播娘蒿花部综合征与传粉模式的适应性研究
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摘要
被子植物中花部特征和形态多变性显示出对多种传粉模式的适应性。前期观测发现,早春短命植物播娘蒿(Descurainia sophia)因花小、不够醒目表现出与自花传粉相适应的花部特征,而花药状的花瓣、宿存的花萼似乎与吸引传粉者有一定的关系。为了探讨其花部综合征与传粉模式的相互关系,通过野外观测、人工控制试验等对其开花习性、花部特征、花瓣的广告效应和交配系统等进行了研究,结果表明:播娘蒿为兼性自交的交配系统类型,自交亲和但保持异交能力。晴朗天气,10:00左右开放,17:00左右闭合,花期1～2 d;开放后的单花,雌蕊不断伸长,经历了低于雄蕊、与雄蕊等高和高于雄蕊3个阶段。播娘蒿有传粉者,但访花频率极低,分别为(0. 062±0. 027) flower·h~(-1)(2015年)和(0. 01±0. 005) flower·h~(-1)(2016年)。控制试验显示花药状花瓣和宿存花萼不具有增强吸引传粉者的广告效应。播娘蒿柱头花粉主要来源于自花粉,自花粉的传递发生在雌蕊与两轮雄蕊分别等高的时期。播娘蒿极小的花、花药状花瓣、较低的胚珠比(P/O值)等是花器官资源投入减少的结果,是与自花传粉模式相适应的繁殖对策。
Floral syndrome and morphological variability show the adaptability to a variety of pollination modes in angiosperms. Descurainia sophia,an early spring ephemeral plant species,possesses the floral design matching with self-pollination due to the small and inconspicuous petals,but the anther-shape-like petals and persistent calyxes seem to have a certain relationship with attracting pollinators. In order to explore the relationship between floral syndromes and pollination patterns,in this study the flowering habits,floral characteristics,advertising effect of petals,mating system and so on of D. sophia,a natural population in Xinjiang,were lucubrated based on the field observation and artificial control test. The results showed that D. sophia is a facultative selfer,self-compatible with a maintained capability to outcross. On a sunny day,the flowers bloom around 10: 00 o'clock and closure around 17: 00,and the florescence is for 1-2 days. After blooming,pistils grow continuously and experience three stages: they are lower than stamens,as high as stamens and higher than stamens. There are pollinators of D. sophia,but their visiting frequency is extremely low,and the visiting frequency was 0. 062 ± 0. 027 flower·h~(-1) in 2015 and 0. 01 ± 0. 005 flower·h~(-1) in 2016. There was no significant difference between the control experiment by removing petals and persistent calyxes and the contrast. Both of the anther-shape-like petals and persistent calyxes have no advertising in attracting pollinators. The pollen loads on stigmas mainly derived from self-pollen in D. sophia. The self-pollen transfer occurs at the two stages when pistils are the same as height of longer stamens and shorter stamens respectively.The tiny flower size,low P/O value and anther-shape-like petals may be the result of decrease of resources allocation invested in floral show and rewards,and also show the reproductive strategies of being compatible with self-pollination pattern.
Floral syndrome and morphological variability show the adaptability to a variety of pollination modes in angiosperms. Descurainia sophia,an early spring ephemeral plant species,possesses the floral design matching with self-pollination due to the small and inconspicuous petals,but the anther-shape-like petals and persistent calyxes seem to have a certain relationship with attracting pollinators. In order to explore the relationship between floral syndromes and pollination patterns,in this study the flowering habits,floral characteristics,advertising effect of petals,mating system and so on of D. sophia,a natural population in Xinjiang,were lucubrated based on the field observation and artificial control test. The results showed that D. sophia is a facultative selfer,self-compatible with a maintained capability to outcross. On a sunny day,the flowers bloom around 10: 00 o'clock and closure around 17: 00,and the florescence is for 1-2 days. After blooming,pistils grow continuously and experience three stages: they are lower than stamens,as high as stamens and higher than stamens. There are pollinators of D. sophia,but their visiting frequency is extremely low,and the visiting frequency was 0. 062 ± 0. 027 flower·h~(-1) in 2015 and 0. 01 ± 0. 005 flower·h~(-1) in 2016. There was no significant difference between the control experiment by removing petals and persistent calyxes and the contrast. Both of the anther-shape-like petals and persistent calyxes have no advertising in attracting pollinators. The pollen loads on stigmas mainly derived from self-pollen in D. sophia. The self-pollen transfer occurs at the two stages when pistils are the same as height of longer stamens and shorter stamens respectively.The tiny flower size,low P/O value and anther-shape-like petals may be the result of decrease of resources allocation invested in floral show and rewards,and also show the reproductive strategies of being compatible with self-pollination pattern.
