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PubMed2000

Effects of population structures and selection strategies on the purging of inbreeding depression due to deleterious mutations.

Stochastic simulations were run to compare the effects of nine breeding schemes, using full-sib mating, on the rate of purging of inbreeding depression due to mutations with equal deleterious effect on viability at unlinked loci in an outbred population. A number of full-sib mating lines were initiated from a large outbred population and maintained for 20 generations (if not extinct). Selection against deleterious mutations was allowed to occur within lines only, between lines or equal within and between lines, and surviving lines were either not crossed or crossed following every one or three generations of full-sib mating. The effectiveness of purging was indicated by the decreased number of lethal equivalents and the increased fitness of the purged population formed from crossing surviving lines after 20 generations under a given breeding scheme. The results show that the effectiveness of purging, the survival of the inbred lines and the inbreeding level attained are generally highest with between-line selection and lowest with within-line selection. Compared with no crossing, line crossing could lower the risk of extinction and the inbreeding coefficient of the purged population substantially with little loss of the effectiveness of purging. Compromising between the effectiveness of purging, and the risk of extinction and inbreeding coefficient, the breeding scheme with equal within- and between-line selection and crossing alternatively with full-sib mating is generally the most desirable scheme for purging deleterious mutations. Unless most deleterious mutations have relatively large effects on fitness in species with reproductive ability high enough to cope with the depressed fitness and thus increased risk of extinction with inbreeding, it is not justified to apply a breeding programme aimed at purging inbreeding depression by inbreeding and selection to a population of conservation concern.

Alleles↗
PubMed2005

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.

Analysis of Variance↗
PubMed2005

Environmental dependence of inbreeding depression in cultured Coho salmon (Oncorhynchus kisutch): aggressiveness, dominance and intraspecific competition.

We evaluated the effects of inbreeding on traits related to territorial dominance and tested whether the magnitude of inbreeding depression (ID) was modified by social environment in Coho salmon. Evaluation of behaviour in paired contests between juvenile salmon with different inbreeding (low, LI=9.5%; medium, MI=29.6%), did not show significant differences between their capacities for establishing territorial dominance (mean aggressiveness score, LI=20.0+/-22; MI=16.7+/-23 or for feeding attempts, LI=18.3+/-12; MI=21.1+/-12). However, fish with low inbreeding (LI) showed almost twice the aggressive pursuit of fish with medium inbreeding (MI), and had a higher specific growth rate (SGR) in culture (SGR(MI)=1.83+/-0.58; SGR(LI)=2.20+/-0.67). Additionally, we found evidence that the magnitude of ID was modified by social environment: (1) Masking: In small groups of fish (N=20), large dominant fish of MI, cultivated with small subordinate fish of LI, showed the same SGR as dominant fish of LI cultivated with small subordinate fish of MI. (2) Magnifying: A significant effect of ID on juvenile survival was detected only in high-density competitive environments. Thus, the number of lethal equivalents was 2.70 at high-density, and only 0.24 in a low-density environment. Our results show that differences in size associated with territorial dominance may mask deleterious effects of inbreeding under certain conditions, and support the concept that intraspecific competition usually magnifies the deleterious effects of inbreeding.

Animals↗
PubMed2006

Marker-based investigation of inbreeding depression in the endangered species Brassica insularis.

Various methods have been proposed to estimate inbreeding depression and to assess its consequences for natural populations. As an alternative to controlled crosses, the use of molecular markers has allowed direct investigation of inbreeding depression in natural populations, but usually suffers from low statistical power. Here, we investigated the effect of inbreeding depression on survival in two populations of the rare species Brassica insularis, using both controlled crosses and a marker-based approach. We compare the respective merits of the two approaches for studying inbreeding depression. We also use information from the molecular markers to dissect in detail patterns of inbreeding depression in this species. A posteriori, we find that combining the approaches was not necessary to obtain simple point estimates of inbreeding depression. However, using molecular markers may give insight into the genetic basis of inbreeding depression, such as the occurrence of epistatic interactions among deleterious alleles or purging.

Alleles↗
PubMed2006

An investigation of inbreeding depression and purging in captive pedigreed populations.

We use regression models to investigate the effects of inbreeding in 119 zoo populations, encompassing 88 species of mammals, birds, reptiles and amphibians. Meta-analyses show that inbreeding depression for neonatal survival was significant across the 119 populations although the severity of inbreeding depression appears to vary among taxa. However, few predictors of a population's response to inbreeding are found reliable. The models are most likely to detect inbreeding depression in large populations, that is, in populations in which their statistical power is maximised. Purging was found to be significant in 14 populations and a significant trend of purging was found across populations. The change in inbreeding depression due to purging averaged across the 119 populations is <1%, however, suggesting that the fitness benefits of purging are rarely appreciable. The study re-emphasises the necessity to avoid inbreeding in captive breeding programmes and shows that purging cannot be relied upon to remove deleterious alleles from zoo populations.

