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1.
An analysis is made of the variation among individuals in finite populations of the proportion of their genes which are identical by descent. There are two causes of this variability: variation in pedigree among individuals, and linkage which causes whole blocks of genes to be identical or nonidentical by descent. The variation between and within populations is analyzed in detail for several mating systems: monoecious populations with and without random selfing, and dioecious populations with and without a hierarchical mating structure. Transition matrices for two-locus descent measures are given for each system. Total variability is obtained by integrating these measures over the distribution of map distances over whole chromosomes. Approximate methods are also developed for unlinked loci. Unless populations have a very small effective size (Ne) there is little variation in inbreeding between populations. For unlinked loci, the coefficient of variation in nonidentity within populations approaches about 1/√3Ne for random selfing, 1/√6Ne for monogamous matings and 1/√12Ne for monoecy with selfing excluded or dioecy with random pairing. If there is no association between map distance and initial heterozygosity or effect on quantitative traits, the coefficient of variation in mean heterozygosity over the genome is related to that in nonidentity, and the additional variation in a quantitative trait due to dominant genes equals the product of the square of the initial inbreeding depression and the squared coefficient of variation of nonidentity.  相似文献   

2.
Golding GB  Strobeck C 《Genetics》1980,94(3):777-789
The linkage disequilibrium expected in a finite, partially selfing population is analyzed, assuming the infinite allele model. Formulas for the expected sum of squares of the linkage disequilibria and the squared standard linkage disequilibrium are derived from the equilibrium values of sixteen inbreeding coefficients required to describe the behavior of the system. These formulas are identical to those obtained with random mating if the effective population size Ne = (1-½S)N and the effective recombination value re = (1-S)r/(1-½S), where S is the proportion of selfing, are substituted for the population size and the recombination value. Therefore, the effect of partial selfing at equilibrium is to reduce the population size by a factor 1-½S and the recombination value by a factor (1-S)/(1-½S).  相似文献   

3.
Effective population size (N e) quantifies the effects of micro-evolutionary processes and the rate of loss of genetic diversity in a population. Several demographic and mating parameters reduce N e. Theoretical studies elucidate the impacts of various demographic and mating system parameters on N e, while empirical studies illustrate realized N e for species with differing life histories and mating systems. However, effect of intra-specific variation in mating system on effective size remains largely unexplored. In this paper we investigated the effect of promiscuous and polygynous mating on N e in two wild populations of the short-nosed fruit bat, Cynopterus sphinx. N e/N (ratio of effective population size to census size) was lower than unity in both populations, and much lower for the polygynous population compared to promiscuous population. Elasticity analyses reveal that N e/N was sensitive to deviations in the sex ratio. Variance in female reproductive success had a higher impact on N e compared to variance in male reproductive success in the promiscuous population. However, for the polygynous population, impact of variance in male reproductive success on N e was higher than that of variance in female reproductive success. Our results suggest that depending on mating system, different populations of the same species could have alternate evolutionary trajectories. The rate of loss of genetic diversity would be lower for the promiscuous population compared to the polygynous population. Our study is the first to highlight which parameters would most significantly impact population specific N e under different mating systems.  相似文献   

4.
Use of genetic methods to estimate effective population size (Ne) is rapidly increasing, but all approaches make simplifying assumptions unlikely to be met in real populations. In particular, all assume a single, unstructured population, and none has been evaluated for use with continuously distributed species. We simulated continuous populations with local mating structure, as envisioned by Wright''s concept of neighborhood size (NS), and evaluated performance of a single-sample estimator based on linkage disequilibrium (LD), which provides an estimate of the effective number of parents that produced the sample (Nb). Results illustrate the interacting effects of two phenomena, drift and mixture, that contribute to LD. Samples from areas equal to or smaller than a breeding window produced estimates close to the NS. As the sampling window increased in size to encompass multiple genetic neighborhoods, mixture LD from a two-locus Wahlund effect overwhelmed the reduction in drift LD from incorporating offspring from more parents. As a consequence, never approached the global Ne, even when the geographic scale of sampling was large. Results indicate that caution is needed in applying standard methods for estimating effective size to continuously distributed populations.  相似文献   

