Is inbreeding depression more severe in a stressful environment?

Successful reintroduction of endangered species depends in part on their ability to respond to changing environmental conditions. Population genetics theory suggests that inbred populations lacking genetic variability may be unable to respond effectively to environmental stress. There have been very few studies designed explicitly to investigate the phenomenon of inbreeding depression under environmental stress, particularly in the context of conservation genetics. Three separate experiments using Drosophila melanogaster were designed to explore this issue. No increase in the magnitude of inbreeding depression was detected in laboratory lines subjected to three generations of continuous full-sib mating under temperature stress (28°C), lead stress (medium contaminated with 400 ppm Pb), or a combination of these stresses. Individual isofemale lines from a different population, however, did show significant increases in inbreeding depression when exposed to temperature stress for one generation of full-sib mating following three generations of full-sib inbreeding at 25°C. Further, chromosome-2 homozygotes showed, on average, a significant increase in inbreeding depression under lead stress when in competition with corresponding chromosome-2 heterozygotes compared to the same lines in a benign environment. Taken together, these results suggest that inbreeding depression is more severe under conditions of environmental stress and is more likely to be realized in an inter- or intraspecific competitive situation as can be experienced in the wild. Therefore, it is likely that reduced genetic variability through inbreeding is a much greater problem for recently reintroduced populations than it is for populations in a relatively benign zoo environment. © 1994 Wiley-Liss, Inc.     

Selection for non-diapause in the bumblebee Bombus terrestris, with notes on the effect of inbreeding

Bumblebees are generally believed to be annual insects. However, here we will show that under laboratory conditions the bumblebee Bombus terrestris (L.) can produce a second generation without a period of cold storage (diapause) or CO₂ narcosis (a method to break diapause). It is also shown that this so-called non-diapause trait can be selected for. The percentage of non-diapausing queens increased from 8% (minimum) to 97% (maximum) in two generations of selection. However, it was not possible to maintain isofemale non-diapause lines. Colonies of the fourth and fifth generation remained small (expressed in worker number) and produced a small number of queens. Also the percentage of queens that started laying eggs (defined as the percentage non-diapause) decreased in the fourth and fifth generations. To study whether this decline of the non-diapause lines was caused by inbreeding, a control experiment was conducted. In this control experiment queens were mated with their brothers (full-sib mating) for several generations and the number of queens that start egg laying was measured. This revealed that inbreeding can have a negative effect on the egg-laying capacities of queens thus causing the decline of inbred (non-diapause) lines.     

Heterosis, sex and maternal interactions in crosses among inbred lines of mice.

All possible crosses and reciprocals were made among four inbred lines (F = 92%) developed from 12 generations of full-sib mating. All lines originated from a common outbred base population of ICR-albino mice. Data were obtained from 356 litter containing 2,734 mice to evaluate heterosis, reciprocal effects, sex effects and their interactions as they affect body weight and weight gain. Heterosis was significant for most of the postweaning traits (42- and 56-day weight and gain from 21 to 42 days). Nonadditive gene action may have included overdominance and epistasis since both reciprocal linecrosses were generally heavier than those of the better inbred lines. Although significant differences in reciprocals and inbred lines were not frequent, there were sufficient differences to indicate that lines varied in the fixation of loci during inbreeding. Sex-heterosis interactions were significant for 12 of 30 possible cases. However, eight of the 12 significant interactions occurred in crosses involving only one of the lines. The interactions were of the divergent type and arose from males exhibiting more heterosis than females. Overdominance in genes on the sex chromosomes modified by other loci (epistasis) was proposed as a possible explanation for these results. Some sex-linkage affecting growth was evident from the interaction of sex with reciprocal effects.     

