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入侵能力

外来植物的入侵能力与其性状之间的关系是入侵生态学中的基本问题之一。成功的入侵种常常能占据多样化的生境,并以广幅的环境耐受性为特征。遗传分化(包括生态型分化)和表型可塑性是广布性物种适应变化、异质性生境的两种不同但并不矛盾和排斥的策略。越来越多的实验证据表明,表型可塑性具有确定的遗传基础,本身是一种可以独立进化的性状。许多入侵种遗传多样性比较低,但同时又占据了广阔的地理分布区和多样化的生境,表型可塑性可能在这些物种的入侵成功和随后的扩散中起到了关键作用。本文首先介绍表型可塑性的含义,简述表型可塑性和生物适应的关系,然后从理论分析和实验证据两个方面论述了表型可塑性与外来植物入侵能力的相关性,最后针对进一步的研究工作进行了讨论。当然,并非所有入侵种的成功都能归因于表型可塑性,作者认为对于那些遗传多样性比较低同时又占据多样化生境的入侵种,表型可塑性和入侵能力的正相关可能是一条普遍法则,而非特例。     

群体表型性状研究揭示环境与遗传因素对霍山石斛表型及物种分类的影响

石斛属(Dendrobium)种类繁多, 属内物种具有丰富的表型多样性。霍山石斛(D. huoshanense)为我国特有物种, 其与河南石斛(D. henanense)和细茎石斛(D. moniliforme)以及铁皮石斛(D. catenatum)等近缘种表型相似, 在分类处理中存在争议。这种争议很大程度上与植物普遍存在的表型可塑性和代际共存有关。为探究环境和代际间遗传因素对霍山石斛表型性状的影响以及霍山石斛与近缘种的物种边界问题, 本研究观测了安徽省霍山县霍山石斛(野生、林间和温室F₁代、林间和温室F₂代)、野生河南石斛、细茎石斛和铁皮石斛, 共计16个群体2,279株植株的假鳞茎茎长等12个表型性状; 在种内层面, 首次借鉴生态学同质园实验和遗传学代际间性状比较的方法, 对霍山石斛群体表型性状进行差异显著性检验和95%置信区间比较以及主成分和变异系数等统计学分析。在种间层面, 对霍山石斛与河南石斛和铁皮石斛等近缘种群体表型性状进行比较和分析。结果表明, 环境因素对霍山石斛假鳞茎茎长和假鳞茎直径等具有显著的影响, 代际间遗传因素对霍山石斛假鳞茎直径具有显著的影响。霍山石斛与铁皮石斛和细茎石斛等近缘种群体在假鳞茎茎长、假鳞茎直径、花瓣长和花瓣宽等表型性状方面均存在显著性差异和间隔, 但与河南石斛仅在假鳞茎表型性状方面有显著性差异。我们的研究明确了环境和代际间遗传因素对霍山石斛表型性状的影响程度, 为霍山石斛与近缘种等争议物种的分类和鉴定提供了表型证据。     

蚜虫翅的响应

最新研究表明,与雄性蚜虫有翅和无翅多态性有关的一个基因的变异也和雌性蚜虫的相同性状的可塑性相联系。这个令人吃惊的发现表明,因遗传因素和环境因素而引发形态变异的生物体,是解开生物体表型可塑性之谜的有用的模式系统。长期以来,我们已经认识到表型可塑性的普遍重要性。作为对环境变异的响应而改变遗传发育过程的能力,表型可塑性是一种应付可预见的环境变异的有效手段,但是我们对其中的机制却不甚了解。英国科学家ChristianBraendle领导的小组正在研究的课题是表型可塑性性状和遗传控制多态性性状之间的关系,是否调控遗传多态性性状…     