5.
Through stochastic simulations, estimates of breeding values accuracies and response to selection were assessed under traditional pedigree-based and genomic-based evaluation methods. More specifically, several key parameters such as the trait’s heritability (0.2 and 0.6), the number of QTLs underlying the trait (100 to 200), and the marker density (1 to 10 SNPs/cM) were evaluated. Additionally, impact of two contrasting mating designs (partial diallel vs. single-pair mating) was investigated. Response to selection was then assessed in a seed production population (seed orchard consisting of unrelated selections) for different effective population sizes (Ne?=?5 to 25). The simulated candidate population comprised a fixed size of 2050 individuals with fast linkage disequilibrium decay, generally found in forest tree populations. Following the genetic/genomic evaluation, top-ranked individuals were selected to meeting the predetermined effective population size in target production population. The combination of low h2, high Ne, and dense marker coverage resulted at maximum relative genomic prediction efficiency and the most efficient exploitation of the Mendelian sampling term (within-family additive genetic variance). Since genomic prediction of breeding values constitutes the methodological foundation of genomic selection, our results can be used to address important questions when similar scenarios are considered.  相似文献   

6.
C. Chevalet  M. Gillois    R. F. Nassar 《Genetics》1977,86(3):697-713
Properties of identity relation between genes are discussed, and a derivation of recurrent equations of identity coefficients in a random mating, diploid dioecious population is presented. Computations are run by repeated matrix multiplication. Results show that for effective population size (Ne) larger than 16 and no mutation, a given identity coefficient at any time t can be expressed approximately as a function of (1—f), (1—f)3 and (1— f)6, where f is the mean inbreeding coefficient at time t. Tables are presented, for small Ne values and extreme sex ratios, showing the pattern of change in the identity coefficients over time. The pattern of evolution of identity coefficients is also presented and discussed with respect to N eu, where u is the mutation rate. Applications of these results to the evolution of genetic variability within and between inbred lines are discussed.  相似文献   

7.
Effective population size (N e) is a central concept in evolutionary biology and conservation genetics. It predicts rates of loss of neutral genetic variation, fixation of deleterious and favourable alleles, and the increase of inbreeding experienced by a population. A method exists for the estimation of N e from the observed linkage disequilibrium between unlinked loci in a population sample. While an increasing number of studies have applied this method in natural and managed populations, its reliability has not yet been evaluated. We developed a computer program to calculate this estimator of N e using the most widely used linkage disequilibrium algorithm and used simulations to show that this estimator is strongly biased when the sample size is small (<‰100) and below the true N e. This is probably due to the linkage disequilibrium generated by the sampling process itself and the inadequate correction for this phenomenon in the method. Results suggest that N e estimates derived using this method should be regarded with caution in many cases. To improve the method’s reliability and usefulness we propose a way to determine whether a given sample size exceeds the population N e and can therefore be used for the computation of an unbiased estimate.  相似文献   

8.
Clegg MT  Kidwell JF  Horch CR 《Genetics》1980,94(1):217-234
The dynamic behavior of four-locus gametic frequency distributions was studied in five replicate cage populations of Drosophila melanogaster for up to 50 generations. The joint frequency distributions were resolved into gene frequencies and various disequilibrium measures. In addition, F statistics for marginal single-locus genotypic frequency distributions were followed through time. The gene frequency, disequilibrium and F statistics were obtained for four chromosome 3 enzyme marker loci [isocitrate dehydrogenase (3–27.1), esterase-6 (3–36.8), phosphoglucomutase (3–43.4) and esterase-C (3–49.0)]. The initial structure of the experimental populations featured random mating proportions, and two complementary gametic types with respect to the marker loci, thus assuring complete pairwise linkage disequilibrium among the markers.——The experimental results indicate: (1) the between-replicate variance in gene frequency varied substantially among loci, with isocitrate dehydrogenase showing the greatest between-replicate variance, and esterase-C the least. (2) The F statistics initially were strongly negative but decayed to the neighborhood of zero for all marker loci except esterase-C. The rate at which the F statistics approached zero varied among the marker loci, indicating substantial differences in the distribution of selective effects along the chromosome. The centromeric region, marked by esterase-C, shows the strongest selective effects. (3) The rate of decay of linkage disequilibrium was much faster than expected for pairs of neutral loci, averaging 1.82 times the neutral rate over all replicates and pairs of loci. This acceleration, which was observed for all six pairwise combinations of loci, was interpreted as resulting from the interaction between selection and recombination. Our experimental results are consistent with many investigations of linkage disequilibrium in natural populations of Drosophila melanogaster that show little or no disequilibrium among enzyme loci. (4) A fortuitous contamination of two cages revealed an apparent regulatory interaction between the migrant and nonmigrant chromosomes at the esterase-C locus. The migrant chromosomes were very rapidly absorbed into the recipient populations, despite this interaction. This result suggests that the dynamics of migration in populations may be phenomenologically richer than anticipated by simple theory.  相似文献   

