白花鬼针草的繁殖特性及入侵性
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摘要
白花鬼针草是一种菊科入侵植物。本研究从单个花序花期的观察、繁育系统、花粉胚珠比、自交亲和性、花粉活力、访花昆虫、种子萌发特性等方面对白花鬼针草的繁殖特性进行了分析,探讨了其与入侵性之间的关系。研究表明:白花鬼针草单个花序全部开放约5～7 d,约第9天全部凋落;花粉活力在1～8 d内维持在较高的水平(87.07%～93.41%);花粉胚珠比为2508,单个花序可自交亲和,亦可兼性异交,为混合交配系统,结实率高;边花(舌状花)的存在有利于结实率的提高;访花昆虫主要有粉蝶科、蛱蝶科、茧蜂科和蚁科的昆虫;对萌发率、萌发速率、萌发开始时间和萌发持续时间做因子及聚类分析,白花鬼针草种子的萌发特性可以表现为爆发型、过渡型、缓萌型、低萌型4种萌发类型,根据不同的环境条件选择不同的萌发类型。总之,白花鬼针草具有自交亲和,结实率高,结实量大,种子萌发率高、萌发速率快且具有不同的萌发策略和边花等繁殖特点,是其成功入侵的主要原因。
Bidens pilosa var. radiata( Asteraceae) is an invasive plant species. Here,we examined its reproductive characteristics from aspects of flowering phenology of single inflorescence,breeding system,pollen ovule ratio,self-compatibility,pollen vigor,pollinators,and seed germination. The effects of these aspects on its invasiveness were analyzed. The results showed that the flowering duration of a single inflorescence was 5-7 days. The pollen could maintain a high vigor( 87.07%-93.41%) within 1-8 days. The pollen ovule ratio was 2508. Single inflorescence can be self-compatible or facultative heterosynthesis. B. pilosa var. radiata had a mixed mating system with high seed setting rate. The existence of edge flower( tongue flower) was beneficial to the increases of seed setting rate. The main pollinators were Phaeidae,Nymphalidae,Cochonidae,and Formicidae. By factors analysis and cluster analysis on germination percentage,germination rate,germination start time,and germination duration,the germination strategies of seeds of B. pilosa var. radiata could be classified into four types: outbreak type,transition type,slow budding type,and low sprouting type. Different germination types were selected by different environmental conditions. In conclusion,the self-compatibility,high seed setting rate,high seed germination percentage,fast germination rate,and diverse germinating strategies work together for the successful invasion of B. pilosa var. radiata.
Bidens pilosa var. radiata( Asteraceae) is an invasive plant species. Here,we examined its reproductive characteristics from aspects of flowering phenology of single inflorescence,breeding system,pollen ovule ratio,self-compatibility,pollen vigor,pollinators,and seed germination. The effects of these aspects on its invasiveness were analyzed. The results showed that the flowering duration of a single inflorescence was 5-7 days. The pollen could maintain a high vigor( 87.07%-93.41%) within 1-8 days. The pollen ovule ratio was 2508. Single inflorescence can be self-compatible or facultative heterosynthesis. B. pilosa var. radiata had a mixed mating system with high seed setting rate. The existence of edge flower( tongue flower) was beneficial to the increases of seed setting rate. The main pollinators were Phaeidae,Nymphalidae,Cochonidae,and Formicidae. By factors analysis and cluster analysis on germination percentage,germination rate,germination start time,and germination duration,the germination strategies of seeds of B. pilosa var. radiata could be classified into four types: outbreak type,transition type,slow budding type,and low sprouting type. Different germination types were selected by different environmental conditions. In conclusion,the self-compatibility,high seed setting rate,high seed germination percentage,fast germination rate,and diverse germinating strategies work together for the successful invasion of B. pilosa var. radiata.
引文
曾建军,肖宜安,孙敏.2010.入侵植物剑叶金鸡菊的繁殖特征及其与入侵性之间的关系.植物生态学报,34(8):966-972.
陈君,韦建华,蔡少芳,等.2013.白花鬼针草化学成分研究.中药材,36(3):410-413.
陈文,王桔红,陈丹生,等.2015.五种菊科植物种子萌发对温度的响应及其入侵性.生态学杂志,34(2):420-424.
陈志云,梁水凤,李东文,等.2011.假臭草等12种植物对白花鬼针草幼苗的化感作用.热带亚热带植物学报,19(5):454-462.
崔现亮,罗娅婷,蒋智林,等.2014.光照和冷藏时间对青藏高原东缘三种灌木种子萌发的影响.生态学杂志,33(9):2330-2335.
崔现亮,王桔红,齐威,等.2008.青藏高原东缘灌木种子的萌发特性.生态学报,28(11):5294-5302.
