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PubMed2006

Ancestral inbreeding reduces the magnitude of inbreeding depression in Drosophila melanogaster.

The influence of natural selection on the magnitude of inbreeding depression is an important issue in conservation biology and the study of evolution. It is generally expected that the magnitude of inbreeding depression in small populations will depend upon the average homozygosity of individuals, as measured by the coefficient of inbreeding (F). However, if deleterious recessive alleles are selectively purged from populations during inbreeding, then inbreeding depression may differ among populations in which individuals have the same inbreeding coefficient. In such cases, the magnitude of inbreeding depression will partly depend on the ancestral inbreeding coefficient (fa), which measures the cumulative proportion of loci that have historically been homozygous and therefore exposed to natural selection. We examined the inbreeding depression that occurred in lineages of Drosophila melanogaster maintained under pedigrees that led to the same inbreeding coefficient (F = 0.375) but different levels of ancestral inbreeding (fa = 0.250 or 0.531). Although inbreeding depression varied substantially among individual lineages, we observed a significant 40% decrease in the median level of inbreeding depression in the treatment with higher ancestral inbreeding. Our results demonstrate that high levels of ancestral inbreeding are associated with greater purging effects, which reduces the inbreeding depression that occurs in isolated populations of small size.

Alleles↗

PubMed2000

Whole-genome characterization of embryonic stage inbreeding depression in a selfed loblolly pine family.

Inbreeding depression is important in the evolution of plant populations and mating systems. Previous studies have suggested that early-acting inbreeding depression in plants is primarily due to lethal alleles and possibly epistatic interactions. Recent advances in molecular markers now make genetic mapping a powerful tool to study the genetic architecture of inbreeding depression. We describe a genome-wide evaluation of embryonic viability loci in a selfed family of loblolly pine (Pinus taeda L.), using data from AFLP markers from an essentially complete genome map. Locus positions and effects were estimated from segregation ratios using a maximum-likelihood interval mapping procedure. We identified 19 loci showing moderately deleterious to lethal embryonic effects. These loci account for >13 lethal equivalents, greater than the average of 8.5 lethal equivalents reported for loblolly pine. Viability alleles show predominantly recessive action, although potential overdominance occurs at 3 loci. We found no evidence for epistasis in the distribution of pairwise marker correlations or in the regression of fitness on the number of markers linked to deleterious alleles. The predominant role of semilethal alleles in embryonic inbreeding depression has implications for the evolution of isolated populations and for genetic conservation and breeding programs in conifers.

Crosses, Genetic↗

PubMed2005

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.

Alabama↗

PubMed2007

Inbreeding depression in an autotetraploid herb: a three cohort field study.

Autotetraploids are predicted to have reduced inbreeding depression relative to diploids. However, recent theory and information on genomic changes following autopolyploidy suggest that inbreeding depression may be closer to diploids. In three consecutive years, self and outcross pollinations were conducted on autotetraploid Campanulastrum americanum, seeds were planted into native sites, and biennial offspring were followed through seed production. Inbred individuals had lower germination rates, reduced survival, were smaller, and flowered later, producing fewer fruits with fewer seeds. Inbred offspring had 6% of the cumulative fitness of outcross offspring. Although performance varied substantially among cohorts, inbreeding depression for cumulative fitness was relatively constant, with delta ranging only from 0.92 to 0.95. C. americanum, like many outcrossing species, expressed very high amounts of inbreeding depression. This supports the hypothesis that inbreeding depression of some autotetraploids may be similar to that of diploids. Furthermore, few studies have measured temporal variation in inbreeding depression. Constant inbreeding depression given a sixfold range in cohort performance suggests that inbreeding depression may be relatively robust to environmental variation experienced by natural populations.

Campanulaceae↗

PubMed2006

Life history correlates of inbreeding depression in mandrills (Mandrillus sphinx).

