克隆植物的表型可塑性与等级选择

表型可塑性是指生物个体生长发育过程中遭受不同环境条件作用时产生不同表型的能力。进化的发生有赖于自然选择对种群遗传可变性产生的效力以及各基因型的表型可塑性。有足够的证据说明表型可塑性的可遗传性,它实际上是进化改变的一个成分。一般通过优化模型、数量遗传模型和配子模型来研究表型可塑性的进化。植物的构型是相对固定的,并未完全抑制表型可塑性。克隆植物因其双构件性而具有更广泛的、具有重要生态适应意义的表型可塑性。构件性使克隆植物具有以分株为基本单位的等级结构,从而使克隆植物的表型选择也具有等级性。构件等级一般包含基株、克隆片段或分株系统以及分株3个典型水平。目前认为克隆植物的自然选择有两种模式,分别以等级选择模型和基因型选择模型表征。等级选择模型认为:不同的等级水平同时也是表型选择水平,环境对各水平具有作用,各水平之间也有相互作用,多重表型选择水平的净效应最终通过繁殖水平——分株传递到随后的世代中。基因型选择模型指出:克隆生长引起分株的遗传变异,并通过基株内分株间以及基株间的非随机交配引起种子库等位基因频率的改变,产生微进化。这两种选择模式均突出强调了分株水平在自然选择过程中的变异性以及在进化中的重要性,强调了克隆生长和种子繁殖对基株适合度的贡献。基因型选择模型包含等级选择模型的观点,是对等级选择模型的重要补充。克隆植物的表型可塑性表现在3个典型等级层次上,由于各层次对自然选择压力具有不同的反应,其表型变异程度一般表现出“分株层次>分株片段层次>基株层次”的等级性反应模式。很多证据表明,在构件有机体中构件具有最大的表型可塑性,植物的表型可塑性实际上是构件而非整个遗传个体的反应。这说明克隆植物的等级反应模式可能具有普适性。如果该反应模式同时还是构件等级中不同“个体”适应性可塑性反应的模式,那么可以预测:1)在克隆植物中,分株层次受到的自然选择强度也最大,并首先发生适应性可塑性变化,最终引起克隆植物微进化;2)由于较弱的有性繁殖能力,克隆植物在进化过程中的保守性可能大于非克隆植物。克隆植物等级反应模式的普适性亟待验证。     

四种菊科植物开花期构件生物量及表型可塑性比较

本文对粤东地区菊科2种入侵植物三叶鬼针草(Bidens pilosa)、加拿大蓬(Conyza canadensis)以及2种本土植物宽叶鼠曲草(Gnaphalium adnatum)、夜香牛(Vernonia cinerea)3种生境下的构件生物量、生物量分配及变异趋势进行了研究,对株高与各构件之间的关系建立了定量化描述模型,以揭示4种菊科植物在不同小生境中各构件的变异大小及生殖分配策略;对4种植物繁殖与营养构件之间的关系进行相关性、回归分析和生长趋势分析,以确定入侵种是否存在较高的表型可塑性。结果表明:三叶鬼针草在贫瘠、干燥的生境中,茎生物量分配显著增大,叶和花序生物量分配有减小的趋势;加拿大蓬在肥沃、潮湿的生境中其茎、叶、花序及总生物量显著大于其他2个生境(P0.01);宽叶鼠曲草和夜香牛在肥力较差、干燥的环境中,根生物量分配增大,茎和花序生物量分配有减小的趋势;三叶鬼针草、加拿大蓬和夜香牛各构件生物量的变异系数宽叶鼠曲草;相关性检验和生长趋势分析显示,三叶鬼针草、加拿大蓬和夜香牛繁殖与营养构件之间为异速生长。因此,三叶鬼针草、加拿大蓬和夜香牛具有较高的表型可塑性,以增强对多样环境的耐受性和适应性。     

表型可塑性变异的生态-发育机制及其进化意义

表型可塑性赋予生物个体在不同环境条件下通过产生不同表型来维持其适合度的能力.研究结果显示多数可塑性变异的产生是基于对环境变异信号的响应、改变基因表达式样并调整发育轨迹的结果,表观遗传调控体系在基因选择性表达和可塑性变异的跨世代传递过程中发挥了重要作用.不同物种和种群对环境变化的敏感性、发生可塑性变异的能力以及可塑性反应模式不尽相同,预示着控制可塑性能力并独立于控制性状的可塑性基凶的存在,这些基因是直接响应环境信号并控制表型表达的调控基因.表型可塑性不仅是物种适应性进化的一个重要方面,也是选择进化的产物,物种的表型可塑性变异对其生态适应和进化模式有深远的影响.     

