新疆党参的花部综合征与次级花粉呈现

doi:10.3724/SP.J.1003.2011.10167

生物多样性 ›› 2011, Vol. 19 ›› Issue (1): 24-33. DOI: 10.3724/SP.J.1003.2011.10167 cstr: 32101.14.SP.J.1003.2011.10167

新疆党参的花部综合征与次级花粉呈现

王丽¹^,², 谭敦炎¹^,^*()

1 新疆农业大学草业与环境科学学院, 新疆草地资源与生态重点实验室及西部干旱荒漠区草地资源与生态教育部重点实验室, 乌鲁木齐 830052
2 新疆生产建设兵团农六师青格达湖自然保护区管理站, 新疆五家渠 831300

收稿日期:2010-06-30 接受日期:2010-10-14 出版日期:2011-01-20 发布日期:2011-04-01
通讯作者: 谭敦炎
作者简介:*E-mail: tandunyan@163.com
基金资助:
新疆维吾尔自治区高技术研究发展计划(200810102);新疆维吾尔自治区重大科技专项(200731138-1)

Floral syndrome and secondary pollen presentation in Codonopsis clematidea(Campanulaceae)

Li Wang¹^,², Dunyan Tan¹^,^*()

1 Xinjiang Key Laboratory of Grassland Resources and Ecology & Ministry of Education Key Laboratory for Western Arid Region Grassland Resource and Ecology, College of Grassland and Environment Sciences, Xinjiang Agricultural University, Urümqi 830052
2 Qinggeda Lake Nature Reserve Administration Station, Xinjiang Production and Construction Corps Construction of the Sixth Agricultural Division, Wujiaqu, Xinjiang 831300

Received:2010-06-30 Accepted:2010-10-14 Online:2011-01-20 Published:2011-04-01
Contact: Dunyan Tan

摘要/Abstract

摘要：

次级花粉呈现是植物提高花粉输出和接受的效率与准确性以及促进异交的一种繁殖策略。新疆党参(Codonopsis clematidea)具有典型的次级花粉呈现特性。本文采用野外观测、授粉实验与室内显微观测等方法, 对该物种的花部综合征及其次级花粉呈现过程进行了详细观测, 并对这些特征的适应意义进行了探讨。结果表明: 新疆党参钟状下垂的蓝色花冠及黑黄色蜜腺, 不仅避免了雨水冲刷雌雄蕊、稀释花蜜以及风移出花粉等不利影响, 还增加了对传粉者的吸引, 延长了花粉活力及柱头可授性持续时间。次级花粉呈现属于亚顶端花柱呈现者, 呈现机制为沉积机制。该特性限制了单个传粉者带走的花粉量, 延长了花粉呈现时间, 使更多传粉者参与传粉过程, 增加了雄性适合度。其雄性先熟及雌雄蕊空间位置变化, 避免了雌雄功能干扰, 为传粉者取食花蜜、输出花粉以及柱头接受异源花粉提供了通道和机会。其泛化传粉系统的主要传粉昆虫为林野熊蜂(Bombus silvarum)、草地熊蜂(B. paradoxus)和树长黄胡蜂(Dolichovespula sylvestris), 其较长的单花花期(6 d左右), 昆虫访花高峰期与花粉活力最高时期及柱头最佳授粉期相吻合, 以及较大的泌蜜量、较高的花蜜糖浓度和较长的泌蜜时间等特征, 可提高传粉者的访花频率, 增加成功传粉的机会和传粉效率。在气候多变的天山山区,该物种特殊的花部综合征和次级花粉呈现机制对提高其传粉效率并促进异交繁殖成功具有重要意义。

