入侵植物银胶菊的繁殖生物学与生态适应性研究
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摘要
生物入侵破坏了本地生态系统的结构和功能,造成生物多样性丧失,带来了巨大的经济损失,并且威胁人类健康。作为全球变化的重要因素之一,生物入侵在过去10多年时间里引起了广泛的关注和重视。然而,外来物种成为入侵物种的机制还没有得到比较合理的解释,影响了有效地预测和控制入侵物种。因此,需要对能够影响外来物种入侵的因素开展更多的研究工作。
银胶菊(Parthenium hysterophorus)为菊科入侵植物,原产北美南部、中美洲、西印度洋群岛、南美中部;是国际性的大毒草。其疯狂生长、快速传播和产生有毒的化感物质,对作物产量、生物多样性和人类健康具有极大的威胁。该杂草具有广泛的生态适应性和很强的繁殖能力,在最近几年已发展成为我国危害较为严重的外来入侵植物。
本文通过野外定点观察、人工控制实验,对银胶菊在不同生境中的开花物候、种群开花结实数量特征、生物量及其分配、生殖过程进行了研究,比较银胶菊种群在不同生境不同季节的繁殖能力。另外,本文还研究了银胶菊对不同N、P养分水平的生态适应性以及在不同光照水平下的适应性。旨在揭示银胶菊的入侵能力和入侵机制,为银胶菊的预防和控制提供依据。主要研究结果如下:
(1)银胶菊繁殖期长,使其能产生大量的种子和幼苗,在较短时间内就能爆发和入侵。在草地、疏林和路旁生境中,银胶菊从3月至10月都有植株生长,从4月底5月初至10月均有开花结实。在耕地中,银胶菊从3至12月均有植株生长,从5月至12月均有开花结实。
(2)银胶菊在不同生境不同季节种群开花结实数量特征不同,繁殖能力不同。
在草地、疏林和路旁生境,秋季开花植株数、分枝数、头状花序数较夏季和春季多,产生的种子数也多,秋季是这三个生境银胶菊的繁殖能力最强的时期。在耕地中,冬季的分枝数和头状花序数比夏季和秋季多,种子产量相应也多。因此,耕地中冬季的繁殖能力最强。
秋季,银胶菊在4个生境中繁殖能力的大小依次为:耕地>草地>路旁>疏林。耕地中银胶菊种群密度、分枝数和头状花序数较小,但结实率和种子千粒重大,种子产量最多,繁殖能力最强;草地种群密度大,产生的分枝数和头状花序数多,因此种子产量高,繁殖能力次之;路旁开花植株数、分枝数、头状花序数虽多,但结实率和种子千粒重小,种子产量也较草地少;疏林种群密度小,产生的分枝数和头状花序数少,种子产量最低,但其种子质量高。总的来说,银胶菊在所研究的生境中繁殖能力均极强大。1
银胶菊在同一生境中在2008年和2009年的结实率和种子千粒重无显著差异。在疏林和公路旁生境中,不同年份的头状花序、种子产量无明显区别,可能这两个生境银胶菊受人为影响较少,繁殖能力变化较小。但在开阔草地、耕地生境中,不同年份的头状花序数、种子产量均有显著差异,这两个生境由于放牧和耕作活动,银胶菊受影响大,繁殖能力变化较大。
(3)银胶菊通过调节各构件的生物量分配比例来适应不同生境条件。在草地和路旁中,提高根生物量比例来竞争地下资源;在疏林地中,提高茎和叶生物量比例来提高对光的获取率;在耕地中,既提高根生物量比例来竞争地下资源,又提高叶生物量比例来竞争光。繁殖期各构件生物量与环境密切相关,并受植株密度和高度的制约。在不同生境不同季节各构件生物量分配表现出较高的形态可塑性,与该植物具有强大的入侵性有关。
(4)银胶菊繁殖方式为有性繁殖,花果期较短,6月份时从出现花蕾到种子完全成熟仅需22天。花粉具锥状尖刺突起,极易粘附在昆虫上传播,组织化学主要含淀粉,散粉时间主要在9:00-13:00,此间活性较强,花粉/胚珠比率(P/O)高。柱头可授期为5天,最佳授粉期为柱头伸出后的第二和第三天的9:00-13:00。自交不亲和,不存在无融合生殖现象,繁育系统属于异交类型,需要传粉媒介,入侵生境中一些普通的昆虫能为其传粉,自然结实率较高。主要传粉昆虫有5种,包括膜翅目的蜜蜂、小蜜蜂、黑小蜜蜂和双翅目的黑边家蝇、丝光绿蝇,访花高峰期基本集中在10:00-13:00。
(5)银胶菊对不同土壤养分水平表现出不同的适应性。在一定范围内,银胶菊根生物量比随氮的增加而降低,总生物量、叶生物量比(LMR)、花果生物量比(FMR)、叶面积指数(LAI)、总叶面积(TLA)、叶面积比(LAR)、相对生长速率(RGR)、分枝数、头状花序数等随氮养分水平的增加而增加;而对于磷养分来说,叶生物量比、叶面积比、总叶面积、相对生长速率随磷水平的增加而升高;总生物量、叶面积指数、分枝数、头状花序数等在各磷水平下无明显变化。土壤N的增加提高了银胶菊的种子产量,土壤P的增加提高了种子质量。对不同N、P养分具有不同的适应性。
(6)银胶菊不能在光强低于5%的生境中生存,但能适应变幅较大的光环境(22%-100%),通过调节总生物量、根生物量比、叶生物量比、花生物量比、总叶面积、叶面积比来适应不同光强。
上述研究结果表明,银胶菊具有强大的繁殖能力,并且在不同生境不同季节繁殖能力不同,此外,该杂草还具有广泛的生态适应性。应该注意其在不同生境不同季节的入侵和控制策略。
Biological invasions destroy the structure and function of native ecosystems, make the loss of the biodiversity, and cause huge economic losses, as well as threats health of human being. It is an important element in global changes and has received considerable attention during the last decades. However, the mechanisms that underlie invasiveness are still not well explained. Researches, which identify the factors that influence invasions by alien plants species are needed to be carried out. This will help us improve our ability to predict and control potentially invasive species.
Parthenium hysterophorus (Asteraceae) is an invasive weed native to southern North America, Central America, the West Indies, and central South America. It is a famous noxious weed in the world. This weed has posed serious threats to crop production, natural biodiversity and human health, because of its prolific growth, rapid spreading, and production of toxic allelochemicals. It has a wide range of ecological adaptation and great reproductive potential. The populations of this invasive species have spread rapidly during recent years in China.
