榕–传粉榕小蜂非一对一共生关系的研究进展

榕–传粉榕小蜂系统是研究动植物专性互惠共生关系的经典体系。早期形态学研究认为它们之间遵循一对一原则,即1种榕树只有1种榕小蜂为其传粉,而1种传粉榕小蜂也只能在1种榕树上繁殖后代。然而随着研究的深入,报道了越来越多非一对一共生关系的案例,尤其是隐存种的发现,打破了一对一原则在榕–传粉榕小蜂系统中的普适性。非一对一共生关系包括多种传粉者共现于1种宿主榕树和多种榕树共享1种传粉者两种情况,有各自不同的发生机制和规律。本文从形态学描述阶段、多学科综合证据阶段和传粉榕小蜂隐存种的发现3个阶段综述了榕–传粉榕小蜂非一对一共生关系的研究进展,并对其所引发的科学问题进行了探讨。多种传粉者共现于1种宿主榕树的情况,使得榕小蜂的种间杂交成为可能;而多种榕树共享1种传粉者,为宿主榕树的种间杂交提供了机会。然而,杂交事件的检出率却很低,暗示存在明显的生殖隔离。另外,传粉者共现和共享传粉者两种情况在不同性系统榕树上的发生率和发生机制存在明显差异,暗示不同性系统榕树的宿主专一性不同。大量小蜂隐存种的发现,引发了对其成种机制、共现机制的研究,以及对小蜂生物多样性的重新评估,并给人工控制实验的开展带来了新的问题。每个榕–传粉榕小蜂组合都具有其独特的协同进化历史,为研究动植物间的协同进化机制和规律提供了丰富素材。对榕–传粉榕小蜂专性互惠共生关系的形成、维持和打破机制开展综合研究,才能充分认识该系统,也有助于认识动植物间的协同进化过程和规律。     

榕-传粉小蜂化学通讯机制研究进展

榕属植物与其传粉小蜂组成了高度专一的专性共生关系(榕-蜂共生系统),如此高度紧密的互作关系被认为是驱动两者多样化的关键因素。榕-蜂共生系统主要依靠化学通讯完成相互识别,但目前仍不清楚化学通讯是如何维系现有共生关系并促进物种形成的。结合已有研究,系统梳理了榕-蜂共生系统化学通讯的基础与两者特异性识别的机制,阐述化学通讯在物种和种群层次对维持这一专性传粉关系的重要贡献,进而探讨化学通讯如何在协同成种和宿主转移成种两种模式中介导物种形成。最后,结合生理与多组学等技术展望榕-蜂共生系统的未来研究方向,为深入解析植物与昆虫协同进化的机制以及全球变化下物种的潜在响应模式提供重要参考。     

传粉榕小蜂与榕树的繁衍

榕树传粉现象被广泛地作为研究协同进化特别是互惠共生的重要模式之一。本文总结了榕果与传粉榕小蜂的有关研究,试图解释其形态结构之间的相互适应,总结传粉榕小蜂的传粉行为,探讨传粉榕小蜂在雌雄同株及雌雄异株榕树上的传粉模式,讨论传粉榕小蜂的寄主专一性,并展望中国在榕小蜂方面的研究前景。     

榕树——传粉者共生体系的研究

榕属（Ｆｉｃｕｓ）植物与它的传粉昆虫榕小蜂之间有丰专一性的互惠共生关系,它们是动植物间历史悠久关系最密切的共生伙伴,它们在形态结构,生理功能,生活史同步上构成了榕树－传粉者共生体系并协同进化至今,本文论述了榕属植物和榕小蜂的生物学特征以及二者的相互作用,阐述了榕树－传粉者共生体系的构成,维持及其生物学意义。     

榕树-榕小蜂互惠合作系统中的非对称性博弈

榕树及其传粉榕小蜂是自然界中目前所知道的关系最为紧密的互利共生系统之一。随着研究的深入, 越来越多的证据发现榕树-传粉榕小蜂之间互惠合作的过程中存在着复杂的竞争和对抗关系, 例如榕树与传粉榕小蜂之间对公共资源的竞争、传粉欺骗与宿主对传粉者的惩罚、榕树与传粉小蜂之间的“军备竞赛”等。在相互竞争或者对抗关系中, 双方表现出非对称性相互作用。其非对称性关系主要表现出如下3个特征: (1)收益不对称, 即榕树(宿主)与传粉榕小蜂(共生体)之间在资源利用等方面的实力不对称; (2)榕树与传粉榕小蜂之间的信息不对称; (3)进化速率不对称。这些非对称的相互作用可能导致种群的波动、榕树与传粉榕小蜂相互适应和进化策略的变化。因此, 理解榕树与传粉榕小蜂之间的非对称交互作用有助于理解为什么合作和冲突在互利共生关系中经常能同时存在, 也将有助于解释榕树-传粉榕小蜂种间相互关系和物种的多样性。     

