入侵植物美洲商陆的传粉生态学特性
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摘要
从开花物候、花朵数量性状、花粉活力、花粉组织化学、柱头可授性、花粉在柱头上的萌发状况、杂交指数、花粉胚珠比、传粉媒介以及繁育系统等方面,研究了外来入侵植物美洲商陆(Phytolacca americana)的传粉生态学特性,为揭示该物种成功入侵的机理提供依据。结果表明:美洲商陆种群花期可达45～70 d,单花花期为2～3 d,可进一步分为杯状花期、花瓣平展期、花瓣下垂期、花瓣反卷期;花冠直径在下垂期最大且不超过1 cm,花药与柱头间距离在1～2.5 mm,有利于自交的发生;花瓣平展期与花瓣下垂期花粉活力差异显著(P<0.05),杯状花期与花瓣平展期花粉活力差异不显著(P>0.05),花瓣平展期花粉活力可达85%;雌蕊柱头可授性在花瓣平展期最强;杯状花期合生柱头开裂,花瓣平展期柱头9～10裂,接受花粉的表面积增大;花药散粉时间与合生柱头开裂时间相吻合,有助于柱头接受花粉;扫描电镜下观察到花粉在花瓣平展期的柱头上已萌发;每朵花可提供1667～3222粒花粉,胚珠9～10个,花粉胚珠比(P/O)为340.88±34.99,表现为兼性自交;美洲商陆杂交指数(OCI)为3,花粉粒属于含脂型,主要传粉昆虫有蜜蜂、胡蜂、食蚜蝇、细腰蜂。综上所述,美洲商陆自交亲和,花粉活力高、柱头可授性强,且高花粉活力与强柱头可授性出现在相同的时期,增加了花粉落置柱头及在柱头上萌发的几率;虫媒花,访花者行为与花朵相适应,有利于提高坐果率,这样的繁殖机制与传粉特点是美洲商陆成功入侵的主要因素之一。
Pollination ecological characteristics of pokeweed( Phytolacca americana) were systematically studied by measuring flowering phenology,flower quantitative characters,pollen vitality and histochemistry,stigma receptivity,pollen germination on stigmata,out-crossing index,pollen-ovule ratio,pollinators,and mating system. The aim of this study is to understand the mechanism underlying successful invasion of pokeweed. The results showed that the flowering span of pokeweed was 45-70 d. The life span of single flowers was 2-3 d,including goblet flowering,petal flattening,petal sagging and petal rewinding stage. The corolla was no larger than 1 cm in diameter in petal sagging stage. The distance between the anther and the stigma was 1-2.5 mm,promoting self-pollination. There was a significant difference in pollen viability between petal flattening stage and petal sagging stage( P<0.05),and no significant difference between goblet flowering stage and petal flattening stage( P>0.05). The pollen viability during petal flattening stage was higher than 85%. In the petal flattening stage,the stigma receptivity was the highest and the synthetic stigmata cracked into 9 to 10,which increased the surface area of stigma for receiving more pollen. Results from the scanning electron microscope revealed that pollen had been germinated on the stigma of this stage. Therefore,the timing of anther loose pollen coincided with the timing of synthetic stigma cracking,which might be beneficial to self-pollination. Each flower contained 1667-3222 pollen grains and 9-10 ovules. The pollen ovule ratio( P/O) was340.88±34.99 and hybridization index( OCI) was 3. The P/O ratio suggests that pokeweed is facultative autogamous. Pokeweed has fat-containing pollen. At least four different species of insect taxa,such as bees,wasps,scorpion flies,and thin-wax bees,were the"principal pollinators". In conclusion,pokeweed is self-compatible with high pollen viability and strong stigma receptivity. The co-occurrence of high pollen viability and strong stigma receptivity is conducive to improve fruit setting. Those reproductive mechanism and pollination characteristics are the main factors for the successful invasion of pokeweed.