引文
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[10]Pierre J,Mesquida J,Marilleau R,et al.Nectar secretion in winter oil seed rape,Brassica napus-quantitative and qualitative variability among 71 genotypes[J].Plant Breeding,2010,118(6):471-476.
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[14]范惠玲,罗芳芳,黄雪莲,等.芸芥自交亲和性变异的初步研究[J].植物学报,2015,50(5):598-604.[Fan Huiling,Luo Fangfang,Huang Xuelian,et al.Preliminary studies on the variation in self-compatibility among 52 cultivars of Eruca sativa[J].Chinese Bulletin of Botany,2015,50(5):598-604.]
[15]Holsinger K E.Reproductive systems and evolution in vascular plants[J].Proceedings of the National Academy of Science,2000,97(13):7 037-7 042.
[16]Foxe J P,Slotte T,Stahl E A,et al.Recent speciation associated with the evolution of selfing in Capsella[J].Proceedings of the National Academy of Sciences of the United States of America,2009,106(13):5 241.
[17]Wright S I,Kalisz S,Slotte T.Evolutionary consequences of selffertilization in plants[J].Proceedings Biological Sciences,2013,280(1 760):1-10.
[18]Sicard A,Lenhard M.The selfing syndrome:A model for studying the genetic and evolutionary basis of morphological adaptation in plants[J].Annals of Botany,2011,107(9):1 433-1 433.
[19]Takahata Y,Konno N,Hinata K.Genotypic variation for floral characters in Brassica and allied genera with special reference to breeding system[J].Breeding Science,2008,58(4):385-392.
[20]Duncan T M,Rausher M D.Evolution of the selfing syndrome in Ipomoea[J].Frontiers in Plant Science,2013,4(1):301.
[21]Stebbins C L J.Variation and evolution in plants[J].Kew Bulletin,1971,26(1):187.
[22]Vogler D W,Kalisz S.Sex among the flowers:The distribution of plant mating systems[J].Evolution,International Journal of Organic Evolution,2001,55(1):202-204.
[23]Motten A F.Autogamy and competition for pollinators in Hepatica americana(ranunculaceae)[J].American Journal of Botany,1982,69(8):1 296-1 305.
[24]Dole J A.Role of corolla abscission in delayed self-pollination of Mimulus guttatus(scrophulariaceae)[J].American Journal of Botany,1990,77(11):1 505-1 507.
[25]Tedder A,Ansell S W,Lao X,et al.Sporophytic self-incompatibility genes and mating system variation in Arabis alpina[J].Annals of Botany,2011,108(4):699-713.
[26]Buehler D,Graf R,Holderegger R,et al.Contemporary gene flow and mating system of Arabis alpina in a Central European alpine landscape[J].Annals of Botany,2012,109(7):1 359-1 367.
[27]Lobréaux S,Manel S,Melodelima C.Development of an Arabis alpina genomic contig sequence data set and application to single nucleotide polymorphisms discovery[J].Molecular Ecology Resources,2014,14(2):411-418.
[28]Amasino R.Floral induction and monocarpic versus polycarpic life histories[J].Genome Biology,2009,10(7):228-230.
[29]Holsinger K E.Pollination biology and the evolution of mating systems in flowering plants[J].Evolutionary Biology,1996,29:107-149.
