Within-population craniometric variability of insular populations of deer mice, Peromyscus maniculatus, elucidated by landscape configuration

Within-population genetic variability of twelve insular and four mainland populations of deer mice ( Peromyscus maniculatus ) was assessed using craniometric characters, and compared to results previously obtained from RAPD data. An index of Craniometric Variance ( CVar ) was computed from pairwise distances among all specimens. Variations in CVar measures were then compared to landscape variables using a linear regression approach. Our results suggest that CVar decreases in presence of large number of a competitive species (the boreal redback vole, Clethrionomys gapperi ; r =−0.527, p <0.037) in deer mouse populations. Island remoteness ( r =−0.251, p <0.220) and the geometry of the bank opposite to each island ( r =−0.459, p <0.067) were marginally correlated with CVar , but the linear combination of these two variables, forming a composite isolation index, represented the major factor explaining the observed CVar ( r =−0.648, p <0.011). Using a multiple regression model, 76.3% of the CVar was explained by a combination of this isolation index and the competitors' abundance. These results suggest that taking into account landscape barriers as well as the dispersal behavior of small mammals might provide sounder ecological variables than geographical distances alone for predicting within-population genetic variability in a network of habitat patches.     

Inbreeding depression in self-incompatible and self-compatible populations of Leavenworthia alabamica

Inbreeding depression is one of the leading factors preventing the evolution of self-fertilization in plants. In populations where self-fertilization evolves, theory suggests that natural selection against partially recessive deleterious alleles will reduce inbreeding depression. The purpose of this study was to evaluate this hypothesis by comparing the magnitude of inbreeding depression in self-incompatible and self-compatible populations of Leavenworthia alabamica. Within-population crosses were conducted to compare the quantity and quality of offspring produced by outcrossing and self-fertilization. These progeny were grown in a common greenhouse and inbreeding depression was measured in germination, survival, biomass, transition rate to flowering, flower number, petal length, pollen grains/anther, pollen viability, and ovule number. In comparison to outcrossing, self-fertilization led to the production of fewer and smaller seeds within self-incompatible populations. Moreover, inbreeding depression was observed in eight of 11 offspring traits within self-incompatible populations of L. alabamica. In contrast, there was significant inbreeding depression only in flower number within self-compatible populations. The results of this study are consistent with the idea that self-fertilization selectively removes partially recessive deleterious alleles causing inbreeding depression in natural plant populations. However, in plant species such as L. alabamica where self-compatibility may evolve in small populations following long-distance dispersal, declines in inbreeding depression may also be facilitated by genetic drift.     

Inbreeding depression in small populations of self-incompatible plants.

Self-incompatibility (SI) is a widespread mechanism that prevents inbreeding in flowering plants. In many species, SI is controlled by a single locus (the S locus) where numerous alleles are maintained by negative frequency-dependent selection. Inbreeding depression, the decline in fitness of selfed individuals compared to outcrossed ones, is an essential factor in the evolution of SI systems. Conversely, breeding systems influence levels of inbreeding depression. Little is known about the joint effect of SI and drift on inbreeding depression. Here we studied, using a two-locus model, the effect of SI (frequency-dependent selection) on a locus subject to recurrent deleterious mutations causing inbreeding depression. Simulations were performed to assess the effect of population size and linkage between the two loci on the level of inbreeding depression and genetic load. We show that the sheltering of deleterious alleles linked to the S locus strengthens inbreeding depression in small populations. We discuss the implications of our results for the evolution of SI systems.     

