Gene flow maintains a large genetic difference in clutch size at a small spatial scale

Understanding the capacity of natural populations to adapt to their local environment is a central topic in evolutionary biology. Phenotypic differences between populations may have a genetic basis, but showing that they reflect different adaptive optima requires the quantification of both gene flow and selection. Good empirical data are rare. Using data on a spatially structured island population of great tits (Parus major), we show here that a persistent difference in mean clutch size between two subpopulations only a few kilometres apart has a major genetic component. We also show that immigrants from outside the island carry genes for large clutches. But gene flow into one subpopulation is low, as a result of a low immigration rate together with strong selection against immigrant genes. This has allowed for adaptation to the island environment and the maintenance of small clutches. In the other area, however, higher gene flow prevents local adaptation and maintains larger clutches. We show that the observed small-scale genetic difference in clutch size is not due to divergent selection on the island, but to different levels of gene flow from outside the island. Our findings illustrate the large effect of immigration on the evolution of local adaptations and on genetic population structure.     

Non-adaptive origins of interactome complexity

The boundaries between prokaryotes, unicellular eukaryotes and multicellular eukaryotes are accompanied by orders-of-magnitude reductions in effective population size, with concurrent amplifications of the effects of random genetic drift and mutation. The resultant decline in the efficiency of selection seems to be sufficient to influence a wide range of attributes at the genomic level in a non-adaptive manner. A key remaining question concerns the extent to which variation in the power of random genetic drift is capable of influencing phylogenetic diversity at the subcellular and cellular levels. Should this be the case, population size would have to be considered as a potential determinant of the mechanistic pathways underlying long-term phenotypic evolution. Here we demonstrate a phylogenetically broad inverse relation between the power of drift and the structural integrity of protein subunits. This leads to the hypothesis that the accumulation of mildly deleterious mutations in populations of small size induces secondary selection for protein-protein interactions that stabilize key gene functions. By this means, the complex protein architectures and interactions essential to the genesis of phenotypic diversity may initially emerge by non-adaptive mechanisms.     

Host-plant adaptation drives the parallel evolution of reproductive isolation

Parallel evolution of similar traits in independent populations that inhabit ecologically similar environments strongly implicates natural selection as the cause of evolution. Parallel speciation is a special form of parallel evolution where traits that determine reproductive isolation evolve repeatedly, in closely related populations, as by-products of adaptation to ecological conditions. The outcome of such parallel evolution is that ecologically divergent pairs of populations exhibit greater levels of reproductive isolation than ecologically similar pairs of populations of a similar or younger age. The parallel evolution of reproductive isolation provides strong evidence for natural selection in the process of speciation, but only one conclusive example from nature is known. Populations of the walking-stick insect Timema cristinae that use different host-plant species have diverged in body size and shape, host preference, behaviour and the relative frequency of two highly cryptic colour-pattern morphs. Here we report that divergent selection for host adaptation, and not genetic drift, has promoted the parallel evolution of sexual isolation in this species. Our findings represent a clear demonstration that host-plant adaptation can play a crucial and repeatable role in the early stages of speciation.     

Stage-structured cycles promote genetic diversity in a predator-prey system of Daphnia and algae

Competition theory predicts that population fluctuations can promote genetic diversity when combined with density-dependent selection. However, this stabilizing mechanism has rarely been tested, and was recently rejected as an explanation for maintaining diversity in natural populations of the freshwater herbivore Daphnia pulex. The primary limitation of competition theory is its failure to account for the alternative types of population cycles that are caused by size- or stage-dependent population vital rates--even though such structure both explains the fluctuating dynamics of many species and may alter the outcome of competition. Here we provide the first experimental test of whether alternative types of cycles affect natural selection in predator-prey systems. Using competing Daphnia genotypes, we show that internally generated, stage-structured cycles substantially reduce the magnitude of selection (thereby contributing to the maintenance of genetic diversity), whereas externally forced cycles show rapid competitive exclusion. The change in selection is ecologically significant, spanning the observed range in natural populations. We argue that structured cycles reduce selection through a combination of stalled juvenile development and stage-specific mortality. This potentially general fitness-equalizing mechanism may reduce the need for strong stabilizing mechanisms to explain the maintenance of genetic diversity in natural systems.     