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。     

果蝇中顺式调控序列的进化

顺式调控序列(cis-regularory sequences)是与基因表达调控相关、能够被调控因子特异性识别和结合的非编码DNA序列。顺式调控序列可以通过增减所含转录因子结合位点的数目,重构转录调控网络,以精准调控基因的时空表达模式,从而调控动物的生理和形态表型。顺式调控假说认为顺式调控序列进化是自然界丰富的动物表型进化的主要遗传机制。本文首先简述了顺式调控序列的结构和功能,然后重点讨论了近年来顺式调控序列进化调控果蝇表型进化如刚毛表型进化、色素沉积表型进化和胚胎发育方面的研究进展,阐释了顺式调控序列进化是动物表型进化的主要遗传机制之一。最后本文展望了顺式调控序列未来研究方向,例如应用功能基因组研究、开展ENCODE计划等,为进化发育生物学中顺式调控序列的研究提供了参考。     

蚜虫种群遗传多样性的影响因素及分子基础

蚜虫是一个复杂的类群 ,不同种群之间常常表现遗传多样性 ,特别是同种蚜虫的不同种群 ,这种多样性与环境因素 (寄主植物、地理气候条件等 )的影响密切相关 ,而且蚜虫种群多样性无论在细胞学水平 ,还是分子生物学水平均表现明显的遗传分化。该文在分析了蚜虫种群遗传多样性影响因素的基础上 ,从蚜虫核型变化、核DNA和线粒体DNA遗传分化和多样性方面总结了导致蚜虫种群遗传多样性的内在分子基础 ,并讨论了研究蚜虫种群遗传多样性的重要意义和前景     

牛、绵羊角的遗传定位及遗传机制研究进展

角属于动物颅骨附属物,为反刍动物所特有。牛(Bos taurus)、绵羊(Ovis aries)角的表型包括野生型两角表型、人工驯化的无角表型、畸形角等多种。牛和绵羊是阐明角的质量性状和数量性状之间的关系以及质量性状的多基因调控机制等方面的理想动物模型。近年来,对角性状研究不断深入,在阐明新器官起源进化、自然选择、性别选择和人工选择对角表型的影响等方面取得了一系列进展。本文详细介绍了角的研究概况、多角表型遗传定位、无角位点基因遗传定位和畸形角等,并对目前牛和绵羊角的遗传机制及存在的问题进行了分析,以期为反刍动物角性状和其他特异性性状遗传机制研究提供参考。     

克隆和有性亲体效应及其调控机制

植物表型受自身基因型、所处环境及其亲体所经历环境的共同影响;其中,亲体环境对子代表型的影响被称为亲体效应。亲体效应不仅可通过有性繁殖产生的种子传递给后代(即有性亲体效应),也可以通过克隆生长等无性繁殖产生的分株传递给后代(即克隆亲体效应)。亲体效应对植物种群,特别是对有性繁殖受限、缺乏遗传变异的克隆植物种群的长期进化可能发挥着极其重要的作用,因此,对亲体效应研究进展的梳理非常必要。对克隆亲体效应和有性亲体效应的内涵进行了阐释,并论述了克隆和有性亲体效应对子代表型、适合度、种内/种间竞争能力以及种群/群落结构和功能的潜在影响;阐述了亲体效应的潜在调控机制,包括供给机制、代谢物质调控机制、表观遗传机制等;论述了克隆亲体效应在克隆植物适应进化中的作用。未来可以就克隆亲体效应的遗传稳定性及其对克隆生活史性状变异的贡献程度,以及克隆和有性亲体效应引起的表型多样性对种内/种间关系、种群/群落多样性及生态系统结构、功能和稳定性的影响开展深入研究。     

胚胎孵化温度和初孵仔鱼发育温度对斑马鱼仔鱼热适应性的影响

发育可塑性是引起生物新性状产生与多样化的关键因素。在气候变化的背景下,探究鱼类热适应性的发育可塑性,有助于理解早期生活史阶段鱼类热适应表型的进化,具有重要的生态学意义。为探究胚胎孵化温度和初孵仔鱼发育温度对鱼类热适应表型的影响,本研究选取卵生鱼类斑马鱼(Danio rerio)为实验对象,采用2×2双因素研究设计,测定了不同胚胎孵化温度(低温组22℃、高温组28℃)和不同初孵仔鱼发育温度(低温组22℃、高温组28℃)及其交互作用对斑马鱼热偏好(偏好温度)、热回避(冷回避温度、热回避温度)、非回避温度范围以及热耐受(临界低温、临界高温、致死低温、致死高温)的影响。结果表明：胚胎孵化温度和初孵仔鱼发育温度对斑马鱼偏好温度、回避温度、临界温度以及致死温度等均有显著影响(P<0.05),胚胎孵化温度和初孵仔鱼发育温度交互作用对非回避温度范围影响显著(P<0.05);胚胎孵化热环境对早期生活史阶段斑马鱼热适应表型特征的塑造具有重要的调节作用,变动的发育温度环境可能扩展鱼类的适宜温度区间,从而提升其在未来变动气候环境中的存活力。     

