入侵种喜旱莲子草对光照强度的表型可塑性反应

对外来人侵种喜旱莲子草[Alternanthera philoxeroides（Mart．）Griseb．]在不同强度光照处理中的表型可塑性反应进行了研究。结果表明：在高光照（100％）、中等光照（60．4％）、低光照（35．4％）和弱光照（16．5％）条件下,随光照强度的降低,分枝强度、基株株长、茎节长度随之下降,总生物量及根、茎、叶生物量显著减少;中等光强度处理喜旱莲子草根冠比显著低于其他处理;弱光条件下喜旱莲子草生长迟缓,呈直立状。研究结果表明,光照强度是影响喜旱莲子草种群生存与维持、生长和成功入侵、扩散的重要因子,同时也说明光照强度较低的生态系统喜旱莲子草入侵成功的机会很小。     

喜旱莲子草茎叶解剖结构从原产地到入侵地的变异式样

长期以来人们一直认为,外来种入侵及其危害是由于一个物种从原产地到入侵地其环境因子改变(如天敌压力的减弱等)而导致的。近年来,越来越多的研究者开始认识到,生物入侵过程实际上是一个现代人类活动影响下的物种的快速进化过程,生物入侵的进化遗传学已成为入侵生物学研究中最活跃的分支之一。作者比较了来自原产地(阿根廷)和入侵地(中国和美国)的喜旱莲子草(Alternantheraphiloxeroides)的11个种群在茎、叶解剖结构方面的变异式样,发现所研究的19个性状在原产地(阿根廷)和入侵地(中国和美国)的变异情况明显不同:在原产地种群中,共有9个性状指标存在显著差异,遗传率在49–89%之间,这9个性状是气孔密度、气孔指数、茎直径、髓腔直径、维管柱直径、皮层厚度、维管柱面积比、髓腔面积比和叶形指数;而在入侵地种群间,19个性状指标均无明显差异。这表明喜旱莲子草从原产地到入侵地其遗传多样性降低;入侵地喜旱莲子草种群间的形态变异主要为表型可塑性。根据19个形态指标对喜旱莲子草11个种群进行主成分分析和聚类,结果显示:所有入侵地种群和原产地的Ar1种群(SantaFé,59°49′W,29°16′S)聚为一类,原产地的Ar4(Tandil,59°03′W,37°11′S)单独聚为一类,原产地的其他4个种群聚为一类。表明Ar1种群可能与入侵中国的喜旱莲子草在基因型上更为接近。这一结果为进一步揭示喜旱莲子草入侵机理(如杂交适应性)和在原产地寻求对应天敌的生物防治工作提供了基础数据。     

喜旱莲子草对模拟全天增温的可塑性: 引入地和原产地种群的比较

植物可以通过关键功能性状的表型可塑性来适应气候变暖背景下环境温度的增加。表型可塑性增强进化假说(evolution of increased phenotypic plasticity hypothesis)认为外来植物在引入地进化出了更强的表型可塑性。以往对该假说的验证多集中于外来植物对光照、水分、养分、邻体以及天敌等的可塑性进化, 而对增温条件下植物生长和功能性状可塑性进化的研究相对较少。仅有的几项研究多集中在温带地区, 且多集中于研究植物生长相关的性状, 而对植物的抗性和草食作用对增温的响应的关注相对较少。本研究采用同质园实验比较了喜旱莲子草(Alternanthera philoxeroides)引入地(中国)和原产地(阿根廷)各8个种群的生物量、功能性状和草食作用在热带地区(广州市增城区)对模拟全天增温2℃的响应差异。结果表明: (1)模拟全天增温显著降低了喜旱莲子草总生物量(-7.8%)、贮藏根生物量(-12.8%)、分枝强度(-11.6%)和茎端取食率(-34.4%)。(2)模拟全天增温造成的引入地种群总生物量降低幅度大于原产地种群; 模拟全天增温使引入地种群的比茎长和茎端取食率降低, 而原产地种群则相反。(3)无论是否模拟全天增温, 引入地种群的贮藏根生物量(+31.5%)、分枝强度(+38.5%)、比茎长(+30.2%)、根冠比(+24.5%)和比叶面积(+20.0%)均高于原产地种群, 茎端取食率则低于原产地种群(-35.8%)。这些结果表明, 热带地区全天增温2℃对喜旱莲子草是一种胁迫; 引入地种群的生物量对模拟全天增温2℃的响应更强, 而其株形相关性状(比茎长)和草食作用(茎端取食率)对模拟全天增温的可塑性方向与原产地种群相反。由于引入地种群在热带地区模拟全天增温条件下生物量的下降和草食作用的增加明显高于原产地种群, 因此在未来全球气候变暖的背景下, 热带地区温度升高可能不利于喜旱莲子草种群多度的增加。     

