Random and nonrandom inbreeding.
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The Y and the Z strains of Praomys natalensis (Mastomys) are being sucessfully inbred, as far as is known for the first time. Spontaneous tumours of the glandular stomach have been found to develop in these rodents. Evidence is provided of the great potential of the laboratory Mastomys for use in medical research on cancer, plague, bilhaziasis, and more recently in the research on the Lassa fever virus.
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Inbreeding depression, the reduction in fitness that accompanies inbreeding, is one of the most important topics of research in evolutionary and conservation genetics. In the recent literature, much attention has been paid to the possibility of purging the genetic load. If inbreeding depression is due to deleterious alleles, whose effect on fitness are negative when in a homozygous state, then successive generations of inbreeding may result in a rebound in fitness due to the selective decrease in frequency of deleterious alleles. Here we examine the experimental evidence for purging of the genetic load by collating empirical tests of rebounds in fitness-related traits with inbreeding in animals and plants. We gathered data from 28 studies including five mammal, three insect, one mollusc, and 13 plant species. We tested for purging by examining three measures of fitness-component variation with serial generations of inbreeding: (1) changes in inbreeding depression, (2) changes in fitness components of inbred lines relative to the original outbred line, and (3) purged population (outcrossed inbred lines) trait means as a function of ancestral outbred trait means. Frequent and substantial purging was found using all three measures, but was particularly pronounced when tracking changes in inbreeding depression. Despite this, we found little correspondence between the three measures of purging within individual studies, indicating that the manner in which a researcher chooses to estimate purging will affect interpretation of the results obtained. The discrepancy suggests an alternative hypothesis: rebounds in fitness with inbreeding may have resulted from adaptation to laboratory conditions and not to purging when using outcrossed inbred lines. However, the pronounced reduction in inbreeding depression for a number of studies provides evidence for purging, as the measure is likely less affected by selection for laboratory conditions. Unlike other taxon-specific reviews on this topic, our results provide support for the purging hypothesis, but firm predictions about the situations in which purging is likely or the magnitude of fitness rebound possible when populations are inbred remain difficult. Further research is required to resolve the discrepancy between the results obtained using different experimental approaches.
Mating patterns in plant populations are influenced by interactions between reproductive traits and ecological conditions, both factors that are likely to vary geographically. Narcissus triandrus, a wide-ranging heterostylous herb, exhibits populations with either two (dimorphic) or three (trimorphic) style morphs and displays substantial geographical variation in demographic attributes and floral morphology. Here, we investigate this variation to determine if demography, morphology, and mating system differ between the two sexual systems. Our surveys in Portugal and NW Spain indicated that dimorphic populations were less dense, of smaller size, and had larger plants and flowers compared to trimorphic populations. Outcrossing rates estimated using allozyme markers revealed similar outcrossing rates in dimorphic and trimorphic populations (t(m) dimorphic=0.759; t(m) trimorphic=0.710). All populations experienced significant inbreeding in progeny (mean F=0.143). In contrast, parental estimates of inbreeding were not significantly different from zero (mean F=0.062), implying that few inbred offspring survive to reproductive maturity due to inbreeding depression. Although the majority of inbreeding results from selfing, significant levels of biparental inbreeding were also detected in eight of the nine populations (mean s(s)-s(m)=0.081). Density was negatively associated with levels of selfing but positively associated with biparental inbreeding. Population size was positively associated with outcrossing but not biparental inbreeding. There were no consistent differences among the style morphs in outcrossing or biparental inbreeding indicating that the maintenance of trimorphism vs dimorphism is unlikely to be associated with inbreeding of maternal parents.