Predator-induced morphological shift in the pea aphid

Aphids exhibit a polymorphism whereby individual aphids are either winged or unwinged. The winged dispersal morph is mainly responsible for the colonization of new plants and, in many species, is produced in response to adverse environmental conditions. Aphids are attacked by a wide range of specialized predators and predation has been shown to strongly influence the growth and persistence of aphid colonies. In two experiments, we reared two clones of pea aphid (Acyrthosiphon pisum) in the presence and absence of predatory ladybirds (Coccinella septempunctata or Adalia bipunctata). In both experiments, the presence of a predator enhanced the proportion of winged morphs among the offspring produced by the aphids. The aphid clones differed in their reaction to the presence of a ladybird, suggesting the presence of genetic variation for this trait. A treatment that simulated disturbance caused by predators did not enhance winged offspring production. The experiments indicate that aphids respond to the presence of a predator by producing the dispersal morph which can escape by flight to colonize other plants. In contrast to previous examples of predator-induced defence this shift in prey morphology does not lead to better protection against predator attack, but enables aphids to leave plants when mortality risks are high.     

Body colour and genetic variation in winged morph production in the pea aphid

Aphids (Homoptera: Aphidoidea) produce a number of different phenotypes in their life-cycle, among which are winged (alate) and wingless (apterous) morphs. Lowe & Taylor (1964) and Sutherland (1969a, b) were the first to suggest that aphid clones differ in their propensity to produce the winged morph and that in the pea aphid (Acyrthosiphon pisum Harris), this propensity is linked to the colour of the phenotype. We tested for the occurrence of genetic variation in winged morph production by rearing individuals from red and green clones of pea aphid under wing-inducing (crowding) and control conditions, and scored the phenotypes of their offspring. Clones differed significantly in alate production and red clones produced on average a higher proportion of winged morphs than green clones. Importantly, however, there was considerable variation between clones of the same colour. Broad-sense heritabilities of winged morph production were 0.69 (crowding treatment) and 0.63 (control). Clones also differed in the number of offspring they produced. When exposed to the crowding stimulus, aphids deferred offspring production, resulting in a higher number of offspring produced in the crowding treatment than in the control.     

Mechanisms of species‐sorting: effect of habitat occupancy on aphids' host plant selection

相似文献   

Climate effects on life cycle variation and population genetic architecture of the black bean aphid, Aphis fabae

Aphid species may exhibit different reproductive modes ranging from cyclical to obligate parthenogenesis. The distribution of life cycle variation in aphids is generally determined by ecological forces, mainly climate, because only sexually produced diapausing eggs can survive harsh winters or periods of absence of suitable host plants. Aphids are thus interesting models to investigate intrinsic and environmental factors shaping the competition among sexual and asexual lineages. We conducted a Europe-wide sampling of black bean aphids, Aphis fabae, and combined population genetic analyses based on microsatellite data with an experimental determination of life cycle strategies. Aphids were collected from broad beans (Vicia faba) as well as some Chenopodiaceae, but we detected no genetic differentiation between aphids from different host plants. Consistent with model predictions, life cycle variation was related to climate, with aphids from areas with cold winters investing more in sexual reproduction than aphids from areas with mild winters. Accordingly, only populations from mild areas exhibited a clear genetic signature of clonal reproduction. These differences arise despite substantial gene flow over large distances, which was evident from a very low geographic population structure and a lack of isolation-by-distance among 18 sites across distances of more than 1000 km. There was virtually no genetic differentiation between aphids with different reproductive modes, suggesting that new asexual lineages are formed continuously. Indeed, a surprising number of A. fabae genotypes even from colder climates produced some parthenogenetic offspring under simulated winter conditions. From this we predict that a shift to predominantly asexual reproduction could take place rapidly under climate warming.     

