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Genetic and environmental stress, and the persistence of populations.

Many populations of endangered species have to cope both with stressful and deteriorating environmental conditions (mostly the primary cause of the endangerment) and with an increase in homozygosity due to genetic drift and/or inbreeding in small isolated populations. The latter will result in genetic stress often accompanied by a decrease in fitness (inbreeding depression). We have studied the consequences of genetic stress, under optimal as well as stressful environmental conditions, for the fitness and persistence of small populations using Drosophila melanogaster as a model system. The results show that, already under optimal environmental conditions, an increase in homozygosity or inbreeding both impairs fitness and increases the extinction risk of populations significantly. Under environmental stress, however, these effects become greatly enhanced. More important, the results show that the impact of environmental stress becomes significantly greater for higher inbreeding levels. This explicitly demonstrates that genetic and environmental stress are not independent but can act synergistically. This apparent interaction may have important consequences for the conservation of endangered species.

Animals↗

Genome-wide deleterious mutation favors dispersal and species integrity.

Here I develop the idea that ubiquitous harmful genome-wide mutation with local differentiation favors dispersal, even though migration reduces average fitness. Historical contingency of the mutational process means that demes (sub-populations) differentiate from one another. Deleterious or lethal partially recessive mutations carried by migrants then do not encounter similar mutations in the recipient deme. Migrant offspring have higher fitness than offspring of residents, because migrant offspring are heterozygous rather than homozygous for harmful mutations. The advantage is inversely related to local inbreeding depression. Genome-wide deleterious mutation favors the evolution of dispersal, which in turn enhances the genetic integrity of the species.

Animals↗

Changes in mortality patterns and temperature dependence of lifespan in Drosophila melanogaster caused by inbreeding.

After an inbreeding event, lifespan can be curtailed through the expression of deleterious alleles. This will impact on both mortality patterns and interactions with the environment as visualised in reaction norms. We have established the effects of inbreeding on the temperature dependence of lifespan and on mortality patterns in Drosophila melanogaster. Four inbred lines displaying severely decreased lifespan and five outbred controls were assessed for male adult survival at three temperatures. As expected, all inbred lines showed a shorter lifespan than noninbred lines. The mechanisms behind this, however, appeared to be very diverse. Two inbred lines showed a significantly decreased temperature dependence of lifespan compared to the control lines. Analysis of variance on the mortality parameters over all lines showed that inbreeding changes the age-independent mortality but not the age-dependent mortality, whereas temperature does the opposite. This suggests that gene-by-environment interaction caused by inbreeding is the result of changes in the processes of lifespan determination. Importantly, for the two other inbred lines, a particular temperature regime triggered the expression of conditional lethal alleles. Mortality was concentrated in short lethal phases early in adult life. These conditionally expressed lethal alleles affecting lifespan demonstrate line specificity for inbreeding depression and will help ageing studies as such alleles may serve as candidate genes for ageing processes and age-related pathologies in humans.

Animals↗

Do female great reed warblers seek extra-pair fertilizations to avoid inbreeding?

Females of many species mate with several males. According to a recent hypothesis, female promiscuity serves to avoid inbreeding. We tested this hypothesis in a polygynous bird, the great reed warbler (Acrocephalus arundinaceus), in which extra-pair fertilization, inbreeding and inbreeding depression occur. However, the extra-pair males and social males did not differ in relatedness to the promiscuous females, nor did the least related males sire most of the females' chicks. Thus, contrary to recent findings in some insects, birds and reptiles, we found no evidence for inbreeding avoidance among the promiscuous females. Instead, female great reed warblers may seek other potential benefits when cuckolding, such as good gene effects at particular functional genes.

Animals↗

Decline in heterozygosity under full-sib and double first-cousin inbreeding in Drosophila melanogaster.

The effects of inbreeding on heterozygosities and reproductive fitness were determined by carrying out full-sib and double first-cousin inbreeding in Drosophila melanogaster populations for up to 18 generations. Parents were scored each generation for five or six polymorphic enzyme loci, and progeny numbers per pair were recorded. Inbreeding depression, in the form of significant reductions in progeny numbers and significant extinction of lines, was observed. Heterozygosity decreased at a significantly slower rate than predicted, being about 80% of expected. The full-sib and double first-cousin treatments showed similar disagreement with expectations over comparable ranges of inbreeding. Natural selection was shown to favor heterozygotes in the inbred lines. Associative overdominance was the most probable explanation for the slower than expected decline in heterozygosity.