喜旱莲子草在青藏高原对模拟增温的可塑性: 引入地和原产地种群的比较

气候变暖背景下植物可通过关键性状的表型可塑性来适应环境温度的增加。表型可塑性增强进化假说预测定植到新环境中的入侵植物种群具有演化出更强表型可塑性的潜力。此前对可塑性进化的研究涵盖了外来植物性状对水分条件、光照变化、土壤养分、邻体根系以及天敌防御等的响应, 而较少有研究关注增温条件下植物重要性状的可塑性进化。已有的部分研究多集中在温带和热带地区, 而较少关注入侵植物在高寒地区对增温的响应; 且研究多集中在植物生长相关性状, 较少关注功能性状和防御性状。本研究采用同质园实验比较了喜旱莲子草6个引入地(中国)种群和6个原产地(阿根廷)种群, 在西藏拉萨模拟全天增温2℃处理下的适合度性状、功能性状和防御性状的响应差异。结果表明: (1)高寒地区模拟全天增温显著提高了喜旱莲子草总生物量(+36.4%)、地上生物量(+34.5%)、贮藏根生物量(+51.4%)和毛根生物量(+33.6%), 降低了分枝强度(-19.8%)和比茎长(-30.2%); (2)模拟全天增温使引入地种群的比叶面积和黄酮含量增加, 而原产地种群则相反。这些结果表明高寒地区全天增温2℃对喜旱莲子草可能是一种有利条件。引入地种群的适合度性状对模拟全天增温2℃的响应比原产地种群更强, 而其光能利用相关性状和防御性状的响应可能提升了其在高寒地区的适合度。因此, 在未来全球气候变暖的背景下, 高寒地区温度升高可能更有利于喜旱莲子草引入地种群的定植和扩散。     

喜旱莲子草对同基因型邻体根系的表型可塑性: 入侵地和原产地的比较

植物对邻体根系的表型可塑性是指与无邻体对照相比, 即使个体平均可获取土壤资源相同, 在有邻体根系存在时植物也会改变根系生物量分配, 并影响其他功能性状和适合度。表型可塑性进化假说(evolution of plasticity hypothesis)认为外来植物在入侵地进化出了更强的表型可塑性。对该假说的验证多集中于外来植物对光照、水分、养分以及天敌等的可塑性进化, 但对邻体根系的可塑性在入侵植物中是否发生进化尚未见报道。我们采用同质园实验比较了喜旱莲子草(Alternanthera philoxeroides)入侵地(美国)和原产地(阿根廷)各5个基因型的适合度与功能性状对同基因型邻体根系的可塑性。结果表明: 喜旱莲子草的根冠比(P = 0.088)和比叶面积(P = 0.007)对同基因型邻体根系的可塑性在入侵地和原产地基因型间存在差异: 入侵地基因型在有邻体根系时根冠比和比叶面积增加, 而原产地基因型则相反。但是, 总生物量、贮藏根生物量、比茎长和分枝强度对邻体根系的可塑性在入侵地和原产地间没有显著差异。此外, 与分隔邻体根系相比, 同基因型邻体根系存在时总生物量(+9.9%)和贮藏根生物量(+13.9%)显著增加, 比茎长(-9.5%)显著降低。最后, 与原产地基因型相比, 总体上入侵地基因型的总生物量(+62.0%)和贮藏根生物量(+58.9%)增加, 比茎长(-28.5%)和分枝强度(-42.8%)降低。这些结果表明喜旱莲子草入侵地基因型与资源利用相关功能性状(如根冠比和比叶面积)对邻体根系的可塑性方向与原产地基因型相反; 但适合度和株型相关性状(如比茎长和分枝强度)对同基因型邻体根系的可塑性与原产地没有差异。     