9.
Several Peronospora species are carried by wind over short and long distances, from warmer climates where they survive on living plants to cooler climates. In eastern Canada, this annual flow of sporangia was thought to be the main source of Peronospora destructor responsible for onion downy mildew. However, the results of a recent study showed that the increasing frequency of onion downy mildew epidemics in eastern Canada is associated with warmer autumns, milder winters, and previous year disease severity, suggesting overwintering of the inoculum in an area where the pathogen is not known to be endogenous. In this study, genotyping by sequencing was used to investigate the population structure of P. destructor at the landscape scale. The study focused on a particular region of southwestern Québec—Les Jardins de Napierville—to determine if the populations were clonal and regionally differentiated. The data were characterized by a high level of linkage disequilibrium, characteristic of clonal organisms. Consequently, the null hypothesis of random mating was rejected when tested on predefined or nonpredefined populations, indicating that linkage disequilibrium was not a function of population structure and suggesting a mixed reproduction mode. Discriminant analysis of principal components performed with predefined population assignment allowed grouping P. destructor isolates by geographical regions, while analysis of molecular variance confirmed that this genetic differentiation was significant at the regional level. Without using a priori population assignment, isolates were clustered into four genetic clusters. These results represent a baseline estimate of the genetic diversity and population structure of P. destructor.  相似文献   

10.
The variances of actual inbreeding and coancestry in terms of their corresponding identities by descent were studied for finite populations. For inbreeding at a single locus, the total variance σ2 = F(1 ? F) (F is the inbreeding coefficient) is comprised of a component σw2 within populations and a component σb2 between replicate populations. These variances increase in time to a maximum at about 1.1Ne generations for σw2, about 2.3Ne generations for σb2, and about 1.4Ne generations for σ2, and decrease thereafter (Ne is effective population size). The ratio σb2σ2 is ever increasing to an asymptote in the range 0.4-0.5 depending on Ne and the mating system. For finite populations with variation in pedigree F's, there are contributions σwF2 within and σbF2 between populations. The component σbF2 is insignificant except for very small populations, and σwF2 is largest in the early generations and then decreases roughly as (1 ? F)2KNewhere K is formulated in terms of the mating strategy and the degree of avoidance of mating relatives. An additional degree of avoidance increases K by a factor of 4. In a large population at equilibrium with respect to mixed self and random mating, σwF2 accounts for onehalf to two-thirds of σw2. Bringing in more loci leads to the decomposition of the total variance into four components whose values are affected by linkages among the loci. The relationships between these components and σw2, σb2, σwF2, and σbF2, are elaborated in terms of tight and loose linkage. The exact computations of σwF2 and σbF2 require the use of two locus descent measures without linkage. The variances of various averages of actual identities by descent, such as the proportions for individuals or populations, are formulated for a sample of individuals.  相似文献   

11.
The use of dense SNPs to predict the genetic value of an individual for a complex trait is often referred to as “genomic selection” in livestock and crops, but is also relevant to human genetics to predict, for example, complex genetic disease risk. The accuracy of prediction depends on the strength of linkage disequilibrium (LD) between SNPs and causal mutations. If sequence data were used instead of dense SNPs, accuracy should increase because causal mutations are present, but demographic history and long-term negative selection also influence accuracy. We therefore evaluated genomic prediction, using simulated sequence in two contrasting populations: one reducing from an ancestrally large effective population size (Ne) to a small one, with high LD common in domestic livestock, while the second had a large constant-sized Ne with low LD similar to that in some human or outbred plant populations. There were two scenarios in each population; causal variants were either neutral or under long-term negative selection. For large Ne, sequence data led to a 22% increase in accuracy relative to ∼600K SNP chip data with a Bayesian analysis and a more modest advantage with a BLUP analysis. This advantage increased when causal variants were influenced by negative selection, and accuracy persisted when 10 generations separated reference and validation populations. However, in the reducing Ne population, there was little advantage for sequence even with negative selection. This study demonstrates the joint influence of demography and selection on accuracy of prediction and improves our understanding of how best to exploit sequence for genomic prediction.  相似文献   

