6种铁线莲属植物的柱头特征对传粉的适应
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摘要
为了解传粉过程中柱头对花粉的捕获策略,对6种铁线莲属植物(甘青铁线莲、灌木铁线莲、粉绿铁线莲、薄叶铁线莲、粗齿铁线莲和短尾铁线莲)的柱头显微结构进行扫描电镜观察,发现6种铁线莲属植物的柱头均位于花柱的腹缝面,由花柱腹缝两侧细胞发育成柱头乳突,乳突形状随花期逐步从球状到指状甚至长指状过度,并伴随着柱头受粉面从花柱顶端朝花柱基部渐次发育成熟的特殊发育式样。观察分析6种铁线莲的花部综合征及花粉胚珠比(P/O)发现,柱头的此类发育式样与其他花部构成存在功能上的协同一致。为理解铁线莲属植物花部的进化提供了新的思路和视野,对观赏用铁线莲属植物的育种栽培具参考价值。
An unusual developmental pattern of stigma was observed in six Clematis species(Clematis tangutica(Maxim.) Korsh.,Clematis fruticosa Turcz.,Clematis glauca Willd.,Clematis gracilifolia Rehd. et Wils.,Clematis argenfilucida(Levl. et Vant.) W. T. Wang, Clematis brevicaudata DC.) by scanning electron microscopy. The stigmas of Clematis flowers were on the ventral sides of the styles,and developed into stigmatic papillae by cells along the ventral slits. The shape of papillae change along with the development of stigma.During anthesis,the receptive surfaces matured in a sequence from the apical towards the basal ends of the styles. Stigma development pattern is functionally concerted with other floral design. Our study will be a new field of vision to understand the evolution of stigma in pollen capture with a reference value in breeding and cultivation of ornamental Clematis species.
An unusual developmental pattern of stigma was observed in six Clematis species(Clematis tangutica(Maxim.) Korsh.,Clematis fruticosa Turcz.,Clematis glauca Willd.,Clematis gracilifolia Rehd. et Wils.,Clematis argenfilucida(Levl. et Vant.) W. T. Wang, Clematis brevicaudata DC.) by scanning electron microscopy. The stigmas of Clematis flowers were on the ventral sides of the styles,and developed into stigmatic papillae by cells along the ventral slits. The shape of papillae change along with the development of stigma.During anthesis,the receptive surfaces matured in a sequence from the apical towards the basal ends of the styles. Stigma development pattern is functionally concerted with other floral design. Our study will be a new field of vision to understand the evolution of stigma in pollen capture with a reference value in breeding and cultivation of ornamental Clematis species.
引文
1.黄双全.花部特征演化的最有效传粉者原则:证据与疑问[J].生命科学,2014,26(2):118-124.Huang S Q.Most effective pollinator principle of floral evolution:evidence and query[J].Chinese Bulletin of Life Sciences,2014,26(2):118-124.
2 .Baker H G.Evolutionary mechanisms in pollination biology:Origins and functions of floral systems are being elucidated by genetical and ecological studies[J].Science,1963,139(3558):877-883.
3 .Stebbins G L.Adaptive radiation of reproductive characteristics in angiosperms,Ⅱ:seeds and seedlings[J].Annual Review of Ecology and Systematics,1971,2:237-260.
4 .Faegri K,Van Der Pijl L.The principles of pollination ecology[J].The Quarterly Review of Biology,1967,42(2):310-311.
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9 .蔡艳飞,李世峰,李涵,等.中国铁线莲属植物研究进展[J].中国农学通报,2009,25(4):195-198.Cai Y F,Li S F,Li H,et al.Advance in the research of Clematis L.[J]Chinese Agricultural Science Bulletin,2009,25(4):195-198.
10 .Kunze H.Evolution of the translator in Periplocaceae and Asclepiadaceae[J].Plant Systematics and Evolution,1993,185(1-2):99-122.
11 .黄双全,郭友好.传粉生物学的研究进展[J].科学通报,2000,45(3):225-237.Huang S Q,Guo Y H.New advances in pollination biology and the studies in China[J].Chinese Science Bulletin,2000,45(1):1441-1447.
12 .吴海燕.毛茛科部分植物花器官发育及柱头式样对传粉的适应[D].兰州:西北师范大学,2008.Wu H Y.Floral development and patterns of stigmas for pollination adaptation in some species of Ranunculaceae[D].Lanzhou:Northwest Normal University,2008.
13 .蔡正旺.几种铁线莲属植物的花部综合征与访花昆虫多样性及甘青铁线莲传粉生物学研究[D].兰州:西北师范大学,2008.Cai Z W.Study on floral characters and diversity of flowervisiting insects in 6 species of clematis and pollination biology of Clematis tangutica(Ranunculaceae)[D].Lanzhou:Northwest Normal University,2008.
14 .Lloyd D G,Yates J M A.Intrasexual selection and the segregation of pollen and stigmas in hermaphrodite plants,exemplified by Wahlenbergia albomarginata(Campanulaceae)[J].Evolution,1982,36(5):903-913.
