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Udder health shows inbreeding depression in Danish Holsteins.

The purpose of this study was to estimate inbreeding depression for udder health using records on mastitis incidence and somatic cell count (SCC). Data were selected based on a pedigree completeness index so that inbreeding coefficients were reliable. Average inbreeding was 3.3%. The presence or absence of mastitis was considered in first (CM1), second (CM2), and third (CM3) lactations, and was recorded as a binary trait. Somatic cell count was also included and analyzed on the log scale. A minimum of 140,000 cows with records were included per trait. Linear sire models were used and (co)variance components were estimated. Linear and curvilinear effects of the coefficient of inbreeding were included. Inbreeding significantly affected all traits investigated. Three traits, CM1, CM3, and SCC, showed a nonlinear relationship between phenotype and inbreeding coefficient. A cow with 5% inbreeding compared with a cow with 2% inbreeding had a higher SCC and a higher incidence of mastitis. The SCC increased by 1,500 cells/mL in first lactation and the incidence of mastitis increased by 1.08, 0.55, and 0.98% in first, second, and third lactation, respectively. The corresponding reduction in net return from these traits over 3 lactations amounted to 11.00 US dollars under Danish production circumstances. These results are a step on the way to estimating the total cost of inbreeding depression to determine an acceptable rate of inbreeding.

Animals↗

Inbreeding depression and heterosis in a subdivided population: influence of the mating system.

We investigate the joint effects of gene flow and selfing on the level of inbreeding depression, heterosis and genetic load in a subdivided population at equilibrium. Low gene flow reduces inbreeding depression and substantially increases heterosis. However, in highly self-fertilizing populations, inbreeding depression is independent of the amount of gene flow. When migration occurs via pollen, consanguinity of the reproductive system could have a negative influence on subpopulation persistence, in contrast to the case of isolated populations. However, with only seed migration, genetic load and heterosis depend mildly on the mating system. From an evolutionary point of view, we reach two main conclusions: first, outcrossing is selected for if gene flow is low; second, intermediate levels of gene flow could promote mixed mating systems, especially when migration occurs through pollen.

Biological Evolution↗

Modes and rates of selfing and associated inbreeding depression in the self-incompatible plant Senecio squalidus (Asteraceae): a successful colonizing species in the British Isles.

The strength of the self-incompatibility (SI) response in Senecio squalidus was measured across its British range. Geographic variation in SI was investigated and the extent and inheritance of pseudo-self-compatibility (PSC) and inbreeding depression were determined. Mean self-fruit-set per capitulum was calculated for individuals and sample populations. The heritability of PSC and the magnitude of inbreeding depression were assessed by comparing selfing rates and fitness trait values between SI and PSC parent-progeny lines. SI was found to be strongly expressed in S. squalidus throughout its British range, with only 3.1% of the individuals sampled showing PSC. This PSC had relatively low heritability with stronger expression of SI in selfed progeny relative to PSC parents. Inbreeding depression was shown to be great in S. squalidus, with mean life history stage values ranging from 0.18 to 0.25. The strength of SI in S. squalidus appears not to have weakened in response to its rapid colonization of Britain. The avoidance of inbreeding depression is likely to be the primary factor maintaining strong SI in this successful colonizing species.

Ecosystem↗

Inbreeding depression and genetic load of sexually selected traits: how the guppy lost its spots.

To date, few studies have investigated the effects of inbreeding on sexually selected traits, although inbreeding depression on such traits can play an important role in the evolution and ecology of wild populations. Sexually selected traits such as ornamentation and courtship behaviour may not be primary fitness characters, but selection and dominance coefficients of their mutations will resemble those of traits under natural selection. Strong directional selection, for instance, through female mate-choice, purges all but the most recessive deleterious mutations, and the remaining dominance variation will result in inbreeding depression once populations undergo bottlenecks. We analysed the effects of inbreeding on sexually selected traits (colour pattern and courtship behaviour) in the male guppy, Poecilia reticulata, from Trinidad, and found a significant decline in the frequency of mating behaviour and colour spots. Such effects occurred although the genetic basis of these traits, many of which are Y-linked and hemizygous, would be expected to leave relatively little scope for inbreeding depression. Findings suggest that these sexually selected traits could reflect the genetic condition or health of males, and thus may be informative mate-cue characters for female choice as suggested by the 'good genes' model.