喜旱莲子草对喀斯特三种不同生境的可塑性反应

喜旱莲子草自入侵以来在我国各地广为传播蔓延,对人们生产生活产生了重要影响。该研究通过野外采样和室内理化指标分析,探讨喀斯特不同生境中喜旱莲子草的可塑性。结果表明:(1)在三种生境中喜旱莲子草的节间长度、叶面积、单株重和叶干重差异显著。(2)在岩石环境中其无性繁殖器官——茎的投入比例占其生物量的比例最大,其次是行道生境和湘江河道。(3)节间距随环境因子从岩石环境、绿化行道和水生生境的变化,依次增加,叶质比(叶面积/质量)也随水分环境的增加而增加;(4)在岩石环境中,喜旱莲子草通过缩短节间距并增加无性繁殖器官——茎的质量来增加其在水分匮乏生境的繁殖能力。该研究结果为喜旱莲子草今后的防治对策提供了科学依据。     

不同土壤水分和养分条件下喜旱莲子草与同属种生长状况的比较研究

喜旱莲子草(Alternanthera philoxeroides)入侵已在中国造成巨大的生态和经济损失。为揭示喜旱莲子草成功入侵的生态机制并预测其种群扩张趋势及其与环境因子的关系, 作者比较了喜旱莲子草与其同属的外来弱入侵种刺花莲子草(A. pungens)以及土著种莲子草(A. sessilis)在不同土壤水分、养分条件下的生长状况。结果显示: 在高水高肥条件下, 喜旱莲子草的生物量要高于刺花莲子草和莲子草, 而在低水低肥条件下却不如这两个同属种; 弱入侵种刺花莲子草在低水条件下的生物量要高于强入侵种喜旱莲子草和土著种莲子草, 说明植物的入侵性受环境条件的影响。另外, 强入侵种喜旱莲子草形态学性状的可塑性较高, 在各种条件下都具有较高的比叶面积, 暗示这两个指标可作为莲子草属外来植物入侵性的预测指标。     

光照和氮素对喜旱莲子草形态特征和生物量分配的影响

研究了两个光照梯度和3个土壤氮素水平交互作用对喜旱莲子草(Alternan thera philoxeroides(Mart.)Griseb.)形态特征和生物量分配的影响。结果表明,全光照促进喜旱莲子草总生物量的积累,但在遮荫条件下,喜旱莲子草可以通过增加株高、光合叶面积和改变生物量分配来适应弱光生境。土壤中氮素含量对喜旱莲子草生长有明显影响,总生物量、株高、叶面积、茎生物量比和叶生物量比等随土壤氮素水平增加而增加。光照和氮素的交互作用对总生物量、根生物量比、茎生物量比和叶生物量比也有显著影响。随着氮素水平的增高,遮荫和高光照处理下喜旱莲子草的叶面积、总生物量和叶生物量比之间的差异减小,而株高和根生物量比之间的差异增大。此外,光照强度对茎生物量比的影响具有明显的氮素浓度依赖性,低氮条件下,茎生物量比在高光照处理下显著高于遮荫处理,而在中氮条件下,遮荫处理却显著高于高光照处理,且在高氮处理下其差异进一步加大。这些结果表明喜旱莲子草在高氮素环境下能够通过形态可塑性和生物量分配模式的改变来适应弱光环境所带来的不利影响。研究结果不但可为研究喜旱莲子草对异质生境的入侵机制提供资料,也可为进一步研究喜旱莲子草的入侵和扩散与农业等生态系统中土壤氮素残留的关系提供参考。       

完全水淹环境中光照和溶氧对喜旱莲子草表型可塑性的影响

光照和溶氧是水环境和陆地环境间差异显著的两个环境因子,对水淹植物的生长和存活具有重要的意义。以三峡库区常见外来入侵植物喜旱莲子草(Alternanthera philoxeroides)为研究对象,考察了水体中的光照(L)和溶氧(DO)对完全水淹环境中喜旱莲子草的形态特征和生物量分配等表型可塑性的影响。实验设置水淹和非水淹对照两组处理,对水淹组的光照和溶氧两个环境因子再分别设置有(+)、无(-)以及高(+)、低(-)两种水平,共计4个处理。实验结果表明:(1)水淹可促进喜旱莲子草主茎和叶片发生可塑性反应,引发伸长生长。水淹后,其细长的主茎以及长而薄的直立叶更有利于植株早日出露水面。(2)完全水淹条件下,喜旱莲子草主茎和叶片的表型可塑性受光照和溶氧的复合影响,其中主茎的伸长生长主要受溶氧的影响,而叶片的形态变化则主要受光照影响。高溶氧处理下喜旱莲子草的主茎伸长生长显著(P0.05)。在相同光照条件下,高溶氧处理下喜旱莲子草的主茎长、节间数、节间长以及主茎长/主茎直径均明显高于低溶氧处理。不论有光还是无光,高溶氧处理下喜旱莲子草主茎长以及节间数的平均增长率均处于最高水平,分别为61.8%、34.2%。喜旱莲子草叶片的形态变化在有光处理下表现得尤为显著,其平均叶片长宽比、比叶面积以及叶倾角分别较水淹前增加了39.65%、28.3%、45.9°。(3)光照和溶氧对于喜旱莲子草不定根和分枝的发生及发展存在影响差异。有光条件下可促进植株抽枝,而高溶氧处理时更有利于植株生根。这些形态变化有助于喜旱莲子草扩大株型占据有利生境,进一步提高植株的水下存活能力。     