云南松的种群遗传与进化

本文从种群遗传学的角度研究了云南松（pinus yunnanensis)的遗传体制，种群内的多态现象，种群的遗传结构，演化潜力，生态小种和地理小种以及共交种等问题，得出的若干结论如下：（1）种内存在巨大的遗传变异，如在自然 群中表明的，几乎所有可见的形态性状都具有多态现象，这种巨大的遗传变异是自然选择进化的一个必要条件。（2）自然种群中，大多数个体的基因型很多座位都是杂合的，它们常表现出杂种优势，并且也可能表明增强了生理上发育上的体内平衡。（3）由于种群大，而且又分割成许多相对独立的亚群，各种进化要素都有能同时起作用，并已分化出生态种和地理小种，（4）云南松具有拟态种子，其起源和适应意义作了初步讨论。     

Reinforcement drives rapid allopatric speciation

Allopatric speciation results from geographic isolation between populations. In the absence of gene flow, reproductive isolation arises gradually and incidentally as a result of mutation, genetic drift and the indirect effects of natural selection driving local adaptation. In contrast, speciation by reinforcement is driven directly by natural selection against maladaptive hybridization. This gives individuals that choose the traits of their own lineage greater fitness, potentially leading to rapid speciation between the lineages. Reinforcing natural selection on a population of one of the lineages in a mosaic contact zone could also result in divergence of the population from the allopatric range of its own lineage outside the zone. Here we test this with molecular data, experimental crosses, field measurements and mate choice experiments in a mosaic contact zone between two lineages of a rainforest frog. We show that reinforcing natural selection has resulted in significant premating isolation of a population in the contact zone not only from the other lineage but also, incidentally, from the closely related main range of its own lineage. Thus we show the potential for reinforcement to drive rapid allopatric speciation.     

多辐溲疏居群过氧化物酶同工酶分化的研究

运用聚丙烯酰胺凝胶电泳，测试了采自四川南川县金佛山不同生境的多辐溲疏５个居群５０株个体的过氧化物酶同工酶，对酶谱带位置进行了二态编码，然后采用Ｒｏｇｅｒｓ－Ｔａｎｉｍｏｔｏ，Ｓｏｋａｌ－Ｍｉｃｈ－ｅｎｅｒ，Ｓｏｋａｌ－Ｓｎｅａｔｈ，Ｊａｃｃａｒｄ等结合系数用组平均法（ＵＰＧＭＡ）对各谱带位置进行了聚类分析．结果表明，相同生境同一居群内的个体间表现出较大的遗传同质性，而来源于不同生境的不同居群个体间却呈现出较大的遗传趋异性；同时，５个居群不但具有许多共同谱带，而且整体相聚的结合系数大多超过０．５，这表明５个居群在很早以前可能是属于同一大居群，在长期进化过程中，由于微生境的地理与气候因子选择，使得多辐溲疏现仅在金佛山海拔６００～１２００ｍ的约ｋｍ２范围内呈岛状分布；又由于其特殊的繁殖方式——居群内个体虽能杂交，但因该种仅能正常开花传粉，大部分种子败育，使得居群内个体遗传变异性不能通过有性生殖方式固定下来，仅能通过营养繁殖方式繁殖后代，而正是营养繁殖导致了居群内较大遗传同质性以及居群间的遗传趋异性．在整个进化过程中生境只不过是对那些遗传变异起到了选择固定作用     

Cryptic evolution in a wild bird population.