Effect of initial size on subsequent growth and carcass characteristics of divergently selected channel catfish

Summary Two experiments were conducted simultaneously to determine (1) if fast-growing fingerlings of channel catfish, Ictalurus punctatus, could be identified by simple visual selection of body size and (2) if initial size advantages influenced subsequent growth and carcass traits of divergently selected channel catfish. Exp. 1 included large (L), medium (M), and small (S) fingerling sizes from each of the control (C), selected upward (+) and selected downward (–) lines for body weight. Exp. 2 included all fmgerlings of the same size (25±5 g) from the 3 lines. Catfish from the L size-class, within each full-sib family in each line, were consistently heavier and longer than M and S size-classes throughout the 53-week experimental period. Fingerlings from the M size-class were also superior in growth to those from the S size-class. Catfish from the + line exceeded those from the C and –lines in body weight and total length at the conclusion of Exp. 1 but not in Exp. 2. This was attributed to the selection of equal size fmgerlings in Exp. 2 which may have excluded fingerlings with the best growth potential from the + body weight line. Results of the two experiments combined indicated that one generation of divergent selection has created genetic differences among lines of channel catfish.Supported by State and Hatch funds allocated to the Georgia Agricultural Experiment Station     

Inbreeding avoidance through kin recognition: choosy females boost male dispersal

Inbreeding avoidance is predicted to induce sex biases in dispersal. But which sex should disperse? In polygynous species, females pay higher costs to inbreeding and thus might be expected to disperse more, but empirical evidence consistently reveals male biases. Here, we show that theoretical expectations change drastically if females are allowed to avoid inbreeding via kin recognition. At high inbreeding loads, females should prefer immigrants over residents, thereby boosting male dispersal. At lower inbreeding loads, by contrast, inclusive fitness benefits should induce females to prefer relatives, thereby promoting male philopatry. This result points to disruptive effects of sexual selection. The inbreeding load that females are ready to accept is surprisingly high. In absence of search costs, females should prefer related partners as long as delta相似文献   

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

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

Effect of rate of inbreeding on inbreeding depression in Drosophila melanogaster

Summary This experiment was designed to study the relationship between rate of inbreeding and observed inbreeding depression of larval viability, adult fecundity and cold shock mortality in Drosophila melanogaster. Rates of inbreeding used were full-sib mating and closed lines of N=4 and N=20. Eight generations of mating in the N=20 lines, three generations in the N=4 lines and one generation of full-sib mating were synchronised to simultaneously produce individuals with an expected level of inbreeding coefficient (F) of approximately 0.25. Inbreeding depression for the three traits was significant at F=0.25. N=20 lines showed significantly less inbreeding depression than full-sib mated lines for larval viability at approximately the same level of F. A similar trend was observed for fecundity. No effect of rate of inbreeding depression was found for cold shock mortality, but this trait was measured with less precision than the other two. Natural selection acting on loci influencing larval viability and fecundity during the process of inbreeding could explain these results. Selection is expected to be more effective with slow rates of inbreeding because there are more generations and greater opportunity for selection to act before F=0.25 is reached. Selection intensities seem to have been different in the three traits measured. Selection was most intense for larval viability, less intense for fecundity and, perhaps, negligible at loci influencing cold shock mortality.     

The Effect of Inbreeding on Competitive Male-Mating Ability in DROSOPHILA MELANOGASTER

The effect of full-sib inbreeding on competitive male-mating ability (CI♂) in Drosophila melanogaster was investigated in two experiments. In the first, five inbred lines (with reserves) were assessed up to 18 generations. Linear inbreeding depression, of 5.9% per 10% increase in homozygosity, was observed. In a second experiment, 21 inbred lines were tested after three generations of full-sib mating (without reserves), and the decline with inbreeding was more severe, the male competitive index (CI♂) decreasing by 10.7% per 10% increase in F. The difference between these results is attributed to natural selection acting on variation within the inbred lines in extent of homozygosity, which can arise because of the peculiarly strong influence of linkage in Drosophila. Furthermore, differentiation between the lines may have reflected this variation rather than the various effects of different alleles fixed.—These results imply that the genetic variation in male-mating ability is largely due to dominance (no epistasis was detected) and are consonant with the proposition that intermale sexual selection is a very important component of fitness in D. melanogaster . There was no evidence of a positive correlation between male body size and competitive mating ability.     

