菊科12种外来植物的有性繁殖特征和入侵风险研究

菊科植物具有极强的有性繁殖能力,对外来菊科观赏植物的引进存在极高的入侵风险。本研究以武汉常见的菊科12种外来观赏植物为对象,通过有性繁殖特征调查,研究植株的花部特征、花粉活力、传粉系统、种子产量,分析各物种有性繁殖能力的差异,评估其入侵能力。结果显示,多数物种可在隔绝传粉者情况下结实;访花昆虫类型多样,共观察到26种访花昆虫,其中膜翅目蜂类是主要的传粉者;结实情况具有一定差异。综合考虑花粉活力、传粉情况和结实情况,认为黑心金光菊(Rudbeckia hirta L.)、松果菊(Echinacea purpurea(L.)Moench)、大花金鸡菊(Coreopsis grandiflora Hogg.)、蛇目菊(Sanvitalia procumbens Lam.)的有性繁殖竞争力较高,对本地传粉环境可能有较大影响。本研究对近缘物种有性繁殖特征的综合分析结果可为评估外来物种的入侵风险提供一定的参考。     

群落组成及物种间相互作用对外来植物入侵的影响

外来植物入侵已成为严重的环境和社会问题,了解外来植物的入侵机制是有效控制其入侵的前提。生物阻抗假说认为,入侵地本地植物群落中的许多生物因子及生物过程能够抵御外来植物入侵。但关于群落抵抗外来种入侵的主要机制,目前还没有确定的结论。本文综述了群落中物种功能特征的多样性以及与外来种功能特征的相似度、植物与动物、植物与植物以及植物与土壤微生物间的相互作用等因素对外来植物入侵的影响,以及以前研究存在的不足。未来研究应该注重不同条件下植物与植物间的相互作用;不同竞争强度下,植物与食草动物的相互作用;植物、动物及土壤微生物三者之间的相互作用对外来植物入侵的影响。这些研究不仅能够丰富和完善入侵生态学理论,而且对于预测外来植物未来的扩散范围,合理有效地管理生态系统,防止外来植物入侵,保护本地生物多样性具有重要的实际意义。     

基于功能性状的外来植物入侵预测模型框架构建

预测外来植物的潜在入侵性已成为生物多样性保护研究的重要内容,外来植物与乡土物种间的亲缘关系是预测外来植物能否成功入侵的一个重要途径。然而,达尔文归化难题却预测了两种截然不同的结果(即达尔文归化假说和预适应假说)。该研究解析了达尔文归化难题的内涵,提出了基于功能性状的外来植物与乡土群落间的相似性关系应该是进行外来植物入侵预测的重要切入点,而功能性状的种间分化与种内变异可能是外来植物成功入侵的两种不同生态策略。在此基础上,该研究还通过物种功能性状的多维超体积构建了外来植物与乡土群落间的相似性,提出了基于这种相似性的外来植物入侵预测的研究框架和基本流程。该模型框架的建立有助于理解外来植物的入侵机制,对外来植物的潜在入侵性预测提供了理论依据。当然,要实现外来植物能否成功入侵的准确预测,不仅依赖于功能性状的选择,还要考虑入侵的生境依赖性、空间尺度的重要性以及乡土群落的可入侵性等,未来的研究重点是通过控制实验对该模型进行验证和进一步完善。     

海南岛外来植物入侵风险评价指标体系

基于海南岛生物入侵防控及外来物种管理的需求,对海南外来入侵植物的种类、分布情况和危害现状进行了调查。调查发现,目前海南受外来植物入侵影响较重,外来入侵植物共约160种,隶属于38个科。构建了适应于海南当地生态环境特点与经济社会发展的外来植物风险评估指标体系,该指标体系共分3级,包括5个一级指标,和相应的12个二级指标,42个三级指标,采用层次分析法确定了体系各级指标的权重。对海南50种常见的主要外来入侵植物进行了风险评估,并按照评估结果将外来入侵植物划分为不同风险等级,提出相应的管理策略。外来入侵生物风险评估体系可被作为管理部门制定早期预警和决策措施的重要工具,通过风险评估与风险管理,建立一套应对海南外来生物入侵的快速反应体系,为外来生物入侵的防范和生态安全提供保障。     

