡秀珍.中国凤仙花属Impatiens L一些系统学问题的研究[D].湖南,湖南师范大学.2008
[20]C.Grex-wilson.Impatiens bicornuta and its allies Studies in Balsaminaceae:Ⅶ [J]. Kew Bull, 1988.227(35):61-72
[21]C.Grey-wilson.Impatiens cymbifera and its allies Studies in Balsaminaceae:ⅩⅠ [J]. Kew Bull,1989, 122(44):711-714
[22]V.Bhaskar. Impatiens clavata Bhaskar sp.nov.-a new scapigerous balsam (Balsaminaceae) from Bisle Ghat, Western Ghats, South India[J].Current Science,2006,91(9):1138-1140
[23]C.Grey-wilson.New taxa in African Impatiens(Balsaminaceae)[J].Kew Bull,1978,5(3):641
[24]C.Grey-wilson.Impatiens in Papuasia Studies in Balsaminaceae:I[J] Kew Bull,1979,9(12):661
[25]C.Grey-wilson.Impatiens of Africa[M].Balkema,Rotterdam.1980
[26]Fisher E.Contributions to the Flora of Central Africa V:Two new species of Impatiens (Balsamin-aceae) from Eastern Zaire [J]. Bull. Jard. Bot. Nat. Belg,1997,66:63-71
[27]Cheer M and Fischer E.A tuberous and epiphytic new species of Impatiens Balsaminaceae from southwest Cameroon[J].Kew Bull,1999,54:471-475
[28]Frimodt-mler C and C.Grey-wilson.Two new taxa of Impatiens (Balsaminaceae)from the Udzung-wa Mountains,Tanzania[J].Kew Bull,1999,54:179-184.
[29]Martin Cheek and Laszlo Csiba.A new epiphytic species of Impatiens (Balsaminaceae) from west-ern Cameroon.Kew Bulletin,2002,57:669-674
[30]Fischer E and Dhetchuvi J.P, Ntaganda C.A new species of Impatiens Balsaminaceae from Nyyun-gwe Forest,Rwanda[J].Syst.Geography PI,2003,73:91-101
[31]Fischer E. New taxa of Impatiens (Balsaminaceae) from Madagascar. Ⅱ.A collection from Masoala Peninsula[J].Adansonia,ser.3,2003,25(1):17-31
[32]Fischer E and Rahelivololona M. E.New taxa of Impatiens (Balsaminaceae) from Madagascar. Ⅲ [J].Adansonia, ser.3.2004,26(1):37-52
[33]Fischer E and Rahelivololona M. E. New taxa of Impatiens (Balsaminaceae) from Madagascar. Ⅴ. New species of Impatiens from Masoala Peninsula. Adansonia, ser.3,2007,29 (2):317-332
[34]S.B. Janssens and E.B. Knox,S.Dessein et.Impaties msisimwanensis (Balsaminaceae):description, pollen morphology and phylogenetic position of a new East African species[J].South African Journal of Botany,2009,75:104-109
[35]刘克明,湖南凤仙花属一新种[J].植物分类学报,1999,37(2)：201-203
[36]黄素华,税玉民,陈文红.云南凤仙花属新类群[J].云南植物研究,2003,25(3)：261-280
[37]胡光万,刘克明,刘林愉.湖南的新记录植物(五)[J].植物研究,2003,23(4)：399-401
[38]黄文新,刘克明,蔡秀珍,田建平,胡光万.湖南的新记录植物(六)[J],植物研究,2004,24 (4)：397-399
[39]陈艺林.中国植物志第47卷第二分册[M].北京：科学出版社,2001
[40]陈艺林.国产凤仙花属六新种[J].植物分类学报,1999,37(1)：88-99
[41]陈艺林.国产凤仙花属三新种[J].植物分类学报.2000.38(3)：557-562
[42]陈艺林,浙江凤仙花属的订止--国产凤仙花属植物的研究(二).植物分类学报,1988,8(2)：1-15
[43]于胜祥,陈艺林,覃海宁.广西凤仙花属(凤仙花科)一新变种—瑶山凤仙花.植物分类学报,2007,45(5)：708-712
[44]熊源新.贵州凤仙花科植物的地理分布与区系[J].贵州农学院学报,1990,9(1)：51-57
[45]熊源新.贵州凤仙花属二新种[J].植物分类学报,1996,34(1),98-101
[46]金孝锋,丁炳扬.浙江凤仙花属野生花卉资源及开发.中国野生植物资源,2000,19(4)：27-29
[47]丛义艳.云南高黎贡山凤仙花属(Impatiens L.)植物区系研究[D].湖南：湖南师范大学.2007
[48]Yi-Yan CONG, Ke-Ming LIU et.Impatiens fugongensis (Balsaminaceae),a new species from Yunnan, China,Botanical Studies,2008,49:161-165
[49]王越,温放.“明黄复嫩紫,淡粉换轻红.熠熠还生辉,灼灼又成丛.”--记原产我国的凤仙花(一)[J].中国花卉盆景,2007,4：8-10
[50]王越,温放.“明黄复嫩紫,淡粉换轻红.熠熠还生辉,灼灼又成丛.”--记原产我国的凤仙花(二)[J].中国花卉盆景,2007,5：2-4
[51]新疆八一农学院.植物分类学[M].新疆.农业出版社.1980：5
[52]王开发,王宪曾.孢粉学概论[M].北京.北京大学出版社,1983：4-5
[53]陈学森,郭延奎等.不同制样方法对几种落叶果树花粉形态的影响[J].果树科学,1992,9(4)：198-202.