INBREEDING: ITS EFFECT ON RESPONSE TO SELECTION FOR PUPAL WEIGHT AND THE HERITABLE VARIANCE IN FITNESS IN THE FLOUR BEETLE,TRIBOLIUM CASTANEUM

We report our studies of the effect of inbreeding on the response to selection for increased pupal weight in the flour beetle, Tribolium castaneum. We also report the effects of inbreeding and selection for pupal weight on the heritable variation in fitness and fitness components. We created replicate and independent inbred lines with F-values of 0.00, 0.375, and 0.672, by 0, 2, and 5 generations, respectively, of brother-sister mating of adult beetles from an outbred stock population. Subsequently, we imposed artificial within-family selection for increased pupal weight in each of 15 inbred lines for eight generations; each line had its own paired, unselected control. We compared the response to selection across the three levels of inbreeding with theoretical expectation, and investigated the effects of inbreeding and selection on fitness variation among families within all 30 selected and control lines. Among-line variation in pupal weight increased with increased inbreeding prior to selection but diminished with directional selection. Inbreeding reduced the realized heritability of pupal weight concordant with quantitative predictions of additive theory. Mean fitness, measured in several ways, declined with inbreeding and declined further with selection. In contrast, the genetic variation for fitness in the inbred and selected lines lines equalled or exceeded that of the outbred controls. Our results suggest that inbreeding and selection may affect traits in different ways depending on the relative amounts of additive and nonadditive genetic variation.     

Trait mean depression for second-generation inbred strawberry populations with and without parent selection

 Strawberry genotypes selected for superior fruit yield or chosen at random from first-generation self, full-sib, and half-sib populations were crossed to provide second-generation inbred progenies and composite cross-fertilized control populations. Mean yields for inbred offspring from crosses among selected parents exceeded those from the offspring of unselected parents by 87%, 23%, and 37% for self, full-sib, and half-sib populations, respectively; yields for offspring from unrelated crosses among selected parents were 54% larger than those for crosses among unselected parents. Selection for yield also resulted in significant correlated response for fruit number and plant diameter. Mean yields for second-generation half-sib and full-sib offspring from selected parents were greater than those for offspring from the unselected but non-inbred control population. This suggests that selection can be a powerful force in counteracting most of the inbreeding depression expected in cross-fertilized strawberry breeding programs. Selection treatment× inbreeding rate interactions were non-significant for all traits; thus, selection among partially inbred offspring did not have a large effect on the rate of genetic progress. Differential realized selection intensity among individuals with differing levels of homozygosity accumulated due to inbreeding is suggested as the most likely explanation for the absence of association between pedigree inbreeding coefficients and cross performance detected previously in strawberry. Received: 21 July 1996 / Accepted: 7 March 1997     

Effect of recombination in the parent populations on the means and combining ability variances in hybrid populations of maize (<Emphasis Type="Italic">Zea mays</Emphasis> L.)

Recombination of selected genotypes plays a key role in plant breeding for generating new base populations. We investigated the influence of recombination in two parent populations on the means and combining ability variances of their hybrid population by (1) quantitative genetic theory and (2) experiments with maize. The two parent populations were founded by four early flint and four early dent inbred lines, respectively. Each population was studied in three generations: Syn-0, the four inbred lines themselves; Syn*-1, the six intrapool single crosses (SC); and Syn*-2, the three intrapool double crosses (DC). Four interpool hybrid populations were created: (1) all 16 SC and (2) all 36 DC were produced from generations Syn-0 and Syn*-1, respectively, (3) 168 biparental progenies (BIP) of type flint x dent (female x male), and (4) 168 BIP of type dent x flint were produced according to NC-design I with randomly sampled plants of generation Syn*-2. The half-sib and full-sib families obtained in this manner were evaluated for grain yield, dry matter concentration and plant height. According to theoretical results, differences in the population means of these hybrid populations indicate the presence of various types of epistasis. Changes in combining ability variances from SC to DC reflect different levels of parental inbreeding (F = 1 vs F = 0), whereas changes from DC to BIP only reflect the effects of recombination and are attributable to covariances between additive and dominance effects caused by linkage disequilibrium in the Syn-0 generations. The experimental results showed a significant decline in yield from DC to BIP due to a loss of gene combinations with favourable epistatic effects. Estimates of sigma(2)(GCA) attributable to flint or dent lines decreased or remained unchanged from SC to DC, but generally increased in the BIP populations. The consequences of these trends for developing improved interpool hybrids are discussed.     

