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PubMed2003

Selfing results in inbreeding depression of growth but not of gas exchange of surviving adult black spruce trees.

In most tree species, inbreeding greatly reduces seed production, seed viability, survival and growth. In a previous large-scale quantitative analysis of a black spruce (Picea mariana (Mill.) B.S.P.) diallel experiment, selfing had large deleterious effects on growth but no impact on stable carbon isotope discrimination (an indirect measure of the ratio of net photosynthesis (A) to stomatal conductance (gwv)). It was hypothesized that selfing has no effect on carbon (C) fixation at the leaf level but impairs subsequent utilization of C. Alternatively, A and gwv may be impacted by selfing to the same extent. However, no gas exchange data were collected to test these hypotheses. We have now obtained photosynthetic gas exchange data from three selfed families and three outcrossed families (all the result of controlled pollination) from the same diallel experiment. Photosynthetic responses to intercellular CO2 concentration (A-Ci curves) were generated on four replicates per family, one block per day, over a 4-day period in July. There were no differences between selfed and outcrossed families in maximum carboxylation rate, maximum electron transport, A or gwv (both estimated at 370 ppm CO2), or the ratio A/gwv. Because selfed trees had higher mortality than outcrossed trees during the experiment, we cannot exclude the possibility that previously living selfed progeny had low A. Nevertheless, the data indicate that inbreeding can result in trees that have low productivity despite high A, supporting our hypothesis that gas exchange is similar between selfed and outcrossed progeny trees. We conclude that utilization of fixed C is modified in the surviving selfed progeny.

Inbreeding↗
PubMed2002

Inbreeding depression and male fitness in black grouse.

The male lifetime lekking performance was studied, and related to inbreeding-outbreeding in a wild population of black grouse (Tetrao tetrix) in central Finland between 1989 and 1995. Inbreeding was measured as the mean heterozygosity and mean d(2) of 15 microsatellite loci. We found a significantly positive relationship between mean d(2) and lifetime copulation success (LCS), while the relationship between heterozygosity and LCS was close to significant. We also found that males that never obtained a lek territory had significantly lower mean heterozygosity than males that were observed on a territory at least during one mating season in their life. Furthermore, among males that were successful in obtaining a lek territory, LCS and mean d(2) were highest for those males that held central territories. We suggest that inbred males have a disadvantage (or outbred males have an advantage) in the competition for territories that may explain the relationships with LCS and inbreeding. Furthermore, the fact that mean d(2) was positively correlated with LCS whereas heterozygosity was not when we restricted the analysis to territorial males, suggests that mean d(2) provides more information about levels of inbreeding-outbreeding than heterozygosity alone, and potentially highlights the effects of heterosis. To our knowledge, this is the first time that measures of inbreeding and lifetime fitness have been linked in a non-isolated population. This is important in establishing that the relationships found in previous studies are not artefacts of low gene flow created by limited dispersal but a general feature of wild vertebrate populations.

Animals↗
PubMed1999

Inbreeding and inbreeding depression on reproduction and production traits of White Leghorn lines selected for egg production traits.

1. The rate of inbreeding and its effect on reproduction and egg production traits were studied in White Leghorn lines selected for egg production traits. The experiment was carried out for 10 generations in a control line (C) and in lines selected for increased egg number (EN), egg weight (EW) and egg mass (EM). 2. Data were available on reproduction traits, such as percent fertile eggs (PF), percent hatched of fertile eggs (PHF) and percent hatched of total eggs set (PHT), and on egg production traits such as age at 1st egg (AFE), egg number and egg weight. 3. The rate of increase in average inbreeding per generation was 1.50, 1.24, 1.14 and 0.18% for the line EN, EW, EM and C, respectively. The effect of inbreeding on reproduction and production traits was estimated by including the inbreeding coefficient of the hen (Fh), embryo (Fe) and mate (Fs) as a partial linear regression in the model. 4. There was a significant effect of inbreeding on reproduction traits in line EW attributable to the inbreeding of the hen, embryo and mate. No such effect was observed in the other lines. 5. In all lines inbreeding tended to reduce egg number and delay sexual maturity. In general, all lines reacted differently to inbreeding.

Animals↗
PubMed1993

Inbreeding depression and male-mating behavior in Drosophila melanogaster.