Inbreeding depression and drift load in small populations at demographic disequilibrium

Inbreeding depression is a major driver of mating system evolution and has critical implications for population viability. Theoretical and empirical attention has been paid to predicting how inbreeding depression varies with population size. Lower inbreeding depression is predicted in small populations at equilibrium, primarily due to higher inbreeding rates facilitating purging and/or fixation of deleterious alleles (drift load), but predictions at demographic and genetic disequilibrium are less clear. In this study, we experimentally evaluate how lifetime inbreeding depression and drift load, estimated by heterosis, vary with census (N_c) and effective (estimated as genetic diversity, H_e) population size across six populations of the biennial Sabatia angularis as well as present novel models of inbreeding depression and heterosis under varying demographic scenarios at disequilibrium (fragmentation, bottlenecks, disturbances). Our experimental study reveals high average inbreeding depression and heterosis across populations. Across our small sample, heterosis declined with H_e, as predicted, whereas inbreeding depression did not vary with H_e and actually decreased with N_c. Our theoretical results demonstrate that inbreeding depression and heterosis levels can vary widely across populations at disequilibrium despite similar H_e and highlight that joint demographic and genetic dynamics are key to predicting patterns of genetic load in nonequilibrium systems.     

The genetic heterogeneity of deer mouse populations (Peromyscus maniculatus) in an insular landscape

A survey of the genetic variability in deer mouse populations was performed using specimens collected from six different islands on a lake covering approximately 50 km². Random amplified polymorphic DNA (RAPD) was used to measure the extent of the genetic differences in this insular system. An analysis of molecular variance (AMOVA) revealed that populations are clearly separated at this microgeographic scale (F _st = 0.13863; P < 0.001). The homogeneity of molecular variance test (HOMOVA) indicated that within-population levels vary greatly (B _p = 0.76831; P < 0.001). The within-population molecular variance was found to be mainly correlated with the accessibility of the islands, computed as the inverse of the geographic distance separating an island from the lakeshore (r = 0.916; P < 0.003). Received: March 5, 1999 / Accepted: July 16, 1999     

Inbreeding depression due to overdominance in partially self-fertilizing plant populations

The effect of the rate of partial self-fertilization and viability selection on the magnitude of inbreeding depression was investigated for the overdominance genetic model. The influence of these factors was determined for populations with equilibrium genotypic frequencies. Inbreeding depression was measured as the normalized disadvantage in mean viability of selfed progeny as compared to outcrossed progeny. When caused by symmetric homozygous disadvantage at a single locus it is shown always to be less than one-third. Moreover, for fixed rates of self-fertilization, its maximum value is found at intermediate levels of homozygous disadvantage. As the rate of self-fertilization increases, inbreeding depression increases and the homozygote viability that results in maximum depression tends toward one-half the heterozygote viability. Symmetric selection against homozygotes at multiple loci can lead to substantially higher values than selection at a single-locus. As the number of independent loci involved increases, inbreeding depression can reach high levels even though the selfing rate is low. Viability distributions for progenies produced from both random mating and self-fertilization were derived for the case of symmetric selection at independently assorting multiple loci. Distributions of viabilities in progenies resulting from mixtures of selfing and outcrossing were shown to be bimodal when inbreeding depression is high.     

Genetic divergence of insular populations of deer mice

Electrophoretic variants at 28 genetic loci were analyzed in subspecies of Peromyscus maniculatus endemic to the Channel Islands off the California coast. The genetic variability within insular populations was calculated. These deer mice have relatively high levels of polymorphism for insular populations. The mean heterozygosity per individual varies in the populations, being much higher on one of the islands than the others. Nei's measure of genetic distance between groups compared on the basis of electrophoretic variants was used. His estimate of time of divergence of these groups, based on genetic distance, is applicable particularly to closely related groups. The length of time each island population has been isolated from the others was calculated and found to be consistent with the periods of isolation estimated on the basis of geological data.     

Inbreeding depression and heterosis in populations of Schiedea viscosa, a highly selfing species

Progeny from self-pollinations and intrapopulation crosses were examined in Schiedea viscosa to determine the extent of inbreeding depression in this highly selfing species. Progeny of interpopulation crosses were also investigated to determine whether deleterious alleles have been fixed in populations of S. viscosa. There was no inbreeding depression at early life history stages, including seeds per capsule, seed mass, or germination. Inbreeding depression was detected for the later life history stage of fruit biomass, although not for survival or life span. Heterosis for vegetative biomass and fruit biomass was detected in progeny from crosses between populations. Levels of inbreeding depression in S. viscosa are low relative to out-crossing species of Schiedea, especially when early life history stages are compared.     