We investigated inbreeding depression and selfing in hermaphroditic Schiedea menziesii to assess the stability of the breeding system. A combination of high selfing rates and strong inbreeding depression suggests that the mating system is unstable. The population-level selfing rate measured in three years ranged considerably from 0.425 (SE = 0.138) to 0.704 (0.048); family measures of selfing rate varied from zero to one in all three years. Inbreeding coefficients did not differ from zero, suggesting that inbred plants do not survive to reproduction in the field. Average inbreeding depression measured in two greenhouse experiments was 0.608-0.870, with values for individual plants ranging from -0.170 to 0.940. The magnitude of inbreeding depression expressed at different life-history stages depended on experimental conditions. When plants were grown during the winter, inbreeding depression was expressed at early and late life-history stages. When plants were grown during the summer, inbreeding depression was detected for germination but not for later life-history stages. Inbreeding depression for vegetative and inflorescence biomass was also measured using field-collected seeds where cross status was assigned using genotypes determined electrophoretically. We did not detect a relation between inbreeding depression and the selfing rate at the level of the individual plant. We saw no evidence for intrafloral selfing, suggesting that the evolution of increased selfing through autogamy is unlikely, despite high selfing rates. A more likely outcome of breeding system instability is the evolution of gynodioecy, which occurs in species of Schiedea closely related to S. menziesii. Females have been detected in progeny of S. menziesii that have been raised in the greenhouse. In the absence of biotic pollen vectors, the failure of these females to establish in the natural population may result from the absence of adaptations for wind pollination.
The present study investigated two rarely measured aspects of inbreeding depression: the relationship between inbreeding and behaviour, and the possibility that inbred individuals that survive infancy may still suffer from inbreeding depression by failing to breed or failing to show appropriate mating or parental behaviours. Specifically, the relationship between (1) behaviour at pairing and reproductive success, (2) inbreeding and reproductive success and (3) inbreeding and pairing behaviour, was examined in two subspecies of the oldfield mouse, Peromyscus polionotusEffects of parental and offspring inbreeding were separated through experimental design and analysis. Activity level during the first 25 days after pairing predicted future reproductive success: pairs that remained less active during the nocturnal (active) period were significantly less likely to breed than pairs that remained more active. Inbred females took significantly longer to produce their first litters and were less likely to produce litters than were outbred females, independently of whether females were related to their mates (i.e. whether their offspring would be inbred). Inactive pairs averaged fewer surviving pups than did active pairs. Inbreeding coefficient of female was a significant predictor of activity level in one of the two subspecies, suggesting that inbreeding may affect behaviour. Inbred adult females showed inbreeding depression in the form of lower conception rates and fewer surviving offspring, although the specific traits affected differed for the two subspecies. The implications for captive breeding programs, and likely causes of the subspecific differences, are discussed.
In order to estimate the isonymy structure of Spain, we studied surname distribution in 283 Spanish towns based on 3.625 million telephone users selected from 6.328 million users, downloaded from a commercial CD-ROM which contains all 13 million users in the country. Since in Spain the surname is made by the paternal and the maternal surname, it was possible to classify surnames according to parental origin. Two matrices of isonymy distances, one for paternal and one for maternal surnames, were constructed and tested for correlation with geographic distance. For the whole of Spain, Euclidean distance was significantly but weakly correlated with geographic distance both for paternal and maternal surnames, with r = 0.205 +/- 0.013 and r = 0.263 +/- 0.012, respectively. Two dendrograms of the 283 sampled towns were built from the two matrices of Euclidean distance. They are largely colinear. Four main clusters identified by the dendrograms are correlated with geography. Given the surname structure of Spain, we were able to calculate from isonymy and for each town 1). total or expressed inbreeding, 2). random or expected inbreeding, and 3). local inbreeding. Total inbreeding, F(IT), was highest in the North Atlantic regions and lowest along the Mediterranean Coast. The lowest levels were found in Andalusia, Catalunyia, Valencia, and Navarra. Random inbreeding, F(ST), had a similar geographical pattern. Local inbreeding, F(IS), was relatively uniform in the whole of Spain. In towns, random inbreeding dominates over local inbreeding. From the analysis, it emerges that the northwestern area of Spain is the most inbred.