表型可塑性与外来植物的成功入侵

生物入侵是导致生物多样性丧失的第二大因素,掌握入侵机制可提高外来入侵物种防控管理能力。表型可塑性被认为是外来植物成功入侵的重要机制之一,具有确定的遗传基础。近30年来表型可塑性的研究发展迅猛,野外调查与移植实验相结合,宏观生态与分子生态交叉等多种方法被用来探索外来植物成功入侵的表型可塑性机制。本文从表型可塑性概念、研究发展历程及其在外来植物入侵中的作用等方面进行了综述,并对今后外来入侵植物表型可塑性的研究方向进行了展望。表型可塑性的生态作用是目前研究的关注重点,表型可塑性增强了外来入侵植物的适应性和入侵能力,与遗传分化一起共同促进了外来植物的成功入侵。未来多学科相结合的综合研究方法将成为表型可塑性研究发展趋势,气候变化、土地利用等因素与入侵植物表型可塑性相结合,以及运用基因组学和分子生物学等方法,定量分析比较个体、种群或物种间的表型可塑性是未来研究外来植物入侵机制的方向之一。     

环境信号诱导发育的可塑性

有关环境因素作为影响发育的信号导致产生表观多型性的研究,属于遗传学、发育生物学、进化生物学和生态学研究领域的热点问题,在长期的积淀和拓展中形成了一门新的交叉学科——生态发育生物学。该学科以发育的可塑性为理论基础,研究多种环境因子诱导机体在发育中产生表观多型性的机制,包括非遗传多型性和应激性多型性。对于环境、发育和进化三者关系的研究尤为重视。本文介绍了该学科形成的背景,并对其研究主题进行分析和归纳,重点讨论了不同环境因子导致动物表观多型性的机制,包括季节和捕食者诱导的非遗传多型性,营养和激素调节社会性昆虫的品级分化,温度依赖型性别决定中的基因、酶与激素,动物对环境的适应与进化,机体的免疫应答与神经元的可塑性、环境污染物的致畸作用等。并对生态学与发育生物学结合的未来研究前景做了展望。     

入侵植物银胶菊在不同生境下表型可塑性和构件生物量

研究了入侵植物银胶菊在4种不同小生境间花果期形态特征变化和构件生物量特征。结果显示:在植株密度小但土壤肥沃的小生境中,植株各形态指标如茎长、茎直径和花序直径等都明显高于其它小生境,在生物量结构特征上则表现为总生物量和花果生物量所占比例的升高。随着植株密度的增加以及土壤肥力下降,上述各形态指标都发生了较明显的变化,生物量投资也进行了优化配置,银胶菊表现出了较高的形态可塑性。银胶菊与觅光和竞争相关的几个指标如叶和根的比例都增加,但用于生殖构件的比例却减少了。相关分析显示,银胶菊花果期各构件生物量与高度成正相关,与密度为负相关,并受环境因素的制约。表明,较高的形态可塑性和较强的生殖配置策略可能是银胶菊成功入侵我国的重要特征。     

昆虫翅型分化的表型可塑性机制

翅多型现象在昆虫中广泛存在,是昆虫在飞行扩散和繁殖能力之间权衡的一种策略,对种群的环境适应性进化具有重要的意义。目前在植食性昆虫中研究较多,有关寄生蜂的翅型分化鲜见报道。综述了昆虫翅型分化的表型可塑性机制。遗传因素和环境因素均对昆虫翅的发育产生影响,基因型对翅型的决定具有显著作用,外界环境条件,包括温度、光周期、食物质量、自身密度、外源激素等因素对昆虫翅的发育也产生重要的调节作用,从而产生翅的非遗传多型性现象。此外,天敌的寄生或捕食作用可能会诱导某些昆虫的翅型产生隔代表型变化。对昆虫产生翅多型现象的生态学意义及其在生物进化过程中的作用进行了讨论,并探讨了寄生性昆虫翅型分化机制在生物防治上的可能应用途径。功能基因组学和表观遗传学的进一步发展可望为彻底揭示昆虫翅型分化机制提供新的机遇和技术手段。     

