二化螟寄主种群及种群间生殖隔离形成机制研究

水稻和茭白是二化螟的两种主要寄主植物。经过长期的适应进化,二化螟已分化出水稻种群和茭白种群。国内外的大量研究表明,二化螟水稻种群和茭白种群在形态特征、交配行为、寄主选择性、寄主适应性、田间种群动态、越冬生物学、生理生化及遗传多样性等方面均存在一定差异。二化螟水稻种群和茭白种群的发生期差异、交配节律差异和交配后配子不亲合等可能造成了寄主种群的部分生殖隔离。目前认为,二化螟两寄主种群间仍存在基因交流,处于物种形成的早期阶段,但尚需从寄主种群鉴定、表型差异区分和种群间生殖隔离机制等方面开展深入研究来进一步明确,为理解昆虫同域物种形成机制提供参考。     

生态物种形成及其研究进展

物种形成是基本的进化过程, 也是生物多样性形成的基础。自然选择可以导致新物种的产生。生态物种形成是指以生态为基础的歧化选择使不同群体分化产生生殖隔离的物种形成过程。本文首先回顾了生态物种形成的研究历史, 并详细介绍了生态物种形成的3个要素, 即歧化选择的来源、生殖隔离的形式以及关联歧化选择与生殖隔离的遗传机制。歧化选择的来源主要包括不同的环境或生态位、不同形式的性选择, 以及群体间的相互作用。生殖隔离的形式多种多样, 我们总结了合子前和合子后隔离的遗传学机制以及在生态物种形成中起到的作用。控制适应性性状的基因与导致生殖隔离的基因可以通过基因多效性或连锁不平衡相互关联起来。借助于第二代测序技术, 研究者可以对生态物种形成的遗传学与基因组学基础进行研究。此外, 本文还总结了生态物种形成领域最新的研究进展, 包括平行进化的全基因组基础, 以及基因流影响群体分化的理论基础。通过归纳比较由下至上和由上至下这两种不同的研究思路, 作者认为这两种思路的结合可以为生态物种形成基因的筛选提供更有力也更精确的方法。同时, 作者还提出生态物种形成的研究应该基于更好的表型描述以及更完整的基因组信息, 研究的物种也应该具有更广泛的代表性。     

基于生物学物种定义探讨物种形成理论与验证的研究进展

物种形成是进化生物学研究的一个永恒主题, 由于生物群体进化是连续和动态的, 物种界限变得难于界定。本文首先讨论了3种地理物种形成模式(同域、邻域及异域), 并分析了近期报道的研究证据。其次, 评述了合子后生殖隔离机制的分子遗传基础和应用群体基因组数据分析的证据, 包括BDMI模型(Bateson-Dobzhansky-Muller incompatibility)、QTLs (quantitative trait loci)、霍尔丹法则及大X染色体效应。最后, 探讨了交配系统作为合子前隔离机制之一与物种形成的关系, 认为近交或自交通过扩大种群遗传结构分化, 增强不同交配系统的种群间不对称基因渐渗, 或种群间无基因渐渗等途径, 促进新物种形成。已知植物交配系统的演化更倾向于从异交(或自交不亲和)向自交(或近交亲和)方式, 花性状和基因组的分化推动形成所谓的自交综合征, 研究交配系统驱动或强化物种形成模式对认识植物物种形成机制有重要意义。     

物种形成过程中的分化基因组岛及其形成机制

理解物种形成机制是生态和进化领域的重要任务。得益于测序技术的快速发展, 越来越多研究发现分化种群(亚种、物种)间的基因组常呈现异质性分化景观, 存在分化基因组岛, 这被认为是基因流存在下的歧化选择引起的, 支持基因流存在下的成种假说。然而, 基因渐渗、祖先多态性的差异分选、连锁选择等其他进化过程也可导致分化基因组岛的形成。现有实证研究在解析分化基因组岛的形成机制时, 往往忽略了上述其他进化过程的作用。为此, 本文在辨析分化基因组岛相关概念的基础上, 总结了利用种群基因组数据鉴定分化基因组岛的方法, 对比了不同进化过程形成分化基因组岛的特征, 指出在区分不同机制时联用基因渐渗程度、绝对分化指数(d_XY)、相对节点深度(RND)、重组率等多个指标的必要性, 归纳了物种形成过程中分化基因组岛形成机制解析的研究思路, 并对未来在生殖隔离机制上的深入探索以及实证研究的整合分析等方面进行了展望。     