Parasitoids induce production of the dispersal morph of the pea aphid, Acyrthosiphon pisum

In animals, inducible morphological defences against natural enemies mostly involve structures that are protective or make the individual invulnerable to future attack. In the majority of such examples, predators are the selecting agent while examples involving parasites are much less common. Aphids produce a winged dispersal morph under adverse conditions, such as crowding or poor plant quality. It has recently been demonstrated that pea aphids, Acyrthosiphon pisum, also produce winged offspring when exposed to predatory ladybirds, the first example of an enemy‐induced morphological change facilitating dispersal. We examined the response of A. pisum to another important natural enemy, the parasitoid Aphidius ervi, in two sets of experiments. In the first set of experiments, two aphid clones both produced the highest proportion of winged offspring when exposed as colonies on plants to parasitoid females. In all cases, aphids exposed to male parasitoids produced a higher mean proportion of winged offspring than controls, but not significantly so. Aphid disturbance by parasitoids was greatest in female treatments, much less in male treatments and least in controls, tending to match the pattern of winged offspring production. In a second set of experiments, directly parasitised aphids produced no greater proportion of winged offspring than unparasitised controls, thus being parasitised itself is not used by aphids for induction of the winged morph. The induction of wing development by parasitoids shows that host defences against parasites may also include an increased rate of dispersal away from infected habitats. While previous work has shown that parasitism suppresses wing development in parasitised individuals, our experiments are the first to demonstrate a more indirect influence of parasites on insect polyphenism. Because predators and parasites differ fundamentally in a variety of attributes, our finding suggests that the wing production in response to natural enemies is of general occurrence in A. pisum and, perhaps, in other aphids.     

Geographic and clonal variation in the milkweed-oleander aphid,Aphis nerii (Homoptera: Aphididae), for winged morph production,life history,and morphology in relation to host plant permanence

Summary Populations of the milkweed-oleander aphid,Aphis nerii, were sampled in California, Iowa and Puerto Rico. Among these localities the aphid's host plants differ greatly in permanence. I compared populations for migratory potential, measured as the proportion of winged offspring produced in response to being crowded, and for life history and morphometric traits of the subsequent adult winged aphids. I predicted a negative correlation between degree of host plant permanence and migratory potential. As predicted, aphids from Iowa, where migration on to temporary hosts must occur each year, produce a greater proportion of winged offspring (37.7%) than those from California (25.7%) or Puerto Rico (31.6%) where hosts are more permanent. However, hosts in Puerto Rico appear to be more permanent than those in California, yet the difference between populations for migratory potential was opposite to that predicted. Within California the prediction again held: aphids collected from the most impermanent sites produce the greatest proportion of winged offspring. There were no population differences for any life history or morphometric traits of winged aphids that are important contributors to fitness or migratory ability such as time to reproductive maturity, fecundity or wing length. Nor did any traits covary with migratory potential. Thus, there does not appear to be an association of life history and morphology with migratory potential that could enhance the colonizing ability of migrant aphids. I was unable to detect population differentiation for life history and morphology even though there is ample genetic variation within populations on which selection could act and an absence of constraints arising from genetic correlations that could prevent appropriate evolution of traits within populations. The exploitation of temporary host plants therefore occurs by an increase in the number of colonists produced and not by change in life history or morphology of those colonists.     

EVOLUTION OF AN APHID-PARASITOID INTERACTION: VARIATION IN RESISTANCE TO PARASITISM AMONG APHID POPULATIONS SPECIALIZED ON DIFFERENT PLANTS

