THE EFFECTS OF POPULATION BOTTLENECKS ON CLONAL INTERFERENCE, AND THE ADAPTATION EFFECTIVE POPULATION SIZE

Clonal interference refers to the competition that arises in asexual populations when multiple beneficial mutations segregate simultaneously. A large body of theoretical and experimental work now addresses this issue. Although much of the experimental work is performed in populations that grow exponentially between periodic population bottlenecks, the theoretical work to date has addressed only populations of a constant size. We derive an analytical approximation for the rate of adaptation in the presence of both clonal interference and bottlenecks, and compare this prediction to the results of an individual-based simulation, showing excellent agreement in the parameter regime in which clonal interference prevails. We also derive an appropriate definition for the effective population size for adaptive evolution experiments in the presence of population bottlenecks. This "adaptation effective population size" allows for a good approximation of the expected rate of adaptation, either in the strong-selection weak-mutation regime, or when clonal interference comes into play. In the multiple mutation regime, when the product of the population size and mutation rate is extremely large, these results no longer hold.     

THE SIGNIFICANCE OF GENETIC EROSION IN THE PROCESS OF EXTINCTION. IV. INBREEDING DEPRESSION AND HETEROSIS EFFECTS CAUSED BY SELFING AND OUTCROSSING IN SCABIOSA COLUMBARIA

The effects of self-fertilization, within-population crosses (WPC) and between-population crosses (BPC) on progeny fitness were investigated in the greenhouse for Scabiosa columbaria populations of varying size. Plants grown from field collected seeds were hand pollinated to produce selfed, WPC, and BPC progeny. The performance of these progenies was examined throughout the entire life cycle. The different pollination treatments did not significantly affect germination, seedling-to-adult survival, flowering percentage and the number of flower heads. But severe inbreeding depression was demonstrated for biomass production, root development, adult survival, and seed set. Additionally, multiplicative fitness functions were calculated to compare relative fitnesses for progeny. On average, WPC progeny showed a more than 4-fold, and BPC progeny an almost 10-fold, advantage over selfed progeny, indicating that S. columbaria is highly susceptible to inbreeding. No clear relationship was found between population size and level of inbreeding depression, suggesting that the genetic load has not yet been reduced substantially in the small populations. A significant positive correlation was found between plant dry weight and total fitness. In two out of six populations, the differences between the effects of the pollination treatments on dry weight increased significantly when seedlings were grown under competitive conditions. This result is interpreted as an enhancement of inbreeding depression under these conditions. It is argued that improvement of the genetic exchange between populations may lower the probability of population extinction.     

影响禾谷缢管蚜种群密度的因子分析

运用回归分析、通径分析和网络理论探讨了小麦植株体内氨基酸、蔗糖、水分和温度、光照等因子对禾谷缢管蚜种群量及其胚胎数的单个影响和综合作用。结果表明,单个因子对麦蚜的作用符合二次多项式ｙ２＝ａ＋ｂ１ｘ＋ｂ２ｘ＾２或三次多项式ｙ３＝ａ＋ｂ１ｘ＋ｂ２ｘ＾２＋ｂ３ｘ＾３关系,５个因子的共同作用以多元非线性回归模型ｙ＝ａ＋ΣΣ（ｂｉｊｘｉ＾ｊ）＋ΣΣ（ｂｉｊｘｉ＇ｊ＇ｘｉ＾ｊｘｉ＇＾ｊ＇）拟合比较合理。计算结     

MUTATION AND EXTINCTION: THE ROLE OF VARIABLE MUTATIONAL EFFECTS,SYNERGISTIC EPISTASIS,BENEFICIAL MUTATIONS,AND DEGREE OF OUTCROSSING