蚜虫的表型可塑性及其遗传基础

表型可塑性(phenotypic plasticity)是有机体在适应生物或非生物环境时呈现不同表型的能力,并且有遗传基础。蚜虫是农林业的重要经济害虫,易受外部环境因素和自身遗传因素的影响而表现出表型的可塑性。本文综述了外部环境因素（如寄主植物、温度、光照、天敌等）的变异对蚜虫表型的影响。总体来说,蚜虫表型会因寄主植物的种类、品系以及发育阶段和营养状况的不同而有所差异; 温度变化对不同蚜虫种类的生殖力、生存力以及有翅蚜产生与否有极大影响。研究人员利用RAPD-PCR、微卫星等分子遗传标记确认寄主植物和温度是造成蚜虫种群遗传分化的重要因素。就内部因素而言,不同的蚜虫种类以及同一种蚜虫的不同克隆系在表型和遗传进化上也有不同程度的差异,在蚜虫受外界条件影响的不同虫态以及不同体色克隆系、不同生活周期的类群之间, 其生物学、生态学和遗传学都有所差异。分析上述各个因素对蚜虫表型可塑性的影响,对于蚜虫的生态进化研究和有效治理蚜害均有重要意义。本文在最后讨论了还有必要深入研究的诸多问题,如表观遗传调控,包括DNA甲基化、基因所在的核小体上的组蛋白的共价修饰和染色质重塑、siRNA介导的基因沉默以及微RNA（microRNA 或 miRNA）调控的基因表达变化等,又如有翅蚜的表型和遗传学研究,以及全球气候变化对蚜虫的生态进化的影响等问题。     

鸟鸣及其鸣唱控制系统发育可塑性研究进展

鸟类的鸣唱控制系统已成为研究神经系统与学习、行为和发育相关的一个重要模型。鸣禽鸣唱学习行为的神经基础为脊椎动物复杂习得行为的解剖学功能定位提供了一个范例。它也可为我们研究人类语言学习记忆提供借鉴,对近年来在鸟类鸣唱及其呜唱控制系统发育可塑性方面的研究进展进行了综述。     

河北木蓝的叶表型可塑性研究

以中国分布最广、形态变异复杂且分类上存在争议的木蓝属植物河北木蓝(Indigofera bungeana Walp.)为研究对象,运用GIS技术从较大尺度上(17省28县29个居群)进行叶表型可塑性分析,利用表型可塑性指数和变异系数对叶表型可塑性进行评价,并对叶表型性状与环境因子的相关性进行分析.结果显示:河北木蓝叶表...     

Adaptive developmental plasticity: what is it,how can we recognize it and when can it evolve?

Developmental plasticity describes situations where a specific input during an individual''s development produces a lasting alteration in phenotype. Some instances of developmental plasticity may be adaptive, meaning that the tendency to produce the phenotype conditional on having experienced the developmental input has been under positive selection. We discuss the necessary assumptions and predictions of hypotheses concerning adaptive developmental plasticity (ADP) and develop guidelines for how to test empirically whether a particular example is adaptive. Central to our analysis is the distinction between two kinds of ADP: informational, where the developmental input provides information about the future environment, and somatic state-based, where the developmental input enduringly alters some aspect of the individual''s somatic state. Both types are likely to exist in nature, but evolve under different conditions. In all cases of ADP, the expected fitness of individuals who experience the input and develop the phenotype should be higher than that of those who experience the input and do not develop the phenotype, while the expected fitness of those who do not experience the input and do not develop the phenotype should be higher than those who do not experience the input and do develop the phenotype. We describe ancillary predictions that are specific to just one of the two types of ADP and thus distinguish between them.     

