生物入侵: 中国学者面临的转化生态学机遇与挑战
Biological invasions: opportunities and challenges facing Chinese ecologists in the era of translational ecology
通讯作者: *E-mail:bool@fudan.edu.cn
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Corresponding authors: *E-mail:bool@fudan.edu.cn
Online: 2010-11-20
本文引用格式
李博, 马克平.
Bo Li, Keping Ma.
今天, 乘坐飞机长途旅行已是一件太普通不过的事情了, 我们可以在不到一天的时间内到达世界的任何大城市。在我们的孩提时代, 世界大城市是那么的遥不可及, 而今只不过是邻近的“小村落”而已, 世界变得越来越小。近代技术革新所带来的空间上的相对变小不仅给我们人类的各种交往和商业活动带来了便捷, 而且也有助于生物物种越过自然的物理屏障, 进行快速和低风险的长距离扩散, 从而重塑物种的分布区。从这种意义上来说, 全球化就像地质事件一样, 对物种的分布区产生着深刻的影响。就全球范围来看, 由人类有意或无意“嫁到”新的地理分布区的物种总数可能已达到50万种之多(Pimentel et al., 2001)。
尽管引入一个区域的物种中只有极少数会最终成为有害的外来入侵性生物, 即外来入侵种(invasive alien species), 但正是这少数的物种被全世界公认为是新千年最严重的生态、人类健康、经济威胁之一(Pimentel, 2002), 已给土著生态系统的生物多样性带来了灭顶之灾, 给人类健康造成了严重的威胁, 给所入侵区域的经济带来了巨大的损失。所以, 生物入侵的问题已成为全球性的重大生态和环境问题, 引起了各国政府、社会和学术界的普遍关注, 常常与全球变暖相提并论, 并被视为全球变化的重要组成部分(Vitousek et al., 1997)。
生物入侵过程压缩了传统生态学和进化生物学所涉及的时间和空间尺度, 如外来入侵种中的进化现象可以发生在数十个至数百个世代(而非百万代)的时间尺度内(Cox, 2004), 所以外来入侵种成为研究生态学和进化生物学重大问题的理想材料或模式生物。然而, 生物入侵问题不仅仅是科学家发表科学论文、满足自身研究兴趣的“宠物”, 而且由于生物入侵问题已成为政府和社会的心病, 所以它是联系科学家、政策制定者和公众的转化生态学(translational ecology)问题(Schlesinger, 2010), 要求生态学家以一种便于理解的方式将生物入侵研究的成果向公众普及, 或成为制定政策的依据和有意义的管理行动。
我国的大部分疆域处在中纬度地区, 景观、气候、生态系统类型多样, 因而是一个生物多样性的大国; 也正由于此, 来自世界各地的物种都很容易在我国的领土上安家落户, 也就是说, 我国疆土的可入侵性(invasibility)高。我国有着悠久的文明史, 引种在西汉时期就已开始, 引入了大量的经济物种及其相关的有害生物; 譬如, 石榴就是由张骞出使西域时从原产地中亚细亚带回。我国又是一个人口大国, 尤其是生产性人口巨大, 高强度的生产和经济活动给外来种的引入和散布创造了有利条件。更为重要的是, 我国目前正处在高速经济发展的时期, 进入了一个前所未有的国际物流和人流的高峰。尽管相关部门采取了许多积极有效的防御措施, 但外来物种进入我国的数量仍是空前的。如此等等, 很多因素使得我国成为了生物入侵的重灾国, 而且生物入侵问题还有日趋严重的趋势。
有鉴于此, 我国政府也采取了一系列积极的措施, 如环保部于2003年和2010年分别发布了第一、二批入侵物种清单, 共36种; 科技部设立了有关生物入侵的“973”项目, 专门研究生物入侵中的重大基础理论问题以及管理中的应用转化问题, 该项目现已进入滚动期。近10年来, 我国学术界对生物入侵的研究可谓异常火爆, 许多大学和科研单位均设有生物入侵研究团队。当然, 相对于欧美国家对生物入侵问题的重视程度, 相对于我国生物入侵问题的严重程度, 无论是我们政府的行动还是科学家的热情都来得比较晚。尽管如此, 我们也很高兴地看到, 我国在近10年内已在生物入侵研究领域中取得了可喜的进展, 尤其是有关互花米草(Spartina alterniflora)、紫茎泽兰(Eupatorium adenophorum)、喜旱莲子草(Alternanthera philoxeroides)、松材线虫(Bursaphelenchus xylophilus)、B型烟粉虱(Bemisia tabaci)等我国重要入侵种的研究已取得了国际认可的成果。
为了让更多的有志者了解生物入侵研究的最新进展, 加入到“外来种入侵”这一关系到国计民生的热点问题的研究领域中来, 推动国内这一领域研究的更快发展, 《生物多样性》杂志继2001年和2002年的“生物多样性与外来物种管理”专栏后, 愿再一次推出有关生物入侵的专辑。经过《生物多样性》编辑部、相关编委以及作者近半年的努力, 这一愿望终于在“第九届生物多样性保护与持续利用研讨会”之时得以实现。本专辑由15篇论文组成, 涉及生物入侵研究领域中三大核心问题, 即外来生物入侵的机制、后果以及控制和管理。