植物边缘种群遗传多样性研究进展

边缘种群指地理分布边缘可检测到的一定数量的同种个体集合, 准确评价其遗传多样性对于理解第四纪冰期后气候变化对物种边缘扩展或收缩、遗传资源保护与利用以及物种形成等有重要意义。该文探讨了维持植物边缘种群遗传多样性的进化机制, 分析交配系统对物种边缘及其遗传多样性的影响, 比较了边缘与中心种群遗传多样性的差异及其形成的生态与进化过程, 并探讨了边缘种群遗传多样性与其所在的群落物种多样性的关系及理论基础。该文提出今后研究的重点是应用全基因组序列或转录组基因序列研究前缘-后缘种群之间或边缘-中心种群之间的适应性差异, 边缘种群与所在群落其他物种之间相互作用的分子机制, 深入解析边缘种群对环境的适应及边缘种群遗传多样性与群落物种多样性关系的生态与进化过程。     

植物边缘种群遗传多样性研究进展

边缘种群指地理分布边缘可检测到的一定数量的同种个体集合,准确评价其遗传多样性对于理解第四纪冰期后气候变化对物种边缘扩展或收缩、遗传资源保护与利用以及物种形成等有重要意义。该文探讨了维持植物边缘种群遗传多样性的进化机制,分析交配系统对物种边缘及其遗传多样性的影响,比较了边缘与中心种群遗传多样性的差异及其形成的生态与进化过程,并探讨了边缘种群遗传多样性与其所在的群落物种多样性的关系及理论基础。该文提出今后研究的重点是应用全基因组序列或转录组基因序列研究前缘-后缘种群之间或边缘-中心种群之间的适应性差异,边缘种群与所在群落其他物种之间相互作用的分子机制,深入解析边缘种群对环境的适应及边缘种群遗传多样性与群落物种多样性关系的生态与进化过程。     

基因流存在条件下的物种形成研究述评:生殖隔离机制进化

物种形成过程是生物多样性形成的基础, 长期以来一直是进化生物学的中心议题之一。传统的异域物种形成理论认为, 地理隔离是物种分化的主要决定因子, 物种形成只有在种群之间存在地理隔离的情况下才能发生。近年来, 随着种群基因组学的发展和溯祖理论分析方法的完善, 种群间存在基因流情况下的物种形成成为进化生物学领域新的研究焦点。物种形成过程中是否有基因流的发生?基因流如何影响物种的形成与分化?基因流存在条件下物种形成的生殖隔离机制是什么?根据已发表的相关文献资料, 作者综述了当前物种形成研究中基因流的时间和空间分布模式、基因流对物种分化的影响以及生殖隔离机制形成等问题, 指出基因流存在条件下的物种形成可能是自然界普遍发生的一种模式。     

协同物种形成及其生物学意义

自达尔文与华莱士以来的每位生物学家都承认种间生态学作用对进化的意义。1964年,埃利希和雷文提出了协同进化理论。它反映了生态学相关的不同物种之间的相互作用引起的相互进化。这一理论引起了研究者的广泛重视。近年来,随着进化生态研究的深入发展,学者们对进化过程的生态机制以及生态特性与生态关系进化规律、种间生态学作用引起的物种形成有了进一步的认识。本文仅从协同物种形成的概念出发,对协同物种形成的生态机制以及生物学意义作一简要概述。协同物种形成的概念进化在本质上是一种生态过程,是生物与环境、生物与生物之间相互作用的结果。种群分化的生态机制和物种形成的本质在于对生存条件的逐步适应。按照协同进化理论,物种间的相互作用引起的协同适应     

生殖隔离浅析

现代进化论认为,物种形成大致要经过3个阶段:1)由于基因突变、染色体变异和基因重组,使种群中产生可遗传的变异;2)自然选择等因素作用于可遗传的变异,使种群的遗传结构(基因型频率和基因频率)发生了适应性的改变;3)不同种群由于地理隔离和生态隔离而加深了性状分歧,逐渐形成亚种,一旦出现了生殖隔离,亚种就变成了新种。此外,在植物界异源多倍体的产生亦可导致新种形成。由于各方面的原因,使亲缘关系接近的类群之间在自然条件下不交配,即使能交配也不能产生后代或不能产生可育性后代的隔离机制,便称为生殖隔离。若隔离发生在受精以前,就称为…     