Animals↗

Inbreeding depression in anthropometric traits among Telaga boys of Kharagpur, West Bengal, India.

Inbreeding depression in phenotypic variations as well as reproductive capacity was reported in animals. Depression of physical measurements in human beings especially among infants and adults due to inbreeding has also been reported all over the world. The present study is an attempt to examine the nature of inbreeding effect on growing children at each yearly age with increase of inbreeding intensity in a local endogamous population based on pedigree data. In general, mean values of measurements appear to be smaller in higher levels of inbreeding from that of lower levels of inbreeding in each yearly age. These differences are even more marked when grouped as low and high inbreeding level. The percent of inbreeding depression tends to be larger in post adolescent boys than in younger children.

Adolescent↗

Frequency-dependent inbreeding depression in Amsinckia.

If the competitive ability of plants produced by self-pollination differs from that of plants derived by outcrossing, then the magnitude of inbreeding depression may be influenced by the composition of the competitive environment (i.e., the frequency of plants that have arisen from selfing and outcrossing in the neighborhood of "target" plants in which inbreeding depression is expressed). Here, we report the results of experiments designed to examine whether inbreeding depression is influenced by the frequency of inbred plants in the competitive neighborhood. Two species of the annual plant genus Amsinckia were studied, one a near-complete selfer (Amsinckia gloriosa) and the other a partial outcrosser (Amsinckia douglasiana). Competition experiments were conducted in artificial stands composed of different mixtures of inbred and outbred progeny. The fitnesses of progeny were found to be significantly influenced by the composition of the competing neighborhood. The fitness of target plants, however, did not vary monotonically with the frequency of inbred plants in the neighborhood. Rather, for A. gloriosa, maximum performance was observed when there was an intermediate frequency of inbred neighbors. For A. douglasiana, the opposite pattern was found. The results suggest that competition among progeny has the potential to play a role in the selection of self-fertilization and possibly in the maintenance of mixed mating systems.

Amsinckia↗

Levels of inbreeding depression over seven generations of selfing in the androdioecious clam shrimp, Eulimnadia texana.

Androdioecy (mixtures of males and hermaphrodites) is a rare mating system in both plants and animals. Theory suggests that high levels of inbreeding depression can maintain males in androdioecious populations if hermaphrodites commonly self-fertilize. However, if inbreeding depression (delta) can be 'purged' from selfing populations, maintaining males is more difficult. In the androdioecious clam shrimp, Eulimnadia texana, delta is estimated to be as high as 0.7. Previous work suggests that this high level is maintained in the face of high levels of inbreeding due to an associative overdominance of fitness-related loci with the sex-determining locus. Such associative overdominance would make purging of inbreeding depression difficult to impossible. The current experiment was designed to determine if delta can be purged in these shrimp by tracking fitness across seven generations in selfing and outcrossing treatments. Evidence of purging was found in one of four populations, but the remaining populations demonstrated a consistent pattern of delta across generations. Although the experimental design allowed ample opportunity for purging, the majority of populations were unable to purge their genetic load. Therefore, delta in this species is likely due to associative overdominance caused by deleterious recessive alleles linked to the sex determining locus.

Analysis of Variance↗

Genomic inbreeding coefficients and inbreeding depression of semen production traits at genome-wide and chromosomal levels in Japanese Holstein bulls.

We aimed to estimate inbreeding coefficients and the effects of inbreeding depression on semen production traits at both the genome-wide and chromosomal levels. We utilized pedigree data for 19,921 animals, single nucleotide polymorphism (SNP) data on 5700 Japanese Holstein bulls, and 52,193 semen collection records from 775 bulls. We estimated 4 different inbreeding coefficients, namely a pedigree-based coefficient (FPED) and 3 genomic coefficients derived from SNP data. The genomic coefficients consisted of one based on the genomic relationship matrix (FGRM), one based on runs of homozygosity (ROH), and one based on homozygous-by-descent (HBD) segments (FHBD). These genomic coefficients were estimated at both the genome-wide and chromosomal levels. Furthermore, we investigated the effects of these coefficients on semen production traits: semen volume (VOL), sperm concentration (CON), sperm number (NUM), and sperm motility (MOT). In the genome-wide-level analysis, inbreeding coefficients increased markedly in bulls born after 2009, coinciding with the introduction of genomic selection. Significant inbreeding depression of VOL was found. At the chromosomal level, the inbreeding coefficients for most chromosomes showed a similar trend to the genome-wide metrics, although some (e.g., chr10 and chr20) exhibited a more pronounced trend. Suggestive inbreeding effects were detected on specific chromosomes for all traits (chr1 and chr22 for VOL, chr24 and chr29 for CON, chr1, chr12, and chr27 for NUM, chr10 and chr18 for MOT), including the traits that were not significant at the genome-wide level. Our results highlight that chromosomal-level analysis provides information complementary to whole-genome metrics, offering a more detailed perspective for managing inbreeding effects. To mitigate the adverse effects of inbreeding on semen production traits, future breeding programs would benefit from the control of inbreeding effects on high-risk chromosomal regions.