Alleles↗

Inbreeding and relatedness in Scandinavian grey wolves Canis lupus.

Management of small and threatened populations may require detailed knowledge about the genetic status of individuals and the genetic relatedness between individuals. I show here that individual heterozygosity at a set of 29 microsatellite loci correlates closely to the degree of inbreeding in a captive grey wolf population. Microsatellite allele sharing similarly correlates closely to known relatedness between pairs of individuals. Genotyping the same markers in a small (60-70 individuals) natural population of grey wolves in Sweden, low individual heterozygosities and high values of allele sharing between some animals were found. Since inbreeding depression has been documented in a captive grey wolf population of Scandinavian origin, the results point out an additional risk for the small Swedish wild population.

Alleles↗

Inbreeding alters resistance to insect herbivory and host plant quality in Mimulus guttatus (Scrophulariaceae).

Previous studies have demonstrated genetic variation for resistance to insect herbivores and host plant quality. The effect of plant mating system, an important determinant of the distribution of genetic variation, on host plant characteristics has received almost no attention. This study used a controlled greenhouse experiment to examine the effect of self- and cross-pollination in Mimulus guttatus (Scrophulariaceae) on resistance to and host plant quality for the xylem-feeding spittlebug Philaenus spumarius (Homoptera: Cercopidae). Spittlebugs were found to have a negative effect on two important fitness components in M. guttatus, flower production and above ground biomass. One of two M. guttatus populations examined showed a significant interaction between the pollination and herbivore treatments. In this case, the detrimental effects of herbivores on biomass and flower production were much more pronounced in inbred (self) plants. The presence of spittlebug nymphs increased inbreeding depression by as much as three times. Pollination treatments also had significant effects on important components of herbivore fitness, but these effects were in opposite directions in our two host plant populations. Spittlebug nymphs maturing on self plants emerged as significantly larger adults in one of our host plant populations, indicating that inbreeding increased host plant quality. In our second host plant population, spittlebug nymphs took significantly longer to develop to adulthood on self plants, indicating that inbreeding decreased host plant quality. Taken together these results suggest that the degree of inbreeding in host plant populations can have important and perhaps complex effects on the dynamics of plant-herbivore interactions and on mating-system evolution in the host.

Animals↗

Effects of inbreeding and heterosis in Hereford females on fertility, calf survival and preweaning growth.

Effects of inbreeding and heterosis and the difference between them were estimated by comparing linecross (L), topcross (T), inbred (I) and control line (C) Hereford females for reproductive and preweaning growth traits of their progeny. Inbred females (average inbreeding coefficient = 26.5%) originated from four single-sire inbred lines. Control females (average inbreeding coefficient = 6.9%) were produced by a four-sire, 60-cow line. Linecross females were produced from all possible reciprocal crosses of the I lines. Topcross females were produced by mating I bulls to C cows. Differences in pregnancy rate among these lines were not detected. Effects of maternal heterosis were positive for both prenatal and postnatal survival. The weaning rate by L females exceeded the weaning rate by I females as a result. Prenatal survival was reduced in calves from I females relative to those from C females, resulting in corresponding differences in birth and weaning rates. Differences in the magnitudes of maternal heterosis and inbreeding effects were not detected, except for birth weight. For birth weight the effect of maternal inbreeding was of greater magnitude than the effect of maternal heterosis. Within the I and C lines regressions of pregnancy rate, prenatal survival, birth rate, postnatal survival, weaning rate, weaning weight/cow exposed, birth weight and weaning weight on the inbreeding coefficients of the cows indicated significant inbreeding depression on all traits except pregnancy rate and postnatal survival.

Animals↗

The effect of synergistic epistasis on the inbreeding load.

The magnitude of inbreeding depression in Drosophila melanogaster appears too large to be accounted for by mutational load with multiplicative fitness interactions among loci, if current estimates of mutation and selection parameters are valid. One possible explanation for this discrepancy is synergistic epistasis among the fitness effects of deleterious mutations. A simple model of the effect of synergistic epistasis on the inbreeding load is developed. This model is used to show that deleterious mutations could account for the Drosophila data on the effects of inbreeding on components of fitness such as viability.