植物繁育系统研究的最新进展和评述

 植物繁育系统是当今进化生物学研究中最为活跃的领域。繁育系统是指代表所有影响后代遗传组成的有性特征的总和,主要包括花形态特征、花的开放式样、花各部位的寿命、传粉者种类和频率、自交亲和程度和交配系统,其中交配系统是核心。我们重点综述了植物 有性繁育系统研究中1）传粉模式的多样性,2）从历史发生角度利用系统发育方法检验花性状的演化和繁育系统的生态转变过程,3）近交衰退对植物生殖史的影响及其机制和4）混和交配系统的时空动态、维持机理以及进化趋势的最新研究结果。总结了繁育系统在研究花适应性、物种形成机制和濒危植物     

Frequency-dependent selection and the evolution of assortative mating

A long-standing goal in evolutionary biology is to identify the conditions that promote the evolution of reproductive isolation and speciation. The factors promoting sympatric speciation have been of particular interest, both because it is notoriously difficult to prove empirically and because theoretical models have generated conflicting results, depending on the assumptions made. Here, we analyze the conditions under which selection favors the evolution of assortative mating, thereby reducing gene flow between sympatric groups, using a general model of selection, which allows fitness to be frequency dependent. Our analytical results are based on a two-locus diploid model, with one locus altering the trait under selection and the other locus controlling the strength of assortment (a "one-allele" model). Examining both equilibrium and nonequilibrium scenarios, we demonstrate that whenever heterozygotes are less fit, on average, than homozygotes at the trait locus, indirect selection for assortative mating is generated. While costs of assortative mating hinder the evolution of reproductive isolation, they do not prevent it unless they are sufficiently great. Assortative mating that arises because individuals mate within groups (formed in time or space) is most conducive to the evolution of complete assortative mating from random mating. Assortative mating based on female preferences is more restrictive, because the resulting sexual selection can lead to loss of the trait polymorphism and cause the relative fitness of heterozygotes to rise above homozygotes, eliminating the force favoring assortment. When assortative mating is already prevalent, however, sexual selection can itself cause low heterozygous fitness, promoting the evolution of complete reproductive isolation (akin to "reinforcement") regardless of the form of natural selection.     

HABITAT AVOIDANCE: OVERLOOKING AN IMPORTANT ASPECT OF HOST‐SPECIFIC MATING AND SYMPATRIC SPECIATION?

Understanding speciation requires discerning how reproductive barriers to gene flow evolve between previously interbreeding populations. Models of sympatric speciation for phytophagous insects posit that reproductive isolation can evolve in the absence of geographic isolation as a consequence of an insect shifting and ecologically adapting to a new host plant. One important adaptation contributing to sympatric differentiation is host-specific mating. When organisms mate in preferred habitats, a system of positive assortative mating is established that facilitates sympatric divergence. Models of host fidelity generally assume that host choice is determined by the aggregate effect of alleles imparting positive preferences for different plant species. But negative effect genes for avoiding nonnatal plants may also influence host use. Previous studies have shown that apple and hawthorn-infesting races of Rhagoletis pomonella flies use volatile compounds emitted from the surface of fruit as key chemosensory cues to recognize and distinguish between their host plants. Here, we report results from field trials indicating that in addition to preferring the odor of their natal fruit, apple and hawthorn flies, and their undescribed sister species infesting flowering dogwood (Cornus florida), also avoid the odors of nonnatal fruit. We discuss the implications of nonnatal fruit avoidance for the evolutionary dynamics and genetics of sympatric speciation. Our findings reveal an underappreciated role for habitat avoidance as a potential postmating, as well as prezygotic, barrier to gene flow.     

Vibrational communication and reproductive isolation in the Enchenopa binotata species complex of treehoppers (Hemiptera: Membracidae)

Sexual communication can contribute to population divergence and speciation because of its effect on assortative mating. We examined the role of communication in assortative mating in the Enchenopa binotata species complex of treehoppers. These plant-feeding insects are a well studied case of sympatric speciation resulting from shifts to novel host-plant species. Shifting to hosts with different phenologies causes changes in life-history timing. In concert with high host fidelity, these changes reduce gene flow between populations on ancestral and novel hosts and facilitate a rapid response to divergent natural selection. However, some interbreeding can still occur because of partial overlap of mating periods. Additional behavioral mechanisms resulting in reproductive isolation may thus be important for divergence. In E. binotata, mating pairs form after an exchange of plant-borne vibrational signals. We used playback experiments to examine the relevance of inter- and intraspecific variation in male advertisement signals for female mate choice in a member of the E. binotata species complex. Female signals given in response to male signals provided a simple and reliable assay. Male species and male individual identity were important determinants of female responses. Females failed to respond to the signals of the two most closely related species in the complex, but they responded strongly to the signals of conspecific males, as well as to those of the most basal species in the complex. Communication systems in the E. binotata species complex can therefore play a role in reproductive isolation. Female responses were influenced by among-individual variation in male signals and females, suggesting the involvement of sexual selection in the evolution of these communication systems.     