12.
The Effects of Overdominance on Linkage in a Multilocus System   总被引:3,自引:2,他引:1       下载免费PDF全文
Computer simulations were performed with overdominant multiple alleles among tightly linked multiple loci under a multiplicative fitness model. The quantity X2/N(n — 1) was introduced as a new measure of linkage disequilibrium which, unlike previously available measures, can be applied to multiple allele models, where N is the sample size, and n is the number of alleles at the locus possessing fewest alleles. Simulations showed that (1) With multiple (three or four) alleles, the approach to stable disequilibrium is slower and the amount of disequilibrium established is weaker than in a two allele system. (2) The number of complementary chromosomes is a function of number of alleles and of population size. (3) As population size increases, the rate of the approach to stable disequilibrium is slower. (4) There is an optimum selection coefficient which minimizes the transient fixation probability of alleles when linkage is present. (5) The absence of linkage disequilibrium is in most cases not a practical method of testing the hypothesis of balancing selection of genetic polymorphisms because it depends strongly on population size in determining linkage disequilibria.  相似文献   

13.
The effective population size (Ne) is proportional to the loss of genetic diversity and the rate of inbreeding, and its accurate estimation is crucial for the monitoring of small populations. Here, we integrate temporal studies of the gecko Oedura reticulata, to compare genetic and demographic estimators of Ne. Because geckos have overlapping generations, our goal was to demographically estimate NbI, the inbreeding effective number of breeders and to calculate the NbI/Na ratio (Na = number of adults) for four populations. Demographically estimated NbI ranged from 1 to 65 individuals. The mean reduction in the effective number of breeders relative to census size (NbI/Na) was 0.1 to 1.1. We identified the variance in reproductive success as the most important variable contributing to reduction of this ratio. We used four methods to estimate the genetic based inbreeding effective number of breeders NbI(gen) and the variance effective populations size NeV(gen) estimates from the genotype data. Two of these methods - a temporal moment-based (MBT) and a likelihood-based approach (TM3) require at least two samples in time, while the other two were single-sample estimators - the linkage disequilibrium method with bias correction LDNe and the program ONeSAMP. The genetic based estimates were fairly similar across methods and also similar to the demographic estimates excluding those estimates, in which upper confidence interval boundaries were uninformative. For example, LDNe and ONeSAMP estimates ranged from 14–55 and 24–48 individuals, respectively. However, temporal methods suffered from a large variation in confidence intervals and concerns about the prior information. We conclude that the single-sample estimators are an acceptable short-cut to estimate NbI for species such as geckos and will be of great importance for the monitoring of species in fragmented landscapes.  相似文献   

14.
Strains of Fusarium species belonging to section Liseola cause stalk and ear rot of maize and produce important mycotoxins, such as fumonisins. We isolated two species, Fusarium verticillioides (Gibberella fujikuroi mating population A) and Fusarium proliferatum (G. fujikuroi mating population D) from maize cultivated under no-till conditions at five locations in the Córdoba province of Argentina. We determined the effective population number for mating population A (Ne) and found that the Ne for mating type was 89% of the count (total population) and that the Ne for male or hermaphrodite status was 36%. Thus, the number of strains that can function as the female parent limits Ne, and sexual reproduction needs to occur only once every 54 to 220 asexual generations to maintain this level of sexual fertility. Our results indicate that the fungal populations isolated from no-till maize are similar to those recovered from maize managed with conventional tillage. We placed 36 strains from mating population A into 28 vegetative compatibility groups (VCGs). Of the 13 strains belonging to five multimember VCGs, only 2 isolates belonging to one VCG were clones based on amplified fragment length polymorphism (AFLP) fingerprints. Members of the other four multimember VCGs had an average similarity index of 0.89, and members of one VCG were no more closely related to other members of the same VCG than they were to other members of the population as a whole. This finding suggests that the common assumption that strains in the same VCG are either clonal or very closely related needs to be examined in more detail. The variability observed with AFLPs and VCGs suggests that sexual reproduction may occur more frequently than estimated by Ne.  相似文献   