青藏高原及周边高山地区的植物繁殖生态学研究进展

青藏高原及周边高山地区孕育了极为丰富的植物多样性资源, 研究该地区植物如何顺利完成繁殖过程有助于我们理解植物对典型高山环境的进化和适应机制。该文综述了青藏高原地区高山植物在资源分配、繁殖方式、花部特征演化等方面的研究进展, 包括全球气候变化对植物繁殖特征的影响, 以及一些新技术和新方法在本研究领域的应用。在高山地区限制性环境中, 随海拔升高, 繁殖分配通常表现出增大的趋势, 其中投入到雄性资源的比例上升, 但具体的资源分配模式还要取决于植株的交配系统、个体大小、生活史特征、遗传特性以及环境中的资源有效性等。面对资源和传粉的双重限制, 植物在不同繁殖方式之间存在权衡, 当传粉者稀少时, 克隆繁殖和自交有利于繁殖保障; 而有性繁殖和异交能够提高种子的质量和后代的遗传多样性, 从而在复杂多变的气候条件下有利于种群的维持。因此, 不同繁殖方式的结合以及泛化的传粉互作网络可能是应对高山限制性环境的最优选择。花部特征的演化主要受到当地传粉者的选择压力, 但是外来传粉者、植食者、盗蜜者以及非生物环境(如温度、雨水和紫外辐射等)对花部性状的影响越来越受到重视。近年来, 青藏高原因其脆弱性和对气候变化的高度敏感性而在全球气候变化研究中备受关注, 以全球变暖和氮沉降增加为显著特征的全球气候变化正在直接或间接地影响着该地区高山植物的繁殖特征。气候变化影响植物和传粉者的物候并引起物种的迁移, 最终将导致植物与传粉者的时空不匹配。植物通过改变花部特征(花展示、花冠结构、花报酬的数量和质量)来响应气候变化, 这可能会改变其传粉者的类型、数量和访花行为, 从而最终影响植物的繁殖成功。3D打印和高通量测序等新技术和新方法的应用有助于促进植物繁殖生态学研究的进一步发展。3D打印的花能够精确控制其形态构造, 可以用于研究精细的花部特征变化对于传粉者行为的影响, 在此基础上与人工饲养的传粉者结合使用, 有助于进一步研究传粉者介导的花部特征演化。随着高通量测序技术的发展, 植物繁殖生态学领域, 尤其是花部特征演化的许多重要问题的潜在机制得以深入研究。该文最后提出了目前研究中需要注意的问题以及值得深入研究的发展方向。     

外来植物入侵的化感作用机制探讨

在最近的几十年,在全球范围内的外来植物入侵的频率达到了前所未有的水平,所以对其入侵机制的研究显得尤为重要。目前,已有几种假说从不同的侧面解释外来植物的成功入侵。本文在综述前人工作基础上,从化感作用的角度解释外来植物入侵,并给出解释模式。由于本土动植物对某些外来植物释放的化感物质比较敏感,外来植物可以利用它的“化学武器”与本土植物竞争,抵御植食性动物的取食和病源菌的感染,从而在与本土物种的相互干扰中占据优势,实现成功入侵。进一步从理论和应用的角度探讨了化感作用作为外来植物入侵的机制给我们的启示,并对今后这方面研究提出几点建议和看法。     