In this paper, we studied the phenology and quantitative characters of flowering and seed-set, biomass allocation as well as reproductive process of P. hysterophorus population in different environments by field investigation and control trial. We compared the reproductive potential of this weed in different environments and in different seasons according to above research. We also studied its ecological adaptability under different nitrogen and phosphorus levels and light levels. The objective of this research was to explore the reproductive capacity and mechanism of invasion of Parthenium weed, and provide some basic information for preventing and controlling this weed. The results are as follows:
(1) Parthenium weed has long reproductive period, which made this weed continuously produce abundant seeds and seedlings. It grew from March to October and propagated from early May to October in grassland, sparse forest, and roadside. It propagated until to December in farmland.
(2) The quantitative characters of flowering and seed set were different under different environments and in different seasons. Therefore, the reproductive capacity was also different. In grassland, sparse forest, and roadside, flowering plants, branch numbers and capitulum’s number as well as seed production were more in autumn than in summer and in spring. The reproductive capacity of the weed in the three habitats was the greatest in autumn. In farmland, branch number and capitulum’s number were the most in winter. The reproductive capacity was also the greatest.
In autumn, the sequence of reproductive capacity in four habitats was farmland > grassland > roadside > sparse forest. In farmland, the population density, branch number and capitulum’s number were less, but the seed production was the most and the reproductive capacity was the greatest because of high seed set and 1000-seed weigh. In grassland, the population density, branch number and capitulum’s number were more, the seed production was high, and the productive capacity was great. In roadside, the flowering plants, branch numbers and capitulum’s number were more than those in former habitats, but the seed set and1000-seed weigh were less, so the seed production was less than that in grassland. In sparse forest, the population density, branch number, capitulum’s number, and seed production were less than those in former habitats were, but the quality of seed was high. In a word, the reproductive capacity was very great in the studied sites.
The seed set and 1000-seed weigh in 2008 had no significant difference with those in 2009 in the same habitat. In sparse forest and roadside, the capitulum’s number and seed production in 2008 had no significant difference with those in 2009. The reproductive capacity had no much change in different years in the two habitats because of fewer disturbances. In grassland and farmland, the capitulum’s number and seed production in 2008 had significant difference with that in 2009. The reproductive capacity was various because of the interferences by grazing and farming.