Amphibians↗
PubMed1998

Allozyme diversity in Leavenworthia populations with different inbreeding levels.

Levels of allozyme diversity were studied in five species of the annual plant genus Leavenworthia. The breeding systems of the populations were also characterized in terms of measures of autogamous seed-set and self-fertility. The populations that appeared more inbreeding by these criteria also tended to have lower within-population gene diversity (Hs) values, but there was nevertheless considerable allozyme variability in two of the four sets of inbred populations studied. Comparing the outcrosser L. stylosa and the related inbreeders L. uniflora and L. torulosa the reduction in diversity associated with inbreeding was very high, as no variants were seen within populations of the inbreeders. In L. crassa and L. alabamica, the reduction was lower, but correcting for the fact that this comparison is over less extreme selfing rate differences, it is estimated that the difference between extreme inbreeding and outcrossing populations of L. crassa considerably exceeds twofold. These results are compared with published studies within closely related taxa with different inbreeding levels. The effect of inbreeding on allozyme diversity is consistently larger than has been estimated from comparisons of unrelated species.

Brassicaceae↗
PubMed2003

Environment-dependent inbreeding depression in a hermaphroditic freshwater snail.

Inbreeding depression was simultaneously studied under contrasted environments, laboratory and natural conditions, using individuals originating from 14 families of the freshwater snail Physa acuta. Both survival and growth of juveniles showed inbreeding depression under laboratory conditions. The same fitness components were monitored with mature snails either kept under laboratory conditions or released at a natural site and analysed using capture-mark-recapture models. Genetic composition of both samples was similar. Inbreeding depression on survival was highest in the laboratory while strong outbreeding depression was revealed in the field. Thus inbreeding depression may not be always higher under natural conditions, at the opposite of what is commonly assumed. We suggest that inbreeding depression is dependent on metabolic requirements imposed by the environment. Other evidences showing that inbreeding depression is environment-dependent are reviewed. We conclude that genetic models should include both genetic and environmental variance in inbreeding depression for studying mating system evolution.

Adaptation, Physiological↗
PubMed1999

Inbreeding of bottlenecked butterfly populations. Estimation using the likelihood of changes in marker allele frequencies.

Polymorphic enzyme and minisatellite loci were used to estimate the degree of inbreeding in experimentally bottlenecked populations of the butterfly, Bicyclus anynana (Satyridae), three generations after founding events of 2, 6, 20, or 300 individuals, each bottleneck size being replicated at least four times. Heterozygosity fell more than expected, though not significantly so, but this traditional measure of the degree of inbreeding did not make full use of the information from genetic markers. It proved more informative to estimate directly the probability distribution of a measure of inbreeding, sigma2, the variance in the number of descendants left per gene. In all bottlenecked lines, sigma2 was significantly larger than in control lines (300 founders). We demonstrate that this excess inbreeding was brought about both by an increase in the variance of reproductive success of individuals, but also by another process. We argue that in bottlenecked lines linkage disequilibrium generated by the small number of haplotypes passing through the bottleneck resulted in hitchhiking of particular marker alleles with those haplotypes favored by selection. In control lines, linkage disequilibrium was minimal. Our result, indicating more inbreeding than expected from demographic parameters, contrasts with the findings of previous (Drosophila) experiments in which the decline in observed heterozygosity was slower than expected and attributed to associative overdominance. The different outcomes may both be explained as a consequence of linkage disequilibrium under different regimes of inbreeding. The likelihood-based method to estimate inbreeding should be of wide applicability. It was, for example, able to resolve small differences in sigma2 among replicate lines within bottleneck-size treatments, which could be related to the observed variation in reproductive viability.

Alleles↗
PubMed2001

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.

Alleles↗
PubMed2006

Inbreeding ancestors: the role of sibmating in the social evolution of gall thrips.

We used microsatellite data to estimate levels of inbreeding in four species of solitary gall thrips that are in the same clade as the six species with soldier castes. Three of the four species were highly inbred (Fis 0.54-0.68), and the other apparently mated randomly (Fis near zero). These estimates, combined with previous data from species with soldiers, suggest that inbreeding is a pervasive life-history feature of the gall-inducing thrips on Australian Acacia. Mapping of inbreeding estimates onto the phylogeny of the gall inducers showed that the ancestral lineage that gave rise to soldiers was apparently highly inbred, and therefore, inbreeding could have played a role in the origin of sociality within this group. Moreover, there was a trend from high levels of inbreeding at the origin of soldiers to low levels in the most derived species with soldiers, which exhibits the highest levels of reproductive division of labor and soldier altruism. These patterns are consistent with considerations from population genetics, which show that the likelihood of the origin of soldier altruism is higher in inbreeding populations but that, once soldiers have evolved, a reduction in inbreeding levels may facilitate the evolution of enhanced division of labor and reproductive skew.