Increase of the spontaneous mutation rate in a long-term experiment with Drosophila melanogaster

In a previous experiment, the effect of 255 generations of mutation accumulation (MA) on the second chromosome viability of Drosophila melanogaster was studied using 200 full-sib MA1 lines and a large C1 control, both derived from a genetically homogeneous base population. At generation 265, one of those MA1 lines was expanded to start 150 new full-sib MA2 lines and a new C2 large control. After 46 generations, the rate of decline in mean viability in MA2 was approximately 2.5 times that estimated in MA1, while the average degree of dominance of mutations was small and nonsignificant by generation 40 and moderate by generation 80. In parallel, the inbreeding depression rate for viability and the amount of additive variance for two bristle traits in C2 were 2-3 times larger than those in C1. The results are consistent with a mutation rate in the line from which MA2 and C2 were derived about 2.5 times larger than that in MA1. The mean viability of C2 remained roughly similar to that of C1, but the rate of MA2 line extinction increased progressively, leading to mutational collapse, which can be ascribed to accelerated mutation and/or synergy after important deleterious accumulation.     

Isozyme polymorphisms maintained by lethal loci in inbred strains of Aedes triseriatus

We analyzed the progeny of brother-sister matings of two inbred strains of the treehole mosquito, Aedes triseriatus, at several isozyme loci to determine which factors were maintaining heterozygosity at these loci after prolonged inbreeding. The triseriatus Kramer (TK) lines were polymorphic for the Odh and Hbd loci after 12 generations of full-sib mating, and the triseriatus Vero Beach (TV) lines were polymorphic for the Odh, Hk-4, Pgm, and Hbd loci after 14 generations of full-sib mating. Genetic analysis of 22 TK F13 matings and nine TV F15 matings showed that heterozygosity was enforced by lethal loci to which all the polymorphic isozyme loci were linked. The lethal loci formed a balanced lethal system. We determined the relative positions and map distances of lethals and enzyme loci.     

Effects on heterozygosity and reproductive fitness of inbreeding with and without selection on fitness in Drosophila melanogaster

The effects of inbreeding, with (IS) and without selection (IO) for reproductive fitness, on inbreeding depression and heterozygosity were evaluated in 20 lines of each treatment inbred over seven generations using full-sib mating. The survival of lines was significantly greater in IS (20/20) than in IO (15/20). The competitive index measure of reproductive fitness was significantly lower in the inbred lines than in the outbred base population, but not significantly different in surviving IS and IO lines. There was a trend for higher fitness in the IS treatment as relative fitnesses were 19% higher in IS than IO for surviving lines and 59% higher for all lines. Heterozygosities were lower in the inbred lines than in the base population, and significantly higher in the IS than the IO lines. Consequently, the reduction of inbreeding depression in IS has been achieved, at least in part, by slowing the rate of fixation.     

Variability of localization of retrotransposon copia ant its effect on adaptation in inbred strains of Drosophila melanogaster with the different rate of transposition

Three sublines of an inbred laboratory line of Drosophila melanogaster with the initial copia transposition rate 2 x 10(-2), 2 x 10(-3), and 5 x 10(-4) per copy per generation were reared for several dozen generations under conditions of low effective population size (by full-sib crosses or in a small mass culture of 10 females x 10 males). All six lines were tested for the transposition rate, location pattern, and copy number of copia in euchromatic genome regions and for fitness inferred from the intraspecific competition index. The copia transposition rate remained constant in both versions of the lines with an initially lower rate and decreased by an order of magnitude in both versions of the line with an initially higher rate. New copia insertions behaved as selectively neutral and were accumulated in the genome. Each new copy decreased fitness by less than 1% on average. Some of the existing unfixed insertions remained segregating after long-term inbreeding and were assumed to provide a selective advantage to heterozygotes.     