Inbreeding depression and mixed mating in Leptosiphon jepsonii: a comparison of three populations

BACKGROUND AND AIMS: Inbreeding depression is thought to play a central role in the evolution and maintenance of cross-fertilization. Theory indicates that inbreeding depression can be purged with self-fertilization, resulting in positive feedback for the selection of selfing. Variation among populations of Leptosiphon jepsonii in the timing and rate of self-fertilization provides an opportunity to study the evolution of inbreeding depression and mating systems. In addition, the hypothesis that differences in inbreeding depression for male and female fitness can stabilize mixed mating in L. jepsonii is tested. METHODS: In a growth room experiment, inbreeding depression was measured in three populations with mean outcrossing rates ranging from 0.06 to 0.69. The performance of selfed and outcrossed progeny is compared at five life history stages. To distinguish between self-incompatibility and early inbreeding depression, aborted seeds and unfertilized ovules were counted in selfed and outcrossed fruits. In one population, pollen and ovule production was quantified to estimate inbreeding depression for male and female fitness. KEY RESULTS: Both prezygotic barriers and inbreeding depression limited self seed set in the most outcrossing population. Cumulative inbreeding depression ranged from 0.297 to 0.501, with the lowest value found in the most selfing population. Significant inbreeding depression for early life stages was found only in the more outcrossing populations. Inbreeding depression was not significant for pollen or ovule production. CONCLUSIONS: The results provide modest support for the hypothesized relationship between inbreeding depression and mating systems. The absence of early inbreeding depression in the more selfing populations is consistent with theory on purging. Differences in male and female expression of inbreeding depression do not appear to stabilize mixed mating in L. jepsonii. The current estimates of inbreeding depression for L. jepsonii differ from those of previous studies, underscoring the effects of environmental variation on its expression.     

Age structure of six populations of old-field mice,Peromyscus polionotus

An accurate age-estimating technique, based on biochemical changes in eye lens protein, was used to study age structures of six populations of the old-field mouse, Peromyscus polionotus. A new mathematical procedure permitted quantitative comparisons of these populations. Four inland populations had essentially the same median ages (75–84 days), maximum ages (248–297 days) and relative production rates (56–58% of the surviving population had been born in the 100 days prior to sampling). Approximately 50% of the females were old enough to have weaned a litter. One inland population had a lower median age (49 days), a lower maximum age (181 days) and a higher relative production rate (73%). Relatively fewer females (38%) could have weaned a litter. The single beach population was also different, with higher median age (180 days), and lower relative production rate (2%). All females were old enough to have weaned a litter. Maximum age was essentially the same as for the four similar, inland populations. Factors which may have contributed to observed differences in these populations are discussed.     

Effects of forest edge on populations of white-footed mice Peromyscus leucopus

Several studies have reported higher densities of white-footed mice in small fragments than in large fragments of eastern deciduous forests. The edge hypothesis states that higher densities in smaller fragments reflect an increase in relative amount of edge habitat, which supports higher densities of mice because of its higher quality. To test this hypothesis we live trapped white-footed mice along edge-to-interior gradients in forest fragments of east-central Illinois. Our results indicated a greater abundance of mice in the forest interior than near the edge, which did not support the edge hypothesis. This pattern could occur because dominant adults hold larger territories of higher quality habitat, thereby reducing density and increasing fitness near the edge (an ideal despotic distribution). We found some evidence of increased reproductive success (juveniles per female) at the edge, but this could reflect density-dependent demographic processes rather than habitat quality. Furthermore, other indicators of dominance (body weight, and reproductive activity) did not show an increase at the edge, and other studies indicate higher prevalence of natural enemies at edges, which could account for lower densities there. Reduced competition from larger rodents and reduced predation could cause higher densities in small fragments but the distributions of competitors and predators do not strongly support these hypotheses. We suggest two additional hypotheses that could account for greater densities in smaller fragments: 1) estimates of high densities could be artifacts of the large effect that a few captures can have on density estimates for very small fragments, and 2) densities in smaller fragments are overestimated because mice use a relatively larger area of surrounding habitat as fragment size decreases.     