Genomic inbreeding coefficient↗

Group living and inbreeding depression in a subsocial spider.

Social spiders are unusual among social organisms in being highly inbred-males and females mature within their natal nest and mate with each other to produce successive generations. Several lines of evidence suggest that in spiders inbred social species originated from outbred subsocial ancestors, a transition expected to have been hindered by inbreeding depression. As a window into this transition, we examined the fitness consequences of artificially imposed inbreeding in the naturally outbred subsocial spider Anelosimus cf. jucundus. Subsocial spiders alternate periods of solitary and social living and are thought to resemble the ancestral system from which the inbred social species originated. We found that inbreeding depression in this subsocial spider only becomes evident in spiders raised individually following the end of their social phase and that ecological and demographic factors such as eclosion date, number of siblings in the group and mother's persistence are more powerful determinants of fitness during the social phase. A potential explanation for this pattern is that maternal care and group living provide a buffer against inbreeding depression, a possibility that may help explain the repeated origin of inbred social systems in spiders and shed light on the origin of other systems involving regular inbreeding.

Animals↗

Influence of parentage upon growth in Ostrea edulis: evidence for inbreeding depression.

Genetic variability for growth was analysed in three populations of Ostrea edulis, selected for resistance to the protozoan parasite Bonamia ostreae. This study was undertaken first to determine the potential for selection for growth in populations that have never been selected for this character, and second to estimate heterosis versus inbreeding depression. Growth was monitored in culture for 10 months. The selected populations (namely S85-G3, S891-G2 and S89W-G2), their crossbred population and a control population were composed of full-sib families whose parents were already genotyped using five microsatellite markers. This genotyping allowed the estimation of genetic relatedness among pairs of parents. The parents' relatedness was then correlated with the growth performance of their offspring within each of the three populations, and inbreeding depression was estimated. The population effect for growth was highly significant, with the crossbred population having the highest growth rate, followed by S891-G2 and S89W-G2, S85-G3 and the control population. The within-populations family effect was also highly significant, indicating, as well as the high value for heritability at the family level (between 0.57 and 0.92), that a potential for a further selection for growth still exists within the three populations. Estimates of inbreeding depression (relative to the mean, for complete inbreeding) were high (1 for S891-G2, 0.44 for S89W-G2 and between 0.02 and 0.43 for S85-G3), which correlates with the apparent heterosis for growth observed in the crossbred population. These results are discussed in the context of the future management of the selected populations.

Animals↗

A model of the evolution of dichogamy incorporating sex-ratio selection, anther-stigma interference, and inbreeding depression.

Historically, explanations for the evolution of floral traits that reduce self-fertilization have tended to focus on selection to avoid inbreeding depression. However, there is growing support for the hypothesis that such traits also play a role in promoting efficient pollen dispersal by reducing anther-stigma interference. The relative importance of these two selective pressures is currently a popular topic of investigation. To date, there has been no theoretical exploration of the relative contributions of selection to avoid the genetic costs of self-fertilization and selection to promote efficient pollen dispersal on the evolution of floral traits. We developed a population genetic model to examine the influence of these factors on the evolution of dichogamy: the temporal separation of anther maturation and stigma receptivity. Our analysis indicates that anther-stigma interference can favor dichogamy even in the absence of in-breeding depression. Although anther-stigma interference and inbreeding depression are the key forces driving the initial evolution of dichogamy, selection to match the timing of pollen dispersal to the availability of ovules at the population level becomes a more potent force opposing the further evolution of dichogamy as the extent of temporal separation increases. This result may help to explain otherwise puzzling phenomena such as why dichogamy is rarely complete in nature and why dichogamy tends to be associated with asynchronous flower presentation.