The fitness consequences of inbreeding and outbreeding are poorly understood in natural populations. We explore two microsatellite-based variables, individual heterozygosity (likely to correlate with recent inbreeding) and a new individual-specific internal distance measure, mean d2 (focusing on events deeper in the pedigree), in relation to two measures of fitness expressed early in life, birth weight and neonatal survival, in 670 red deer calves (Cervus elaphus) born on the Isle of Rum between 1982 and 1996. For comparison, we also analyse inbreeding coefficients derived from pedigrees in which paternity was inferred by molecular methods. Only 14 out of 231 calves (6.1%) had non-zero inbreeding coefficients, and neither inbreeding coefficient nor individual heterozygosity was consistently related to birth weight or neonatal survival. However, mean d2 was consistently related to both fitness measures. Low mean d2 was associated with low birth weight, especially following cold Aprils, in which foetal growth is reduced. Low mean d2 was also associated with low neonatal survival, but this effect was probably mediated by birth weight because fitting birth weight to the neonatal survival model displaced mean d2 as an explanatory variable. We conclude that in the deer population fitness measures expressed early in life do not show evidence of inbreeding depression, but they do show evidence of heterosis, possibly as a result of population mixing. We also demonstrate the practical problems of estimating inbreeding via pedigrees compared with a direct marker-based estimate of individual heterozygosity. We suggest that, together, individual heterozygosity and mean d2, estimated using microsatellites, are useful tools for exploring inbreeding and outbreeding in natural population.
Fecundity is usually considered as a trait closely connected to fitness and is expected to exhibit substantial nonadditive genetic variation and inbreeding depression. However, two independent experiments, using populations of different geographical origin, indicate that early fecundity in Drosophila melanogaster behaves as a typical additive trait of low heritability. The first experiment involved artificial selection in inbred and non-inbred lines, all of them started from a common base population previously maintained in the laboratory for about 35 generations. The realized heritability estimate was 0.151 +/- 0.075 and the inbreeding depression was very small and nonsignificant (0.09 +/- 0.09% of the non-inbred mean per 1% increase in inbreeding coefficient). With inbreeding, the observed decrease in the within-line additive genetic variance and the corresponding increase of the between-line variance were very close to their expected values for pure additive gene action. This result is at odds with previous studies showing inbreeding depression and, therefore, directional dominance for the same trait and species. All experiments, however, used laboratory populations, and it is possible that the original genetic architecture of the trait in nature was subsequently altered by the joint action of random drift and adaptation to captivity. Thus, we carried out a second experiment, involving inbreeding without artificial selection in a population recently collected from the wild. In this case we obtained, again, a maximum-likelihood heritability estimate of 0.210 +/- 0.027 and very little nonsignificant inbreeding depression (0.06 +/- 0.12%). The results suggest that, for fitness-component traits, low levels of additive genetic variance are not necessarily associated with large inbreeding depression or high levels of nonadditive genetic variance.
The objectives of this study were to 1) evaluate the genetic diversity of Navajo-Churro sheep using pedigree information; 2) examine the distribution of the Navajo-Churro population; and 3) evaluate the effect of breeder dynamics on genetic conservation of the breed. Pedigree data and breeder information (city and state) were obtained from the Navajo-Churro Sheep Breed Association. Inbreeding coefficients were calculated for each individual animal using pedigree information. A geographic information system program was used to divide the United States into four regions and overlay breeder locations, flock size, and flock inbreeding level. The small correlation between level of inbreeding and flock size (r = -0.07, P = 0.07) indicated that inbreeding levels are not different across flock sizes. The mean flock inbreeding levels ranged from 0 to 11% across regions. The level of inbreeding did not differ among regions (P = 0.15), except for Region 4 (Kansas and Missouri; P = 0.001). The number of breeders registering sheep averaged 34 per year. Most of the breeders were transient, with only eight breeders maintaining ownership for more than 7 yr. Average inbreeding level for 2000 was found to be 1.2%, with a linear increase in inbreeding of 0.1%/yr over the period studied, suggesting a minimal loss of genetic diversity for the Navajo-Churro. However, given the relatively small effective population size (92) and the transient nature of the breeders, development of an ex situ cryo-preserved germplasm bank may be the best long-term strategy for maintaining this breed's genetic diversity.