郝建华,刘倩倩,强胜.2009b.菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系.植物学报,44(6):656-665.
郝建华,钱海军,姜雯,等.2009a.加拿大一枝黄花有性生殖特征研究.生态环境学报,18(6):2278-2282.
郝建华.2008.部分菊科入侵种的有性繁殖特征与入侵性的关系(博士学位论文).南京:南京农业大学.
何报作,曾静,韦郃,等.2009.鬼针草与易淆品白花鬼针草的叶形态---脉序图谱鉴别特征.中国中药杂志,34(20):2559-2563.
何田华,葛颂.2001.植物种群交配系统、亲本分析以及基因流动研究.植物生态学报,25(2):144-154.
贺俊英,谢彩琴.2009.繁殖生物学特性在外来入侵植物入侵性中的意义.内蒙古师大学报:自然汉文版,38(2):217-221.
洪岚,沈浩,杨期和,等.2004.外来入侵植物三叶鬼针草种子萌发与贮藏特性研究.植物科学学报,22(5):433-437.
姜勇,李艳红,王文杰,等.2013.光和不同打破种子休眠方法对紫茎泽兰种子萌发及幼苗状态的影响.生态学报,33(1):302-309.
刘明超,韦春强,唐赛春,等.2012.不同土壤养分水平下2种外来鬼针草和近缘本地种的比较研究.生物安全学报,21(1):32-40.
田兴山,岳茂峰,冯莉,等.2010.外来入侵杂草白花鬼针草的特征特性.江苏农业科学,(5):174-175.
万方浩,侯有明,蒋明星.2015.入侵生物学.北京:科学出版社.
王桔红,崔现亮,陈学林,等.2007.中、旱生植物萌发特性及其与种子大小关系的比较.植物生态学报,31(6):1037-1045.
王四海,孙卫邦,成晓,等.2008.外来植物肿柄菊(Tithonia diversifolia)的繁殖特性及其地理扩散.生态学报,28(3):1307-1313.
韦春强,潘玉梅,唐赛春,等.2013.不同光照和温度下白花鬼针草和金盏银盘种子萌发研究.杂草科学,31(4):1-4.
谢田朋,杜国祯,张格非,等.2010.黄帚橐吾种子生产的花序位置效应及其对幼苗建植的影响.植物生态学报,34(4):418-426.
许慧男,王文杰,于兴洋,等.2010.菊科几种入侵和非入侵植物种子需光发芽特性差异.生态学报,30(13):3433-3440.
张志强,李庆军.2009.花寿命的进化生态学意义.植物生态学报,33(3):598-606.
钟军弟,周贤熙,李晓琳,等.2017.不同埋藏深度对白花鬼针草、假臭草和胜红蓟种子出苗及生长的影响.热带农业科学,37(1):1-6.
周兵,闫小红,肖宜安,等.2013.外来入侵植物美洲商陆的繁殖生物学特性及其与入侵性的关系.生态环境学报,22(4):567-574.
Adam NM,McDonald MB,Henderlong PR.1989.The influence of seed position,planting and harvesting dates on soybean seed quality.Seed Science and Technology,17:143-152.
Ashman TL,Schoen DJ.1997.The cost of floral longevity in Clarkia tembloriensis:An experimental investigation.Evolutionary Ecology,11:289-300.
Baskin CC,Baskin JM.1998.Seeds:Ecology,Biogeography,and Evolution of Dormancy and Germination.San Diego:Academic Press.
Brunet J.1996.Male reproductive success and variation in fruit and seed set in Aquilegia caerulea(Ranunculaceae).Ecology,77:2458-2471.
Buide ML.2004.Intra-inflorescence variation in floral traits and reproductive success of the hermaphrodite Silene acutifolia.Annals of Botany,94:441-448.
Buide ML.2008.Disentangling the causes of intra-inflorescence variation in floral traits and fecundity in the hermaphrodite Silene acutifolia.American Journal of Botany,95:490-497.
Burtt BL.1961.Compositae and the study of functional evolution.Transactions of the Botanical Society of Edinburgh,39:216-232.
Canossa RS,Oliveira JR,RS,Constantin J,et al.2008.Effect of temperature and light on joyweed(Alternanthera tenella)seed germination.Planta Daninha,26:745-750.
Cruden RW.1977.Pollen-Ovule Ratios:A conservative indicator of breeding systems in flowering plants.Evolution,31:32-46.
Curnutt JL.2000.Host-area specific climatic-matching:Similarity bread plant,the varying success of invaders.Ecology,77:1661-1666.
Daws MI,Hall J,Flynn S,et al.2007.Do invasive species have bigger seeds?Evidence from intra-and inter-specific comparisons.South African Journal of Botany,73:138-143.