(3) Parthenium weed adapted to different environments by adjusting each organ biomass ratio. In grassland and roadside, it competed underground resource by improving the root biomass ratio; in sparse forest, it captured more light by improving the shoot and leave biomass ratio. In farmland, it increased root biomass ratio to compete underground and improved leaf biomass ratio to capture light. The biomass allocations of Parthenium weed closely related with environments during the reproductive period, and were restricted by plant height and density. It had high plasticity to environment. The high plasticity under different environments in different seasons showed the powerful ability to invasion.
(4) Parthenium weed was fast-maturated weed, which propagated by seeds. It took only 22 days from the beginning of flower bud to seed maturity in June. Pollen grain with spine prominence was easy to attach the body of pollinators. Pollen-ovule (P/O) ratio was11556.6±323.2. Pollen was starchy and shed from 9:00 to 13:00. Pollen viability was high during this period. Stigma acceptability lasted 5 days. The optimal time for accepting pollen is the second and third day after stigma protruded. There was no apomixes. The breeding system of this weed was xenogamy in China and dependent on pollen vector. The pollinators were local general insects, such as Apis cerana, A. frorea, A. andreiformis, Musca hervei, Lucillia sericata. The peak for visiting flower was 10:00-13:00.
(5) Parthenium weed exhibited considerable nutrition-acclimation abilities. In a certain nitrogen range, root mass ratio (RMR) reduced with increasing nitrogen levels; total biomass, leaf mass ratio relative (LMR), flower and fruit mass ratio (FMR), leaf area index (LAI), total leaf area (TLA) and leaf area ratio (LAR), relative growth rates (RGR), branch number, as well as capitulum’s number increased with increasing nitrogen levels. To phosphorus nutrient, LMR, LAR, TLA, RGR increased with rising phosphorus levels. Total biomass, LAI, branch number, capitulum’s number had no difference among different phosphorus levels. Parthenium weed improved seed production with the increase of nitrogen, while improve seed quality with the increase of phosphorus. It could adjust the reproductive traits by environment nutrient which promote its invasiveness in rich nitrogen and phosphorus environments.
(6) Parthenium weed could not survive under low light condictions (5% of light intensity). However, it was able to acclimate to different light condiction (22-100% of light intensity) by adjusting total biomass, RMR, LMR, FMR, TLA, LAR.
The above research results show that the productive capacity of Parthenium weed,which exhibited various under different environments in different seasons, was very great. It has a broad ecological adaption under different nutrient levels and light levels. We should pay attention to the invasion of Parthenium weed under different environments in different seasons and take suitable controlling measures.
银胶菊(Parthenium hysterophorus)为菊科入侵植物,原产北美南部、中美洲、西印度洋群岛、南美中部;是国际性的大毒草。其疯狂生长、快速传播和产生有毒的化感物质,对作物产量、生物多样性和人类健康具有极大的威胁。该杂草具有广泛的生态适应性和很强的繁殖能力,在最近几年已发展成为我国危害较为严重的外来入侵植物。
本文通过野外定点观察、人工控制实验,对银胶菊在不同生境中的开花物候、种群开花结实数量特征、生物量及其分配、生殖过程进行了研究,比较银胶菊种群在不同生境不同季节的繁殖能力。另外,本文还研究了银胶菊对不同N、P养分水平的生态适应性以及在不同光照水平下的适应性。旨在揭示银胶菊的入侵能力和入侵机制,为银胶菊的预防和控制提供依据。主要研究结果如下:
(1)银胶菊繁殖期长,使其能产生大量的种子和幼苗,在较短时间内就能爆发和入侵。在草地、疏林和路旁生境中,银胶菊从3月至10月都有植株生长,从4月底5月初至10月均有开花结实。在耕地中,银胶菊从3至12月均有植株生长,从5月至12月均有开花结实。
(2)银胶菊在不同生境不同季节种群开花结实数量特征不同,繁殖能力不同。