Animals↗

Host inbreeding increases susceptibility to ectoparasitism.

Inbreeding, which increases homozygosity throughout the genome by increasing the proportion of alleles that are identical by descent, is expected to compromise resistance against parasitism. Here, we demonstrate that host inbreeding increases susceptibility to ectoparasitism in a natural fruit fly (Drosophila nigrospiracula) - mite (Macrocheles subbadius) association, and that this effect depends on host genetic background. Moreover, flies generated from reciprocal crosses between susceptible inbred lines exhibited elevated levels of resistance similar to that in the mass-bred base population, confirming in reverse direction the causative link between expected heterozygosity and resistance. We also show that inbreeding reduces the host's ability to sustain energetically expensive behaviours, and that host exhaustion dramatically increases susceptibility. These findings suggest that inbreeding depression for resistance results from an inability to sustain defensive behaviours because of compromised physiological competence.

Analysis of Variance↗

Effects of bottlenecks on quantitative genetic variation in the butterfly Bicyclus anynana.

The effects of a single population bottleneck of differing severity on heritability and additive genetic variance was investigated experimentally using a butterfly. An outbred laboratory stock was used to found replicate lines with one pair, three pairs and 10 pairs of adults, as well as control lines with approximately 75 effective pairs. Heritability and additive genetic variance of eight wing pattern characters and wing size were estimated using parent-offspring covariances in the base population and in all daughter lines. Individual morphological characters and principal components of the nine characters showed a consistent pattern of treatment effects in which average heritability and additive genetic variance was lower in one pair and three pair lines than in 10 pair and control lines. Observed losses in heritability and additive genetic variance were significantly greater than predicted by the neutral additive model when calculated with coefficients of inbreeding estimated from demographic parameters alone. However, use of molecular markers revealed substantially more inbreeding, generated by increased variance in family size and background selection. Conservative interpretation of a statistical analysis incorporating this previously undetected inbreeding led to the conclusion that the response to inbreeding of the morphological traits studied showed no significant departure from the neutral additive model. This result is consistent with the evidence for minimal directional dominance for these traits. In contrast, egg hatching rate in the same experimental lines showed strong inbreeding depression, increased phenotypic variance and rapid response to selection, highly indicative of an increase in additive genetic variance due to dominance variance conversion.

Analysis of Variance↗

Establishment of inbred strains of chicken and Japanese quail and their potential as animal models.

We started establishing inbred strains of chicken and Japanese quail in 1970. In class Aves, full sib mating is highly difficult due to inbreeding depression. In the chicken, we attempted to establish some inbred strains in three breeds, Black Minorca, White Leghorn and Fayoumi by fixing all the characters that differentiate individuals homozygously. In this paper, we describe some marker genes and characters fixed in the inbred strains of chicken and Japanese quail as well as a calculation of a putative coefficient of inbreeding in 8 chicken inbred strains using band sharing values detected by AFLP analysis. We established generalized glycogenosis type II quail, myotonic dystrophy quail, neurofilament deficient quail, visually impaired chicken, double oviduct chicken with partial kidney deficiency, chicken showing spontaneous lymphocytic thyroiditis with feathered amelanosis, and chicken with a hereditary nervous disorder. The processes of establishment and characteristics of these animal models are described with some interesting information obtained from these animal models. In generalized glycogenosis type II quail, the results of enzyme replacement therapy and gene therapy are described.

Animals↗

Inbreeding effects on lifetime in David's deer (Elaphurus davidianus, Milne Edwards 1866) population.

The current population of David's deer is derived from 18 individuals kept in Woburn Abbey Park (England). The aim of this study is to evaluate the inbreeding rate as well as inbreeding depression in longevity. The recorded data have been extracted from the International Species Information System (ISIS). Complete records of 2042 individuals (born in 1947-2000) from zoological gardens were studied. The following four subsets of data were formed: all individuals, individuals over 31 days of life, sexually mature individuals (above 450 days old) and individuals with identified sex. Two models (including inbreeding coefficient as linear and quadratic covariables, respectively) have been employed. These computations were performed by the use of the DFREML package programs. This study has shown that average levels of inbreeding in the David's deer are relatively low (no exceed 0.028). The highest level of inbreeding was registered for mature individuals. In general, the average inbreeding on length of life was small for the above mentioned the inbreeding level (from 24 days to 77 days). However, on the basis estimated regression coefficients it can be suggested that an increase of inbreeding could lead to a drastic reduction of longevity. Relationships between inbreeding level and longevity are usually better described by quadratic partial regression (except for the oldest individuals). On the other hand, from a statistical point of view, a relatively low inbreeding level of the population studied is not suitable to derive the slope of the dependencies.