Finch-Savage WE,Leubner-Metzger G.2006.Seed dormancy and the control of germination.New Phytologist,171:501-523.
Kleemann SGL,Chauhan BS,Gill GS.2007.Factors affecting seed germination of perennial wall rocket(Diplotaxis tenuifolia)in Southern Australia.Weed Science,55:481-485.
Koltunow AM.1993.Apomixis:Embryo scas and embryos formed without meiosis or fertilization in ovules.Plant Cell,5:1425-1437.
Liu K,Baskin JM,Baskin CC.2011.Effect of storage conditions on germination of 489 species from high elevation grassland of the eastern Tibet Plateau and implications for climate change.American Journal of Botany,98:12-19.
Marques AR,Apf A,Fao S,et al.2014.Are seed germination and ecological breadth associated?Testing the regeneration niche hypothesis with bromeliads in a heterogeneous neotropical montane vegetation.Plant Ecology,215:517-529.
Mason RAB,Cooke J,Moles AT,et al.2008.Reproductive output of invasive versus native plants.Global Ecology&Biogeography,17:633-640.
Primack RB.2003.Longevity of individual flowers.Annual Review of Ecology&Systematics,16:15-37.
Rambuda TD,Johnson SD.2004.Breeding systems of invasive alien plants in South Africa:Does Baker’s rule apply?Diversity&Distributions,10:409-416.
Sun M,Ganders FR.1988.Mixed mating systems in Hawaiian Bidens(Asteraceae).Evolution,42:516-527.
Tuckeet RE,Merritt DJ,Hay FR.2010.Dormancy,germination and seed bank storage:A study on support of ex situ conservation of macrophytes of southwest Australian temporary pools.Freshwater Biology,55:1118-1129.
Wulff RD.1986.Seed size variation in Desmodium paniculatum.I.Factors affecting seed size.Journal of Ecology,74:87-97.
陈君,韦建华,蔡少芳,等.2013.白花鬼针草化学成分研究.中药材,36(3):410-413.
陈文,王桔红,陈丹生,等.2015.五种菊科植物种子萌发对温度的响应及其入侵性.生态学杂志,34(2):420-424.
陈志云,梁水凤,李东文,等.2011.假臭草等12种植物对白花鬼针草幼苗的化感作用.热带亚热带植物学报,19(5):454-462.
崔现亮,罗娅婷,蒋智林,等.2014.光照和冷藏时间对青藏高原东缘三种灌木种子萌发的影响.生态学杂志,33(9):2330-2335.
崔现亮,王桔红,齐威,等.2008.青藏高原东缘灌木种子的萌发特性.生态学报,28(11):5294-5302.
郝建华,刘倩倩,强胜.2009b.菊科入侵植物三叶鬼针草的繁殖特征及其与入侵性的关系.植物学报,44(6):656-665.
郝建华,钱海军,姜雯,等.2009a.加拿大一枝黄花有性生殖特征研究.生态环境学报,18(6):2278-2282.
郝建华.2008.部分菊科入侵种的有性繁殖特征与入侵性的关系(博士学位论文).南京:南京农业大学.
何报作,曾静,韦郃,等.2009.鬼针草与易淆品白花鬼针草的叶形态---脉序图谱鉴别特征.中国中药杂志,34(20):2559-2563.
何田华,葛颂.2001.植物种群交配系统、亲本分析以及基因流动研究.植物生态学报,25(2):144-154.
贺俊英,谢彩琴.2009.繁殖生物学特性在外来入侵植物入侵性中的意义.内蒙古师大学报:自然汉文版,38(2):217-221.
洪岚,沈浩,杨期和,等.2004.外来入侵植物三叶鬼针草种子萌发与贮藏特性研究.植物科学学报,22(5):433-437.
姜勇,李艳红,王文杰,等.2013.光和不同打破种子休眠方法对紫茎泽兰种子萌发及幼苗状态的影响.生态学报,33(1):302-309.
刘明超,韦春强,唐赛春,等.2012.不同土壤养分水平下2种外来鬼针草和近缘本地种的比较研究.生物安全学报,21(1):32-40.
田兴山,岳茂峰,冯莉,等.2010.外来入侵杂草白花鬼针草的特征特性.江苏农业科学,(5):174-175.
万方浩,侯有明,蒋明星.2015.入侵生物学.北京:科学出版社.
王桔红,崔现亮,陈学林,等.2007.中、旱生植物萌发特性及其与种子大小关系的比较.植物生态学报,31(6):1037-1045.
王四海,孙卫邦,成晓,等.2008.外来植物肿柄菊(Tithonia diversifolia)的繁殖特性及其地理扩散.生态学报,28(3):1307-1313.