青藏高原及周边高山地区的植物繁殖生态学研究进展

青藏高原及周边高山地区孕育了极为丰富的植物多样性资源,研究该地区植物如何顺利完成繁殖过程有助于我们理解植物对典型高山环境的进化和适应机制。该文综述了青藏高原地区高山植物在资源分配、繁殖方式、花部特征演化等方面的研究进展,包括全球气候变化对植物繁殖特征的影响,以及一些新技术和新方法在本研究领域的应用。在高山地区限制性环境中,随海拔升高,繁殖分配通常表现出增大的趋势,其中投入到雄性资源的比例上升,但具体的资源分配模式还要取决于植株的交配系统、个体大小、生活史特征、遗传特性以及环境中的资源有效性等。面对资源和传粉的双重限制,植物在不同繁殖方式之间存在权衡,当传粉者稀少时,克隆繁殖和自交有利于繁殖保障;而有性繁殖和异交能够提高种子的质量和后代的遗传多样性,从而在复杂多变的气候条件下有利于种群的维持。因此,不同繁殖方式的结合以及泛化的传粉互作网络可能是应对高山限制性环境的最优选择。花部特征的演化主要受到当地传粉者的选择压力,但是外来传粉者、植食者、盗蜜者以及非生物环境(如温度、雨水和紫外辐射等)对花部性状的影响越来越受到重视。近年来,青藏高原因其脆弱性和对气候变化的高度敏感性而在全球气候变化研究中备受关注,以全球变暖和氮沉降增加为显著特征的全球气候变化正在直接或间接地影响着该地区高山植物的繁殖特征。气候变化影响植物和传粉者的物候并引起物种的迁移,最终将导致植物与传粉者的时空不匹配。植物通过改变花部特征(花展示、花冠结构、花报酬的数量和质量)来响应气候变化,这可能会改变其传粉者的类型、数量和访花行为,从而最终影响植物的繁殖成功。3D打印和高通量测序等新技术和新方法的应用有助于促进植物繁殖生态学研究的进一步发展。3D打印的花能够精确控制其形态构造,可以用于研究精细的花部特征变化对于传粉者行为的影响,在此基础上与人工饲养的传粉者结合使用,有助于进一步研究传粉者介导的花部特征演化。随着高通量测序技术的发展,植物繁殖生态学领域,尤其是花部特征演化的许多重要问题的潜在机制得以深入研究。该文最后提出了目前研究中需要注意的问题以及值得深入研究的发展方向。     

植物物种入侵研究

详细阐述了生物入侵对生态系统的危害、对社会经济造成的严重损失,重点论述了植物外来种入侵的机制：化学介导机制、植物基因交流机制和干扰机制等;影响入侵的因素：外因、内因,其中外因主要是人为原因和环境因素。总结了植物生态入侵的预防、控制、去除策略及方法,包括机械、化学和生物方法,探讨了今后植物生态入侵的研究重点。     

丛枝菌根真菌（AMF）对外来植物入侵反馈机制的研究进展

丛枝菌根真菌（AMF）在植物群落竞争演替、物种多样性的形成及群落空间分布格局、植物群落对全球变化的响应中均起着重要的调节作用;同样也能影响外来植物与本地植物的互作,影响外来植物入侵过程中植物群落演替进程,甚至决定入侵的成败。因此,AMF与外来植物共生及其对外来植物入侵的反馈已成为国际上外来植物入侵机制研究的一个热点。本文基于外来植物的入侵过程,从AMF对外来植物生长、外来植物与本地植物竞争关系的影响,以及外来植物入侵对AMF的影响及AMF对入侵的反馈3个方面综述了AMF对外来植物入侵的反馈机制。外来植物可以通过多种途径改变土著AMF的群落结构和功能,而土著AMF也能直接或间接地改变甚至逆转外来植物与入侵地植物的互作关系。未来的研究不仅需要考虑AMF与外来植物共生的菌根特性和对竞争关系的影响,还需要通过大尺度条件下的野外试验及室内补充试验深入探究影响AMF在外来植物与本地植物竞争演替中的作用的生物和非生物因子,以全面解释AMF影响外来植物入侵的反馈机制。     

中国公布的3 批外来入侵植物种类特征与入侵现状分析

生物入侵是当今世界各国发展的一大挑战。外来入侵物种已对各国经济造成了不同程度的影响, 中国也未能幸免。为了防止外来物种入侵, 减轻其对我国造成的影响, 国家环保总局先后于2003 年、2010 年及2014 年公布了3 批中国外来入侵物种名单。据统计, 中国公布的3 批外来物种名单中的外来入侵植物共29 种, 占半数以上, 其中菊科和禾本科植物合计20 种, 约占3 批外来入侵植物总种数的68.96%; 生活型为草本的植物最多, 占96.55%; 原产地为美洲的占93.10%。3 批外来入侵植物中62.07%的植物是非有意引入的, 现今已全部建立种群, 且在全国各地大范围扩散。     