关键词: Codonopsis clematidea, 花部综合征, 异交, 雄性先熟, 次级花粉呈现

Abstract

Secondary pollen presentation has been described as a reproductive strategy that enhances the efficiency and accuracy of pollen exportation and pollen reception, thereby promoting outcrossing. Codonopsis clematidea is characterized by secondary pollen presentation. The objective of this research is to study the floral syndrome and the procedure of secondary pollen presentation of this species, with special reference to the adaptive significance of these features based on the field observations, hand-pollination experiments and indoor microscopic observations. The results showed that the blue campanulate pendulous corolla and black and yellow nectary of C. clematidea not only protect the androecium and gynoecium from rain damage and prevent rain water from diluting the nectar and removal of pollen by wind, but they also attract more pollinators and extend the duration of pollen viability and stigma receptivity. The secondary pollen presentation of this species is sub-terminal stylar presenters and belongs to the deposition mechanism. This character limits the amount of pollen removed by the pollinators and prolongs pollen presentation, thus attracting more pollinators and thereby increasing male fitness. The protandry and the change of the spatial position of male and female organs during the flowering stage of C. clematidea not only avoid interference between male and female functions, but also create a passageway for pollinators to forage for nectar and pollen and for stigmas to receive outcross pollen. Bombus silvarum, B. paradoxusandDolichovespula sylvestris are the effective pollinators in the generalist pollination system of this species. Other floral characteristics of this species, such as long flowering duration (approximately 6 d) per flower, overlap of the peak of insects visitation in the male and female phases with the period of highest pollen viability and the optimal stigma receptivity of pollen, lots of nectar that contains a higher sugar concentration and a long period of nectar secretion during the flowering stage, etc, enhance visitation frequencies and the chances of successful pollination. Therefore, these traits help improve pollination efficiency and promote outcrossing. The special floral syndrome and the mechanism of secondary pollen presentation are significant in improving pollination efficiency and promoting the reproductive success of Codonopsis clematidea by outcrossing under changeable weather conditions in Tianshan Mountains.

Key words: Codonopsis clematidea, floral syndrome, outcrossing, prodandry, secondary pollen presentation

王丽, 谭敦炎 (2011) 新疆党参的花部综合征与次级花粉呈现. 生物多样性, 19, 24-33. DOI: 10.3724/SP.J.1003.2011.10167.

Li Wang, Dunyan Tan (2011) Floral syndrome and secondary pollen presentation in Codonopsis clematidea(Campanulaceae). Biodiversity Science, 19, 24-33. DOI: 10.3724/SP.J.1003.2011.10167.

导出引用管理器 EndNote|Ris|BibTeX

链接本文: https://www.biodiversity-science.net/CN/10.3724/SP.J.1003.2011.10167

https://www.biodiversity-science.net/CN/Y2011/V19/I1/24

图/表 7

图1 新疆党参单花开放进程。(A)花苞中与花柱靠合且尚未开裂的花药; (B)花药在柱头裂片背面散粉; (C)花药与花柱分离, 花粉沉积在花柱毛上; (D)花粉释放完毕; (E)柱头裂片展开; (F)柱头和花冠萎蔫, 开花结束。An: 花药; Es: 尚未展开的柱头外表面; Fi: 花丝; Ne: 蜜腺; Po: 花粉; Sti: 柱头; Sty: 花柱。标尺 = 1 mm。

Fig. 1 Codonopsis clematidea flowering stages. (A) Anthers are closed around the style within the unopened bud; (B) Anthers disperse pollen onto the back of stigma lobes; (C) Pollens are deposited onto the stylar hairs via anthers and style separation; (D) End of pollen release; (E) Stigma lobes spreading; (F) Withered stigma and corolla showing end of anthesis. An, Anther; Es, External surface of still unexpanded stigma; Fi, Filament; Ne, Nectary; Po, Pollen; Sti, Stigma; Sty, Style. Scale bar = 1 mm.

图2 新疆党参花柱毛的动态变化。(A)花苞内花柱上的单细胞毛; (B)花柱毛回缩, 花柱毛间的花粉暴露并释放; (C)花柱毛完全回缩形成的点状窝。Dw: 点状窝; Hi: 正在回缩的花柱毛; Sh: 花柱毛; Po: 花粉。标尺 = 0.5 mm。

Fig. 2 The course of successive stylar hairs invaginated in Codonopsis clematidea. (A) Unicellular hairs of the style within the unopened bud; (B) Pollens between hairs are exposed and released by successive invagination of hairs; (C) Dotted with pits formed by withdrawal of stylar hairs. Dw, Dotted with pits; Hi, Hair invaginating; Sh, Stylar hairs; Po, Pollen. Scale bar = 0.5 mm.

图3 新疆党参的花粉活力(Mean±SE, n = 10)。A: 花药刚散粉; B: 花药散粉完毕并开始释放; C-E: 花粉释放24 h (C)、48 h (D)和72 h (E)。

Fig. 3 Pollen viability in Codonopsis clematidea(Mean±SE,n = 10). A, Beginning of dehiscence of anthers; B, End of dehiscence and beginning of pollen release; C-E, Pollen is released for 24 h (C), 48 h (D), and 72 h (E).