15.
Mating systems and patterns of reproductive success in fishes play an important role in ecology and evolution. While information on the reproductive ecology of many anadromous salmonids (Oncorhynchus spp.) is well detailed, there is less information for nonanadromous species including the Yellowstone Cutthroat Trout (O. clarkii bouvieri), a subspecies of recreational angling importance and conservation concern. Using data from a parentage‐based tagging study, we described the genetic mating system of a migratory population of Yellowstone Cutthroat Trout, tested for evidence of sexual selection, and identified predictors of mating and reproductive success. The standardized variance in mating success (i.e., opportunity for sexual selection) was significantly greater for males relative to females, and while the relationship between mating success and reproductive success (i.e., Bateman gradient) was significantly positive for both sexes, a greater proportion of reproductive success was explained by mating success for males (r 2 = 0.80) than females (r 2 = 0.59). Overall, the population displayed a polygynandrous mating system, whereby both sexes experienced variation in mating success due to multiple mating, and sexual selection was variable across sexes. Tests for evidence of sexual selection indicated the interaction between mating success and total length best‐predicted relative reproductive success. We failed to detect a signal of inbreeding avoidance among breeding adults, but the group of parents that produced progeny were on average slightly less related than adults that did not produce progeny. Lastly, we estimated the effective number of breeders (N b) and effective population size (N e) and identified while N b was lower than N e, both are sufficiently high to suggest Yellowstone Cutthroat Trout in Burns Creek represent a genetically stable and diverse population.  相似文献   

16.
Effective population size (Ne) determines the rate of genetic drift and the relative influence of selection over random genetic changes. While free-living protist populations characteristically consist of huge numbers of cells (N), the absence of any estimates of contemporary Ne raises the question whether protist effective population sizes are comparably large. Using microsatellite genotype data of strains derived from revived cysts of the marine dinoflagellate Pentapharsodinium dalei from sections of a sediment record that spanned some 100 years, we present the first estimates of contemporary Ne for a local population in a free-living protist. The estimates of Ne are relatively small, of the order of a few 100 individuals, and thus are similar in magnitude to values of Ne reported for multicellular animals: the implications are that Ne of P. dalei is of many orders of magnitude lower than the number of cells present (Ne/N ∼ 10−12) and that stochastic genetic processes may be more prevalent in protist populations than previously anticipated.  相似文献   

17.
The transition from outcrossing to selfing is predicted to reduce the genome-wide efficacy of selection because of the lower effective population size (Ne) that accompanies this change in mating system. However, strongly recessive deleterious mutations exposed in the homozygous backgrounds of selfers should be under strong purifying selection. Here, we examine estimates of the distribution of fitness effects (DFE) and changes in the magnitude of effective selection coefficients (Nes) acting on mutations during the transition from outcrossing to selfing. Using forward simulations, we investigated the ability of a DFE inference approach to detect the joint influence of mating system and the dominance of deleterious mutations on selection efficacy. We investigated predictions from our simulations in the annual plant Eichhornia paniculata, in which selfing has evolved from outcrossing on multiple occasions. We used range-wide sampling to generate population genomic datasets and identified nonsynonymous and synonymous polymorphisms segregating in outcrossing and selfing populations. We found that the transition to selfing was accompanied by a change in the DFE, with a larger fraction of effectively neutral sites (Nes < 1), a result consistent with the effects of reduced Ne in selfers. Moreover, an increased proportion of sites in selfers were under strong purifying selection (Nes > 100), and simulations suggest that this is due to the exposure of recessive deleterious mutations. We conclude that the transition to selfing has been accompanied by the genome-wide influences of reduced Ne and strong purifying selection against deleterious recessive mutations, an example of purging at the molecular level.  相似文献   