Inbreeding and outbreeding depression in Daphnia

Egg-to-adult viability of sexual offspring in Daphnia magna is lower for selfed (average: 43.0%) than for outcrossed families (average: 74.7%). This suggests that intraclonal mating is not the rule in Daphnia populations. For a given family, hatching rate of eggs resulting from interpopulation crosses is lower than for intrapopulation crosses. This breakdown in hatching responses may result in the effective gene flow between Daphnia populations being severely reduced, offering an explanation for the apparent paradox of genetic differentiation of Daphnia populations in spite of efficient dispersal.     

Inbreeding depression and low between-population heterosis in recently diverged experimental populations of a selfing species

In fragmented populations, genetic drift and selection reduce genetic diversity, which in turn results in a loss of fitness or in a loss of evolvability. Genetic rescue, that is, controlled input of diversity from distant populations, may restore evolutionary potential, whereas outbreeding depression might counteract the positive effect of this strategy. We carried out self-pollination and crosses within and between populations in an experimental subdivided population of a selfing species, Triticum aestivum L., to estimate the magnitude of these two phenomena. Surprisingly, for a self-fertilizing species, we found significant inbreeding depression within each population for four of the six traits studied, indicating that mildly deleterious mutations were still segregating in these populations. The progeny of within- and between-population crosses was very similar, indicating low between-population heterosis and little outbreeding depression. We conclude that relatively large population effective sizes prevented fixation of a high genetic load and that local adaptation was limited in these recently diverged populations. The kinship coefficient estimated between the parents using 20 neutral markers was a poor predictor of the progeny phenotypic values, indicating that there was a weak link between neutral diversity and genes controlling fitness-related traits. These results show that when assessing the viability of natural populations and the need for genetic rescue, the use of neutral markers should be complemented with information about the presence of local adaptation in the subdivided population.     

Inbreeding depression in monarch butterflies

Monarch butterflies and their unique system of multigenerational migration have long fascinated the public, and concerns for the fate of this charismatic insect have grown due to the consistent declines in overwintering colony size over the last 20 years. Risks to this migratory insect have been considered in terms of climate change, habitat and thus population fragmentation, and decreased host plant availability. However, another obvious danger, that of decreased heterozygosity resulting from decreasing population size, has yet to be explored. Here we report experimental evidence for immediate inbreeding depression in individuals from the migratory population. Inbred matings produced less viable eggs and inbred offspring had higher developmental mortality and shorter lifespans. We discuss these results in the context of monarch migration extinction risk and suggest that additional genetic monitoring should be undertaken to protect this iconic animal.     

Inbreeding depression due to mildly deleterious mutations in finite populations: size does matter

We studied the effects of population size on the inbreeding depression and genetic load caused by deleterious mutations at a single locus. Analysis shows how the inbreeding depression decreases as population size becomes smaller and/or the rate of inbreeding increases. This pattern contrasts with that for the load, which increases as population size becomes smaller but decreases as inbreeding rate goes up. The depression and load both approach asymptotic limits when the population size becomes very large or very small. Numerical results show that the transition between the small and the large population regimes is quite rapid, and occurs largely over a range of population sizes that vary by a factor of 10. The effects of drift on inbreeding depression may bias some estimates of the genomic rate of deleterious mutation. These effects could also be important in the evolution of breeding systems in hermaphroditic organisms and in the conservation of endangered populations.     