Low reproductive isolation and highly variable levels of gene flow reveal limited progress towards speciation between European river and brook lampreys

Ecologically based divergent selection is a factor that could drive reproductive isolation even in the presence of gene flow. Population pairs arrayed along a continuum of divergence provide a good opportunity to address this issue. Here, we used a combination of mating trials, experimental crosses and population genetic analyses to investigate the evolution of reproductive isolation between two closely related species of lampreys with distinct life histories. We used microsatellite markers to genotype over 1000 individuals of the migratory parasitic river lamprey (Lampetra fluviatilis) and freshwater‐resident nonparasitic brook lamprey (Lampetra planeri) distributed in 10 sympatric and parapatric population pairs in France. Mating trials, parentage analyses and artificial fertilizations demonstrated a low level of reproductive isolation between species even though size‐assortative mating may contribute to isolation. Most parapatric population pairs were strongly differentiated due to the joint effects of geographic distance and barriers to migration. In contrast, we found variable levels of gene flow between sympatric populations ranging from panmixia to moderate differentiation, which indicates a gradient of divergence with some population pairs that may correspond to alternative morphs or ecotypes of a single species and others that remain partially isolated. Ecologically based divergent selection may explain these variable levels of divergence among sympatric population pairs, but incomplete genome swamping following secondary contact could have also played a role. Overall, this study illustrates how highly differentiated phenotypes can be maintained despite high levels of gene flow that limit the progress towards speciation.     

Positive assortative mating between recently described sympatric morphs of Icelandic sticklebacks

Recently, models of sympatric speciation have suggested that assortative mating can develop between sympatric morphs due to divergence in an ecologically important character. For example, in sympatric pairs of threespine stickleback (Gasterosteus aculeatus L.) size-assortative mating seems to be instrumental in reproductive isolation. Here, we examine courtship behaviour and assortative mating of newly described sympatric stickleback morphs in Lake Thingvallavatn, Iceland. We find that the two morphs show strong positive assortative mating. However, the mechanism involved in mate choice does not seem to be as straightforward as in other similar systems of sympatric stickleback morphs and may involve variation in nest type.     

The concept of effective recombination rate and its application in speciation theory

The goal of this study is to develop a unifying theoretical framework to quantify the strength of reproductive isolation. We propose the use of the "effective recombination rate," which measures how fast associations of genes are broken by interlocus recombination. Applying the well-established theory of the effective migration rate, we derive two techniques to investigate the effective recombination rate in models of speciation: the weak migration approximation for parapatric scenarios and the weak recombination approximation for sympatric scenarios. We illustrate the use of these two methods by two examples each: (1) single-locus genetic incompatibility and (2) two-locus genetic incompatibility for the first method, and (3) assortative mating and (4) assortative mating combined with disruptive selection for the second method. An advantage of the effective recombination rate over previous approaches is that it integrates gene flow in both directions into a single index measuring the strength of isolation. This enables straightforward comparisons of speciation scenarios with the same or different geographic histories. The method also allows us to evaluate the relative contributions of F2 hybrid deficiency or linkage between multiple barriers in reproductive isolation.     

INCIPIENT REPRODUCTIVE ISOLATION BETWEEN TWO SYMPATRIC MORPHS OF THE INTERTIDAL SNAIL LITTORINA SAXATILIS

The study of speciation in recent populations is essentially a study of the evolution of reproductive isolation mechanisms between sub-groups of a species. Prezygotic isolation can be of central importance to models of speciation, either being a consequence of reinforcement of assortative mating in hybrid zones, or a pleiotropic effect of morphological or behavioral adaptation to different environments. To suggest speciation by reinforcement between incipient species one must at least know that gene flow occurs, or have recently occurred, and that assortative mating has been established in the hybrid zone. In Galician populations of the marine snail Littorina saxatilis, two main morphs appear on the same shores, one on the upper-shore barnacle belt and the other in the lower-shore mussel belt. The two morphs overlap in distribution in the midshore where hybrids are found together with pure forms. Allozyme variation indicates that the two parental morphs share a common gene pool, although within shores, gene flow between morphs is less than gene flow within morphs. In this study, we observed mating behavior in the field, and we found that mating was not random in midshore sites, with a deficiency of heterotypic pairs. Habitat selection, assortative mating, and possibly sexual selection among females contributed to the partial reproductive isolation between the pure morphs. Sizes of mates were often positively correlated, in particular, in the upper shore, indicating size-assortative mating too. However, this seemed to be a consequence of nonrandom microdistributions of snails of different sizes. Because we also argue that the hybrid zone is of primary rather than secondary origin, this seems to be an example of sympatric reproductive isolation, either established by means of reinforcement or as a by-product to divergent selection acting on other characters.     