图4 新疆党参的传粉昆虫。A-C: 雄性阶段花; D-F: 雌性阶段花。A和D: 林野熊蜂; B和E: 树长黄胡蜂; C和F: 草地熊蜂。标尺 = 1 mm。

Fig. 4 The pollinators ofCodonopsis clematidea. A-C, Flowers in male phase; D-F, Flowers in female phase. A and D,Bombus silvarum; B and E,Dolichovespula sylvestris; C and F,B. paradoxus. Scale bar = 1 mm.

图5 传粉昆虫的访花频率(Mean±SE, n = 30)

Fig. 5 Visiting frequency of pollinators (Mean±SE,n = 30)

图6 新疆党参花蜜量和糖含量(Mean±SE, n = 30)

Fig. 6 Nectar volume and sugar content in Codonopsis clematidea flowers (Mean±SE,n = 30)

表1 新疆党参不同授粉处理下的座果率和结籽率(Mean± SE)

Table 1 Effect of different pollination treatments on fruit set and seed set in Codonopsis clematidea(Mean ± SE)

授粉处理 Pollination treatment	n	座果率 Fruit-set (%)	结籽率 Seed-set (%)
(1) 不去雄套袋 Bagged without emasculation	30	0^a	0^a
(2) 去雄套网袋 Emasculated and bagged with nylon net	30	0^a	0^a
(3) 人工自花授粉 Artificial autogamy	30	0^a	0^a
(4) 人工异株异花授粉 Artificial xenogamy	30	73.3^b	83.7 ± 4.4 ^b
(5) 自然授粉 Natural pollination	30	66.7^b	80.6 ± 2.4 ^b