Inbreeding depression reflects the negative consequences of increased homozygosity at genes that affect fitness. We investigate inbreeding depression in a semi-free-ranging colony of mandrills (Mandrillus sphinx), using high-quality pedigree data, comprising five maternal generations and 20 years of morphological and demographic data. We examine the relationship between inbreeding coefficients and four fitness correlates: two growth parameters (mass and height for age) and longevity in both sexes, and age at first conception in females. Inbreeding was correlated with both growth parameters, but only in females, with inbred females being smaller than noninbred females. Inbreeding was also correlated significantly with age at first conception, with inbred females giving birth earlier in life than noninbred females. We suggest that sex-biased maternal investment may explain this sex-differential response to inbreeding, although the lack of a significant association between inbreeding and growth in males may also be due to the provisioned nature of the colony. The surprising relationship between age at first conception and inbreeding may be related to smaller adult size in inbred females, or to their being less able to escape from male sexual coercion.

Age Factors↗

PubMed2001

Consequences of life history for inbreeding depression and mating system evolution in plants.

Many plants are perennial, but most studies of inbreeding depression and mating system evolution focus on annuals. This paper extends a population genetic model of inbreeding depression due to recessive deleterious mutations to perennials. The model incorporates life history and mating system variation, and multiplicative selection across many genetic loci. In the absence of substantial mitotic mutation, perennials have higher mean fitness and lower, or even negative, inbreeding depression than annuals with the same mating system. As in annuals, self fertilization exposes deleterious recessive mutations to selection, increasing mean fitness and decreasing inbreeding depression. Including mitotic mutation decreases mean fitness while increasing inbreeding depression. Perenniality introduces a kind of selective sieve, such that strongly recessive mutations contribute disproportionately to mean fitness and inbreeding depression. In the presence of high mitotic mutation, this selective sieve may provide a mechanistic basis for high inbreeding depression observed in some long lived perennials. Without substantial mitotic mutation, it is difficult to reconcile genetically based models of inbreeding depression with the empirical generalization that perennials outcross while related annuals self fertilize.

Biological Evolution↗

PubMed2000

Inbreeding depression and mating systems in two species of Linanthus (Polemoniaceae).

Theory predicts that, at equilibrium, the magnitude of inbreeding depression caused by recessive alleles should decrease as the level of self-fertilization increases, facilitating the evolution of even higher selfing rates. Data are presented on mating systems and inbreeding depression in multiple populations of two annual species in the genus Linanthus (Polemoniaceae). Outcrossing rates were found to be less than 0.075 in populations of a small-flowered species, L. bicolor, and ranged from 0.104 to 0.300 in populations of a larger-flowered species, L. jepsonii. Inbreeding depression for both species was low for all life history stages, with cumulative inbreeding depression less than 0.23 in all populations. Significant inbreeding depression for pollen fertility was found in L. jepsonii populations and may contribute substantially to total inbreeding depression. A competitive treatment had no significant effect on the expression of inbreeding depression for above-ground dry mass. In accordance with theoretical expectations, the magnitude of inbreeding depression was lower in L. bicolor, the more highly selfing species. The magnitude of inbreeding depression in L. jepsonii, although greater than that of L. bicolor, is not sufficient to prevent evolution of increased selfing rates in this species, according to some theoretical models of mating system evolution.

Analysis of Variance↗

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↗

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

Inbreeding depression and the evolution of dispersal rates: a multilocus model.

Inbreeding depression is one of the possible reasons organisms disperse. In this article, we present a two-locus model for the evolution of dispersal in the presence of inbreeding depression. The first locus codes for a modifier of the migration rate, while the second locus is a selected locus generating inbreeding depression. We express the change in frequency of the migration modifier as a function of allele frequencies and genetic associations and then use a quasi-equilibrium assumption to express genetic associations as functions of allele frequencies. Our model disentangles two effects of inbreeding depression: it gives an advantage to migrant individuals because their offspring are on average less homozygous, but it also decreases the degree of population structure, thus decreasing the strength of kin selection for dispersal. We then extend our model to include an infinite number of selected loci. When the cost of dispersal is not too high, the model predictions are confirmed by multilocus simulation results and show that inbreeding depression can have a substantial effect on the dispersal rate. For high costs of dispersal, we observe discrepancies between the model and the simulations, probably caused by associations among selected loci, which are neglected in the analysis.

Animal Migration↗

PubMed1994

Purging inbreeding depression and the probability of extinction: full-sib mating.