Recent theoretical studies have illustrated the potential role of spontaneous deleterious mutation as a cause of extinction in small populations. However, these studies have not addressed several genetic issues, which can in principle have a substantial influence on the risk of extinction. These include the presence of synergistic epistasis, which can reduce the rate of mutation accumulation by progressively magnifying the selective effects of mutations, and the occurrence of beneficial mutations, which can offset the effects of previous deleterious mutations. In stochastic simulations of small populations (effective sizes on the order of 100 or less), we show that both synergistic epistasis and the rate of beneficial mutation must be unrealistically high to substantially reduce the risk of extinction due to random fixation of deleterious mutations. However, in analytical calculations based on diffusion theory, we show that in large, outcrossing populations (effective sizes greater than a few hundred), very low levels of beneficial mutation are sufficient to prevent mutational decay. Further simulation results indicate that in populations small enough to be highly vulnerable to mutational decay, variance in deleterious mutational effects reduces the risk of extinction, assuming that the mean deleterious mutational effect is on the order of a few percent or less. We also examine the magnitude of outcrossing that is necessary to liberate a predominantly selfing population from the threat of long-term mutational deterioration. The critical amount of outcrossing appears to be greater than is common in near-obligately selfing plant species, supporting the contention that such species are generally doomed to extinction via random drift of new mutations. Our results support the hypothesis that a long-term effective population size in the neighborhood of a few hundred individuals defines an approximate threshold, below which outcrossing populations are vulnerable to extinction via fixation of deleterious mutations, and above which immunity is acquired.     

FIVE GENERATIONS OF ENFORCED SELFING AND OUTCROSSING IN MIMULUS GUTTATUS: INBREEDING DEPRESSION VARIATION AT THE POPULATION AND FAMILY LEVEL

The focus of this study was to examine the consequences of five sequential generations of enforced selfing and outcrossing in two annual populations of the mixed-mating Mimulus guttatus. Our primary goal was to determine whether purging of deleterious recessive alleles occurs uniformly between populations and among families, and thus gain insights into the mode of gene action (dominance, overdominance, and/or epistasis) governing the expression of inbreeding depression at both the population and family levels across the life cycle.     

THE INFLUENCE OF POPULATION SIZE AND ISOLATION ON GENE FLOW BY POLLEN IN SILENE ALBA

In a series of experiments conducted over two seasons, we used arrays of experimental populations to examine the effects of flower number and distance between patches on gene flow by pollen. For this study we used the dioecious, short-lived perennial plant Silene alba (Caryophyllaceae). This species lives in disturbed roadside and agricultural habitats and displays a weedy population dynamic with high colonization and extinction rates. The motivation for the study was to understand what factors may be influencing genetic connectedness among newly colonized populations within a regional metapopulation. By using experimental populations composed of genotypes homozygous at a diagnostic locus, it was possible to identify explicitly pollen movement into a focal patch as a function of flower number and distance to the nearest neighboring patch. Overall, the mean immigration rate (measured as the fraction of seeds sired by males outside the focal patch) at 20 m was just over 47%, whereas at 80 m immigration rates were less than 6%. In addition, by knowing the context in which each of these gene-flow events occurred, it was possible to understand some of the factors that influenced the exchange of genes. Both the number of flowers in the focal population (target) and in the neighboring populations (source) had a significant effect on the frequency of gene flow. Our experimental data also demonstrate that factors that influence gene flow at one spatial scale may not act in the same way at another. Specifically, the influence of target size and the relative size of the target and source patches on rates of gene flow depended on whether the patches were separated by 20 m or 80 m. These data suggest that the patterns of gene flow within a metapopulation system can be complex and may vary within a growing season.     

THE SIGNIFICANCE OF GENETIC EROSION IN THE PROCESS OF EXTINCTION. IV. INBREEDING LOAD AND HETEROSIS IN RELATION TO POPULATION SIZE IN THE MINT SALVIA PRATENSIS