Acacia↗
PubMed2006

Inbreeding linked to amphibian survival in the wild but not in the laboratory.

We examined the effects of inbreeding on the performance of wood frog (Rana sylvatica) larvae in the field and in the laboratory. We used microsatellite analysis to establish the parentage and degree of inbreeding of the larvae. Two different estimators of inbreeding were used. The first was based on average multilocus heterozygosity, and the second was based on a molecular relatedness estimator. The estimators were highly correlated, and both showed a significant negative relationship between inbreeding and survival in the wild. However, there was no evidence that inbreeding influenced growth or development in the wild. Neither was there any evidence that inbreeding affected survival, growth, or development in the laboratory. These results suggest that, for wood frogs, inbreeding has a bigger effect on fitness in the wild than in captivity and that measurements of survival are more sensitive than measures of growth or development.

Animals↗
PubMed1999

The variance in inbreeding depression and the recovery of fitness in bottlenecked populations.

Theoretical analyses of inbreeding suggest that following an increased degree of inbreeding there may be a temporary recovery of fitness, because of selection either within or among inbred lineages. This is possible because selection can act more efficiently to remove deleterious alleles given the greater homozygosity of such populations. If common, recovery of fitness following inbreeding may be important for understanding some evolutionary processes and for management strategies of remnant populations, yet empirical evidence for such recovery in animals is scant. Here we describe the effects of single-pair population bottlenecks on a measure of fitness in Drosophila melanogaster. We compared a large number of families from each of 52 inbred lines with many families from the outbred population from which the inbred lineages were derived. Measures were made at the third and the 20th generations after the bottleneck. In both generations there was, on average, substantial inbreeding depression together with a highly significant variance among the inbred lines in the amount of fitness reduction. The average fitness of inbred lines was correlated across generations. Our data provide evidence for the possibility of recovery of fitness at two levels, because (i) the average fitness reduction in the F20 generation was significantly less than in the F3 generation, which implies that selection within lines has occurred, and (ii) the large variance in inbreeding depression among inbred lines implies that selection among them is possible. The high variance in inbreeding depression among replicate lines implies that modes of evolution which require a low level of inbreeding depression can function at least in a fraction of inbred populations within a species and that results from studies with low levels of replication should be treated with caution.

Animals↗
PubMed2002

Anther-stigma separation is associated with inbreeding depression in Datura stramonium, a predominantly self-fertilizing annual.

Genetically based variation in outcrossing rate generates lineages within populations that differ in their history of inbreeding. According to some models, mating-system modifiers in such populations will demonstrate both linkage and identity disequilibrium with fitness loci, resulting in lineage-specific inbreeding depression. Other models assert that differences among families in levels of inbreeding depression are mainly attributable to random accumulation of genetic load, unrelated to variation at mating-system loci. We measured female reproductive success of selfed and outcrossed progeny from naturally occurring lineages of Datura stramonium, a predominantly self-fertilizing annual weed that has heritable variation in stigma-anther separation, a trait that influences selfing rates. Progeny from inbred lineages (as identified by high degree of anther-stigma overlap) showed equal levels of seed production, regardless of cross type. Progeny from mixed lineages (as identified by relatively high separation between anthers and stigma) showed moderate levels of inbreeding depression. We found a significant correlation between anther-stigma separation and relative fitness of selfed and outcrossed progeny, suggesting that family-level inbreeding depression may be related to differences among lineages in inbreeding history in this population. Negative inbreeding depression in putatively inbred lineages may be due in part to additive effects or to epistatic interactions among loci.

Datura stramonium↗
PubMed2003

Experimental analysis of biparental inbreeding in a self-fertilizing plant.

Localized dispersal and mating may genetically structure plant populations, resulting in matings among related individuals. This biparental inbreeding has significant consequences for the evolution of mating systems, yet is difficult to estimate in natural populations. We estimated biparental inbreeding in two populations of the largely self-fertilizing plant Aquilegia canadensis using standard inference as well as a novel experiment comparing apparent selfing between plants that were randomly relocated within populations to experimental control plants. Using two allozyme markers, biparental inbreeding (b) inferred from the difference between single-locus and multilocus estimates of selfing (b = s(s) - s(m)) was low. Less than 3% of matings involved close relatives (mean b = 0.029). In contrast, randomly relocating plants greatly reduced apparent selfing (mean s(s) = 0.674) compared to control plants that had been dug up and replanted in their original locations (s(s) = 0.953, P = 0.002). Based on this difference in s(s), we estimated that approximately 30% of all matings involved close relatives (mean b = 0.279, 95% CL = 0.072-0.428). Inference from s(s) - s(m) underestimated b in these populations by more than an order of magnitude. Biparental inbreeding is thought to influence the evolution of self-fertilization primarily through reducing the genetic cost of outcrossing. This is unlikely to be of much significance in A. canadensis because inbreeding depression (a major cost of selfing) is much stronger than the cost of outcrossing. However, biparental inbreeding combined with strong inbreeding depression may influence selection on dispersal.