[30]轩俊伟,郑江华,刘志辉,等.基于SPEI的新疆干旱时空变化特征[J].干旱区研究,2016,33(2):338-344.[Xuan Junwei,Zheng Jianghua,Liu Zhihui,et al.SPEI-based spatiotemporal variation of drought in Xinjiang[J]Arid Zone Research,2016,33(2):338-344.]
[31]常兆丰,马中华,王大为,等.民勤荒漠气候变化的不稳定性分析[J].干旱区研究,2016,33(3):601-608.[Chang Zhaofeng,Ma Zhonghua,Wang Dawei,et al.Instability of climate change in the Minqin desert area[J]Arid Zone Research,2016,33(3):601-608.]
[32]许聪聪,崔洪霞,石雷,等.荚蒾属植物花期物候对春季异常气象事件的响应[J].植物学报,2017,52(3):297-306.[Xu Congcong,Cui Hongxia,Shi Lei,et al.Response of flowering phenology of Viburnum to abnormal meteorological events[J].Chinese Bulletin of Botany,2017,52(3):297-306.]
[33]Hiroshi T,Masashi O.Evolution of hierarchical floral resource allocation associated with mating system in an animal-pollinated hermaphroditic herb,Trillium camschatcense(Trilliaceae)[J].American Journal of Botany,2006,93(1):134-141.
[2]郁文彬,蔡杰,王红,等.马先蒿属植物花冠分化与繁殖适应的研究进展[J].植物学报,2008,25(4):392-400.[Yu Wenbin,Cai Jie,Wang Hong,et al.Advances in floral divergence and reproductive adaptation in Pedicularis L.(Orobanchaceae)[J].Chinese Bulletin of Botany,2008,25(4):392-400.]
[3]Cruden R W.Pollen grains:Why so many?[J].Plant Systematics&Evolution,2000,222(1-4):143-165.
[4]Barrett S C H.The evolution of plant sexual diversity[J].Nature Reviews Genetics,2002,3(4):274-84.
[5]Ornduff R.Reproductive biology in relation to systematics[J].Taxon,1969,18(2):121-133.
[6]Cruden R W.Pollen-ovule ratios:A conservative indicator of breeding systems in flowering plants[J].Evolution,1977,31(1):32-46.
[7]Takahata Y.Floral Variation in the Subtribe Brassicinae with Special Reference to Pollination Strategies and Pollen-Ovule Ratios[M].Biology and Breeding of Crucifers,2009.
[8]Goukon K.The nectary of Cruciferae[J].Saihu to Shiiku,1979,41:428-431.
[9]Free J B,Nuttall P M.The pollination of oilseed rape(Brassica napus)and the behaviour of bees on the crop[J].Journal of Agricultural Science,1968,71(1):91-94.
[10]Pierre J,Mesquida J,Marilleau R,et al.Nectar secretion in winter oil seed rape,Brassica napus-quantitative and qualitative variability among 71 genotypes[J].Plant Breeding,2010,118(6):471-476.
[11]Hinata K,Nishio T.Self-incompatiblity in Crucifers[J].Brassica Crops and Wild Allies,1980:223-234.
[12]Conner J K,Rush S.Effects of flower size and number on pollinator visitation to wild radish,Raphanus raphanistrum[J].Oecologia,1996,105(4):509-516.
[13]中国科学院中国植物志编辑委员会.中国植物志[M].北京:科学出版社,1987:33,448.[Chinese Academy of Sciences,Chinese Ethnography Editorial Board.Flora of China[M].Beijing:Science Press,1987:33,448.]
[14]范惠玲,罗芳芳,黄雪莲,等.芸芥自交亲和性变异的初步研究[J].植物学报,2015,50(5):598-604.[Fan Huiling,Luo Fangfang,Huang Xuelian,et al.Preliminary studies on the variation in self-compatibility among 52 cultivars of Eruca sativa[J].Chinese Bulletin of Botany,2015,50(5):598-604.]