Inbreeding depression has been a topic of interest in recent years from a number of perspectives, particularly in the captive breeding of endangered species. Generally, the goal of captive breeding is to avoid the detrimental effects of inbreeding depression and to retain genetic variation for future adaptation. However, an important component of another suggested approach to captive breeding is to purge rapidly the population of its genetic load so that its long-term fitness is not compromised. I have examined the effectiveness of purging the genetic load by documenting both the reduction in inbreeding depression and the increase of the probability of extinction when there is continuous full-sib mating. When the genetic load is the result of lethals, the inbreeding depression is quickly purged without a high probability of extinction, except when the total genetic load is high. On the other hand, if the load is due to detrimentals of relatively small effect, the genetic load becomes fixed, the mean fitness is reduced, and the probability of extinction may be greatly increased. In other words, the success of such a programme to purge genetic load without an increase in the probability of extinction is highly dependent upon the genetic basis of inbreeding depression, information that is not readily available for most species.

Adaptation, Physiological↗

PubMed1999

Late-acting inbreeding depression in both male and female function of echium vulgare (Boraginaceae)

We present data on late-acting inbreeding depression in pollen performance, siring success and seed production in Echium vulgare. Pollen viability and rate of pollen-tube growth were both lower for pollen from plants derived from selfing than for pollen from plants derived from outcrossing. Pollen tube numbers within the styles did not differ for pollen from plants derived from selfing or outcrossing. A pollination experiment with two mixtures of pollen from plants derived from selfing or outcrossing, revealed a significant decline of 55% in siring success for pollen from plants derived from selfing. A second experiment with a complete diallel design revealed inbreeding depression for both siring success of the offspring (32.8%) and a decline in seed production of the offspring (34.8%-40.6%). In addition, results indicated a heritable component for seed number per flower. Offspring fitness, measured as seed production and siring ability, can be severely affected by late-acting inbreeding depression. Inbreeding depression values for male and female functions were not correlated. Both functions must therefore be considered when calculating inbreeding depression.

Journal Article↗

PubMed2002

Strong inbreeding depression in a Daphnia metapopulation.

The deleterious effects of inbreeding have long been known, and inbreeding can increase the risk of extinction for local populations in metapopulations. However, other consequences of inbreeding in metapopulations are still not well understood. Here we show the presence of strong inbreeding depression in a rockpool metapopulation of the planktonic freshwater crustacean Daphnia magna, which reproduces by cyclical parthenogenesis. We conducted three experiments in real and artificial rockpools to quantify components of inbreeding depression in the presence and the absence of competition between clonal lines of selfed and outcrossed genotypes. In replicated asexual populations, we recorded strong selection against clones produced by selfing in competition with clones produced by outcrossing. In contrast, inbreeding depression was much weaker in single-clone populations, that is, in the absence of competition between inbred and outbred clones. The finding of a competitive advantage of the outbred genotypes in this metapopulation suggests that if rockpool populations are inbred, hybrid offspring resulting from crosses between immigrants and local genotypes might have a strong selective advantage. This would increase the effective gene flow in the metapopulation. However, the finding of low inbreeding depression in the monoclonal populations suggests that inbred and outbred genotypes might have about equal chances of establishing new populations.

Animals↗

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↗

PubMed2001

Overdominant epistatic loci are the primary genetic basis of inbreeding depression and heterosis in rice. I. Biomass and grain yield.

To understand the genetic basis of inbreeding depression and heterosis in rice, main-effect and epistatic QTL associated with inbreeding depression and heterosis for grain yield and biomass in five related rice mapping populations were investigated using a complete RFLP linkage map of 182 markers, replicated phenotyping experiments, and the mixed model approach. The mapping populations included 254 F(10) recombinant inbred lines derived from a cross between Lemont (japonica) and Teqing (indica) and two BC and two testcross hybrid populations derived from crosses between the RILs and their parents plus two testers (Zhong 413 and IR64). For both BY and GY, there was significant inbreeding depression detected in the RI population and a high level of heterosis in each of the BC and testcross hybrid populations. The mean performance of the BC or testcross hybrids was largely determined by their heterosis measurements. The hybrid breakdown (part of inbreeding depression) values of individual RILs were negatively associated with the heterosis measurements of their BC or testcross hybrids, indicating the partial genetic overlap of genes causing hybrid breakdown and heterosis in rice. A large number of epistatic QTL pairs and a few main-effect QTL were identified, which were responsible for >65% of the phenotypic variation of BY and GY in each of the populations with the former explaining a much greater portion of the variation. Two conclusions concerning the loci associated with inbreeding depression and heterosis in rice were reached from our results. First, most QTL associated with inbreeding depression and heterosis in rice appeared to be involved in epistasis. Second, most ( approximately 90%) QTL contributing to heterosis appeared to be overdominant. These observations tend to implicate epistasis and overdominance, rather than dominance, as the major genetic basis of heterosis in rice. The implications of our results in rice evolution and improvement are discussed.