在草地、疏林和路旁生境,秋季开花植株数、分枝数、头状花序数较夏季和春季多,产生的种子数也多,秋季是这三个生境银胶菊的繁殖能力最强的时期。在耕地中,冬季的分枝数和头状花序数比夏季和秋季多,种子产量相应也多。因此,耕地中冬季的繁殖能力最强。
秋季,银胶菊在4个生境中繁殖能力的大小依次为:耕地>草地>路旁>疏林。耕地中银胶菊种群密度、分枝数和头状花序数较小,但结实率和种子千粒重大,种子产量最多,繁殖能力最强;草地种群密度大,产生的分枝数和头状花序数多,因此种子产量高,繁殖能力次之;路旁开花植株数、分枝数、头状花序数虽多,但结实率和种子千粒重小,种子产量也较草地少;疏林种群密度小,产生的分枝数和头状花序数少,种子产量最低,但其种子质量高。总的来说,银胶菊在所研究的生境中繁殖能力均极强大。1
银胶菊在同一生境中在2008年和2009年的结实率和种子千粒重无显著差异。在疏林和公路旁生境中,不同年份的头状花序、种子产量无明显区别,可能这两个生境银胶菊受人为影响较少,繁殖能力变化较小。但在开阔草地、耕地生境中,不同年份的头状花序数、种子产量均有显著差异,这两个生境由于放牧和耕作活动,银胶菊受影响大,繁殖能力变化较大。
(3)银胶菊通过调节各构件的生物量分配比例来适应不同生境条件。在草地和路旁中,提高根生物量比例来竞争地下资源;在疏林地中,提高茎和叶生物量比例来提高对光的获取率;在耕地中,既提高根生物量比例来竞争地下资源,又提高叶生物量比例来竞争光。繁殖期各构件生物量与环境密切相关,并受植株密度和高度的制约。在不同生境不同季节各构件生物量分配表现出较高的形态可塑性,与该植物具有强大的入侵性有关。
(4)银胶菊繁殖方式为有性繁殖,花果期较短,6月份时从出现花蕾到种子完全成熟仅需22天。花粉具锥状尖刺突起,极易粘附在昆虫上传播,组织化学主要含淀粉,散粉时间主要在9:00-13:00,此间活性较强,花粉/胚珠比率(P/O)高。柱头可授期为5天,最佳授粉期为柱头伸出后的第二和第三天的9:00-13:00。自交不亲和,不存在无融合生殖现象,繁育系统属于异交类型,需要传粉媒介,入侵生境中一些普通的昆虫能为其传粉,自然结实率较高。主要传粉昆虫有5种,包括膜翅目的蜜蜂、小蜜蜂、黑小蜜蜂和双翅目的黑边家蝇、丝光绿蝇,访花高峰期基本集中在10:00-13:00。
(5)银胶菊对不同土壤养分水平表现出不同的适应性。在一定范围内,银胶菊根生物量比随氮的增加而降低,总生物量、叶生物量比(LMR)、花果生物量比(FMR)、叶面积指数(LAI)、总叶面积(TLA)、叶面积比(LAR)、相对生长速率(RGR)、分枝数、头状花序数等随氮养分水平的增加而增加;而对于磷养分来说,叶生物量比、叶面积比、总叶面积、相对生长速率随磷水平的增加而升高;总生物量、叶面积指数、分枝数、头状花序数等在各磷水平下无明显变化。土壤N的增加提高了银胶菊的种子产量,土壤P的增加提高了种子质量。对不同N、P养分具有不同的适应性。
(6)银胶菊不能在光强低于5%的生境中生存,但能适应变幅较大的光环境(22%-100%),通过调节总生物量、根生物量比、叶生物量比、花生物量比、总叶面积、叶面积比来适应不同光强。
上述研究结果表明,银胶菊具有强大的繁殖能力,并且在不同生境不同季节繁殖能力不同,此外,该杂草还具有广泛的生态适应性。应该注意其在不同生境不同季节的入侵和控制策略。
Biological invasions destroy the structure and function of native ecosystems, make the loss of the biodiversity, and cause huge economic losses, as well as threats health of human being. It is an important element in global changes and has received considerable attention during the last decades. However, the mechanisms that underlie invasiveness are still not well explained. Researches, which identify the factors that influence invasions by alien plants species are needed to be carried out. This will help us improve our ability to predict and control potentially invasive species.
Parthenium hysterophorus (Asteraceae) is an invasive weed native to southern North America, Central America, the West Indies, and central South America. It is a famous noxious weed in the world. This weed has posed serious threats to crop production, natural biodiversity and human health, because of its prolific growth, rapid spreading, and production of toxic allelochemicals. It has a wide range of ecological adaptation and great reproductive potential. The populations of this invasive species have spread rapidly during recent years in China.
In this paper, we studied the phenology and quantitative characters of flowering and seed-set, biomass allocation as well as reproductive process of P. hysterophorus population in different environments by field investigation and control trial. We compared the reproductive potential of this weed in different environments and in different seasons according to above research. We also studied its ecological adaptability under different nitrogen and phosphorus levels and light levels. The objective of this research was to explore the reproductive capacity and mechanism of invasion of Parthenium weed, and provide some basic information for preventing and controlling this weed. The results are as follows:
(1) Parthenium weed has long reproductive period, which made this weed continuously produce abundant seeds and seedlings. It grew from March to October and propagated from early May to October in grassland, sparse forest, and roadside. It propagated until to December in farmland.