榕树-榕小蜂的物种共形成

榕树-榕小蜂的互惠共生是植物与其传粉者间相互作用的最极端的特例,几乎每一种榕树均由榕小蜂科的一种特定昆虫传粉,而它们的小蜂也只能在此特定榕树种的花内养育它们的幼虫,构成了高度的种专一性(一对一原则).因此榕树及其传粉昆虫间的相互作用被看作是专性互惠共生的最极端特例,也引起人们对它们共趋异及物种形成的诸多推测.对目前榕树一榕小蜂的物种共形成的研究进行了回顾与阐述,希望能对全面认识这对完美的共生伙伴及物种共形成机制提供一些启示.     

榕树及其传粉榕小蜂的系统发育和协同进化研究现状及展望

榕属(Ficus)是有花植物中最大的木本属,全世界有750多种.榕树及其传粉榕小蜂形成的种类专一的互惠共生系统,长期以来被作为研究共生系统比较生物学和协同进化的模式材料.虽然从20世纪90年代才开始对榕树一榕小蜂体系开展分子系统发育研究,但由于这个体系的特殊性和分子技术的快速发展,越来越多的学者开始利用分子学手段来研究榕一蜂共生系统的一系列生物学问题.本文总结了近年来对榕树及其传粉榕小蜂开展的系统发育及协同进化方面的研究,并分析了中国此方面的研究现状,对未来的研究趋势和前景进行了展望.     

薜荔榕小蜂对薜荔和爱玉子雌花期榕果挥发物的行为反应

薜荔和爱玉子均属雌雄异株桑科榕属植物,两者互为原变种与变种的关系,分别与薜荔传粉小蜂和爱玉子传粉小蜂(二者互为隐存种)建立了专性共生关系,榕树榕果挥发物在维系传粉小蜂与其寄主的共生关系上起着重要作用。利用Y型嗅觉仪测定薜荔榕小蜂(薜荔和爱玉子的传粉小蜂)对薜荔和爱玉子雌花期榕果挥发物的行为反应。结果表明:(1)雌花期果型的大小对薜荔榕小蜂行为反应无显著影响,薜荔大、小果型雌花期雌(雄)榕果挥发物对其传粉小蜂均具有强烈的吸引作用;(2)榕果挥发物浓度影响薜荔榕小蜂行为反应,薜荔、爱玉子雌花期雌(雄)榕果挥发物对其传粉小蜂的吸引作用均可能存在阈值反应,即榕果挥发物浓度未超过阈值时,雌花期榕果挥发物对传粉小蜂的吸引作用与挥发物浓度成正相关关系,而一旦超过阈值,榕果挥发物对传粉蜂的吸引作用显著下降,表明寄主榕果挥发物浓度影响传粉小蜂的寄主定位;(3)薜荔传粉小蜂对低浓度爱玉子雌花期雌(雄)榕果挥发物、爱玉子传粉小蜂对低浓度薜荔雌花期雌(雄)榕果挥发物均既无趋向也无驱避行为;薜荔传粉小蜂对高浓度的爱玉子雌花期雌(雄)榕果挥发物表现为显著的驱避行为,而爱玉子传粉小蜂对高浓度薜荔雌(雄)雌花期榕果挥发物表现为显著的趋向行为,因此,薜荔传粉小蜂与爱玉子传粉小蜂存在寄主专一性不对称现象,爱玉子传粉小蜂进入薜荔雌(雄)果内传粉或产卵的可能性较大,而福州地区的薜荔传粉小蜂可能难以进入爱玉子雌(雄)果内传粉或产卵。本研究结果将为榕-蜂共生体系的化学生态学理论研究以及爱玉子栽培提供科学依据。     