The evolution of associations between herbivorous insects and their parasitoids is likely to be influenced by the relationship between the herbivore and its host plants. If populations of specialized herbivorous insects are structured by their host plants such that populations on different hosts are genetically differentiated, then the traits affecting insect-parasitoid interactions may exhibit an associated structure. The pea aphid (Acyrthosiphon pisum) is a herbivorous insect species comprised of genetically distinct groups that are specialized on different host plants (Via 1991a, 1994). Here, we examine how the genetic differentiation of pea aphid populations on different host plants affects their interaction with a parasitoid wasp, Aphidius ervi. We performed four experiments. (1) By exposing pea aphids from both alfalfa and clover to parasitoids from both crops, we demonstrate that pea aphid populations that are specialized on alfalfa are successfully parasitized less often than are populations specialized on clover. This difference in parasitism rate does not depend upon whether the wasps were collected from alfalfa or clover fields. (2) When we controlled for potential differences in aphid and parasitoid behavior between the two host plants and ensured that aphids were attacked, we found that pea aphids from alfalfa were still parasitized less often than pea aphids from clover. Thus, the difference in parasitism rates is not due to behavior of either aphids or wasps, but appears to be a physiologically based difference in resistance to parasitism. (3) Replicates of pea aphid clones reared on their own host plant and on a common host plant, fava bean, exhibited the same pattern of resistance as above. Thus, there do not appear to be nutritional or secondary chemical effects on the level of physiological resistance in the aphids due to feeding on clover or alfalfa, and therefore the difference in resistance on the two crops appears to be genetically based. (4) We assayed for genetic variation in resistance among individual pea aphid clones collected from clover fields and found no detectable genetic variation for resistance to parasitism within two populations sampled from clover. This is in contrast to Henter and Via's (1995) report of abundant genetic variation in resistance to this parasitoid within a pea aphid population on alfalfa. Low levels of genetic variation may be one factor that constrains the evolution of resistance to parasitism in the populations of pea aphids from clover, leading them to remain more susceptible than populations of the same species from alfalfa.     

Genetic variation for an aphid wing polyphenism is genetically linked to a naturally occurring wing polymorphism

Many polyphenisms are examples of adaptive phenotypic plasticity where a single genotype produces distinct phenotypes in response to environmental cues. Such alternative phenotypes occur as winged and wingless parthenogenetic females in the pea aphid (Acyrthosiphon pisum). However, the proportion of winged females produced in response to a given environmental cue varies between clonal genotypes. Winged and wingless phenotypes also occur in males of the sexual generation. In contrast to parthenogenetic females, wing production in males is environmentally insensitive and controlled by the sex-linked, biallelic locus, aphicarus (api). Hence, environmental or genetic cues induce development of winged and wingless phenotypes at different stages of the pea aphid life cycle. We have tested whether allelic variation at the api locus explains genetic variation in the propensity to produce winged females. We assayed clones from an F2 cross that were heterozygous or homozygous for alternative api alleles for their propensity to produce winged offspring. We found that clones with different api genotypes differed in their propensity to produce winged offspring. The results indicate genetic linkage of factors controlling the female wing polyphenism and male wing polymorphism. This finding is consistent with the hypothesis that genotype by environment interaction at the api locus explains genetic variation in the environmentally cued wing polyphenism.     

Phenotypic plasticity of Brevicoryne brassicae in responses to nutritional quality of two related host plants

Abstract 1. The host species used by a herbivorous insect may impose different selective pressures promoting host race formation, yet the presence of plasticity can potentially constrain host race formation. 2. The goal of this study is to determine if there is phenotypic plasticity in life history traits of Brevicoryne brassicae in response to host and nutritional quality of two host species, Brassica oleraceae and Brassica campestris, and to what extent there are genetic differences among genotypes in plasticity. 3. Plants of B. oleraceae and B. campestris were fertilised with three different nitrogen doses (with nutritive solutions of 50, 200 and 400 ppm of soluble nitrogen) to produce plants with different nutritional qualities. Eight clones of B. brassicae were reared on those plants, and days to reproduction (DTR), number of nymphs, and fitness (r_m) were recorded. 4. A significant genotype × host interaction was detected in days to reproduction. Genotype × nitrogen interaction (plasticity) was detected in the number of nymphs when aphids were raised on B. campestris. Aphids showed plasticity in DTR and marginal plasticity in r_m in reaction to the varying nitrogen content of B. oleraceae. 5. The phenotypic plasticity to fine‐scale variation of host (nutritional quality) documented here may be an important source of phenotypic variation and may potentially constrain host race formation.