为了介绍生物入侵研究方法的动态, 我们首先安排了3篇相关综述。随着基因组学方法在不同领域中应用的深入, 该方法也有望为我们认识植物的入侵性提供一把金钥匙(Stewart, 2009)。尚蕾等(2010)综述了可用于植物入侵生物学研究的主要基因组学方法及其应用前景和面临的挑战, 可望能成为从分子水平认识入侵性的工具。朱丽和马克平(2010)总结了外来植物洲际间入侵与生态位属性研究进展, 牛红玉等(2010)则从外来植物入侵的全境性研究角度综述了相关的进展。这些综述为我们进一步开展相关研究提供了研究方法上的借鉴。
外来生物成功入侵的机制可以简单地归结为外来生物的入侵性特征(invasive traits)、入侵地的生态系统特性(ecosystem properties)以及人类活动相互作用的结果, 然而这又是一个悬而未决的问题。本专辑包括6篇相关论文。刘建等(2010)探讨了植物功能性状与外来植物入侵的关系, 指出植物入侵性是其功能性状和环境因素相互作用的结果。杂交(hybridization)与遗传渐渗(genetic introgression)被认为对植物入侵性有深刻影响, 卢宝荣等(2010)对此进行了全面的综述, 但他们也同时指出, 杂交-渐渗与其他理论的结合, 才能从更深的层次来解释外来种的入侵机制。
一个物种的入侵性是否与其遗传多样性相关呢?关于这一问题, 施雯等(2010)在其综述中予以回答。一个外来物种在其所抵达的新环境中能否成功定居并最终成为入侵种还与土著生态系统中物种间相互作用的类型与强度相关。本专辑中, 栾军波和刘树生(2010)探讨了媒介昆虫-病毒-植物相互作用对生物入侵的影响。显然, 从种间相互作用角度来审视生物入侵的机制具有特殊的意义。
生物入侵是全球环境变化的重要过程之一, 而且外来生物的成功入侵还与全球环境变化的其他过程密切相关。邓自发等(2010)以海滨生态系统为对象, 探讨了该类系统中生物入侵的动态如何受到全球变化主要过程的调节。在这一部分的最后, 我们安排了一篇个案的研究论文: 王坤等(2010)采用比较的方法, 研究了喜旱莲子草及其同属种在不同生长条件下的入侵性表达, 充分显示了外来植物的生物学特征与生长环境相互作用在入侵性表达中的地位。
生物入侵之所以受到广泛关注, 很大程度上是因为其造成的环境、经济和社会影响; 影响的程度常常是有关外来物种管理决策的重要依据之一。本专辑有3篇论文涉及到生物入侵的影响。类延宝等(2010)综述了外来植物入侵对土著生态系统中生物多样性及其进化的影响, 为认识生物入侵的影响提供了重要的借鉴。廖成章等(2010)以互花米草为例, 研究了其对重要生态系统过程——氮循环的影响, 认为互花米草取代土著种芦苇(Phragmites australis)后, 增加了生态系统中氮的输入, 最终可能有利于互花米草的快速扩张。从世界范围来看, 城市是人类活动的中心, 也是外来入侵种最丰富的生态系统之一。鞠瑞亭和李博(2010)对新近入侵我国城市的外来害虫——悬铃木方翅网蝽(Corythucha ciliata)进行了较为系统的介绍, 旨在为其控制与管理提供参考。
从应用层面来看, 生物入侵的研究成果最终要转化为公众的知识、管理行动或政府部门的决策依据, 所以有关入侵种控制和管理的研究显得尤为重要。本专辑仅有强胜等(2010)的论文聚焦于入侵种的管理问题, 该文较全面地分析了中国农业生态系统中外来种入侵及其管理现状, 将为我国该类系统中入侵生物的管理提供重要的参考。
由于前述的自然和社会原因, 我国生态系统作为一个总体, 可入侵性很高。就入侵植物而言, Weber等(2008)的初步研究表明, 我国至少有外来入侵植物270种, 而实际的数量很可能会更大。事实上, 通过与美国的植物入侵情况比较, Weber和 Li(2008)发现, 我国的入侵植物远比美国少(美国为519种), 所以未来的趋势可能是, 随着经济的发展我国将面临更多的外来植物的入侵, 尤其是水生和木本植物。他们的预测也可以通过本专辑中杨博等(2010)的论文得到反映。该论文提供了我国800种外来陆生草本植物多样性与生态学特征的初步分析, 说明我国外来植物的物种库很大, 也许若干年后那些目前尚未列入入侵植物清单中的物种成了难以对付的农林或环境杂草。
我们期待, 这期专辑的出版有助于推动我国在生物入侵研究领域的发展。根据生物入侵研究领域的特点以及目前的研究现状, 展望未来, 我们认为如下三个方面在将来的研究中有待进一步加强。
本专辑的大部分论文与植物入侵种相关, 有关真菌、病毒和其他微生物入侵的论文较少, 当然, 这也许是我们在组稿时未能充分重视这些类群。不过, 值得注意的是, 真菌和病毒中有很多是动、植物以及人类疾病的病原生物, 其入侵所造成的经济和人类健康影响是巨大的。美国生态学会前任主席Ron Pulliam教授在2004年复旦大学举办的高级生态学讲习班上指出, 入侵性传染病的生态和进化规律是生态学家在本世纪里所面临的四大挑战之一(Pulliam & Yu, 2005), 所以, 在未来的研究中值得给以更多的关注。
生物入侵之所以受到全球的重视, 不仅仅是因为它为科学家提供了研究生态学和进化问题的理想系统, 更重要的是, 它威胁着我们赖以生存的自然和人工生态系统。近期的外来入侵种编目工作表明, 中国有多达520多种外来入侵生物(万方浩等, 2009), 但是我们对绝大多数物种影响的认识是模糊不清的, 这给我们的依据管理(evidence-based management)带来了困难。因此, 在将来的研究中应进一步重视入侵种的直接和间接影响、产生影响的机制。 事实上, 生物入侵的生态学影响也是生态学家在本世纪里所面临的又一挑战(Pulliam & Yu, 2005)。