三种增温情景对入侵植物空心莲子草形态可塑性的影响

虽然国内外已对克隆植物的表型可塑性开展了大量研究,但是气候变暖对克隆植物,尤其是入侵性克隆植物形态可塑性的影响及其可能的生态学意义研究仍然有限。通过设置白天增温、夜间增温和全天增温3种方式,通过切断或不切断匍匐茎处理,探讨入侵植物空心莲子草(Alternanthera philoxeroides)形态特征对不同气候变暖情景的响应。研究发现:夜间增温2℃条件下,切断匍匐茎连接显著降低空心莲子草匍匐茎总长度和平均分株长度;白天增温和全天增温2℃对匍匐茎切断和连接组的空心莲子草的匍匐茎总长和平均分株长影响不显著;3种增温方式对空心莲子草的分蘖数均无显著影响。这些结果表明:空心莲子草对增温具有较高的耐受性,并对不同增温方式采取不同的响应策略;夜间增温可能通过光合补偿效应增加其匍匐茎的长度和平均分株长从而促进其水平方向的扩展以占据更加有利的生境,增强其入侵能力;白天和全天增温对整个克隆片段匍匐茎形态几乎无影响。     

Phenotypic and genetic differentiation between native and introduced plant populations

Plant invasions often involve rapid evolutionary change. Founder effects, hybridization, and adaptation to novel environments cause genetic differentiation between native and introduced populations and may contribute to the success of invaders. An influential idea in this context has been the Evolution of Increased Competitive Ability (EICA) hypothesis. It proposes that after enemy release plants rapidly evolve to be less defended but more competitive, thereby increasing plant vigour in introduced populations. To detect evolutionary change in invaders, comparative studies of native versus introduced populations are needed. Here, we review the current empirical evidence from: (1) comparisons of phenotypic variation in natural populations; (2) comparisons of molecular variation with neutral genetic markers; (3) comparisons of quantitative genetic variation in a common environment; and (4) comparisons of phenotypic plasticity across different environments. Field data suggest that increased vigour and reduced herbivory are common in introduced plant populations. In molecular studies, the genetic diversity of introduced populations was not consistently different from that of native populations. Multiple introductions of invasive plants appear to be the rule rather than the exception. In tests of the EICA hypothesis in a common environment, several found increased growth or decreased resistance in introduced populations. However, few provided a full test of the EICA hypothesis by addressing growth and defence in the same species. Overall, there is reasonable empirical evidence to suggest that genetic differentiation through rapid evolutionary change is important in plant invasions. We discuss conceptual and methodological issues associated with cross-continental comparisons and make recommendations for future research. When testing for EICA, greater emphasis should be put on competitive ability and plant tolerance. Moreover, it is important to address evolutionary change in characteristics other than defence and growth that could play a role in plant invasions.     