[54]黄锦岭,刘宪章.开口箭属一新种及其核型和花粉形态[J].植物分类学报,1996,34(6)：592-596
[55]李树林,曲泽洲等.枣品种资源的花粉学研究[J].河北农业大学学报,1987,10(3)：1-9
[56]刘海涛,关佩聪.黄花芥蓝与自花芥蓝的分类学关系[J].华南农业大学学报,1997,18(2)：13-16.
[57]刘玉莲.桂花品种分类与木犀属种质资源的利用[J].植物资源与环境,1993,2(2)：44-48
[58]袁涛,王莲英.几个牡丹野生种的花粉形态及其演化.分类的探讨[J].北京林业大学学报,1999,21(1)：17-21
[59]陈智忠.牡丹花粉形态研究初报[J].林业科技通讯,1999,5：33-34
[60]孙俊娅,万惠敏等.芍药属牡丹组花粉形态的研究进展[J].河北农业科学,2009,13(4)：18-19
[61]魏乐.牡丹种间花粉粒形态差异性比较[J].青海大学学报(自然科学版),2007,25(6)：52-54
[62]何丽霞,李睿等.中国野生牡丹花粉形态的研究[J].兰州大学学报(自然科学版).2005,41(4)：43-49
[63]袁涛,王莲英.根据花粉形态探讨中国栽培牡丹的起源[J].北京林业大学学报,2002,24(1)：5-1]
[64]康素红,包满珠.梅花品种分类的花粉形态学研究[J].园艺学报,1997,24(2)：170-174
[65]方从兵,盛炳成等.花粉形态特征与梅品种分类研究[J].安徽农业大学学报,2002,29(2)：137-142
[66]赵小兰,姚崇怀.桂花花粉形态种内变异的研究[J].湖北民族学院学报(自然版),1999,17(1)：16-20
[67]蔡丹.四川桂花品种分类地花粉形态学研究[D].雅安,四川农业大学.2006
[68]王文莉.玫瑰花粉形态学及生活力研究[D].山东,山东农业大学.2004
[69]冯立国,邵大伟等.野生玫瑰的花粉形态及其起源与演化的探讨[J].林业科学,2007,43(12)：76-80
[70]李玉舒,张启翔.中国玫瑰品种资源花粉形态的观察与比较[J].北方园艺,2009,8：179-182
[71]高连明,张长芹.杜鹃花族花粉形态及其系统学意义[J].云南植物研究,2002,24(4)：471482
[72]王玉国,李光照等.杜鹃花属植物花粉形态及其分类学意义[J].广西植物,2006,26(3)：113-119
[73]周兰英,王永清等.26种杜鹃花属植物花粉形态及分类学研究[J].林业科学,2008,44(2)：55-63
[74]刘永翔,龙毅等.贵州几种杜鹃花属植物的花粉形态[J].贵州农业科学,2009,37(3)：24-26
[75]周兰英,董平等.杜鹃花属植物孢粉学特性种内变异研究[J].安徽农业科学,2009,37(18)：8441-8444,8448
[76]曾小英,赵庆芳等.百合品种的花粉形态研究[J].西北师范大学学报(自然科学版),2004,40(2)：66-68,113
[77]张丽娜,雷家军等.百合属(Lilium)几种植物花粉形态的观察[J].西北农业学报,2006,15(6)：174-178
[78]苏立娟.百合科(Liliaceae)植物花粉形态及花粉发育的研究[D].北京,首都师范大学.2006
[79]吴祝华,施季森等.百合种质资源花粉形态及亲缘关系研究[J].浙江林学院学报,2007,24(4)：406-412
[80]张秀英,王雁等.桃花种质资源花粉形态的观察与比较[J].北京林业大学学报,1997,19(2)：57-62
[81]陈亮,童启庆.山茶属8种1变种花粉形态比较[J],茶叶科学,1997,17(2)：183-188
[82]傅玉兰.寒菊新品种花粉形态特征的研究[J].北京林业大学学报,1998.20(2)：110-113
[83]马燕,陈俊愉.一些蔷薇属植物的花粉形态研究[J].植物研究,1991,11(3)：69-76
[84]方从兵.我国果树孢粉学研究进展[J].安徽农业大学学报,2002,29(2)：154-157.
[85]Huynh K L. Cinq especes D Impatiens au pollen insolte[J].Pollen et Spores,1966,8:455-460
[86]王荷生.植物区系地理[M].北京.科学出版社.1992：32-40
[87]李净.北京绣线菊属种质资源调查及其园林应用研究[D].北京,北京林业大学.2007
[88]埃尔特曼G.孢粉学手册[M].中国科学院植物研究所译.北京.科学出版社.1978：1-45.
[89]Punt W., Hoen P.P., Blackmore S., Nilsson S., Le Thomas A. Glossary of pollen and spore terminology. Review of Palaeobotany and Palynology,2007,143:1-81
[90]陈俊愉主编.中国花卉品种分类学[M].北京.中国林业出版社.2001：101-123
[91]曹宏主.雅安地区自然地理志.雅安.2000：263-268
[92]中共雅安地委政策研究室,雅安地区地方志办公室合编.雅安地区概况.雅安.四川省雅安地区印刷厂.1986：119-240
[93]四川省天全县人民政府.天全年鉴.雅安.2004：5-7
[94]于胜祥.广西凤仙花属植物分类学修订—兼论国产凤仙花属的系统发育[D].北京,中国科学院植物研究所.2008