入侵植物加拿大一枝黄花化学性状的增强及其对竞争的作用
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摘要
生物入侵是当今生态学研究的核心问题。尤其是近几年来,对于入侵生物学中讨论和争议的核心问题——为什么入侵种在入侵地表现得活力更强、更具有竞争力?—一科学家提出了多种理论假说。其中,从化感角度提出的"Novel Weapons Hypothesis"(“新奇武器假说”)被认为是很多入侵植物的入侵机制。该假说指出,入侵植物能产生一些化学物质,而这些物质可以发挥较强的化感作用,成为植物和土壤微生物之间相互作用的调节者。由于不同区域的植物群落共同进化的轨迹不同,被入侵群落的植物对这些化学物质缺乏适应性,而这些化学物质也就成为了外来种入侵的武器。但是,目前关于入侵植物释放的化感物质在入侵地是否增强的研究甚少。
     因此,本论文以入侵欧亚的杂草加拿大一枝黄花(Solidago canadensis L.)为例,通过生物地理比较(中国和美国取样)和同质园种植实验,探讨入侵植物的化感性状在入侵地是否发生了快速适应性增强;中国种群的竞争作用是否大于美国种群的竞争作用;化感作用在入侵植物竞争过程中的贡献大小如何。本研究取得了以下主要研究成果:
     1外来植物加拿大一枝黄花的化学性状
     加拿大一枝黄花在种群扩张过程中,向土壤中释放的化感物质会逐年积累,3类主要化感物质(总黄酮、总酚类和总皂苷)含量显著高于对照组中未被加拿大一枝黄花侵染过的土壤。从化感效应来看,加拿大一枝黄花显著抑制了莴苣(Lactuca tatarica B.)种子的萌发和苜蓿(Medicago sativa L.)的生长,随着提取物浓度的升高或者加拿大一枝黄花种植密度的增加(即化感物含量的增加),化感抑制效应随之显著增强。
     土壤灭菌处理显著影响了加拿大一枝黄花释放的3类主要次生代谢产物在土壤中的存留率。与灭菌处理土壤相比,这3类主要次生代谢产物在不灭菌的土壤表现为刚开始快速降解,随后降解速度变得缓慢,最后维持在一定水平(总黄酮、总酚类和总皂苷的最终浓度分别约为26.04μg/g,17.01μg/g和55.18μg/g)。
     2外来植物加拿大一枝黄花化感性状的增强
     生物地理水平上的比较实验和温室的同质园实验中,中国种群都比美国种群表现有更高的生物量、株高、叶片总数、根冠比、SLA(比叶面积)、可溶性糖含量及蛋白质含量这些表征性状。
     另外,我们还发现,中国种群的地上部分所含的总黄酮、总酚类和总皂苷的含量比美国种群分别高34.22%-34.28%、34.20%-66.53%和75.56%-107.80%;地下部分的总黄酮、总酚类和总皂苷的含量比美国种群分别高184.93%-209.06%、60.20%-67.20%和49.59%一77.33%。中国种群地上部分和地下部分的粗提液对莴苣种子的抑制率分别比美国种群高17.50%-65.98%和21.03%-70.43%,说明加拿大一枝黄花在入侵进程中,显著增强了其“化学防御武器”这个性状,暗示了入侵植物并不总是以降低化学防御的能量分配为代价。
     3外来植物加拿大一枝黄花的竞争比较
     当以鸡眼草为目标植物时,邻株植物加拿大一枝黄花(中国种群根状茎繁殖)密度的增加,显著降低了鸡眼草的生长,即显著增强了对鸡眼草的竞争作用。
     当以加拿大一枝黄花(中国种群和美国种群种子繁殖)为目标植物时,中国种群的株高、生物量和花序干重都显著高于美国种群。而且,在邻体植物鸡眼草不同密度条件下(除4株/m2),中国种群的竞争作用都显著高于美国的种群。
     当加拿大一枝黄花和鸡眼草1:1竞争时,以根状茎繁殖的加拿大一枝黄花中国种群地上部分的竞争作用比分别以种子繁殖的美国种群和中国种群的竞争作用强179.08%和22.71%,而地下部分的竞争作用分别比以种子繁殖的美国种群和中国种群的竞争作用强197.68%和31.89%,说明地下部分的竞争力要大于地上部分。实验结果表明,中国种群在长期的入侵进程中,竞争能力得到增强。
     4化感性状对入侵植物竞争的贡献
     当以鸡眼草为目标植物时,化感作用的贡献率随着邻体植物加拿大一枝黄花密度的增加而减弱,意味着竞争作用逐渐转向以资源竞争为主;当以加拿大一枝黄花为目标植物时,在鸡眼草的任何密度条件下,化感作用只在中国种群的竞争过程中扮演了重要的角色,说明加拿大一枝黄花在入侵的过程中增强的化感性状对竞争的重要作用;当加拿大一枝黄花和鸡眼草1:1竞争时,化感性状对以根状茎繁殖的加拿大一枝黄花的竞争力贡献也最大,地上部分和地下部分分别达到31.09%和46.12%,而化感性状对以种子繁殖为主的美国种群的竞争力的贡献最低,地上部分和地下部分分别只达到0.48%和0.80%,说明对于加拿大一枝黄花来说,还产生了以种子繁殖变为以根状茎繁殖为主的化感性状的转变。
Invasion by exotic plant species is recognized as an important ecological problem. Several hypothesises have been proposed to explain why some invasive plants are more vigorous in their introduced range comparing to in their native range."Novel weapons hypothesis" is one of these hypothesises, which poists that some invaders transform because they possess novel biochemical weapons that function as unusually powerful allelopathic agents, or as mediators of new plant—soil microbial interactions. Root exudates that are relatively ineffective against their natural neighbors because of adaptation, may be highly inhibitory to newly encountered plants in invaded communities. And these chemicals are the "Novel Weapons" of invasive plants. However, whether and how the alleochemicals evolve rapidly in the introduced range is not well known.