The Hanoveranian Scenthound is a purebred hound with a long breeding history and an exceptional high hunting performance. The kennel club Hirschmann e. V. has the goal to conserve the very high level of performance in the small population of the Hanoveranian Scenthound. Therefore, the structure of the current population was analyzed by estimating the gene contributions of the founder breeds of other European kennel clubs, coefficients of relationships, rates of inbreeding and coefficients of inbreeding. All current dogs (n = 334) were included in this study. The pedigrees had a degree completeness of 73.3% for 10 generations. The large influence of the Hanoveranian Scenthound bred in the Austrian kennel club was obvious. The gene contribution of these dog population was with 46.5% much more higher than the gene contribution from the German population with 29.8%. The gene contribution of Czech, Swiss and Hungarian dogs was between 4.3% and 7.6%. The mean coefficient of inbreeding of the current German population was 9.2% with the maximum at 15.6%. For the last 10 generations the increase of the coefficient of inbreeding per generation was 0.71%, regarding the degree of completeness of the pedigrees the coefficient of inbreeding per generation increased by 0.96%. For the last generation the rate of inbreeding amounted to 0.09%. The rate of inbreeding expected for the next generation is 0.02%. Also the effective population size (N(e)) in the current population of the Hanoveranian Scenthound being 70.9 exceeds the critical value of N(e) = 50. The mean coefficient of relationship in the current population was 11.2% with the maximum at 73.6%. The present incrossing of Hanoveranian Scenthounds from other European countries reduced the coefficient of inbreeding.
For a population subdivided into an arbitrary number (s) of subpopulations, each consisting of different numbers of separate sexes, with arbitrary distributions of family size and variable migration rates by males (dm) and females (df), the recurrence equations for inbreeding coefficient and coancestry between individuals within and among subpopulations for a sex-linked locus are derived and the corresponding expressions for asymptotic effective size are obtained by solving the recurrence equations. The usual assumptions are made which are stable population size and structure, discrete generations, the island migration model, and without mutation and selection. The results show that population structure has an important effect on the inbreeding coefficients in any generation, asymptotic effective size, and F-statistics. Gene exchange among subpopulations inhibits inbreeding in initial generations but increases inbreeding in later generations. The larger the migration rate, the greater the final inbreeding coefficients and the smaller the effective size. Thus if the inbreeding coefficient is to be restricted to a specific value within a given number of generations, the appropriate population structure (the values of s, dm, and df) can be obtained by using the recurrence equations. It is shown that the greater the extent of subdivision (large s, small dm and df), the larger the effective size. For a given subdivided population, the effective size for a sex-linked locus may be larger or smaller than that for an autosomal locus, depending on the sex ratio, variance and covariance of family size, and the extend of subdivision. For the special case of a single unsubdivided population, our recurrence equations for inbreeding coefficient and coancestry and formulas for effective size reduce to the simple expressions derived by previous authors.
The genetic and demographic consequences of population subdivision have received considerable attention from conservation biologists. In particular, losses of genetic variability and reduced viability and fecundity due to inbreeding (inbreeding depression) are of concern. Studies of domestic, laboratory and zoo populations have shown inbreeding depression in a variety of traits related to fitness. Consequently, inbreeding depression is widely accepted as a fact. Recently, however, the relative impact of inbreeding on the viability of natural populations has been questioned. Work on the cheetah (Acinonyx jubatus), for example, has emphasized the overwhelming importance of environmental factors on mortality in the wild. Here we report that song sparrows (Melospiza melodia) that survived a severe population bottleneck were a non-random subset of the pre-crash population with respect to inbreeding, and that natural selection favoured outbred individuals. Thus, inbreeding depression was expressed in the face of an environmental challenge. Such challenges are also likely to be faced by inbred populations of endangered species. We suggest that environmental and genetic effects on survival may interact and, as a consequence, that their effects on individuals and populations should not be considered independently.