18.
The advantage or disadvantage of sexual reproduction or recombination for the accumulation of mutant genes in a population is studied under the joint effects of recurrent mutations, selection, and random sampling drift. To obtain the rate at which mutant genes are incorporated three different methods are used; numerical integration of Kolmogorov backward equations, simulation of stochastic difference equations, and Monte Carlo experiments. The first two methods are used in a two-locus system to obtain the fixation probability of double mutants and other related quantities under five different selection models. The third one is conducted for a multiple-locus system and the rate of accumulation of mutant genes per locus is studied. Comparison of the results between sexual and asexual populations shows that the effect of recombination depends on initial linkage disequilibrium, mutation rate v, selection intensity s, and population size Ne. The mode of selection is also an important factor and the large effect of recombination is observed when mutant genes are individually deleterious but collectively favorable. Under a given model of selection, the great advantage or disadvantage of recombination is achieved when a large extent of genetic polymorphism is produced not by mutation but by recombination. Extreme values of Nes and Nev make the effect insignificant. The results of Monte Carlo experiments also reveal the presence of interaction between selection and sampling drift even when the loci segregate independently and selection is multiplicative. Although this interaction is usually small, there are cases in which one locus theory cannot be used freely. In those cases, the effect of recombination is prominent and one locus theory gives an overestimate of the rate.  相似文献   

19.
Quantifying interannual variation in effective adult breeding number (Nb) and relationships between Nb, effective population size (Ne), adult census size (N) and population demographic characteristics are important to predict genetic changes in populations of conservation concern. Such relationships are rarely available for long‐lived iteroparous species like lake sturgeon (Acipenser fulvescens). We estimated annual Nb and generational Ne using genotypes from 12 microsatellite loci for lake sturgeon adults (= 796) captured during ten spawning seasons and offspring (= 3925) collected during larval dispersal in a closed population over 8 years. Inbreeding and variance Nb estimated using mean and variance in individual reproductive success derived from genetically identified parentage and using linkage disequilibrium (LD) were similar within and among years (interannual range of Nb across estimators: 41–205). Variance in reproductive success and unequal sex ratios reduced Nb relative to N on average 36.8% and 16.3%, respectively. Interannual variation in Nb/N ratios (0.27–0.86) resulted from stable N and low standardized variance in reproductive success due to high proportions of adults breeding and the species' polygamous mating system, despite a 40‐fold difference in annual larval production across years (437–16 417). Results indicated environmental conditions and features of the species' reproductive ecology interact to affect demographic parameters and Nb/N. Estimates of Ne based on three single‐sample estimators, including LD, approximate Bayesian computation and sibship assignment, were similar to annual estimates of Nb. Findings have important implications concerning applications of genetic monitoring in conservation planning for lake sturgeon and other species with similar life histories and mating systems.  相似文献   

20.
Malaria vectors in sub-Saharan Africa have proven themselves very difficult adversaries in the global struggle against malaria. Decades of anti-vector interventions have yielded mixed results—with successful reductions in transmission in some areas and limited impacts in others. These varying successes can be ascribed to a lack of universally effective vector control tools, as well as the development of insecticide resistance in mosquito populations. Understanding the impact of vector control on mosquito populations is crucial for planning new interventions and evaluating existing ones. However, estimates of population size changes in response to control efforts are often inaccurate because of limitations and biases in collection methods. Attempts to evaluate the impact of vector control on mosquito effective population size (Ne) have produced inconclusive results thus far. Therefore, we obtained data for 13–15 microsatellite markers for more than 1,500 mosquitoes representing multiple time points for seven populations of three important vector species—Anopheles gambiae, An. melas, and An. moucheti—in Equatorial Guinea. These populations were exposed to indoor residual spraying or long-lasting insecticidal nets in recent years. For comparison, we also analyzed data from two populations that have no history of organized vector control. We used Approximate Bayesian Computation to reconstruct their demographic history, allowing us to evaluate the impact of these interventions on the effective population size. In six of the seven study populations, vector control had a dramatic impact on the effective population size, reducing Ne between 55%–87%, the exception being a single An. melas population. In contrast, the two negative control populations did not experience a reduction in effective population size. This study is the first to conclusively link anti-vector intervention programs in Africa to sharply reduced effective population sizes of malaria vectors.  相似文献   

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