Developmental plasticity in plants: implications of non-cognitive behavior

There has been a surge of interest in phenotypic plasticity in the last two decades. Most studies, however, are being carried out within relatively narrow disciplinary frameworks. Consequently, researchers differ not only in their scientific agenda; they often use different terminologies and conceptual frameworks even when studying the very same phenomena. The diversity of approaches has often generated parallel bodies of theory on subjects that can be best understood in broader interdisciplinary terms. This special issue points out the differences between the concepts and questions that are characteristic of various approaches. Bridging all gulfs may be impossible and not necessarily desirable, yet, awareness of the varied approaches should be instrumental in promoting interdisciplinary advances. It is the contribution to such awareness that is the major purpose of this special issue, and for this reason it deals with molecular, physiological, ecological and evolutionary approaches to the study of developmental plasticity. So as to focus the discussion, six topics have been selected, ranging from the fundamental essence of developmental plasticity to its implications to ecology and evolution. These topics were considered by scholars who were chosen for the diversity of their research, not only their expertise. Rather than a comprehensive body of theory, the current issue thus seeks the diversity of opinions on the discussed topics. It is hoped that the confrontation, in its original Latin sense, which includes bringing together and discussion, of scholars who are studying these phenomena at very different levels and from different points of view will generate new insights and promote future interdisciplinary research.     

Shade avoidance syndrome in Picea omorika seedlings: a growth‐room experiment

The adaptiveness of shade avoidance responses to density was studied in Picea omorika seedlings raised in a growth‐room. Siblings of a synthetic population comprising 117 families from six natural populations were exposed to contrasting density conditions in order to score variation in phenotypic expression of several epicotyl and bud traits included in the shade avoidance syndrome. As predicted for the adaptive plasticity to foliage shade, epicotyl elongation traits tended toward higher, while axillary bud traits toward lower values in high‐density vs. low‐density conditions. Phenotypic selection analysis revealed that the elongated plants had greater relative fitness than the suppressed ones in both density treatments which could be ascribed to the effect of direct selection on epicotyl length. There was no evidence for plasticity costs associated with the expression of the shade avoidance phenotype either under low or under high density, with only a single exception. Estimates of variance component genetic correlations across densities were significantly different from unity for the majority of the seedling traits studied, indicating the existence of heritable variation within reaction norms of these traits. However, since all these correlations were positive in sign and large in magnitude, this conclusively means that the level of the additive genetic variation for plasticity in the shade‐avoidance traits of P. omorika is rather low.     

蚂蚁入侵适应性进化及其机制

入侵种被传至一个新的区域后,在面对当地某些生态因子的胁迫时如何发生适应性变化,是入侵生物学研究的一个重要课题。这方面信息对评价物种的入侵性、制订防控策略具有重要指导意义。入侵性蚂蚁作为无脊椎陆生动物中入侵最成功、危害最严重的一类,其生物学、生态学已得到大量研究,许多报道表明它们在定殖和扩张过程中会发生适应性变化。基于30多年的文献资料,从蚁群社会结构、超级蚁群形成能力、个体大小、觅食行为、种间竞争、天敌防御、适应低温能力、生态位转变等多个角度进行了全面总结,并针对这些方面适应性变化的机制,从表型可塑性、适应性进化、自身遗传学基础、与共生微生物的互利关系等角度进行了剖析。提出了今后有待重点研究的几个问题。将有助于深化对蚂蚁成功入侵机制的认识,为推进我国红火蚁(Solenopsis invicta Buren)、小火蚁(Wasmannia auropunctata Roger)等入侵性蚂蚁的研究与治理提供参考。     

Variation in the degree and costs of adaptive phenotypic plasticity among Rana temporaria populations

Adaptive phenotypic plasticity in the form of capacity to accelerate development as a response to pond drying risk is known from many amphibian species. However, very little is known about factors that might constrain the evolution of this type of plasticity, and few studies have explored to what degree plasticity might be constrained by trade-offs dictated by adaptation to different environmental conditions. We compared the ability of southern and northern Scandinavian common frog (Rana temporaria) larvae originating from 10 different populations to accelerate their development in response to simulated pond drying risk and the resulting costs in metamorphic size in a factorial laboratory experiment. We found that (i) northern larvae developed faster than the southern larvae in all treatments, (ii) a capacity to accelerate the response was present in all five southern and all five northern populations tested, but that the magnitude of the response was much larger (and less variable) in the southern than in the northern populations, and that (iii) significant plasticity costs in metamorphic size were present in the southern populations, the plastic genotypes having smaller metamorphic size in the absence of desiccation risk, but no evidence for plasticity costs was found in the northern populations. We suggest that the weaker response to pond drying risk in the northern populations is due to stronger selection on large metamorphic size as compared with southern populations. In other words, seasonal time constraints that have selected the northern larvae to be fast growing and developing, may also constrain their innate ability for adaptive phenotypic plasticity.