Biological Evolution↗

Marker-based inferences about epistasis for genes influencing inbreeding depression.

We describe a multilocus, marker-based regression method for inferring interactions between genes controlling inbreeding depression in self-fertile organisms. It is based upon selfing a parent heterozygous for several unlinked codominant markers, then analyzing the fitness of progeny marker genotypes. If loci causing inbreeding depression are linked to marker loci, then viability selection is manifested by distorted segregation of markers, and fecundity selection by dependence of the fecundity character upon the marker genotype. To characterize this selection, fitness is regressed on the proportion of loci homozygous for markers linked to deleterious alleles, and epistasis is detected by nonlinearity of the regression. Alternatively, fitness can be regressed on the proportion of heterozygous loci. Other modes of selection can be incorporated with a bivariate regression involving both homozygote and heterozygote marker genotypes. The advantage of this marker-based approach is that "purging" is minimized and specific chromosomal segments are identified; its disadvantage lies in low statistical power when linkage is not strong and/or the linkage phase between marker and selected loci is uncertain. Using this method in the wildflower Mimulus guttatus, we found predominant multiplicative gene interaction determining fecundity and some negative synergistic (nonmultiplicative) interaction for viability.

Epistasis, Genetic↗

Inbreeding depression: tests of the overdominance and partial dominance hypotheses.

The two principal theories of the causal mechanism for inbreeding depression are the partial dominance hypothesis and the overdominance hypothesis. According to the first hypothesis, inbreeding increases the frequency of homozygous combinations of deleterious recessive alleles thereby decreasing fitness, whereas the overdominance hypothesis posits that inbreeding increases homozygosity and thus reduces the frequency of the superior heterozygotes. These two hypotheses make different predictions on the effect of crossing inbred lines: the overdominance hypothesis predicts that trait means will be restored to the outbred means, whereas the partial dominance hypothesis predicts that trait means will exceed those of the outbred population. I tested these predictions using seven inbred lines of the sand cricket, Gryllus firmus. Fourteen generations of brother-sister mating resulted in an inbreeding depression of 20-34% in four traits: nymphal weights at ages 14 days, 21 days, 28 days, and early fecundity. An incomplete diallel cross of these lines showed genetic variation among lines and an increase in all trait means above the outbred means, with three being significantly higher. These results provide support for the partial dominance hypothesis and are inconsistent with the overdominance hypothesis.

Analysis of Variance↗

Point estimation and graphical inference of marginal dominance for two viability loci controlling inbreeding depression.

A deterministic analysis is conducted to examine marginal dominance for two linked viability loci influencing inbreeding depression and its graphical inferences. Four estimators of marginal dominance are derived, assuming a biallelic marker locus completely linked to one of the viability loci, and the biases in expected estimates due to the other deleterious locus are discussed. Three conditions under which apparent partial dominance or underdominance could occur are found, i.e. when two multiplicative, partially recessive loci are linked in coupling phase and when two synergistic, highly overdominant loci are linked in coupling or repulsion phases. Expected frequencies of the three marker genotypes in selfed progeny are derived, considering two linkage phases, two types of marker locus position with respect to the viability loci, and the multiplicative and synergistic fitness models. Segregation ratios are generated for the marker locus linked to either two overdominant or partially recessive loci and plotted in gene action graphs to examine the robustness of the graphical inferences of gene action due to the presence of an additional linked viability locus. Under a multiplicative fitness model, the presence of an additional partially recessive or overdominant locus in the vicinity of the marker locus does not greatly affect the graphical inferences of the relative role of partially recessive or overdominant genes in expression of inbreeding depression. A marker linked to two synergistic, highly overdominant loci can behave as though linked to a partially recessive, partially dominant or underdominant locus, even with relatively weak synergism.

Genes, Dominant↗

Joint effects of self-fertilization and population structure on mutation load, inbreeding depression and heterosis.