Microevolution is expected to be commonplace, yet there are few thoroughly documented cases of microevolution in wild populations. In contrast, it is often observed that apparently heritable traits under strong and consistent directional selection fail to show the expected evolutionary response. One explanation proposed for this paradox is that a genetic response to selection may be masked by opposing changes in the environment. We used data from a 20-year study of collared flycatchers (Ficedula albicollis) to explore selection on, and evolution of, a heritable trait: relative body weight at fledging ('condition'). Despite consistent positive directional selection, on both the phenotypic and the additive genetic component (breeding values, estimated from an animal model) of condition, the mean phenotypic value of this trait in the population has declined, rather than increased, over time. Here we show that, despite this decline, the mean breeding value for condition has increased over time. The mismatch between response to selection at the levels of genotype and phenotype can be explained by environmental deterioration, concealing underlying evolution. This form of cryptic evolution may be common in natural environments.     

Relative fitness can decrease in evolving asexual populations of S. cerevisiae

It is generally accepted from the darwinian theory of evolution that a progressive increase in population adaptation will occur in populations containing genetic variation in fitness, until a stable equilibrium is reached and/or the additive genetic variation is exhausted. However, the theoretical literature of population genetics documents exceptions where mean population fitness may decrease in response to evolutionary changes in gene frequency, due to varying selective coefficients, sexual selection or to epistatic interactions between loci. Until now, no examples of such exceptions have been documented from fitness estimates in either natural or experimental populations. We present here direct evidence that, as a result of epistatic interactions between adaptive mutations, mean population fitness can decrease in asexual evolving populations of the yeast Saccharomyces cerevisiae.     

Genetic variation increases during biological invasion by a Cuban lizard

A genetic paradox exists in invasion biology: how do introduced populations, whose genetic variation has probably been depleted by population bottlenecks, persist and adapt to new conditions? Lessons from conservation genetics show that reduced genetic variation due to genetic drift and founder effects limits the ability of a population to adapt, and small population size increases the risk of extinction. Nonetheless, many introduced species experiencing these same conditions during initial introductions persist, expand their ranges, evolve rapidly and become invasive. To address this issue, we studied the brown anole, a worldwide invasive lizard. Genetic analyses indicate that at least eight introductions have occurred in Florida from across this lizard's native range, blending genetic variation from different geographic source populations and producing populations that contain substantially more, not less, genetic variation than native populations. Moreover, recently introduced brown anole populations around the world originate from Florida, and some have maintained these elevated levels of genetic variation. Here we show that one key to invasion success may be the occurrence of multiple introductions that transform among-population variation in native ranges to within-population variation in introduced areas. Furthermore, these genetically variable populations may be particularly potent sources for introductions elsewhere. The growing problem of invasive species introductions brings considerable economic and biological costs. If these costs are to be mitigated, a greater understanding of the causes, progression and consequences of biological invasions is needed.     

Sexually antagonistic genetic variation for fitness in red deer

Evolutionary theory predicts the depletion of genetic variation in natural populations as a result of the effects of selection, but genetic variation is nevertheless abundant for many traits that are under directional or stabilizing selection. Evolutionary geneticists commonly try to explain this paradox with mechanisms that lead to a balance between mutation and selection. However, theoretical predictions of equilibrium genetic variance under mutation-selection balance are usually lower than the observed values, and the reason for this is unknown. The potential role of sexually antagonistic selection in maintaining genetic variation has received little attention in this debate, surprisingly given its potential ubiquity in dioecious organisms. At fitness-related loci, a given genotype may be selected in opposite directions in the two sexes. Such sexually antagonistic selection will reduce the otherwise-expected positive genetic correlation between male and female fitness. Both theory and experimental data suggest that males and females of the same species may have divergent genetic optima, but supporting data from wild populations are still scarce. Here we present evidence for sexually antagonistic fitness variation in a natural population, using data from a long-term study of red deer (Cervus elaphus). We show that male red deer with relatively high fitness fathered, on average, daughters with relatively low fitness. This was due to a negative genetic correlation between estimates of fitness in males and females. In particular, we show that selection favours males that carry low breeding values for female fitness. Our results demonstrate that sexually antagonistic selection can lead to a trade-off between the optimal genotypes for males and females; this mechanism will have profound effects on the operation of selection and the maintenance of genetic variation in natural populations.     