Animals↗

Selection in Hereford cattle. II. Expected and realized response.

Data from 14 inbred lines and 14 linecross groups of Hereford cattle at the San Juan Basin Research Center, Hesperus, were used to evaluate expected and realized response in birth and weaning traits and postweaning traits in males and females over a 28-yr period. There were large differences in the means and variances of the performance traits among the inbreds and linecrosses, with the inbreds showing inbreeding depression and greater variability among lines, while the linecrosses manifested within-breed heterosis. Except for gain from weaning to 12 mo, in females, genetic progress was expected in all traits studied, mainly due to sire selection. Regressions of annual trait means on years indicated positive phenotypic trends in the inbreds for heart girth circumference at birth, adjusted weaning weight (adjusted for inbreeding), weaning score, final weight, feed consumption and the yearling weights and gains in females. Changes were negative for other traits. In the linecross group phenotypic trends were positive in all traits except heart girth circumference, weaning age, initial test weight and feed efficiency. Estimated genetic progress per generation due to within-line selection was negative in most of the traits in the inbreds but was considerably positive for the linecrosses for most of the traits. As expected, between-line selection yielded greater genetic improvement in the inbred than in the linecross population. The different patterns of response in the two populations are attributed to high rates and levels of inbreeding. Although variable, the actual progress was below prediction in most of the traits studied.

Animals↗

Kin competition, the cost of inbreeding and the evolution of dispersal

Dispersal is often presented as a mechanism to avoid competition among relatives and inbreeding depression. However, the formal analysis of the effects of both these factors on the evolution of dispersal has only been conducted in few studies with strong restrictive assumptions. In this paper, I first derive the evolutionary stable dispersal rate as a function of three parameters: (1) the cost of dispersal, c, (2) the coefficient of relatedness among randomly chosen offspring, R, and (3) the cost of inbreeding, delta. In a second step, relatedness is used as a dynamical variable for the derivation of the evolutionarily stable dispersal rate. Finally, in a third step, relatedness and the cost of inbreeding are assumed to be dynamical variables. This allows to analyse the more realistic situation where dispersal, relatedness and the cost of inbreeding are coevolving simultaneously. Several subcases are considered depending on the genetic determinism (haploid or diploid), the control of the dispersal strategy (parent or offspring control of dispersal) and the plasticity of dispersal with sexes (with or without sex-specific dispersal rates). This analysis clarifies the role of the cost of inbreeding and kin competition on the evolution of dispersal (in particular on the evolution of sex-biased dispersal rates) and leads to quantitative and testable predictions. Copyright 1999 Academic Press.

Journal Article↗

Genetics and conservation biology.

Conservation genetics encompasses genetic management of small populations, resolution of taxonomic uncertainties and management units, and the use of molecular genetic analyses in forensics and to understanding species' biology. The role of genetic factors in extinctions of wild populations has been controversial, but evidence now shows that they make important contributions to extinction risk. Inbreeding has been shown to cause extinctions of wild populations, computer projections indicate that inbreeding depression has important effects on extinction risk, and most threatened species show signs of genetic deterioration. Inappropriate management is likely to result if genetic factors are ignored in threatened species management.

Animals↗

Inbreeding and copulatory behavior in house mice: a further consideration.

Comparisons were made of the copulatory behavior of randomly bred (one population: WRL) and inbred wild (five strains: PAA, ab, ac, ad, and PAE) male house mice. All inbred and randomly bred stocks were derived from a single foundation population. The inbred males tended to have shorter latencies to the first mount and intromission, longer latencies to ejaculation, and more preejaculatory mounts and thrusts than randomly bred males. All these effects parallel those observed in a previous study in which a wild population was compared with various domestic inbred strains. If inbreeding depression is related to adaptive significance, these data suggest that, although rapid initiation of copulation in a novel environment may not be adaptive, it may be adaptive for mice to ejaculate rapidly once copulation is initiated.

Adaptation, Psychological↗