参考文献 43

[1]	Abdusalik N (努尔拜衣·阿不都萨力克) (2004) Codonopsis Wall. In: Flora of Xinjiang, Vol. 4 (新疆植物志, 第4卷) (eds Hudaberdi M (米吉提·胡达拜尔地), Pan XL (潘晓玲)), pp.517-518. Xinjiang Science, Technology and Hygiene Publishing House, Urümqi. (in Chinese)
[2]	Anderson GJ, Bernardello G, Lopez SP, Crawford DJ, Stuessy TF (2000) Reproductive biology of Wahlenbergia (Campanulaceae) endemic to Robinson Crusoe Island (Chile). Plant Systematics and Evolution, 223,109-123. DOI URL
[3]	Barrett SCH (1998) The evolution of mating strategies in flowering plants. Trends in Plant Science, 3,335-341.
[4]	Bingham RA, Orthner AR (1998) Efficient pollination of alpine plants. Nature, 391,238-239.
[5]	Brantjes NBM (1983) Regulated pollen issue in Isotoma, Campanulaceae, and evolution of secondary pollen presentation. Acta Botanica Neerlandica, 32,213-222.
[6]	Castro S, Silveira P, Navarro L (2008a) How does secondary pollen presentation affect the fitness of Polygala vayredae (Polygalaceae)? American Journal of Botany, 95,706-712. DOI URL PMID
[7]	Castro S, Silveira P, Navarro L (2008b) How flower biology and breeding system affect the reproductive success of the narrow endemic Polygala vayredae Costa (Polygalaceae). Botanical Journal of the Linnean Society, 157,67-81.
[8]	Chen X (陈曦), Luo GP (罗格平), Xia J (夏军), Zhou KF (周可法), Lou SP (娄少平), Ye MQ (叶民权) (2004) Climate change in northern slope of Tianshan Mountains of Xinjiang and its impact on eco-environment. Science in China (D)(中国科学D辑), 34,1166-1175. (in Chinese)
[9]	Conner JK, Davis R, Rush S (1995) The effect of wild radish floral morphology on pollination efficiency by four taxa of pollinators. Oecologia, 104,234-245. DOI URL PMID
[10]	Dafni A, Kevan PG, Husband BC (2005) Practical Pollination Biology. Enviroquest Ltd., Cambridge.
[11]	Duan YA (段友爱), Li QJ (李庆军) (2008) The pollination biology of Phrynium oliganthum (Marantaceae). Acta Phytotaxonomica Sinica (植物分类学报), 46,545-553. (in Chinese with English abstract)
[12]	Erbar C, Leins P (1989) On the early floral development and the mechanisms of secondary pollen presentation in Campanula, Jasione and Lobelia. Botanische Jahrbücher für Systematik, 111,29-55.
[13]	Erbar C, Leins P (1995) Portioned pollen release and the syndromes of secondary pollen presentation in the Campanulales-Asterales-complex. Flora, 190,323-328.
[14]	Etcheverry AV (2001) Floral biology and pollination in Crotalaria stipularia (Fabaceae: Papilionoideae). Acta Horticulturae, 561,339-342.
[15]	Feng Y (冯缨), An SZ (安沙舟), Liu CE (刘长娥), Sun ZJ (孙宗玖) (2005) Structure and formation analysis of rangeland complex type in middle zone of northern slop of Tianshan Mountains. Acta Prataculturae Sinica (草业学报), 14(1),18-23. (in Chinese with English abstract)
[16]	Galloni M, Podda L, Vivarelli D, Cristofolini G (2007) Pollen presentation, pollen-ovule ratios, and other reproductive traits in Mediterranean legumes (Fam. Fabaceae-Subfam. Faboideae). Plant Systematics and Evolution, 266,147-164.
[17]	Harder LD, Barrett SCH (1996) Pollen dispersal and mating patterns in animal-pollinated plants. In: Floral Biology: Studies on Floral Evolution in Animal-pollinated Plants (eds Lloyd DG, Barrett SCH), pp.140-190. Chapman & Hall, New York.
[18]	Harder LD, Willson WG (1994) Floral evolution and male reproductive success: optimal dispensing schedules for pollen dispersal by animal-pollinated plants. Evolutionary Ecology, 8,542-559.
[19]	Hong DY (洪德元) (1999) Geographical distribution of the family Campanulaceae with reference to the issue of distribution center. In: The Geography of Spermatophytic Families and Genera (种子植物科属地理) (ed. Lu AM (路安民)), pp.516-529. Science Press, Beijing. (in Chinese)
[20]	Howell GJ, Slater AT, Know RB (1993) Secondary pollen presentation in angiosperms and its biological significance. Australian Journal of Botany, 41,417-438.
[21]	Huang SQ (黄双全), Jin BF (靳宝锋), Wang QF (王青锋), Guo YH (郭友好) (1999) Floral display and pollen flow in a natural population of Sagittaria trifolia. Acta Botanica Sinica (植物学报), 41,726-730. (in Chinese with English abstract)
[22]	Imbert FM, Richards JH (1993) Protandry, incompatibility, and secondary pollen presentation in Cephalanthus occidentalis (Rubiaceae). American Journal of Botany, 80,395-404.
[23]	Johnson LAS, Briggs BG (1984) Myrtales and Myrtaceae: a phylogenetic analysis. Annals of the Missouri Botanical Garden, 71,700-756.
[24]	Ladd PG (1994) Pollen presenters in the flowering plants—form and function. Botanical Journal of the Linnean Society, 115,165-195.
[25]	Lammers TG (1992) Circumscription and phylogeny of the Campanulales. Annals of the Missouri Botanical Garden, 79,388-413.
[26]	Leins P, Erbar C (1990) On the mechanisms of secondary pollen presentation in the Campanulales-Asterales-complex. Botanica Acta, 103,87-92.
[27]	Leins P, Erbar C (2003) Floral developmental features and molecular data in plant systematics. In: Deep Morphology: Toward a Renaissance of Morphology in Plant Systematics (eds Stuessy TF, Mayer V, Hörandl E), pp.81-105. Ruggell, Ganther.
[28]	Leins P, Erbar C (2006) Secondary pollen presentation syndromes of the Asterales—a phylogenetic perspective. Botanische Jahrbücher für Systematik, 127,83-103.
[29]	Lloyd DG, Yates JMA (1982) Intrasexual selection and the segregation of pollen and stigmas in hermaphrodite plants, exemplified by Wahlenbergia albomarginata (Campanulaceae). Evolution, 36,903-913. DOI URL PMID
[30]	Lloyd DG, Webb CJ (1986) The avoidance of interference between the presentation of pollen and stigmas in angiosperms. I. Dichogamy. New Zealand Journal of Botany, 24,135-162.
[31]	Lopes AV, Machado IC (1999) Pollination and reproductive biology of Rauvolfia grandiflora (Apocynaceae): secondary pollen presentation, herkogamy and self-incompatibility. Plant Biology, 1,547-553.
[32]	López J, Rodríguez-Rianño T, Ortega-Olivencia A, Devesa JA, Ruiz T (1999) Pollination mechanisms and pollen-ovule ratios in some Genisteae (Fabaceae) from southwestern Europe. Plant Systematics and Evolution, 216,23-47.
[33]	Lundberg J, Bremer K (2003) A phylogenetic study of the order Asterales using one large morphological and three molecular data sets. International Journal of Plant Sciences, 164,553-578. DOI URL
[34]	Mao YY, Huang SQ (2009) Pollen resistance to water in 80 Angiosperm species: flower structures protect rain-susceptible pollen. New Phytologist, 183,892-899.
[35]	Nilsson LA, Rabakonandrianina E, Perterson B, Ranaivo J (1990) ‘Ixoroid’ secondary pollen presentation by small moths in the Malagasy treeletIxora platythyrsa (Rubiaceae). Plant Systematics and Evolution, 170,161-175.
[36]	Nyman Y (1992) Pollination mechanisms in six Campanula species (Campanulaceae). Plant Systematics and Evolution, 181,97-108.
[37]	Richardson TE, Stephenson AG (1989) Pollen removal and pollen deposition affect the duration of the staminate and pisillate phases in Campanula rapunculoides. American Journal of Botany, 76,532-538.
[38]	Sprengel CK (1793) Das entdeckte Geheimnis der Natur im Bau und in der Befruchtung der Blumen (Reprinted 1972). Wheldon and Wesley, New York.
[39]	Sun JF, Gong YB, Renner SS, Huang SQ (2008) Multifunctional bracts in the dove tree Davidia involucrata (Nyssaceae: Cornales): rain protection and pollinator attraction. The American Naturalist, 171,119-124. DOI URL PMID
[40]	Thomson JD, Thomson BA (1992) Pollen presentation and viability schedules in animal-pollinated plants:consequences for reproductive success. In: Ecology and Evolution of Plant Reproduction(ed. Wyatt R), pp.1-24. Chapman & Hall, New York.
[41]	Westerkamp C (1996) Pollen in bee-flower relations. Some considerations on melittophyly. Acta Botanica, 109,325-333.
[42]	Yeo PF (1993) Secondary pollen presentation—form, function and evolution. Plant Systematics and Evolution, 6 (Suppl.),1-128.
[43]	Zhang DY (张大勇) (2004) The Evolution of Life-History and Reproductive Ecology of Plants (植物生活史进化与繁殖生态学). Science Press, Beijing. (in Chinese)