榕树-传粉者共生体系的协同进化与系统学研究进展及展望

 榕树-传粉者共生体系是目前植物与昆虫协同进化研究中的典型模式之一。国内外已经开展了大量的相关研究,从不同方面探讨了其特殊的一一对应的共生关系。榕树-传粉者的专一性互惠共生关系中蕴含了与系统发育有关的多因子协同进化的机理,因此,进行系统发育研究将有助于更好地揭示榕树-传粉者的协同进化历史和理解二者的专一性互惠共生关系。本文简单地介绍了目前榕树及其传粉者共生体系的研究状况之后,论述了榕树-传粉者协同进化的系统发育分子生物学研究成果。同时针对国际上在榕属植物的传统的系统与分类研究中存在的一些分歧及榕树传粉者亚科分类不匹配等问题,回顾了榕属的分类研究进展及其与传粉者的关系。最后,结合我国榕树与传粉者共生体系的研究状况对我国榕属的重新分类和系统发育研究作了展望。     

榕树–榕小蜂协同进化中的非对称性相互作用及其集合种群效应

在协同进化研究中,传统观点认为物种之间的相互作用是对称性的,在进化过程中将形成一个稳定的均衡状态或进化稳定策略。然而,近来的观测和实验数据表明,物种间的协同进化可能存在非对称性相互作用,而且这种非对称性可能造成集合种群效应(metapopulation effect)或形成非均衡状态(例如混沌,chaos)。本文利用"榕树–榕小蜂"这一经典的协同进化模式系统来介绍协同进化过程中的非对称性相互作用,以及这种非对称性如何产生集合种群效应。在榕–蜂共生系统中,栖身于榕果中的榕小蜂除了传粉小蜂之外,还有一些"投机"的非传粉小蜂。非传粉小蜂比传粉小蜂具有更强的竞争力,而榕树则可惩罚不合作的非传粉蜂,同时奖励合作的传粉小蜂,从而形成了复杂的非对称性种间相互作用。在某个斑块生境中,非传粉小蜂通过竞争作用排斥传粉小蜂,然而随着非传粉小蜂在蜂群中的比例不断升高,榕树惩罚作用(如落果等)常常会导致整个非传粉蜂群的数量急剧下降,甚至局域性灭绝。随后,其他斑块的传粉小蜂则迁移过来填补空白。然而,随着传粉小蜂种群的数量增长,该斑块又会集聚更多的投机性非传粉小蜂,进而诱发榕树再次进行惩罚。这样的非对称性相互作用导致了非传粉小蜂、传粉小蜂以及榕树种群的集合效应,其种群大小呈现周期性的"此消彼长"式的循环。     

Rapid divergent evolution of sexual morphology: comparative tests of antagonistic coevolution and traditional female choice

Male structures specialized to contact females during sexual interactions often diverge relatively rapidly over evolutionary time. Previous explanations for this pattern invoked sexual selection by female choice, but new ideas emphasize possible sexually antagonistic coevolution resulting from male-female conflict over control of fertilization. The two types of selection have often not been carefully distinguished. They do not theoretically exclude one another, but they have not necessarily had equally important roles in producing rapid evolutionary divergence. To date, most recent empirical studies of antagonistic coevolution have emphasized only a few taxa. This study uses the abundant but little-used data in the taxonomic literature on morphology to evaluate the roles of antagonistic coevolution and traditional female choice over a wide taxonomic spectrum (61 families of arthropods, mostly insects and spiders). Groups with species-specific male structures that contact females were checked for coevolution of species-specific female structures that are contacted by the male and that have mechanical properties that could potentially defend her against the male. Facultatively deployable, species-specific female defensive structures, a design that would seem likely to evolve frequently under the sexually antagonistic coevolution hypothesis, were completely absent (0% of 106 structures in 84 taxonomic groups). Although likely cases of sexually antagonistic coevolution exist, using conservative criteria, 79.2% of the 106 structures lacked even potentially defensive female coevolution. A common pattern (53.8% of 106) was a nearly complete absence of female change in areas contacted by species-specific male structures. Post-hoc arguments invoking possible coevolution of defensive female behavior instead of morphology, or of female sensitivities and responses to male sensory traps, could enable the sexually antagonistic coevolution hypothesis to explain these data. No case of such coevolution of female behavior or sensitivities has been demonstrated, and there are additional reasons to doubt that they are general explanations for the data presented here. Detailed studies of female resistance behavior could help illuminate several issues. The possibility of a greater role for antagonistic coevolution in reproductive physiology than in morphology and the possibility that female choice and sexually antagonistic coevolution have both been important in some lineages are discussed.     