韦春强,潘玉梅,唐赛春,等.2013.不同光照和温度下白花鬼针草和金盏银盘种子萌发研究.杂草科学,31(4):1-4.
谢田朋,杜国祯,张格非,等.2010.黄帚橐吾种子生产的花序位置效应及其对幼苗建植的影响.植物生态学报,34(4):418-426.
许慧男,王文杰,于兴洋,等.2010.菊科几种入侵和非入侵植物种子需光发芽特性差异.生态学报,30(13):3433-3440.
张志强,李庆军.2009.花寿命的进化生态学意义.植物生态学报,33(3):598-606.
钟军弟,周贤熙,李晓琳,等.2017.不同埋藏深度对白花鬼针草、假臭草和胜红蓟种子出苗及生长的影响.热带农业科学,37(1):1-6.
周兵,闫小红,肖宜安,等.2013.外来入侵植物美洲商陆的繁殖生物学特性及其与入侵性的关系.生态环境学报,22(4):567-574.
Adam NM,McDonald MB,Henderlong PR.1989.The influence of seed position,planting and harvesting dates on soybean seed quality.Seed Science and Technology,17:143-152.
Ashman TL,Schoen DJ.1997.The cost of floral longevity in Clarkia tembloriensis:An experimental investigation.Evolutionary Ecology,11:289-300.
Baskin CC,Baskin JM.1998.Seeds:Ecology,Biogeography,and Evolution of Dormancy and Germination.San Diego:Academic Press.
Brunet J.1996.Male reproductive success and variation in fruit and seed set in Aquilegia caerulea(Ranunculaceae).Ecology,77:2458-2471.
Buide ML.2004.Intra-inflorescence variation in floral traits and reproductive success of the hermaphrodite Silene acutifolia.Annals of Botany,94:441-448.
Buide ML.2008.Disentangling the causes of intra-inflorescence variation in floral traits and fecundity in the hermaphrodite Silene acutifolia.American Journal of Botany,95:490-497.
Burtt BL.1961.Compositae and the study of functional evolution.Transactions of the Botanical Society of Edinburgh,39:216-232.
Canossa RS,Oliveira JR,RS,Constantin J,et al.2008.Effect of temperature and light on joyweed(Alternanthera tenella)seed germination.Planta Daninha,26:745-750.
Cruden RW.1977.Pollen-Ovule Ratios:A conservative indicator of breeding systems in flowering plants.Evolution,31:32-46.
Curnutt JL.2000.Host-area specific climatic-matching:Similarity bread plant,the varying success of invaders.Ecology,77:1661-1666.
Daws MI,Hall J,Flynn S,et al.2007.Do invasive species have bigger seeds?Evidence from intra-and inter-specific comparisons.South African Journal of Botany,73:138-143.
Finch-Savage WE,Leubner-Metzger G.2006.Seed dormancy and the control of germination.New Phytologist,171:501-523.
Kleemann SGL,Chauhan BS,Gill GS.2007.Factors affecting seed germination of perennial wall rocket(Diplotaxis tenuifolia)in Southern Australia.Weed Science,55:481-485.
Koltunow AM.1993.Apomixis:Embryo scas and embryos formed without meiosis or fertilization in ovules.Plant Cell,5:1425-1437.
Liu K,Baskin JM,Baskin CC.2011.Effect of storage conditions on germination of 489 species from high elevation grassland of the eastern Tibet Plateau and implications for climate change.American Journal of Botany,98:12-19.
Marques AR,Apf A,Fao S,et al.2014.Are seed germination and ecological breadth associated?Testing the regeneration niche hypothesis with bromeliads in a heterogeneous neotropical montane vegetation.Plant Ecology,215:517-529.
Mason RAB,Cooke J,Moles AT,et al.2008.Reproductive output of invasive versus native plants.Global Ecology&Biogeography,17:633-640.
Primack RB.2003.Longevity of individual flowers.Annual Review of Ecology&Systematics,16:15-37.
Rambuda TD,Johnson SD.2004.Breeding systems of invasive alien plants in South Africa:Does Baker’s rule apply?Diversity&Distributions,10:409-416.
Sun M,Ganders FR.1988.Mixed mating systems in Hawaiian Bidens(Asteraceae).Evolution,42:516-527.
Tuckeet RE,Merritt DJ,Hay FR.2010.Dormancy,germination and seed bank storage:A study on support of ex situ conservation of macrophytes of southwest Australian temporary pools.Freshwater Biology,55:1118-1129.
Wulff RD.1986.Seed size variation in Desmodium paniculatum.I.Factors affecting seed size.Journal of Ecology,74:87-97.