新疆党参的花部综合征与次级花粉呈现

Floral syndrome and secondary pollen presentation in Codonopsis clematidea(Campanulaceae)

RichHTML

PDF (PC)

可视化

被引次数

摘要/Abstract

引用本文

使用本文

图/表 7

参考文献 43

相关文章 6

编辑推荐

Metrics

本文评价

[1]	向文倩, 任明迅. 木棉黄花个体的适应意义[J]. 生物多样性, 2019, 27(4): 373-379.
[2]	吉乃提汗·马木提, 成小军, 谭敦炎. 荒漠短命植物异喙菊的小花异形性及繁殖特性[J]. 生物多样性, 2018, 26(5): 498-509.
[3]	李琳, 路宁娜, 樊宝丽, 赵志刚. 雌雄异熟植物露蕊乌头开花时间对雌雄功能期及表型性别的影响[J]. 生物多样性, 2016, 24(6): 665-671.
[4]	任明迅, 姜新华, 张大勇. 植物繁殖生态学的若干重要问题[J]. 生物多样性, 2012, 20(3): 241-249.
[5]	廖万金, 王峥媚, 谢丽娜, 肖雯, 孙岳. 草乌传粉过程中的广告效应与回报物质研究[J]. 生物多样性, 2007, 15(6): 618-625.
[6]	刘敏, 孙杉, 李庆军. 两种姜科花柱卷曲性植物柱头的位置与其可授性的关系[J]. 生物多样性, 2007, 15(6): 639-644.