(2) The quantitative characters of flowering and seed set were different under different environments and in different seasons. Therefore, the reproductive capacity was also different. In grassland, sparse forest, and roadside, flowering plants, branch numbers and capitulum’s number as well as seed production were more in autumn than in summer and in spring. The reproductive capacity of the weed in the three habitats was the greatest in autumn. In farmland, branch number and capitulum’s number were the most in winter. The reproductive capacity was also the greatest.
In autumn, the sequence of reproductive capacity in four habitats was farmland > grassland > roadside > sparse forest. In farmland, the population density, branch number and capitulum’s number were less, but the seed production was the most and the reproductive capacity was the greatest because of high seed set and 1000-seed weigh. In grassland, the population density, branch number and capitulum’s number were more, the seed production was high, and the productive capacity was great. In roadside, the flowering plants, branch numbers and capitulum’s number were more than those in former habitats, but the seed set and1000-seed weigh were less, so the seed production was less than that in grassland. In sparse forest, the population density, branch number, capitulum’s number, and seed production were less than those in former habitats were, but the quality of seed was high. In a word, the reproductive capacity was very great in the studied sites.
The seed set and 1000-seed weigh in 2008 had no significant difference with those in 2009 in the same habitat. In sparse forest and roadside, the capitulum’s number and seed production in 2008 had no significant difference with those in 2009. The reproductive capacity had no much change in different years in the two habitats because of fewer disturbances. In grassland and farmland, the capitulum’s number and seed production in 2008 had significant difference with that in 2009. The reproductive capacity was various because of the interferences by grazing and farming.
(3) Parthenium weed adapted to different environments by adjusting each organ biomass ratio. In grassland and roadside, it competed underground resource by improving the root biomass ratio; in sparse forest, it captured more light by improving the shoot and leave biomass ratio. In farmland, it increased root biomass ratio to compete underground and improved leaf biomass ratio to capture light. The biomass allocations of Parthenium weed closely related with environments during the reproductive period, and were restricted by plant height and density. It had high plasticity to environment. The high plasticity under different environments in different seasons showed the powerful ability to invasion.
(4) Parthenium weed was fast-maturated weed, which propagated by seeds. It took only 22 days from the beginning of flower bud to seed maturity in June. Pollen grain with spine prominence was easy to attach the body of pollinators. Pollen-ovule (P/O) ratio was11556.6±323.2. Pollen was starchy and shed from 9:00 to 13:00. Pollen viability was high during this period. Stigma acceptability lasted 5 days. The optimal time for accepting pollen is the second and third day after stigma protruded. There was no apomixes. The breeding system of this weed was xenogamy in China and dependent on pollen vector. The pollinators were local general insects, such as Apis cerana, A. frorea, A. andreiformis, Musca hervei, Lucillia sericata. The peak for visiting flower was 10:00-13:00.
(5) Parthenium weed exhibited considerable nutrition-acclimation abilities. In a certain nitrogen range, root mass ratio (RMR) reduced with increasing nitrogen levels; total biomass, leaf mass ratio relative (LMR), flower and fruit mass ratio (FMR), leaf area index (LAI), total leaf area (TLA) and leaf area ratio (LAR), relative growth rates (RGR), branch number, as well as capitulum’s number increased with increasing nitrogen levels. To phosphorus nutrient, LMR, LAR, TLA, RGR increased with rising phosphorus levels. Total biomass, LAI, branch number, capitulum’s number had no difference among different phosphorus levels. Parthenium weed improved seed production with the increase of nitrogen, while improve seed quality with the increase of phosphorus. It could adjust the reproductive traits by environment nutrient which promote its invasiveness in rich nitrogen and phosphorus environments.
(6) Parthenium weed could not survive under low light condictions (5% of light intensity). However, it was able to acclimate to different light condiction (22-100% of light intensity) by adjusting total biomass, RMR, LMR, FMR, TLA, LAR.
The above research results show that the productive capacity of Parthenium weed,which exhibited various under different environments in different seasons, was very great. It has a broad ecological adaption under different nutrient levels and light levels. We should pay attention to the invasion of Parthenium weed under different environments in different seasons and take suitable controlling measures.
引文
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