Pollination ecological characteristics of pokeweed( Phytolacca americana) were systematically studied by measuring flowering phenology,flower quantitative characters,pollen vitality and histochemistry,stigma receptivity,pollen germination on stigmata,out-crossing index,pollen-ovule ratio,pollinators,and mating system. The aim of this study is to understand the mechanism underlying successful invasion of pokeweed. The results showed that the flowering span of pokeweed was 45-70 d. The life span of single flowers was 2-3 d,including goblet flowering,petal flattening,petal sagging and petal rewinding stage. The corolla was no larger than 1 cm in diameter in petal sagging stage. The distance between the anther and the stigma was 1-2.5 mm,promoting self-pollination. There was a significant difference in pollen viability between petal flattening stage and petal sagging stage( P<0.05),and no significant difference between goblet flowering stage and petal flattening stage( P>0.05). The pollen viability during petal flattening stage was higher than 85%. In the petal flattening stage,the stigma receptivity was the highest and the synthetic stigmata cracked into 9 to 10,which increased the surface area of stigma for receiving more pollen. Results from the scanning electron microscope revealed that pollen had been germinated on the stigma of this stage. Therefore,the timing of anther loose pollen coincided with the timing of synthetic stigma cracking,which might be beneficial to self-pollination. Each flower contained 1667-3222 pollen grains and 9-10 ovules. The pollen ovule ratio( P/O) was340.88±34.99 and hybridization index( OCI) was 3. The P/O ratio suggests that pokeweed is facultative autogamous. Pokeweed has fat-containing pollen. At least four different species of insect taxa,such as bees,wasps,scorpion flies,and thin-wax bees,were the"principal pollinators". In conclusion,pokeweed is self-compatible with high pollen viability and strong stigma receptivity. The co-occurrence of high pollen viability and strong stigma receptivity is conducive to improve fruit setting. Those reproductive mechanism and pollination characteristics are the main factors for the successful invasion of pokeweed.
引文
陈洁君,王晶.2018.我国外来入侵生物防控科技进展.生物安全学报,27(1):16-19.
程玲,王亚珍,许景伟,等.2015.灵山湾国家森林公园垂序商陆土壤种子库时空动态及其对防除措施的响应.生态学报,35(15):5095-5103.
董周焱,柏新富,侯玉平,等.2015.胶东滨海8种树木叶片热值、建成成本及其适应能力.林业科学,51(3):8-15.
董周焱,柏新富,张靖梓,等.2014.入侵植物美洲商陆对光环境的适应性.生态学杂志,33(2):316-320.
何宝忠,丁建丽,李焕,等.2018.新疆植被物候时空变化特征.生态学报,38(6):2139-2155.
黄利春,金樑,李晶,等.2014.蝶形花亚科植物花部适应机制与传粉系统.生态学报,34(19):5360-5368.
李祖任,晏升禄,廖海民,等.2015.入侵杂草美洲商陆大小孢子的形成和雌雄配子体的发育.华北农学报,30(S1):81-86.
林慧,张明莉,王鹏鹏,等.2018.外来入侵植物意大利苍耳的传粉生态学特性.生态学报,38(5):1810-1816.
刘学良,姚俊修,吴海涛,等.2018.接骨木花粉活力测定、培养基筛选及贮藏研究.山东农业科学,50(3):129-131.
皮华强,权秋梅,高辉,等.2016.锦鸡儿(Caragana sinica(Buchoz)Rehd)传粉生物学研究.生态学报,36(6):1652-1662.
王连喜,陈怀亮,李琪,等.2010.植物物候与气候研究进展.生态学报,30(2):447-454.
王鹏程,王秋红,赵珊,等.2014.商陆化学成分及药理作用和临床应用研究进展.中草药,45(18):2722-2731.
徐宁伟,贾益豪,包志毅.2014.美洲商陆果实和种子组分测定及幼苗形态观察.经济林研究,32(4):175-179.