Invasive and Non-Invasive Congeners Show Similar Trait Shifts between Their Same Native and Non-Native Ranges

Differences in morphological or ecological traits expressed by exotic species between their native and non-native ranges are often interpreted as evidence for adaptation to new conditions in the non-native ranges. In turn this adaptation is often hypothesized to contribute to the successful invasion of these species. There is good evidence for rapid evolution by many exotic invasives, but the extent to which these evolutionary changes actually drive invasiveness is unclear. One approach to resolving the relationship between adaptive responses and successful invasion is to compare traits between populations from the native and non-native ranges for both exotic invaders and congeners that are exotic but not invasive. We compared a suite of morphological traits that are commonly tested in the literature in the context of invasion for three very closely related species of Centaurea, all of which are sympatric in the same native and non-native ranges in Europe and North America. Of these, C. solstitialis is highly invasive whereas C. calcitrapa and C. sulphurea are not. For all three species, plants from non-native populations showed similar shifts in key traits that have been identified in other studies as important putative adaptive responses to post-introduction invasion. For example, for all three species plants from populations in non-native ranges were (i) larger and (ii) produced seeds that germinated at higher rates. In fact, the non-invasive C. calcitrapa showed the strongest trait shift between ranges. Centaurea solstitialis was the only species for which plants from the non-native range increased allocation to defensive spines, and allocated proportionally less resources to reproduction, patterns contrary to what would be predicted by theory and other empirical studies to enhance invasion. Our results suggest caution when interpreting the commonly observed increase in size and reproductive capacity as factors that cause exotics to become invaders.     

Plant reproductive systems and evolution during biological invasion

Recent biological invasions provide opportunities to investigate microevolution during contemporary timescales. The tempo and scope of local adaptation will be determined by the intensity of natural selection and the amounts and kinds of genetic variation within populations. In flowering plants, genetic diversity is strongly affected by interactions between reproductive systems and stochastic forces associated with immigration history and range expansion. Here, we explore the significance of reproductive system diversity for contemporary evolution during plant invasion. We focus in particular on how reproductive modes influence the genetic consequences of long-distance colonization and determine the likelihood of adaptive responses during invasion. In many clonal invaders, strong founder effects and restrictions on sexual reproduction limit opportunities for local adaptation. In contrast, adaptive changes to life-history traits should be a general expectation in both outbreeding and inbreeding species. We provide evidence that evolutionary modifications to reproductive systems promote the colonizing ability of invading populations and that reproductive timing is an important target of selection during range expansion. Knowledge of the likelihood and speed at which local adaptation evolves in invasive plants will be particularly important for management practices when evolutionary changes enhance ecological opportunities and invasive spread.     

细根与根系总质量比的相似性导致了共存的本土植物和入侵植物之间植物-土壤反馈的对比

三维土壤异质性对种子萌发影响的实验研究 土壤生物通过植物-土壤的反馈作用调控植物间相互作用和外来植物入侵。因此,探明植物-土壤反馈作用种间差异的形成原因,对于预测土壤生物在植物入侵过程中的作用具有重要价值。近期的研究发现,植物性状可以用于预测植物-土壤反馈作用。同时,研究发现植物入侵也与植物的一些性状相关联,暗示植物-土壤反馈作用通过植物性状与外来植物入侵之间存在关联,但尚缺乏实验证明。鉴于此,本研究选取了3对近缘入侵和本土植物为对象,比较了其植物-土壤反馈作用,探讨了植物-土壤反馈作用与植物根部性状的关联性。首先,通过种植实验植物3个月,驯化采自于田间的土壤。之后,将实验植物种植于对照和灭菌处理的同种或近缘(同科或同属)种驯化过的土壤中,评价同种或近缘种根际土壤生物对植物生长的净反馈作用(与灭菌土比较),并比较了两类土壤生物对植物的反馈作用。总体而言,同种或近缘种根际土壤生物对入侵与本土植物的净反馈作用无显著差异,两类土壤生物对入侵和本土植物的反馈作用亦无显著差异。土壤反馈作用的强度和种植于对照土壤中植物细根生物量比存在正相关关系,且入侵与本土植物细根生物量比无显著差异。这一发现表明：相似的细根生物量比可能是导致入侵与本土植物间土壤反馈作用无差异的一个重要原因。为提升人们对土壤生物在植物入侵过程中 作用的认识,亟需选取更多入侵与本土植物(尤其是亲缘关系较远的物种)开展实验研究,进一步探明植物性状、土壤反馈作用和外来植物入侵之间的关联性。     