Both the spatial distribution of organisms and their mode of reproduction have important effects on the change in allele frequencies within populations. In this article, we study the combined effect of population structure and the rate of partial selfing of organisms on the efficiency of selection against recurrent deleterious mutations. Assuming an island model of population structure and weak selection, we express the mutation load, the within- and between-deme inbreeding depression, and heterosis as functions of the frequency of deleterious mutants in the metapopulation; we then use a diffusion model to calculate an expression for the equilibrium probability distribution of this frequency of deleterious mutants. This allows us to derive approximations for the average mutant frequency, mutation load, inbreeding depression, and heterosis, the simplest ones being Equations 35-39 in the text. We find that population structure can help to purge recessive deleterious mutations and reduce the load for some parameter values (in particular when the dominance coefficient of these mutations is <0.2-0.3), but that this effect is reversed when the selfing rate is above a given value. Conversely, within-deme inbreeding depression always decreases, while heterosis always increases, with the degree of population subdivision, for all selfing rates.

Crosses, Genetic↗

Genetic adaptation to captivity and inbreeding depression in small laboratory populations of Drosophila melanogaster.

The rate of adaptation to a competitive laboratory environment and the associated inbreeding depression in measures of reproductive fitness have been observed in populations of Drosophila melanogaster with mean effective breeding size of the order of 50 individuals. Two large wild-derived populations and a long-established laboratory cage population were used as base stocks, from which subpopulations were extracted and slowly inbred under crowded conditions over a period of 210 generations. Comparisons have been made of the competitive ability and reproductive fitness of these subpopulations, the panmictic populations produced from them by hybridization and random mating and the wild- or cage-base populations from which they were derived. After an average of approximately 180 generations in the laboratory, the wild-derived panmictic populations exceeded the resampled natural populations by 75% in fitness under competitive conditions. The cage-derived panmictic population, after a total of 17 years in the laboratory, showed a 90% superiority in competitive ability over the corresponding wild population. In the inbred lines derived from the wild-base stocks, the average rate of adaptation was estimated to be 0.33 +/- 0.06% per generation. However, the gain in competitive ability was more than offset by inbreeding depression at an initial rate of approximately 2% per generation. The effects of both adaptation and inbreeding on reproductive ability in a noncompetitive environment were found to be minor by comparison. The maintenance of captive populations under noncompetitive conditions can therefore be expected to minimize adaptive changes due to natural selection in the changed environment.

Adaptation, Biological↗

Analysis of levels of inbreeding and inbreeding depression in Jersey cattle.

A pedigree file of 157,015 male and female Jersey cattle (born after 1955) from the Canadian herdbooks was investigated for the occurrence of inbreeding. A large proportion of Jersey bulls and cows were inbred (32.4 and 36.3% for bulls and cows, respectively). However, average inbreeding coefficients of these inbred cows and of all cows were low. First lactation milk, fat, and fat percentage records for 53,592 Jersey cows were analyzed. Inbreeding was included in the animal model as a linear covariate. The regression coefficients of milk, fat, and fat percentage on inbreeding were -9.84 kg, -.55 kg, and -.0011% per 1% increase of inbreeding. Inbreeding depression was not enough to cause large reductions of milk and fat yield of a cow with average inbreeding. However, when the inbreeding coefficient was greater than 12.5%, the inbreeding depression was significantly higher than expected and such that intentional inbreeding is not justified unless the mating is to an animal with exceptionally high breeding value.

Algorithms↗

Relative contribution of inbreeding depression and eroded adaptive diversity to extinction risk in small populations of shore campion.

To study the relative importance of inbreeding depression and the loss of adaptive diversity in determining the extinction risk of small populations, we carried out an experiment in which we crossed and self-fertilized founder plants from a single, large population of shore campion (Silene littorea Brot.). We used the seeds these plants produced to colonize 18 new locations within the distribution area of the species. The reintroduced populations were of three kinds: inbred and genetically homogeneous, each made up of selfed seed from a single plant; inbred and mixed, made up of a mixture of selfed seeds from all founder plants; and outbred and mixed, made up of a mixture of seeds obtained in outcrosses between the founders. We compared the inbred homogeneous populations with the inbred mixed to measure the effect of genetic diversity among individuals and the inbred mixed with the outbred mixed to measure the effect of inbreeding. Reintroduction success was seriously limited by inbreeding, whereas it was not affected by genetic diversity. This observation and the nonsignificant interaction between family and reintroduction location for individual plant characters suggest that the fixation of overall deleterious genes causing inbreeding depression posed a more serious threat to the short-term survival of the populations than the loss of genes involved in genotype and environment interactions. Thus, reintroduction success was related to adaptive diversity. Preventing such fixation might be the most important consideration in the genetic management and conservation of shore campion populations.

Adaptation, Physiological↗