If, because of genetic erosion, the level of homozygosity in small populations is high, additional selfing will result in small reductions of fitness. In addition, in small populations with a long inbreeding history selection may have purged the population of its genetic load. Therefore, a positive relationship between population size (or level of genetic variation) and level of additional inbreeding depression, here referred to as inbreeding load, may be expected. In a previous study on the rare and threatened perennial Salvia pratensis, a positive correlation between population size and level of allozyme variation has been demonstrated. In the present study, the inbreeding load in six populations of varying size and allozyme variation was investigated. In the greenhouse, significant inbreeding load in mean seed weight, proportion of germination, plant size, regenerative capacity, and survival was demonstrated. In a field experiment with the two largest and the two smallest populations, survival of selfed progeny was 16% to 63% lower than survival of outcrossed progeny. In addition, survival of outcrossed progeny was, with the exception of the largest population, lower (16% to 37%) than of hybrid progeny, resulting from crosses between populations. Effects on plant size were qualitatively similar to the effects on survival, but these effects were variable in time because of differential survival of larger individuals. In all populations the total inbreeding load, that is, the effects on size and survival multiplicated, increased in time. It was demonstrated that inbreeding load in different characters may be independent. At no time and for no character was inbreeding load or the heterosis effect correlated to the mean number of alleles per locus, indicating that allozyme variation is not representative for variation at fitness loci in these populations. Combined with results of previous investigations, these results suggest that the small populations are in an early phase of the genetic erosion process. In this phase, allozyme variation, which is supposed to be (nearly) neutral, has been affected by genetic erosion but the selectively nonneutral variation is only slightly affected. These results stress the need for detailed information about the inbreeding history of small populations. The relative performance of selfed progeny was lowest in all populations, in the greenhouse as well as in the field, and inbreeding depression could still influence the extinction probabilities of the small populations.     

THE INFLUENCE OF MATING SYSTEM AND OVERLAPPING GENERATIONS ON EFFECTIVE POPULATION SIZE

The effective population size (N_e) depends strongly on mating system and generation time. These two factors interact such that, under many circumstances, N_e is close to N/2, where N is the number of adults. This is shown to be the case for both simple and highly polygynous mating systems. The random union of gametes (RUG) and monogamy are two simple systems previously used in estimating N_e, and here a third, lottery polygyny, is added. Lottery polygyny, in which all males compete equally for females, results in a lower N_e than either RUG or monogamy! Given nonoverlapping generations the reduction is 33% for autosomal loci and 25% for sex-linked loci. The highly polygynous mating systems, harem polygyny and dominance polygyny, can give very low values of N_e/N when the generation time (T) is short. However, as T is lengthened, N_e approaches N/2. The influence of a biased sex ratio depends on the mating system and, in general, is not symmetrical. Biases can occur because of sex differences in either survival or recruitment of adults, and the potential for a sex-ratio bias to change N_e is much reduced given a survival bias. The number of juveniles present also has some influence: as the maturation time is lengthened, N_e increases.     

扑虱灵对稻飞虱及其捕食性天敌种群的影响

扑虱灵对稻飞虱的防治效果,及对捕食性天敌的毒性,进行了田间试验,并与叶蝉散,杀虫双的使用结果作比较．试验结果表明,扑虱灵对稻飞虱的防治效果达93％,与叶蝉散,杀虫双相比较。防治效果提高了2．39～32．85％．对黑肩绿盲蝽和蜘蛛种群具有不同程度的选择性,维护较低的稻飞虱与天敌的比值,有利于天敌作用的发挥,维持对稻飞虱较长的残效期,基本上可以控制稻田后期飞虱的增长．     

THE EFFECT OF HOST PLANT PATCH SIZE VARIATION ON THE POPULATION STRUCTURE OF A SPECIALIST HERBIVORE INSECT,TETRAOPES TETRAOPHTHALMUS

The patchy local distribution of the common milkweed, Asclepias syriaca, organizes populations of a beetle that feeds on it, Tetraopes tetraophthalmus, into numerous local demes. Genetic and ecological characteristics of demes of adult milkweed beetles occupying two naturally occurring size classes of patches, defined as large and small, were studied in order to describe the effect of patch size variation on local population structure. Allele frequency variance in two of three protein polymorphisms was significantly greater in collections of beetles from an array of 13 small patches when compared to collections from an array of 11 large populations. A multivariate measure of variance using information from all 3 genetic markers confirmed that the small patches displayed greater overall genetic differentiation. This was further quantified by computing an F_st value, combined across loci, of 0.018 for the small patches and 0.004 for the large patches. No significant difference between patch size classes in mean allele frequency was detected. Mark and recapture studies of the adults found in five small and four large patches showed the residence times of adults in small patches to be less than half of those found in large patches. This was interpreted as resulting from higher emigration rates from small patches. It is proposed that greater genetic differentiation is found among demes from smaller patches because smaller patches support smaller population sizes and further because smaller patches act as net exporters of migrants while larger patches act as net migrant importers.     