闫小红,张蓓玲,周兵,等.2012.外来入侵植物美洲商陆提取物的化感活性.生态与农村环境学报,28(2):139-145.
张斯斯,肖宜安.2013.中国外来入侵植物生活型与性系统多样性.植物研究,33(3):351-359.
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.
周国海,杨美霞,于华忠,等.2004.商陆生物学特性的初步研究.中国野生植物资源,23(4):37-40.
Cruden RW.1977.Pollen-ovule ratio:A conservation indicator of breeding systems in flowering plants.Evolution,31:32-46.
Dafni A.1992.Pollination Ecology.New York:Oxford University Press:1-89.
Dietz H,Edwards PJ.2006.Recognition that causal processes change during plant invasion helps explain conflicts in evidence.Ecology,87:1359-1367.
Doust JL.1989.Plant reproductive strategies and resource allocation.Trends in Ecology&Evolution,4:230-234.
Dumas Y.2011.American grape(Phytolacca americana):An invasive alien species.Rendez-vous,33/34:47-57.
Hargreaves AL,Harder LD,Johnson SD.2010.Native pollen thieves reduce the reproductive success of a hermaphroditic plant,Aloe maculata.Ecology,91:1693-1703.
Hedhly A,Hormaza JI,Herrero M.2003.The effect of temperature on stigmatic receptivity in sweet cherry(Prunus avium L.).Plant,Cell&Environment,26:1673-1680.
Irwin RE.2002.Pollination biology-linking botany and zoology.Ecology,83:1168-1169.
JerskovJ,Jürgens A,milauer P,et al.2012.The evolution of floral mimicry:Identifying traits that visually attract pollinators.Functional Ecology,26:1381-1389.
Jian OU,LU CY,O’toole DK.2008.A risk assessment system for alien plant bio-invasion in Xiamen,China.Journal of Environmental Sciences,20:989-997.
Jordaan LA,Johnson SD,Downs CT.2011.The role of avian frugivores in germination of seeds ofeshy-fruited invasive alien plants.Biological Invasions,13:1917-1930.
Klein AM,Cunningham SA,Bos M,et al.2008.Advances in pollination ecology form tropical plantation crops.Ecology,89:935-943.
Krupnick GA,Weis AE.1999.The effect of floral herbivory on male and female reproductive success in Isomeris arborea.Ecology,80:135-149.
Makrodimos N,Blionis GJ,Krigas N,et al.2008.Flower morphology,phenology and visitor patterns in an alpine community on Mt Olympos,Greece.Flora,203:449-468.
Stohlgren TJ,Binkley D,Chong GW,et al.1999.Exotic plant species invade hot spots of native plant diversity.Ecological Monographs,69:25-46.
Vogelsang KM,Bever JD.2009.Mycorrhizal densities decline in association with nonnative plants and contribute to plant invasion.Ecology,90:399-407.
Weiss MR.1991.Floral changes as cues for pollinators.Nature,354:227-229.
程玲,王亚珍,许景伟,等.2015.灵山湾国家森林公园垂序商陆土壤种子库时空动态及其对防除措施的响应.生态学报,35(15):5095-5103.
董周焱,柏新富,侯玉平,等.2015.胶东滨海8种树木叶片热值、建成成本及其适应能力.林业科学,51(3):8-15.
董周焱,柏新富,张靖梓,等.2014.入侵植物美洲商陆对光环境的适应性.生态学杂志,33(2):316-320.
何宝忠,丁建丽,李焕,等.2018.新疆植被物候时空变化特征.生态学报,38(6):2139-2155.
黄利春,金樑,李晶,等.2014.蝶形花亚科植物花部适应机制与传粉系统.生态学报,34(19):5360-5368.
李祖任,晏升禄,廖海民,等.2015.入侵杂草美洲商陆大小孢子的形成和雌雄配子体的发育.华北农学报,30(S1):81-86.