EVIDENCE FOR NEGATIVE FREQUENCY-DEPENDENT SELECTION DURING EXPERIMENTAL COEVOLUTION OF A FRESHWATER SNAIL AND A STERILIZING TREMATODE

Host–parasite coevolution is often suggested as a mechanism for maintaining genetic diversity, but finding direct evidence has proven difficult. In the present study, we examine the process of coevolution using a freshwater New Zealand snail ( Potamopyrgus antipodarum ) and its common parasite (the sterilizing trematode, Microphallus sp.) Specifically, we test for changes in genotypic composition of clonal host populations in experimental populations evolving either with or without parasites for six generations. As predicted under the Red Queen model of coevolution, the initially most common host genotype decreased in frequency in the presence, but not the absence, of parasitism. Furthermore, the initially most common host genotype became more susceptible to infection by the coevolving parasite populations over the course of the experiment. These results are consistent with parasite-meditated selection leading to a rare advantage, and they indicate rapid coevolution at the genotypic level between a host and its parasite.     

植物与草食动物之间的协同适应及进化

通常协同进化是指一个物种 (或种群 )的遗传结构由于回应于另一个物种 (或种群 )遗传结构的变化而发生的相应改变。广义的理解 ,协同进化是相互作用的物种之间的互惠进化。生物之间、特别是植物与草食动物之间的协同适应与进化 ,已经成为生物进化、生态、遗传等学科十分关注的问题 ,可能成为生物学中各学科研究的交汇点或结点。作者具体阐述了 :(1)生物之间协同进化的研究意义 ,包括对生物学与生态学的价值 ;(2 )生物之间协同进化研究的限制或困难 ,诸如时间、研究对象、进化等级尺度和研究方法的限制 ;(3)植物与草食动物之间协同进化的主要研究对象 (系统 ) ,即昆虫传粉系统、昆虫诱导植物反应系统、种子散布系统、以及大型草食动物采食与植物反应系统 ;(4 )植物与草食动物之间协同进化的主要研究内容 ,包括适应特征 (性状 )——物种的可塑性 ,以及适应机制——物种适应过程与策略两个方面 ;(5 )植物与草食动物之间协同进化研究的存在问题及研究方向     

Genomic Signatures of Coevolution between Nonmodel Mammals and Parasitic Roundworms

Antagonistic coevolution between host and parasite drives species evolution. However, most of the studies only focus on parasitism adaptation and do not explore the coevolution mechanisms from the perspective of both host and parasite. Here, through the de novo sequencing and assembly of the genomes of giant panda roundworm, red panda roundworm, and lion roundworm parasitic on tiger, we investigated the genomic mechanisms of coevolution between nonmodel mammals and their parasitic roundworms and those of roundworm parasitism in general. The genome-wide phylogeny revealed that these parasitic roundworms have not phylogenetically coevolved with their hosts. The CTSZ and prolyl 4-hydroxylase subunit beta (P4HB) immunoregulatory proteins played a central role in protein interaction between mammals and parasitic roundworms. The gene tree comparison identified that seven pairs of interactive proteins had consistent phylogenetic topology, suggesting their coevolution during host–parasite interaction. These coevolutionary proteins were particularly relevant to immune response. In addition, we found that the roundworms of both pandas exhibited higher proportions of metallopeptidase genes, and some positively selected genes were highly related to their larvae’s fast development. Our findings provide novel insights into the genetic mechanisms of coevolution between nonmodel mammals and parasites and offer the valuable genomic resources for scientific ascariasis prevention in both pandas.     

Rapidly fluctuating environments constrain coevolutionary arms races by impeding selective sweeps

Although pervasive, the impact of temporal environmental heterogeneity on coevolutionary processes is poorly understood. Productivity is a key temporally heterogeneous variable, and increasing productivity has been shown to increase rates of antagonistic arms race coevolution, and lead to the evolution of more broadly resistant hosts and more broadly infectious parasites. We investigated the effects of the grain of environmental heterogeneity, in terms of fluctuations in productivity, on bacteria–phage coevolution. Our findings demonstrate that environmental heterogeneity could constrain antagonistic coevolution, but that its effect was dependent upon the grain of heterogeneity, such that both the rate and extent of coevolution were most strongly limited in fine-grained, rapidly fluctuating heterogeneous environments. We further demonstrate that rapid environmental fluctuations were likely to have impeded selective sweeps of resistance alleles, which occurred over longer durations than the fastest, but not the slowest, frequency of fluctuations used. Taken together our results suggest that fine-grained environmental heterogeneity constrained the coevolutionary arms race by impeding selective sweeps.     