A TEST AND REVIEW OF THE ROLE OF EFFECTIVE POPULATION SIZE ON EXPERIMENTAL SEXUAL SELECTION PATTERNS

Experimental evolution, particularly experimental sexual selection in which sexual selection strength is manipulated by altering the mating system, is an increasingly popular method for testing evolutionary theory. Concerns have arisen regarding genetic diversity variation across experimental treatments: differences in the number and sex ratio of breeders (effective population size; N_e ) and the potential for genetic hitchhiking, both of which may cause different levels of genetic variation between treatments. Such differences may affect the selection response and confound interpretation of results. Here we use both census-based estimators and molecular marker-based estimates to empirically test how experimental evolution of sexual selection in Drosophila pseudoobscura impacts N_e and autosomal genetic diversity. We also consider effects of treatment on X-linked N_e s, which have previously been ignored. Molecular autosomal marker-based estimators indicate that neither N_e nor genetic diversity differs between treatments experiencing different sexual selection intensities; thus observed evolutionary responses reflect selection rather than any confounding effects of experimental design. Given the increasing number of studies on experimental sexual selection, we also review the census N_e s of other experimental systems, calculate X-linked N_e , and compare how different studies have dealt with the issues of inbreeding, genetic drift, and genetic hitchhiking to help inform future designs.     

短命植物条叶车前小孢子发生与雄配子体发育研究

条叶车前小孢子发生与雄配子体形成过程如下：花药具４个花粉囊;药壁由表皮、药室内壁、中层、绒毡层等４层细胞组成,发育方式为双子叶型,绒毡层细胞为腺质;小孢子母细胞减数分裂过程中胞质分裂为同时型;小孢子四分体多为四面体型,少数为十字交叉型或两侧对称型;成熟花粉粒为３－细胞,两个精子由原生质丝与营养核联系起来,因此可能存在着“雄性生殖单位”。     

THE EFFECTS OF PREDATION ON THE AGE AND SIZE OF MATURITY OF PREY

The effects of nonselective predation on the optimal age and size of maturity of their prey are investigated using mathematical models of a simple life history with juvenile and adult stages. Fitness is measured by the product of survival to the adult stage and expected adult reproduction, which is usually an increasing function of size at maturity. Size is determined by both age at maturity and the value of costly traits that increase mean growth rate (growth effort). The analysis includes cases with fixed size but flexible time to maturity, fixed time but flexible size, and adaptively flexible values of both variables. In these analyses, growth effort is flexible. For comparison with previous theory, models with a fixed growth effort are analyzed. In each case, there may be indirect effects of predation on the prey's food supply. The effect of increased predation depends on (1) which variables are flexible; (2) whether increased growth effort requires increased exposure to predators; and (3) how increased predator density affects the abundance of food for juvenile prey. If there is no indirect effect of predators on prey food supply, size at maturity will generally decrease in response to increased predation. However, the indirect effect from increased food has the opposite effect, and the net result of predation is often increased size. Age at maturity may either increase or decrease, depending on functional forms and parameter values; this is true regardless of the presence of indirect effects. The results are compared with those of previous theoretical analyses. Observed shifts in life history in response to predation are reviewed, and the role of size-selective predation is reassessed.     

两种珍稀植物群落物种多度分布的核方法研究

首次提出物种多度分布的非参数核密度估计方法,介绍了此方法的构造和主要性质。珍稀濒危植物观光木群落和长苞铁杉群落的乔木层、灌木层、所有木本植物物种多度分布实例拟合结果表明,核方法能很好地描述群落物种多度分布。非参数核估计方法是群落物种多度分布模拟的一种有效方法,它丰富了物种多度分布拟合方法,为珍稀濒危植物的管理与保护提供了理论参考。     