Breeding↗

PubMed2000

The effect of drought stress on inbreeding depression in four populations of the Mediterranean outcrossing plant Crepis sancta (Asteraceae).

The effect of physiological stress on the magnitude of inbreeding depression in plants has been the subject of few studies and is currently controversial because of contradictory results. We measured the inbreeding depression at three drought stress levels, precisely defined by a preliminary physiological experiment. We also tested the hypothesis that more highly self-compatible populations exhibit reduced inbreeding depression due to purging of deleterious mutations. The study was conducted on two populations of the annual and allogamous plant Crepis sancta collected from the French Mediterranean region and two other populations from marginal areas with various self-incompatibility levels. Drought stress did not increase inbreeding depression in terms of plant mortality but significantly increased the inbreeding depression for the date of first flowering, number of heads per plant and relative growth rate. The most self-fertile marginal population showed an absence of inbreeding depression in most of the measured traits indicating that purging could have taken place in this population. The three others populations showed relatively low and similar estimates of inbreeding depression (delta approximately 0.35). The relatively low values obtained compared to the results found in allogamous plants suggests that the absence of competition for C. sancta in our experiment probably underestimated the effects of inbreeding in natural populations where competition occurs.

Analysis of Variance↗

PubMed2002

Severe inbreeding depression in collared flycatchers (Ficedula albicollis).

The causes and magnitude of inbreeding depression are of considerable importance for a wide range of issues in evolutionary and conservation biology, but we have only a limited understanding of inbreeding depression in natural populations. Here, we present a study of inbreeding in a large wild population of collared flycatchers (Ficedula albicollis). Inbreeding was rare, to the extent that we detected only 1.04% of 2139 matings over 18 years that resulted in offspring with a non-zero inbreeding coefficient, f > 0. When it did occur, inbreeding caused a significant reduction in the egg-hatching rate, in fledgling skeletal size and in post-fledging juvenile survival, with the number of offspring being recruited to the breeding population from a nest of f = 0.25 being reduced by 94% relative to a non-inbred nest. A maximum-likelihood estimate of the number of lethal equivalents per gamete was very high at B = 7.47, indicating a substantial genetic load in this population. There was also a non-significant tendency for inbreeding depression to increase with the strength of selection on a trait. The probability of mating between close relatives (f = 0.25) increased throughout the breeding season, possibly reflecting increased costs of inbreeding avoidance. Our results illustrate how severe inbreeding depression and considerable genetic load may exist in natural populations, but detecting them may require extensive long-term datasets.

Animals↗

PubMed2000

The Effects of a Bottleneck on Inbreeding Depression and the Genetic Load.

We study the effects of a population bottleneck on the inbreeding depression and genetic load caused by deleterious mutations in an outcrossing population. The calculations assume that loci have multiplicative fitness effects and that linkage disequilibrium is negligible. Inbreeding depression decreases immediately after a sudden reduction of population size, but the drop is at most only several percentage points, even for severe bottlenecks. Highly recessive mutations experience a purging process that causes inbreeding depression to decline for a number of additional generations. On the basis of available parameter estimates, the absolute fall in inbreeding depression may often be only a few percentage points for bottlenecks of 10 or more individuals. With a very high lethal mutation rate and a very slow population growth, however, the decline may be on the order of 25%. We examine when purging might favor a switch from outbreeding to selfing and find it occurs only under very limited conditions unless population growth is very slow. In contrast to inbreeding depression, a bottleneck causes an immediate increase in the genetic load. Purging causes the load to decline and then overshoot its equilibrium value. The changes are typically modest: the absolute increase in the total genetic load will be at most a few percentage points for bottlenecks of size 10 or more unless the lethal mutation rate is very high and the population growth rate very slow.

bottlenecks↗