     So, I conducted a series of experiments through biogeographical investigation and common garden approaches (eliminating environmental differences) to address the following questions:(1) whether S. canadensis increases its allelopathy in the introduced range compared to in its original range;(2) whether S. canadensis increases its competitive ability in the introduced range compared to in the its original range; and (3) whether allelopathy contributes to the enhanced competititive ability of Solidago. Main results obtained from these experiments are as follows:
     1. Chemical traits of invasive Solidago canadensis
     Concentrations of total flavones, total phenolics, and total saponins in soil accumulated as the population of S. canadensis developed in the field experiment and as density of S. canadensis increased in greenhouse experiment. And the contents of these three allelochemicals in the soil grown Solidago before were significantly higher than that in the soil grown native plants before. Seed germination rate of Lactuca tatarica Breitung and growth of Medicago sativa L. were significantly suppressed as the concentration of allelochemicals released from Solidago increased.
     Total flavones, total phenolics, and total saponins in soil were significantly degraded when the soil was not sterilized. And the contents of three main group compounds declined rapidly6days after incubation, then kept constant at the level of26.04μg/g,17.01μg/g and55.18μg/g respectively.
     2. The evolution of allelopathy in invasive Solidago canadensis
     The results of biogeographical comparisons of Solidago canadensis L. populations and common garden experiments demonstrated that Chinese Solidago have significantly higher shoot height, biomass, the ratio of root to shoot biomass (R/S), leaf numbers, specific leaf area (SLA). Moreover, Solidago of Chinese populations have more34.22%-34.28%total flavones,34.20%-66.53%total phenolics and75.56%-107.80%total saponins than American populations in the aboveground extracts and have more184.93%-209.06%total flavones,60.20%-67.20%total phenolics and49.59%-77.33%total saponins than American populations in the belowground extracts.
     Allelopathic effects of Chinese Solidago on lettuce significantly increased by17.50%-65.98%(aboveground) and21.03%-70.43%(belowground) than that of American Solidago, indicating that during the evolution of Solidago canadensis, the higher performance of S. canadensis in China was not only attributed to the growth and functional traits but also to an increased allelopathy.
     3. The evolution of competition in invasive Solidago canadensis
     When native plant Kummerowia striata was set as the target plant, the growth of K. striata was significantly reduced as the density of neighbor plant S. canadensis (rhizome grown) increased, implying that the competitive abilities of S. canadensis enhanced with the increasing density.
     When Solidago canadensis (seed grown) from different origins (China and America) was set as the target plant, Chinese Solidago had more biomass and height than that of American Solidago. At each density (except for4seedlings/ m2) of neighbor plant K. striata, competitive ability of Chinese Solidago was always higher than American Solidago.
     When Solidago canadensis was grown with Kummerowia striata (1:1), the competitive ability of Chinese Solidago (rhizome grown) were significantly higher by179.08%(aboveground) and197.68%(belowground) than that of seed grown Solidago from America, and by22.71%(aboveground) and31.89%(belowground) than that of seed grown Solidago from China. The results demonstrated that Chinese Solidago have evolved enhanced competitive ability during the invasion.
     4. Contributions of allelopathy to the competition
     Contrary to the results of enhanced competitive abilities of Chinese S. canadensis with target plant Kummerowia striata as the density of S. canadensis increased, allelopathic contributions declined significantly, indicating that resource competition gradually became the dominant form of competition when population of Solidago was very large. But allelopathic contributions were consistent with the results of increased competitive abilities of rhizome grown Chinese Solidago when Solidago grown with Kummerowia striata (1:1), comparing to the seed grown Solidago from China and America. Allelopathy made most contributions to the aboveground (31.09%) and belowground competition (46.12%) of rhizome grown Chinese Solidago. However, allelopathic contributions of seed grown Solidago from America was only up to0.48%(aboveground) and0.80%(belowground), implying that Chinese Solidago had evolved enhanced allelopathic advantage which contributed much more to the enhanced competititive ability.
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