15 .张大勇.植物生活史进化与繁殖生态学[M].北京:科学出版社,2004.Zhang D Y.Plant Life-History Evolution and Reproductive Ecology[M].Beijing:Science Press,2004.
16 .Klips R A,Snow A A.Delayed autonomous self-pollination in Hibiscus laevis(Malvaceae)[J]American Journal of Botany,1997,84(1):48-53.
17 .Barrett S C H,Jesson L K,Baker A M.The evolution and function of stylar polymorphisms in flowering plants[J].Annals of Botany,2000,85:253-265.
18 .Fetscher A E.Resolution of male-female conflict in an hermaphroditic flower[J].Proceeding of the Royal Society B:Biological Sciences,2001,268(1466):525-529.
19 .Stebbins G L.Flowering plants:evolution above the species level[J].Systematic Zoolory,1975,24(4):504-506.
20 .Barrett S C H.The evolution of mating strategies in flowering plants[J].Trends in Plant Science,1998,3(9):335-341.
2 .Baker H G.Evolutionary mechanisms in pollination biology:Origins and functions of floral systems are being elucidated by genetical and ecological studies[J].Science,1963,139(3558):877-883.
3 .Stebbins G L.Adaptive radiation of reproductive characteristics in angiosperms,Ⅱ:seeds and seedlings[J].Annual Review of Ecology and Systematics,1971,2:237-260.
4 .Faegri K,Van Der Pijl L.The principles of pollination ecology[J].The Quarterly Review of Biology,1967,42(2):310-311.
5 .Edlund A F,Swanson R,Preuss D.Pollen and stigma structure and function:the role of diversity in pollination[J].Plant Cell,2004,16(S1):S84-S97.
6 .郭友好,黄双全.茨藻目植物柱头特征与传粉系统的演化[J].植物分类学报,1999,37(2):131-136.Guo Y H,Huang S Q.Evolution of pollination system and characters of stigmas in Najadales[J].Acta Phytotaxonomica Sinica,1999,37(2):131-136.
7 .予茜,郭友好,黄双全.三种耧斗菜属植物柱头的特征[J].植物分类学报,2005,43(6):513-516.Yu Q,Guo Y H,Huang S Q.Characters of stigma in three Aquilegia species[J]Acta Phytotaxonomica Sinica,2005,43(6):513-516.
8 .杨春锋.马先蒿属植物的花部分化与传粉适应[D].武汉:武汉大学,2004.Yang C F.Flower differentiation and pollination adaptation in species of pedicularis(Orobanchaceae)[D].Wuhan:Wuhan University,2004.
9 .蔡艳飞,李世峰,李涵,等.中国铁线莲属植物研究进展[J].中国农学通报,2009,25(4):195-198.Cai Y F,Li S F,Li H,et al.Advance in the research of Clematis L.[J]Chinese Agricultural Science Bulletin,2009,25(4):195-198.
10 .Kunze H.Evolution of the translator in Periplocaceae and Asclepiadaceae[J].Plant Systematics and Evolution,1993,185(1-2):99-122.
11 .黄双全,郭友好.传粉生物学的研究进展[J].科学通报,2000,45(3):225-237.Huang S Q,Guo Y H.New advances in pollination biology and the studies in China[J].Chinese Science Bulletin,2000,45(1):1441-1447.
12 .吴海燕.毛茛科部分植物花器官发育及柱头式样对传粉的适应[D].兰州:西北师范大学,2008.Wu H Y.Floral development and patterns of stigmas for pollination adaptation in some species of Ranunculaceae[D].Lanzhou:Northwest Normal University,2008.
13 .蔡正旺.几种铁线莲属植物的花部综合征与访花昆虫多样性及甘青铁线莲传粉生物学研究[D].兰州:西北师范大学,2008.Cai Z W.Study on floral characters and diversity of flowervisiting insects in 6 species of clematis and pollination biology of Clematis tangutica(Ranunculaceae)[D].Lanzhou:Northwest Normal University,2008.
14 .Lloyd D G,Yates J M A.Intrasexual selection and the segregation of pollen and stigmas in hermaphrodite plants,exemplified by Wahlenbergia albomarginata(Campanulaceae)[J].Evolution,1982,36(5):903-913.
15 .张大勇.植物生活史进化与繁殖生态学[M].北京:科学出版社,2004.Zhang D Y.Plant Life-History Evolution and Reproductive Ecology[M].Beijing:Science Press,2004.
16 .Klips R A,Snow A A.Delayed autonomous self-pollination in Hibiscus laevis(Malvaceae)[J]American Journal of Botany,1997,84(1):48-53.
17 .Barrett S C H,Jesson L K,Baker A M.The evolution and function of stylar polymorphisms in flowering plants[J].Annals of Botany,2000,85:253-265.
18 .Fetscher A E.Resolution of male-female conflict in an hermaphroditic flower[J].Proceeding of the Royal Society B:Biological Sciences,2001,268(1466):525-529.
19 .Stebbins G L.Flowering plants:evolution above the species level[J].Systematic Zoolory,1975,24(4):504-506.
20 .Barrett S C H.The evolution of mating strategies in flowering plants[J].Trends in Plant Science,1998,3(9):335-341.