[15]Holsinger K E.Reproductive systems and evolution in vascular plants[J].Proceedings of the National Academy of Science,2000,97(13):7 037-7 042.
[16]Foxe J P,Slotte T,Stahl E A,et al.Recent speciation associated with the evolution of selfing in Capsella[J].Proceedings of the National Academy of Sciences of the United States of America,2009,106(13):5 241.
[17]Wright S I,Kalisz S,Slotte T.Evolutionary consequences of selffertilization in plants[J].Proceedings Biological Sciences,2013,280(1 760):1-10.
[18]Sicard A,Lenhard M.The selfing syndrome:A model for studying the genetic and evolutionary basis of morphological adaptation in plants[J].Annals of Botany,2011,107(9):1 433-1 433.
[19]Takahata Y,Konno N,Hinata K.Genotypic variation for floral characters in Brassica and allied genera with special reference to breeding system[J].Breeding Science,2008,58(4):385-392.
[20]Duncan T M,Rausher M D.Evolution of the selfing syndrome in Ipomoea[J].Frontiers in Plant Science,2013,4(1):301.
[21]Stebbins C L J.Variation and evolution in plants[J].Kew Bulletin,1971,26(1):187.
[22]Vogler D W,Kalisz S.Sex among the flowers:The distribution of plant mating systems[J].Evolution,International Journal of Organic Evolution,2001,55(1):202-204.
[23]Motten A F.Autogamy and competition for pollinators in Hepatica americana(ranunculaceae)[J].American Journal of Botany,1982,69(8):1 296-1 305.
[24]Dole J A.Role of corolla abscission in delayed self-pollination of Mimulus guttatus(scrophulariaceae)[J].American Journal of Botany,1990,77(11):1 505-1 507.
[25]Tedder A,Ansell S W,Lao X,et al.Sporophytic self-incompatibility genes and mating system variation in Arabis alpina[J].Annals of Botany,2011,108(4):699-713.
[26]Buehler D,Graf R,Holderegger R,et al.Contemporary gene flow and mating system of Arabis alpina in a Central European alpine landscape[J].Annals of Botany,2012,109(7):1 359-1 367.
[27]Lobréaux S,Manel S,Melodelima C.Development of an Arabis alpina genomic contig sequence data set and application to single nucleotide polymorphisms discovery[J].Molecular Ecology Resources,2014,14(2):411-418.
[28]Amasino R.Floral induction and monocarpic versus polycarpic life histories[J].Genome Biology,2009,10(7):228-230.
[29]Holsinger K E.Pollination biology and the evolution of mating systems in flowering plants[J].Evolutionary Biology,1996,29:107-149.
[30]轩俊伟,郑江华,刘志辉,等.基于SPEI的新疆干旱时空变化特征[J].干旱区研究,2016,33(2):338-344.[Xuan Junwei,Zheng Jianghua,Liu Zhihui,et al.SPEI-based spatiotemporal variation of drought in Xinjiang[J]Arid Zone Research,2016,33(2):338-344.]
[31]常兆丰,马中华,王大为,等.民勤荒漠气候变化的不稳定性分析[J].干旱区研究,2016,33(3):601-608.[Chang Zhaofeng,Ma Zhonghua,Wang Dawei,et al.Instability of climate change in the Minqin desert area[J]Arid Zone Research,2016,33(3):601-608.]
[32]许聪聪,崔洪霞,石雷,等.荚蒾属植物花期物候对春季异常气象事件的响应[J].植物学报,2017,52(3):297-306.[Xu Congcong,Cui Hongxia,Shi Lei,et al.Response of flowering phenology of Viburnum to abnormal meteorological events[J].Chinese Bulletin of Botany,2017,52(3):297-306.]
[33]Hiroshi T,Masashi O.Evolution of hierarchical floral resource allocation associated with mating system in an animal-pollinated hermaphroditic herb,Trillium camschatcense(Trilliaceae)[J].American Journal of Botany,2006,93(1):134-141.