喜旱莲子草营养繁殖特征对干扰的响应

养分富集和物理干扰是人类活动影响下外来植物在入侵地所面对的重要生境特征,而克隆生长和营养繁殖是外来植物入侵高资源干扰生境的重要生活史特征。喜旱莲子草似（Alternanthera philoxeroides）是一个世界性的入侵种,它主要分布于暖温带-亚热带气候区各种淡水生态系统的水陆交界区域,这些生境易受养分富集和物理干扰的影响。喜旱莲子草在入侵地有性繁殖缺失,因而贮藏根形成、根芽萌发和分株生长等是其重要的生活史环节和营养繁殖特征。本文中通过种植实验研究了喜旱莲子草的营养繁殖特征对养分增加和物理干扰（贮藏根片段化和埋藏）的响应。结果表明：（1）高养分条件下喜旱莲子草的总生物量增加了约1．5倍,贮藏根分配（营养繁殖分配）增加了约15％,贮藏根平均长度增加了约1．5倍;（2）喜旱莲子草的根芽萌发数随贮藏根段大小的增加而增加,平均每克贮藏根（干重）能够萌发出约15个根芽,萌发出1个根芽只需要不到0．1g的贮藏根,埋藏深度不影响根芽萌发数;（3）贮藏根段大小和埋藏深度对喜旱莲子草分株的主枝长和平均相对生长率均没有显著影响。这些结果暗示,喜旱莲子草的营养繁殖特征对养分富集和物理干扰非常适应,能够促进其局域种群的快速增长和流域范围内集合种群的长期续存。     

Phenotypic plasticity of introduced versus native purple loosestrife: univariate and multivariate reaction norm approaches

The plastic responses to environmental change by Lythrum salicaria (purple loosestrife) were compared between native plants derived from seeds collected in Europe and those introduced into North America. Plants from nine populations each were grown under two levels of water and nutrient conditions. At the end of the growing season, samples were evaluated for eight traits related to their life history, plant size/architecture, and reproduction. Genetic (G), environmental (E), and G × E interactions were assessed by restricted maximum likelihood (REML) analysis of covariance (ANCOVA) and multivariate analysis of covariance (MANCOVA). Both univariate and multivariate reaction norm analyses were used to test for differences in the magnitude and direction of phenotypic plasticity between introduced and native plants. Under high-nutrient conditions, introduced plants were taller and had more branches and greater aboveground biomass. They also exhibited significantly greater amounts of phenotypic plasticity for aboveground biomass than did the natives in response to changing nutrient levels in standing water. This difference in univariate plasticity contributed to the general contrast in multivariate plasticity between introduced and native plants. These results support the idea that introduced plants may successfully invade a habitat and grow better than native plants in response to increased resources.     

Spatial arrangements affect suppression of invasive Alternanthera philoxeroides by native Hemarthria compressa

It has been hypothesized that differences in spatial arrangements change the relative frequency of intra- and interspecific encounters between plant species. Manipulating spatial arrangement may play a role in invasive plant suppression when native species are used as competitors against introduced species. In this study, a replacement series experiment was performed to investigate the effects of intraspecifically random and aggregated spatial arrangements on interactions between the native plant Hemarthria compressa and the invasive plant Alternanthera philoxeroides, to test the possibility and effectiveness of H. compressa in suppressing A. philoxeroides. When both species were planted in intraspecifically random spatial patterns, H. compressa had a competitive advantage over A. philoxeroides at relative densities of 2:2 and 3:1. However, aggregation increased the strength, and therefore the cost, of intraspecific competition in H. compressa, resulting in lower biomass production, which reduced its effectiveness as an interspecific competitor. As the relative density of H. compressa in mixtures decreased, plants allocated more biomass to belowground parts, but fewer interspecific encounters lowered its inhibitory effects on A. philoxeroides. The results not only confirm that the frequency of conspecific and heterospecific encounters can influence competitive outcomes, but also suggest that a reduction in the degree of spatial aggregation in H. compressa and an increase in its relative densities may be essential to increase the suppression of A. philoxeroides.     

Phenotypic plasticity in Cardamine flexuosa: variation among populations in plastic response to chilling treatments and photoperiods

Variability in the plastic responses of seven life history traits to different chilling and photoperiod regimes was studied in four wild populations of Cardamine flexuosa. This species, a winter-green or year-long annual, showed a facultative long-day and chilling requirement for flowering. Considerable variation among populations was noted in plasticity of all traits. Differences in plasticity were greater among three paddy field populations from different climatic areas than between adjacent populations under different disturbance regimes. A paddy field population (OP) and an adjacent orchard population (OG) exhibited similar plasticity, in both amounts and patterns of response. TP, a Japan Sea coast population, was distinct from three other populations, especially in the small amounts of plasticity. Differences in amount of response were much more common than differences in pattern of response. Character expressions of five traits were significantly correlated with the number of days to flowering. Days to flowering and the numbers of inflorescences and siliques showed high negative correlations because the branching ability of meristems decreased with delay of flowering.