生物入侵并非独行, 而是与全球的环境变化问题同时袭来。可以说, 几乎所有全球环境变化的主要过程(如气候变化、生物多样性的丧失、土地利用方式的改变、氮沉降、环境污染等)都与生物入侵过程相互作用(Mooney & Hobbs, 2000; Perrings et al., 2010), 不仅正在催生出一次进化史上的革命(Cox, 2004), 而且将可能产生难以预测的生态、经济和社会后果。仅就气候变化来说, 其与生物入侵相互作用就可能产生五大后果(Hellmann et al., 2008): (1)改变入侵种的迁移(transport)和引入机制; (2)改变气候对入侵种的约束, 从而导致新入侵种的出现; (3)使现有入侵种的分布区发生改变, 扩张到新的区域; (4)改变现有入侵种所造成的影响; (5)用于管理入侵种的现有对策需要作适应性调整, 或者其管理效率会有所改变。所以, 生物入侵作为全球变化的重要过程之一, 应与其他过程整合起来加以研究, 认识这种相互作用的格局、机制与后果。
总之, 生物入侵是关系到国计民生的生态环境问题, 相关的“文章”最终应该写在管理和控制上, 这就要求科学家将研究成果转化成管理的行动、政策制定的依据和公众对生物入侵的认知。所以, 生物入侵生态学所涉及的主要问题正是最近Schlesinger(2010)所倡导的转化生态学问题。因此, 作为生物入侵工作者, 我们所面临的不仅是生物入侵问题本身, 还面临着知识转化的严峻挑战。当然, 机遇与挑战在并行走来。
最后, 我们诚挚地向《生物多样性》的编辑时意专、周玉荣和闫文杰, 编委丁建清、冯玉龙、李振宇、强胜、万方浩、张大勇等以及所有作者表示由衷的感谢, 没有他们的共同努力, 本专辑可能还只是一个美丽的梦想。
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Introduced species: a significant component of human-caused global change
Comparison of morphological traits between alligator weed and two congeners under different water and nutrient conditions
Alternanthera philoxeroides), a worldwide noxious weed, has caused enormous ecological and economic losses in China. To better understand the species’ successful invasion, and to predict relationships between expansion of its population and environmental conditions, we compared morphological traits of alligator weed with two closely related species, i.e. the geographically restricted alien A. pungens and the native A. sessilis, under different water and nutrient conditions in a common garden experiment. Compared to A. pungens and A. sessilis, alligator weed produced more biomass with high availability of water and nutrients, but did not in conditions of low water availability. These results suggest that the invasiveness of alligator weed may be influenced by environmental conditions. In addition, alligator weed had marginally greater specific leaf area (SLA) and phenotypic plasticity than its congeners. Phenotypic plasticity and SLA may thus have predictive value for invasiveness of alien species under multiple environmental conditions.]]>
Plant invasions in China: what is to be expected in the wake of economic development?
Invasive alien plants in China: diversity and ecological insights
Alien terrestrial herbs in China: diversity and ecological insights
On the niche stasis of intercontinental invasive plants
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