Cryptic genetic variation promotes rapid evolutionary adaptation in an RNA enzyme

Cryptic variation is caused by the robustness of phenotypes to mutations. Cryptic variation has no effect on phenotypes in a given genetic or environmental background, but it can have effects after mutations or environmental change. Because evolutionary adaptation by natural selection requires phenotypic variation, phenotypically revealed cryptic genetic variation may facilitate evolutionary adaptation. This is possible if the cryptic variation happens to be pre-adapted, or "exapted", to a new environment, and is thus advantageous once revealed. However, this facilitating role for cryptic variation has not been proven, partly because most pertinent work focuses on complex phenotypes of whole organisms whose genetic basis is incompletely understood. Here we show that populations of RNA enzymes with accumulated cryptic variation adapt more rapidly to a new substrate than a population without cryptic variation. A detailed analysis of our evolving RNA populations in genotype space shows that cryptic variation allows a population to explore new genotypes that become adaptive only in a new environment. Our observations show that cryptic variation contains new genotypes pre-adapted to a changed environment. Our results highlight the positive role that robustness and epistasis can have in adaptive evolution.     

应用小种群远缘杂交提高遗传算法的效率

为提高遗传算法的效率,将作物育种学中远缘杂交策略应用于多种群遗传算法,采用规模较小的多个种群同时进行进化,选择和变异操作在各种群内部独立完成,杂交操作在种群间完成。小种群的采用可以大大提高进化求解的速度,种群间的远缘杂交能够克服由于种群规模小、种群多样性降低导致早熟收敛的弊端,保证算法以较快的速度收敛到全局最优解。研究结果表明:该算法具有高效性。     

Scanning for sisnatures of geosraphically restricted selection based on population genomics analysis

Natural selection, as the driving force of human evolution, has direct impact on population differentiation. However, it is still unclear to what extent the genetic differentiation has been caused by natural selection. To explore this question, we performed a genome-wide scan with single nucleotide polymorphism （SNP） data from the International HapMap Project. Single locus FsTanalysis was applied to assess the frequency difference among populations in autosomes. Based on the empirical distribution of FsT, we identified 12669 SNPs correlating to population differentiation and 1853 candidate genes subjected to geographic restricted natural selection. Further interpretation of gene ontogeny revealed 121 categories of biological process with the enrichments of candidate genes. Our results suggest that natural selection may play an important role in human population differentiation. In addition, our analysis provides new clues as well as research methods for our understanding of population differentiation and natural selection.     

Emergence of cooperation and evolutionary stability in finite populations

To explain the evolution of cooperation by natural selection has been a major goal of biologists since Darwin. Cooperators help others at a cost to themselves, while defectors receive the benefits of altruism without providing any help in return. The standard game dynamical formulation is the 'Prisoner's Dilemma', in which two players have a choice between cooperation and defection. In the repeated game, cooperators using direct reciprocity cannot be exploited by defectors, but it is unclear how such cooperators can arise in the first place. In general, defectors are stable against invasion by cooperators. This understanding is based on traditional concepts of evolutionary stability and dynamics in infinite populations. Here we study evolutionary game dynamics in finite populations. We show that a single cooperator using a strategy like 'tit-for-tat' can invade a population of defectors with a probability that corresponds to a net selective advantage. We specify the conditions required for natural selection to favour the emergence of cooperation and define evolutionary stability in finite populations.     

Detecting recent positive selection in the human genome from haplotype structure

The ability to detect recent natural selection in the human population would have profound implications for the study of human history and for medicine. Here, we introduce a framework for detecting the genetic imprint of recent positive selection by analysing long-range haplotypes in human populations. We first identify haplotypes at a locus of interest (core haplotypes). We then assess the age of each core haplotype by the decay of its association to alleles at various distances from the locus, as measured by extended haplotype homozygosity (EHH). Core haplotypes that have unusually high EHH and a high population frequency indicate the presence of a mutation that rose to prominence in the human gene pool faster than expected under neutral evolution. We applied this approach to investigate selection at two genes carrying common variants implicated in resistance to malaria: G6PD and CD40 ligand. At both loci, the core haplotypes carrying the proposed protective mutation stand out and show significant evidence of selection. More generally, the method could be used to scan the entire genome for evidence of recent positive selection.     