林慧,张明莉,王鹏鹏,等.2018.外来入侵植物意大利苍耳的传粉生态学特性.生态学报,38(5):1810-1816.
刘学良,姚俊修,吴海涛,等.2018.接骨木花粉活力测定、培养基筛选及贮藏研究.山东农业科学,50(3):129-131.
皮华强,权秋梅,高辉,等.2016.锦鸡儿(Caragana sinica(Buchoz)Rehd)传粉生物学研究.生态学报,36(6):1652-1662.
王连喜,陈怀亮,李琪,等.2010.植物物候与气候研究进展.生态学报,30(2):447-454.
王鹏程,王秋红,赵珊,等.2014.商陆化学成分及药理作用和临床应用研究进展.中草药,45(18):2722-2731.
徐宁伟,贾益豪,包志毅.2014.美洲商陆果实和种子组分测定及幼苗形态观察.经济林研究,32(4):175-179.
闫小红,张蓓玲,周兵,等.2012.外来入侵植物美洲商陆提取物的化感活性.生态与农村环境学报,28(2):139-145.
张斯斯,肖宜安.2013.中国外来入侵植物生活型与性系统多样性.植物研究,33(3):351-359.
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.
周国海,杨美霞,于华忠,等.2004.商陆生物学特性的初步研究.中国野生植物资源,23(4):37-40.
Cruden RW.1977.Pollen-ovule ratio:A conservation indicator of breeding systems in flowering plants.Evolution,31:32-46.
Dafni A.1992.Pollination Ecology.New York:Oxford University Press:1-89.
Dietz H,Edwards PJ.2006.Recognition that causal processes change during plant invasion helps explain conflicts in evidence.Ecology,87:1359-1367.
Doust JL.1989.Plant reproductive strategies and resource allocation.Trends in Ecology&Evolution,4:230-234.
Dumas Y.2011.American grape(Phytolacca americana):An invasive alien species.Rendez-vous,33/34:47-57.
Hargreaves AL,Harder LD,Johnson SD.2010.Native pollen thieves reduce the reproductive success of a hermaphroditic plant,Aloe maculata.Ecology,91:1693-1703.
Hedhly A,Hormaza JI,Herrero M.2003.The effect of temperature on stigmatic receptivity in sweet cherry(Prunus avium L.).Plant,Cell&Environment,26:1673-1680.
Irwin RE.2002.Pollination biology-linking botany and zoology.Ecology,83:1168-1169.
JerskovJ,Jürgens A,milauer P,et al.2012.The evolution of floral mimicry:Identifying traits that visually attract pollinators.Functional Ecology,26:1381-1389.
Jian OU,LU CY,O’toole DK.2008.A risk assessment system for alien plant bio-invasion in Xiamen,China.Journal of Environmental Sciences,20:989-997.
Jordaan LA,Johnson SD,Downs CT.2011.The role of avian frugivores in germination of seeds ofeshy-fruited invasive alien plants.Biological Invasions,13:1917-1930.
Klein AM,Cunningham SA,Bos M,et al.2008.Advances in pollination ecology form tropical plantation crops.Ecology,89:935-943.
Krupnick GA,Weis AE.1999.The effect of floral herbivory on male and female reproductive success in Isomeris arborea.Ecology,80:135-149.
Makrodimos N,Blionis GJ,Krigas N,et al.2008.Flower morphology,phenology and visitor patterns in an alpine community on Mt Olympos,Greece.Flora,203:449-468.
Stohlgren TJ,Binkley D,Chong GW,et al.1999.Exotic plant species invade hot spots of native plant diversity.Ecological Monographs,69:25-46.
Vogelsang KM,Bever JD.2009.Mycorrhizal densities decline in association with nonnative plants and contribute to plant invasion.Ecology,90:399-407.
Weiss MR.1991.Floral changes as cues for pollinators.Nature,354:227-229.