Phylogenetic evidence for multiple sympatric ecological diversification in a marine snail

Parallel speciation can occur when traits determining reproductive isolation evolve independently in different populations that experience a similar range of environments. However, a common problem in studies of parallel evolution is to distinguish this hypothesis from an alternative one in which different ecotypes arose only once in allopatry and now share a sympatric scenario with substantial gene flow between them. Here we show that the combination of a phylogenetic approach with life-history data is able to disentangle both hypotheses in the case of the intertidal marine snail Littorina saxatilis on the rocky shores of Galicia in northwestern Spain. In this system, numerous phenotypic and genetic differences have evolved between two sympatric ecotypes spanning a sharp ecological gradient, and as aside effect of the former have produced partial reproductive isolation. A mitochondrial phylogeny of these populations strongly suggests that the two sympatric ecotypes have originated independently several times. Building upon earlier work demonstrating size-based assortative mating as the main contributor to reproductive isolation among ecotypes, our analysis provides strong evidence that divergent selection across a sharp ecological gradient promoted the parallel divergence of body size and shape between two sympatric ecotypes. Thus, divergent selection occurring independently in different populations has produced the marine equivalent of host races, which may represent the first step in speciation.     

Delayed prezygotic isolating mechanisms: evolution with a twist

Assortative mating characterizes the situation wherein reproducing individuals pair according to similarity. Usually, the impetus for this bias is attributed to some type of mate choice conferring benefits (e.g., increased fitness or genetic compatibility) and, thereby, promoting speciation and phenotypic evolution. We investigate, by computer simulation of an evolving deme-structured snail population, the ramifications ensuing from passive assortative mating wherein couples exhibiting opposite shell coil direction phenotypes experience a physical constraint on mating success: putative mating partners inhabiting stout dextral and sinistral shells are unable to exchange sperm. Because shell coil chirality genotype is encoded at a single locus by shell coil alleles that are inherited maternally, snails containing sinistral alleles can present the typical dextral phenotype. Consequently, the incidence of a sinistral allele in as few as one snail can be manifested as prezygotic reproductive isolation within a deme in a subsequent generation. However, because the efficacy of achieving this type of prezygotic reproductive isolation is affected by shell form, the likelihood and product of single-gene speciation should be determined by deme interaction (migration) and composition (morphological distribution). We test this hypothesis and show how stochastic migration interacts with passive assortative mating yielding morphologically induced prezygotic reproductive isolation to produce new species phenotypes. The results show that demes can achieve rapid macroscopic phenotypic transformation and indicate that sympatric speciation might be more plausible than naturalists recognize conventionally.     

Experimental Swap of Anopheles gambiae's Assortative Mating Preferences Demonstrates Key Role of X-Chromosome Divergence Island in Incipient Sympatric Speciation

Although many theoretical models of sympatric speciation propose that genes responsible for assortative mating amongst incipient species should be associated with genomic regions protected from recombination, there are few data to support this theory. The malaria mosquito, Anopheles gambiae, is known for its sympatric cryptic species maintained by pre-mating reproductive isolation and its putative genomic islands of speciation, and is therefore an ideal model system for studying the genomic signature associated with incipient sympatric speciation. Here we selectively introgressed the island of divergence located in the pericentric region of the X chromosome of An. gambiae s.s. into its sister taxon An. coluzzii through 5 generations of backcrossing followed by two generations of crosses within the introgressed strains that resulted in An. coluzzii-like recombinant strains fixed for the M and S marker in the X chromosome island. The mating preference of recombinant strains was then tested by giving virgin recombinant individuals a choice of mates with X-islands matching and non-matching their own island type. We show through genetic analyses of transferred sperm that recombinant females consistently mated with matching island-type males thereby associating assortative mating genes with the X-island of divergence. Furthermore, full-genome sequencing confirmed that protein-coding differences between recombinant strains were limited to the experimentally swapped pericentromeric region. Finally, targeted-genome comparisons showed that a number of these unique differences were conserved in sympatric field populations, thereby revealing candidate speciation genes. The functional demonstration of a close association between speciation genes and the X-island of differentiation lends unprecedented support to island-of-speciation models of sympatric speciation facilitated by pericentric recombination suppression.