实数编码最优子种群遗传算法及结构优化

遗传算法(GA)是利用自然选择和进化思想在高维空间中寻优的方法,其寻优过程始终保持整个种群的进化.本文提出了实数编码最优子种群遗传算法理论,通过从种群中选出适应值最高的若干数量的个体,组成该代最优子种群,将最优子种群中的个体与种群中其它个体进行交叉变异、最优子种群中的个体间也进行交叉变异,从而产生新的种群.该遗传算法使得遗传过程中落入局部最优解的几乎不可能,对于多极值问题非常有效,收敛速度也非常快.     

分子进化中性理论的进展

中性理论与达尔文的自然选择进化论有明显的差别。中性论认为在分子水平上覆盖着多数的进化改变，是在连续产生突变时，由于选择中性（即选择相等），使突变得到随机的固定（即小群体内，样本的随机漂变）。还认为在分子水平上，大多数的种内变异（如蛋白质和ＤＮＡ多态）是选择中性或近于中性，它们能够保持在种内是由于突变的产生和随机消失之间的平衡引起的。近年来，ＤＮＡ顺序的研究资料，都有力地支持中性论，如鼠的拟球蛋白基因、鼹鼠的αＡ－晶状体蛋白、流行性感冒Ａ病毒基因，果蝇的核和线粒体基因等。所以，从地球上生命的历史来看，中性进化的改变至少在数量上已超过达尔文自然选择进化的改变。     

Genetic mechanisms of floral trait correlations in a natural population

Genetic correlations among traits are important in evolution, as they can constrain evolutionary change or reflect past selection for combinations of traits. Constraints and integration depend on whether the correlations are caused by pleiotropy or linkage disequilibrium, but these genetic mechanisms underlying correlations remain largely unknown in natural populations. Quantitative trait locus (QTL) mapping studies do not adequately address the mechanisms of within-population genetic correlations because they rely on crosses between distinct species, inbred lines or selected lines (see ref. 5), and they cannot distinguish moderate linkage disequilibrium from pleiotropy because they commonly rely on only one or two episodes of recombination. Here I report that after nine generations of enforced random mating (nine episodes of recombination), correlations between six floral traits in wild radish plants are unchanged, showing that pleiotropy generates the correlations. There is no evidence for linkage disequilibrium despite previous correlational selection acting on one functionally integrated pair of traits. This study provides direct evidence of the genetic mechanisms underlying correlations between quantitative traits in a natural population and suggests that there may be constraints on the independent evolution of pairs of highly correlated traits.     

MHC II DRB variation and trans-species polymorphism in the golden snub-nosed monkey (Rhinopithecus roxellana)

Genetic variation is generally believed to be important in studying endangered species’ adaptive potential.Early studies assessed genetic diversity using nearly neutral markers,such as microsatellite loci and mitochondrial DNA(mtDNA),which are very informative for phylogenetic and phylogeographic reconstructions.However,the variation at these loci cannot provide direct information on selective processes involving the interaction of individuals with their environment,or on the capability to resist continuously evolving pathogens and parasites.The importance of genetic diversity at informative adaptive markers,such as major histocompatibility complex(MHC) genes,is increasingly being realized,especially in endangered,isolated species.Small population size and isolation make the golden snub-nosed monkey(Rhinopithecus roxellana) particularly susceptible to genetic variation losses through inbreeding and restricted gene flow.In this study,we compared the genetic variation and population structure of microsatellites,mtDNA,and the most relevant adaptive region of the MHC II-DRB genes in the golden snub-nosed monkey.We examined three Chinese R.roxellana populations and found the same variation patterns in all gene regions,with the population from Shennongjia population,Hubei Province,showing the lowest polymorphism among three populations.Genetic drift that outweighed balancing selection and the founder effect in these populations may explain the similar genetic variation pattern found in these neutral and adaptive genes.