Aquilegia↗
PubMed2006

Variation in the intensity of inbreeding depression among successive life-cycle stages and generations in gynodioecious Silene vulgaris (Caryophyllaceae).

Inbreeding depression is one of the hypotheses explaining the maintenance of females within gynodioecious plant populations. However, the measurement of fitness components in selfed and outcrossed progeny depends on life-cycle stage and the history of inbreeding. Comparative data indicate that strong inbreeding depression is more likely to occur at later life-cycle stages. We used hermaphrodite individuals of Silene vulgaris originating from three populations located in different valleys in the Swiss Alps to investigate the effect of two generations of self- and cross-fertilization on fitness components among successive stages of the life cycle in a glasshouse experiment. We detected significant inbreeding depression for most life-cycle stages including: the number of viable and aborted seeds per fruit, probability of germination, above ground biomass, probability of flowering, number of flowers per plant, flower size and pollen viability. Overall, the intensity of inbreeding depression increased among successive stages of the life cycle and cumulative inbreeding depression was significantly stronger in the first generation (delta approximately 0.5) compared with the second generation (delta approximately 0.35). We found no evidence for synergistic epistasis in our experiment. Our finding of more intense inbreeding depression during later stages of the life cycle may help to explain the maintenance of females in gynodioecious populations of S. vulgaris because purging of genetic load is less likely to occur.

Crosses, Genetic↗
PubMedDate not supplied

The effect of fast created inbreeding on litter size and body weights in mice.

This study was designed to reveal any differences in effects of fast created versus total inbreeding on reproduction and body weights in mice. A line selected for large litter size for 124 generations (H) and a control line (K) maintained without selection for the same number of generations were crossed (HK) and used as a basis for the experiment. Within the HK cross, full sib, cousin or random mating were practised for two generations in order to create new inbreeding (IB(F)) at a fast rate. In the first generation of systematic mating, old inbreeding was regenerated in addition to creation of new inbreeding from the mating design giving total inbreeding (IB(T)). The number of pups born alive (NBA) and body weights of the animals were then analysed by a model including both IB(T) and IB(F). The IB(T) of the dam was in the present study found to reduce the mean NBA with -0.48 (+/-0.22) (p < 0.05) pups per 10% increase in the inbreeding coefficient, while the additional effect of IB(F) was -0.42 (+/-0.27). For the trait NBA per female mated, the effect of IB(T) was estimated to be -0.45 (+/-0.29) per 10% increase in the inbreeding coefficient and the effect of IB(F) was -0.90 (+/-0.37) (p < 0.05) pups. In the present study, only small or non-significant effects of IB(F) of the dam could be found on sex-ratio and body weights at three and six weeks of age in a population already adjusted for IB(T).

Animals↗
PubMed2005

A genetic interpretation of the variation in inbreeding depression.

Inbreeding depression is expected to play an important but complicated role in evolution. If we are to understand the evolution of inbreeding depression (i.e., purging), we need quantitative genetic interpretations of its variation. We introduce an experimental design in which sires are mated to multiple dams, some of which are unrelated to the sire but others are genetically related owing to an arbitrary number of prior generations of selfing or sib-mating. In this way we introduce the concept of "inbreeding depression effect variance," a parameter more relevant to selection and the purging of inbreeding depression than previous measures. We develop an approach for interpreting the genetic basis of the variation in inbreeding depression by: (1) predicting the variation in inbreeding depression given arbitrary initial genetic variance and (2) estimating genetic variance components given half-sib covariances estimated by our experimental design. As quantitative predictions of selection depend upon understanding genetic variation, our approach reveals the important difference between how inbreeding depression is measured experimentally and how it is viewed by selection.

Analysis of Variance↗
PubMed2005

Genome-wide analysis on inbreeding effects on gene expression in Drosophila melanogaster.

The deleterious consequences of inbreeding, especially in the form of inbreeding depression, are well known. However, little is known about how inbreeding affects genome-wide gene expression. Here, we show that inbreeding changes transcription levels for a number of genes. Gene expression profiles of Drosophila melanogaster lines inbred to F approximately = 0.67 at different rates changed relative to those of noninbred lines, but the rate of inbreeding did not significantly affect gene expression patterns. Genes being differentially expressed with inbreeding are disproportionately involved in metabolism and stress responses, suggesting that inbreeding acts like an environmental stress factor.

Animals↗