Accurate Simulation and Detection of Coevolution Signals in Multiple Sequence Alignments

Background

While the conserved positions of a multiple sequence alignment (MSA) are clearly of interest, non-conserved positions can also be important because, for example, destabilizing effects at one position can be compensated by stabilizing effects at another position. Different methods have been developed to recognize the evolutionary relationship between amino acid sites, and to disentangle functional/structural dependencies from historical/phylogenetic ones.

Methodology/Principal Findings

We have used two complementary approaches to test the efficacy of these methods. In the first approach, we have used a new program, MSAvolve, for the in silico evolution of MSAs, which records a detailed history of all covarying positions, and builds a global coevolution matrix as the accumulated sum of individual matrices for the positions forced to co-vary, the recombinant coevolution, and the stochastic coevolution. We have simulated over 1600 MSAs for 8 protein families, which reflect sequences of different sizes and proteins with widely different functions. The calculated coevolution matrices were compared with the coevolution matrices obtained for the same evolved MSAs with different coevolution detection methods. In a second approach we have evaluated the capacity of the different methods to predict close contacts in the representative X-ray structures of an additional 150 protein families using only experimental MSAs.

Conclusions/Significance

Methods based on the identification of global correlations between pairs were found to be generally superior to methods based only on local correlations in their capacity to identify coevolving residues using either simulated or experimental MSAs. However, the significant variability in the performance of different methods with different proteins suggests that the simulation of MSAs that replicate the statistical properties of the experimental MSA can be a valuable tool to identify the coevolution detection method that is most effective in each case.     

Coevolution is a short-distance force at the protein interaction level and correlates with the modular organization of protein networks

We investigated what roles coevolution plays in shaping yeast protein interaction network (PIN). We found that the extent of coevolution between two proteins decreases rapidly as their interacting distance on the PIN increases, suggesting coevolutionary constraint is a short-distance force at the molecular level. We also found that protein-protein interactions (PPIs) with strong coevolution tend to be enriched in interconnected clusters, whereas PPIs with weak coevolution are more frequently present at inter-cluster region. The findings indicate the close relationship between coevolution and modular organization of PINs, and may provide insights into evolution and modularity of cellular networks.     

Coevolution Between Food-Rewarding Flowers and Their Pollinators

The concept of coevolution was first developed by Darwin, who used it to explain how pollinators and food-rewarding flowers involved in specialized mutualisms could, over time, develop long tongues and deep tubes, respectively. He famously predicted that Angraecum sesquipedale, a long-spurred Malagasy orchid, must be pollinated by a hawkmoth with an exceptionally long tongue. Darwin’s idea of a coevolutionary “race” was championed by contemporary naturalists, including Alfred Wallace, and a hawkmoth fitting the expected tongue-length profile was eventually discovered in Madagascar during the early twentieth century. However, strong empirical support for the mechanism behind Darwin’s coevolutionary model has been forthcoming only in the past two decades. It is now established that selection often strongly favors plants with floral tubes that exceed the length of their pollinator’s tongues. There is also evidence that pollinators gain an energetic benefit from having tongues that enable them to consume most or all of the nectar in deep tubular flowers. Alternative explanations for the evolution of long pollinator tongues, such as evasion of predators that use flowers as ambush sites, are considered much less compelling and lack quantitative support. Another important advance in coevolution research has been the development of approaches that explicitly predict a geographical mosaic of coevolution. The expectation that coevolution can lead to geographical diversification and trait covariation among strongly interacting organisms is strongly supported by studies of long-proboscid fly and oil-bee pollination systems in South Africa. Macro- and microevolutionary studies of pollination systems suggest that coevolution can operate alongside other one-sided evolutionary processes, such as shifts, in shaping plant and pollinator traits.     

BioPhysConnectoR: Connecting Sequence Information and Biophysical Models

Background  

One of the most challenging aspects of biomolecular systems is the understanding of the coevolution in and among the molecule(s).     

Gene-culture coevolution and the nature of human sociality

Human characteristics are the product of gene-culture coevolution, which is an evolutionary dynamic involving the interaction of genes and culture over long time periods. Gene-culture coevolution is a special case of niche construction. Gene-culture coevolution is responsible for human other-regarding preferences, a taste for fairness, the capacity to empathize and salience of morality and character virtues.