THE EFFECTS OF FIVE GENERATIONS OF ENFORCED SELFING ON POTENTIAL MALE AND FEMALE FUNCTION IN MIMULUS GUTTATUS

In prior work we detected no significant inbreeding depression for pollen and ovule production in the highly selfing Mimulus micranthus, but both characters showed high inbreeding depression in the mixed-mating M. guttatus. The goal of this study was to determine if the genetic load for these traits in M. guttatus could be purged in a program of enforced selfing. These characters should have been under much stronger selection in our artificial breeding program than previously reported characters such as biomass and total flower production because, for example, plants unable to produce viable pollen could not contribute to future generations. Purging of genetic load was investigated at the level of both the population and the individual maternal line within two populations of M. guttatus. Mean ovule number, pollen number, and pollen viability declined significantly as plants became more inbred. The mean performance of outcross progeny generated from crosses between pairs of maternal inbred lines always exceeded that of self progeny and was fairly constant for each trait through all five generations. The consistent performance of outcross progeny and the universally negative relationships between performance and degree of inbreeding are interpreted as evidence for the weakness of selection relative to the quick fixation of deleterious alleles due to drift during the inbreeding process. The selective removal (purging) of deleterious alleles from our population would have been revealed by an increase in performance of outcross progeny or an attenuation of the effects of increasing homozygosity. The relationships between the mean of each of these traits and the expected inbreeding coefficient were linear, but one population displayed a significant negative curvilinear relationship between the log of male fertility (a function of pollen number and viability) and the inbreeding coefficient. The generally linear form of the responses to inbreeding were taken as evidence consistent with an additive model of gene action, but the negative curvilinear relationship between male fertility and the inbreeding coefficient suggested reinforcing epistasis. Within both populations there was significant genetic variation among maternal lineages for the response to inbreeding in all traits. Although all inbred lineages declined at least somewhat in performance, several maternal lines maintained levels of performance just below outcross means even after four or five generations of selfing. We suggest that selection among maternal lines will have a greater effect than selecting within lines in lowering the genetic load of populations.     

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.     

Heterozygosity in inbred strains of the tree-hole mosquitoAedes triseriatus

Inbred stocks of the tree-hole mosquito Aedes triseriatus from four localities were developed using full-sib mating. The progress of inbreeding was followed electrophoretically at eight variable and six less variable enzyme loci. Rates of fixation of several of these loci were substantially lower than expected. Discrepancies between observed and expected fixation values were evident in the early stages of inbreeding and became larger as inbreeding progressed. Odh, Hbd, Pgm, and Hk-4 were usually not fixed. By the F12 and F14 generations of brother-sister mating, most individuals in the two lines were heterozygotes (Odh and Hbd in the TK lines and Odh, Hbd, Pgm, and Hk-4 in the TV lines). The probability of maintaining heterozygosity at several selectively neutral and unlinked loci simultaneously is very low. Odh, Hbd, Pgm, and Hbd loci are linked to lethal recessives on chromosome 2, creating a balanced lethal system which in turn accounts for the heterozygosity in these inbred mosquitoes.     

Effects of Ancestral X Irradiation Followed by Random Mating on Body Weight of Rats

Effects of nine generations of 450r per generation of ancestral spermatogonial X irradiation of inbred rats on body weight were examined. After six generations of random mating (avoiding inbreeding) following the termination of irradiation, descendants of irradiated males (R) were significantly lighter than their controls (C) at 3 and 6 weeks, but not at 10 weeks of age. However, differences in growth between R and C populations were small. Among-litter and within-litter variance estimates were generally larger in the R lines than in the C lines, suggesting that selection responses would be greater in R than in C lines. In conjunction with previous evidence—obtained during the irradiation phase of the experiment—this suggested that more rapid response to selection for 6-week body weight, in particular, might accrue in the R lines.