入侵植物的生理生态特性对碳积累的影响

随着国际贸易的发展和人们交往的增加以及全球环境的变化,生物种类在全球扩散的机会也大大增加,从而为生物入侵创造了机会。生物入侵不仅给农林牧生产造成损失,而且具有长期的生态学效应。外来种的成功入侵不是其自身某一个特性决定的,而是其特性与新的生境综合作用的结果。外来入侵种生理生态特性的研究对其预测和防治具有重要的意义。目前对入侵种生理生态特性的研究较少。已有的研究表明,与本地种相比入侵种可能通过提高光合能力、资源利用率、表型可塑性、化感作用,以及降低繁殖成本等增加植株碳积累,促进其入侵。但并不是所有的入侵种都同时具有这些特性。生境不同限制性资源不同,入侵机制就不同。成功的入侵种应该能够高效地利用生境中的限制性资源,并且能够较快地调节自身的生理特性以适应波动的资源环境。     

土壤养分对菊科一年生入侵种和本地种繁殖性状的影响

入侵植物繁殖性状的研究可为揭示植物入侵机制提供重要的科学依据。为研究土壤养分对入侵植物和本地植物繁殖性状的影响, 并进一步研究养分添加是否更能促进入侵植物的繁殖能力, 我们设置了低、高两个养分水平, 通过同质园实验比较了不同土壤养分对假臭草(Praxelis clematidea)、胜红蓟(Ageratum conyzoides)、三叶鬼针草(Bidens pilosa) 3种菊科一年生入侵种和夜香牛(Vernonia cinerea)、一点红(Emilia sonchifolia)、墨旱莲(Eclipta prostrata) 3种本地种繁殖性状的影响。研究结果显示, 养分添加提高了6种菊科植物的开花株高、株高、地上生物量、单粒种子重量、总花序数、每花序种子数、总种子数量、总种子重量, 并使开花时间提前、花期延长。养分添加对入侵种的开花株高和单粒种子重量的提高幅度要比本地种更显著。相对于部分本地植物(夜香牛、墨旱莲), 养分添加更能促进部分入侵植物(假臭草、胜红蓟)的繁殖能力。三叶鬼针草和一点红的总种子数量和总种子重量在两种土壤养分水平下均较小。本地种墨旱莲的总种子数量和总种子重量在低养分条件下高于3个入侵种。这些结果表明, 高土壤养分仅能促进部分入侵植物相对于部分本地植物的繁殖能力。     

Asexual and sexual reproductive strategies in clonal plants

Most plants can reproduce both sexually and asexually (or vegetatively),and the balance between the two reproductive modes may vary widely between and within species.Extensive clonal growth may affect the evolution of life history traits in many ways.First,in some clonal species,sexual reproduction and sex ratio vary largely among populations.Variation in sexual reproduction may strongly affect plant's adaptation to local environments and the evolution of the geographic range.Second,clonal growth can increase floral display,and thus pollinator attraction,while it may impose serious constraints and evolutionary challenges on plants through geitonogamy that may strongly influence pollen dispersal.Geitonogamous pollination can bring a cost to plant fitness through both female and male functions.Some co-evolutionary interactions,therefore,may exist between the spatial structure and the mating behavior of clonal plants.Finally,a trade-off may exist between sexual reproduction and clonal growth.Resource allocation to the two reproductive modes may depend on environmental conditions,competitive dominance,life span,and genetic factors.If different reproductive modes represent adaptive strategies for plants in different environments,we expect that most of the resources should be allocated to sexual reproduction in habitats with fluctuating environmental conditions and strong competition,while clonal growth should be dominant in stable habitats.Yet we know little about the consequence of natural selection on the two reproductive modes and factors which control the balance of the two reproductive modes.Future studies should investigate the reproductive strategies of clonal plants simultaneously from both sexual and asexual perspectives.     