克隆植物乌菱对底泥磷含量及植株密度的表型可塑性响应

植物,尤其是克隆植物,能够通过表型变化来缓解外界压力,提高对环境的适应能力。该文研究了水生克隆植物乌菱(Trapa bicornis)对底泥磷含量(Sediment phosphorus concentration, SP)、植株密度(Plant density, PD) 及两者间交互作用的可塑性响应,探讨可塑性是否能促进其在富营养化环境中的生长。结果显示,底泥磷含量对乌菱的主菱盘叶数、同化根比根长、吸收根比根长以及叶、茎、同化根、吸收根与植株总磷含量等都有显著影响 (p<0.05),而植株密度对乌菱各生长及生理生态参数均无显著作用 (p>0.05);SP与PD的交互作用弱化了底泥磷含量对乌菱的效应。底泥磷含量和植株密度甚至改变了同化根、吸收根、茎、叶与总生物量之间的异速生长关系。研究结果表明:乌菱的表型可塑性变化主要受底泥磷含量的影响,乌菱通过器官生物量分配、形态结构及生理生态特征的调整来响应底泥磷含量的变化;同时,高的植株密度也可以提高其在富营养化生境下的生态适应性。     

CONSTANCY OF POPULATION PARAMETERS FOR LIFE-HISTORY AND FLORAL TRAITS IN RAPHANUS SATIVUS L. II. EFFECTS OF PLANTING DENSITY ON PHENOTYPE AND HERITABILITY ESTIMATES

To determine the effect of growing conditions on population parameters in wild radish, (Raphanus sativus L.: Brassicaceae), we replicated maternal and paternal half-sib families of seed across three planting densities in an experimental garden. A nested breeding design performed in the greenhouse produced 1,800 F₁ seeds sown in the garden. We recorded survivorship, measured phenotypic correlations among and estimated narrow-sense and broad-sense heritabilities (h²) of: days to germination, days to flowering, petal area, ovule number/flower, pollen production/flower, and modal pollen grain volume. Survivorship declined with increasing density, but the relative abundances of surviving families did not differ significantly among densities. Seeds in high-density plots germinated significantly faster than seeds sown in medium- or low-density plots, but they flowered significantly later. Plants in high-density plots had fewer ovules per flower than those in the other treatments. Petal area and pollen characters did not differ significantly among densities. Densities differed with respect to the number and sign of significant phenotypic correlations. Analyses of variance were conducted to detect additive genetic variance (V_a) of each trait in each density. At low density, there were significant paternal effects on flowering time and modal pollen grain volume; in medium-density plots, germination time, flowering time and ovule number exhibited significant paternal effects; in high-density plots, only pollen grain volume differed among paternal sibships. The ability to detect maternal effects on progeny phenotype also depended on density. Narrow-sense h² estimates differed markedly among density treatments for germination time, flowering time, ovule number and pollen grain volume. Maternal, paternal and error variance components were estimated for each trait and density to examine the sources of variation in narrow-sense h² across densities. Variance components did not change consistently across densities; each trait behaved differently. To provide qualitative estimates of genetic correlations between characters, correlation coefficients were estimated using paternal family means; these correlations also differed among densities. These results demonstrate that: a) planting density influences the magnitude of maternal and paternal effects on progeny phenotype, and of h² estimates, b) traits differ with respect to the density in which heritability is greatest, c) density affects the variance components that comprise heritability, but each trait behaves differently, and d) the response to selection on any target trait should result in different correlated responses of other traits, depending on density.     

松嫩平原碱化草甸天然翅碱蓬种群的密度制约模型

翅碱蓬是一年生藜科植物,耐碱性极强。本文根据在单—种群落随机取样的调查数据,从种群水平分析了松嫩平原碱化草甸天然翅碱蓬种群的密度制约特征。结果表明,翅碱蓬种群在不同生长期及不同数量性状的密度制约模型均可由多种函数形式同时较好地表达出来。但本文仅以相关性最好的拟合方程作为种群某一性状密度制约特征的模型。孕蕾期的平均植株重和单位面积籽实重的密度制约表现为变形双曲线函数y=a+b/x形式;籽实成熟期的平均植株重、平均植株籽实重、平均植株种子数和单位面积种子数均表现为幂函数y=ax-b形式;地上生物量在孕蕾期为变形双曲线函数y=1/(a+bx)形式,在籽实成熟期为对数函数y=a-blnx形式。     

香蕉种植密度对蕉园光能利用与产量的影响

本试验研究了福建沿海地区蕉园种植密度与光能利用效率和产量之间的关系。结果表明,适当提高种植密度,可增加香蕉单位空间的光能利用,充分利用地力,改善蕉园生态条件,显著地提高香蕉亩产量。亩植160～180株是沿海地区达到高产稳产的有效措施。