Inbreeding and outbreeding depression in Caenorhabditis nematodes

The nematode Caenorhabditis elegans reproduces primarily by self-fertilization of hermaphrodites, yet males are present at low frequencies in natural populations (androdioecy). The ancestral state of C. elegans was probably gonochorism (separate males and females), as in its relative C. remanei. Males may be maintained in C. elegans because outcrossed individuals escape inbreeding depression. The level of inbreeding depression is, however, expected to be low in such a highly selfing species, compared with an outcrosser like C. remanei. To investigate these issues, we measured life-history traits in the progeny of inbred versus outcrossed C. elegans and C. remanei individuals derived from recently isolated natural populations. In addition, we maintained inbred lines of C. remanei through 13 generations of full-sibling mating. Highly inbred C. remanei showed dramatic reductions in brood size and relative fitness compared to outcrossed individuals, with evidence of both direct genetic and maternal-effect inbreeding depression. This decline in fitness accumulated over time, causing extinction of nearly 90% of inbred lines, with no evidence of purging of deleterious mutations from the remaining lines. In contrast, pure strains of C. elegans performed better than crosses between strains, indicating outbreeding depression. The results are discussed in relation to the evolution of androdioecy and the effect of mating system on the level of inbreeding depression.     

Inbreeding depression in benign and stressful environments

Understanding the consequences of inbreeding has important implications for a wide variety of topics in population biology. Although it is often stated in the literature that the deleterious effects of inbreeding (inbreeding depression) are expected to be more pronounced under stressful than benign conditions, this issue remains unresolved and controversial. We review the current literature on the relationship between the magnitude of inbreeding depression and environmental stress and calculate haploid lethal equivalents expressed under relatively benign and stressful conditions based on data from 34 studies. Inbreeding depression increases under stress in 76% of cases, although this increase is only significant in 48% of the studies considered. Estimates of lethal equivalents are significantly greater under stressful (mean = 1.45, median = 1.02) than relatively benign (mean = 0.85, median = 0.61) conditions. This amounts to an approximately 69% increase in inbreeding depression in a stressful vs a benign environment. However, we find strong lineage effects to be ubiquitous among studies that examine inbreeding depression in multiple environments, and a prevalence of conditionally expressed deleterious effects within lineages that are uncorrelated across environments. These results have important implications for both evolutionary and conservation biology.     

Inbreeding in populations with overlapping generations

An inbreeding matrix is defined for populations with overlapping generations. In the short term it can be expressed in terms of a matrix specifying the passage of genes between the different age groups (and sexes) and a diagonal matrix whose elements depend on the number of individuals in each age group. Formulae for the inbreeding effective number are derived using matrix theory. A comparison is made between the inbreeding coefficients predicted by this theory and those obtained by assuming a uniform rate of inbreeding from the outset, and these in turn are compared with the exact inbreeding coefficients.     

Inbreeding depression in male gametic performance

One key objective in evolutionary ecology is to understand the magnitude of inbreeding depression expressed across sex‐specific components of fitness. One major component of male fitness is fertilization success, which depends on male gametic performance (sperm and pollen performance in animals and plants, respectively). Inbreeding depression in male gametic performance could create sex‐specific inbreeding depression in fitness, increase the benefit of inbreeding avoidance and reduce the efficacy of artificial insemination and pollination. However, there has been no assessment of the degree to which inbreeding generally depresses male gametic performance and hence post‐copulatory or post‐pollination fertilization success. Because inbreeding depression is understood to be a property of diploid entities, it is not clear what degree of inbreeding depression in haploid gametic performance should be expected. Here, we first summarize how inbreeding depression in male gametic performance could potentially arise through gene expression in associated diploid cells and/or reduced genetic diversity among haploid gametes. We then review published studies that estimate the magnitude of inbreeding depression in traits measuring components of sperm or pollen quantity, quality and competitiveness. Across 51 published studies covering 183 study traits, the grand mean inbreeding load was approximately one haploid lethal equivalent, suggesting that inbreeding depresses male gametic performance across diverse systems and traits. However, there was an almost complete lack of explicit estimates from wild populations. Future studies should quantify inbreeding depression in systematic sets of gametic traits under naturally competitive and noncompetitive conditions and quantify the degree to which gamete phenotypes and performance reflect haploid vs. diploid gene expression.