Biogeographic differences between native and non-native populations of crayfish alter species coexistence and trophic interactions in mesocosms

Biogeographical comparisons of native and non-native populations allow researchers to understand the degree to which traits contributing to invasion success are intrinsic or change during the invasion process. Here, we investigate whether traits underlying interspecific competition change following invasion and whether these alter the impacts of two crayfish congeners that have invaded into each other’s native ranges. Specifically, we compared native and non-native populations of rusty (Faxonius rusticus) and virile crayfish (F. virilis). We compared native and non-native populations of each species using laboratory assays to examine aggression and large mesocosms with the congeners in sympatry to examine growth and survival as well as impacts on lower trophic levels. We found that non-native virile crayfish were more aggressive in response to a threat than native virile crayfish and exhibited greater growth and survival in sympatry with rusty crayfish. These intraspecific differences were large enough to alter coexistence between species in the mesocosm experiment, which is consistent with patterns of coexistence between these species in the field. We did not observe differences in traits between native and non-native rusty crayfish, but rusty crayfish were consistently competitively dominant over virile crayfish in paired laboratory assays. Non-native populations of both species had greater impacts on lower trophic levels than native populations. Taken together, these findings provide new evidence that trait changes during invasions may enhance ecological impacts of invasive animals and their ability to compete with closely related native species.     

Management of invasive plants through ecological resistance

Despite debates on the real impact of plant invasion on native biodiversity, there remain many situations where exotic invasive plants must be managed and habitats restored. Restoration practices that build on plant community assembly principles could be useful to delay or prevent re-invasion after control, but there are still few syntheses of the biodiversity theory, ecological mechanisms and experimental evidence relevant to invasive plant management, possibly delaying applications. To provide such a synthesis, we review current knowledge on three key determinants of invasion success: biotic resistance, abiotic constraints, and propagule pressure. We elaborate on the ecological mechanisms at play for each determinant and emphasize, using case studies, their relevance for invasive plant management and ecological restoration. We find evidence that restoring a plant cover can enhance invasion resistance, but the challenge for both research and field applications is to understand how multiple determinants interact in relation to species traits in the fields. Failure to recognize these interactions and their effect on community assembly processes may explain some of the mixed species responses observed. While we need control and restoration case studies with local species at different sites, the development of a coherent, dynamic and adaptive framework around biotic/ecological resistance will have to go beyond the idiosyncrasy of the many species and systems being tested. Emphasizing the functional diversity of the restored community seems a promising approach when facing potentially multiple invaders and/or fluctuating abiotic conditions.     

Asexual and sexual reproductive strategies in clonal plants

Most plants can reproduce both sexually and asexually (or vegetatively), and the balance between the two reproductive modes may vary widely between and within species. Extensive clonal growth may affect the evolution of life history traits in many ways. First, in some clonal species, sexual reproduction and sex ratio vary largely among populations. Variation in sexual reproduction may strongly affect plant’s adaptation to local environments and the evolution of the geographic range. Second, clonal growth can increase floral display, and thus pollinator attraction, while it may impose serious constraints and evolutionary challenges on plants through geitonogamy that may strongly influence pollen dispersal. Geitonogamous pollination can bring a cost to plant fitness through both female and male functions. Some co-evolutionary interactions, therefore, may exist between the spatial structure and the mating behavior of clonal plants. Finally, a trade-off may exist between sexual reproduction and clonal growth. Resource allocation to the two reproductive modes may depend on environmental conditions, competitive dominance, life span, and genetic factors. If different reproductive modes represent adaptive strategies for plants in different environments, we expect that most of the resources should be allocated to sexual reproduction in habitats with fluctuating environmental conditions and strong competition, while clonal growth should be dominant in stable habitats. Yet we know little about the consequence of natural selection on the two reproductive modes and factors which control the balance of the two reproductive modes. Future studies should investigate the reproductive strategies of clonal plants simultaneously from both sexual and asexual perspectives. Translated from Acta Phytoecologica Sinica, 2006, 20(1): 174–183 [译自: 植物生态学报]