There have been relatively few studies designed to investigate the effects of inbreeding on behavioral traits. To study this phenomenon, five experimental lines of Drosophila melanogaster made isogenic for chromosome 2 were evaluated for their male-mating ability and, subsequently, male courtship behavior. All lines showed significant reductions in overall mating ability, and males from all of these lines displayed impaired mating behavior, with two lines displaying particularly aberrant courtship patterns. Line 16 displayed an inability to successfully initiate copulation following successful courtship, while line 17 displayed significant reduction in locomotor activity, resulting in virtually no successful courtship or copulatory activity. The implications of these findings for competitive mating ability in wild Drosophila populations are presented. Further, the importance of mating success as a fitness component in the management of potentially highly inbred populations of endangered species is discussed.

Animals↗
PubMed2005

Inbreeding depression of female fecundity by genetic factors retained in natural populations of a male-haploid social mite (Acari: Tetranychidae).

We previously determined that certain recessive genes decrease female fecundity in a haplo-diploid spider mite, Stigmaeopsis miscanthi (Saito). However, whether the depression was caused by the breakdown of heterosis or the expression of deleterious genes retained in a population could not be determined, because we had started our inbreeding experiment from a mixture of two isolated populations. In order to answer this basic question, inbreeding effects on survival and fecundity were measured for eight small populations occurring far from the two initial populations. There was little depression of immature survival of inbred lineages in all populations. On the other hand, in two inbred lineages, both originating from the smallest populations, female oviposition decreased significantly with the increase of Wrights f-value, showing that mildly deleterious genes are actually retained even in natural populations of haplo-diploid organisms.

Animals↗
PubMed1996

Age-specific inbreeding depression and components of genetic variance in relation to the evolution of senescence.

Two major theories of the evolution of senescence (mutation accumulation and antagonistic pleiotropy) make different predictions about the relationships between age, inbreeding effects, and the magnitude of genetic variance components of life-history components. We show that, under mutation accumulation, inbreeding decline and three major components of genetic variance are expected to increase with age in randomly mating populations. Under the simplest version of the antagonistic pleiotropy model, no changes in the severity of inbreeding decline, dominance variance, or the genetic variance of chromosomal homozygotes are expected, but additive genetic variance may increase with age. Age-specific survival rates and mating success were measured on virgin males, using lines extracted from a population of Drosophila melanogaster. For both traits, inbreeding decline and several components of genetic variance increase with age. The results are consistent with the mutation accumulation model, but can only be explained by antagonistic pleiotropy if there is a general tendency for an increase with age in the size of allelic effects on these life-history traits.

Aging↗
PubMed1994

The interaction of inbreeding depression and environmental stochasticity in the risk of extinction of small populations.

Current population genetic and population dynamic models are inappropriate to judge the risk of extinction of small populations due to the combined effects of inbreeding, genetic drift, demographic stochasticity, and environmental stochasticity. Instead, a model based on the aggregated fates of individuals is advocated. The unequal distribution of resources over individuals is an essential part of this model. The model allows the incorporation of the mutation-selection dynamics of alleles leading to inbreeding effects and to fixation of slightly deleterious mutations as a result of genetic drift. The slightly deleterious mutations lower the conversion of resources into offspring. Whereas lethal alleles are rapidly eliminated by selection in small populations, the selection against mild deleterious effects depends strongly on effective population size and on the social system, that is, on the division of resources among individuals. The model allows for the study of rates at which processes occur while far away from equilibrium, which is crucial in understanding the extinction risks of threatened populations. One example of the latter is illustrated in simulations in which small populations become extinct between approximately 100 and 200 generations after they became small populations, due to a gradual accumulation of mildly deleterious mutations.

Alleles↗
PubMed1991

Coevolution of self-fertilization and inbreeding depression. III. Homozygous lethal mutations at multiple loci.

We study the evolution of the rate of self-fertilization in response to deleterious mutations at multiple loci. Although partial selfing induces associations among loci even in the absence of linkage, associations among mutations at different loci are of a smaller order of magnitude than the mutation rate. Genotypes that carry homozygous lethal mutations in heterozygous form at i loci occur in frequencies of the order (Ti) mu i, in which T denotes the number of viability loci and mu the mutation rate. While associations between mutations at different loci remain small even under inbreeding, each viability locus develops an association with the modifier of the rate of self-fertilization that substantially affects the evolution of the breeding system. Positive associations between enhancers of selfing and haplotypes carrying multiple wild-type alleles and positive associations in heterozygosity between the modifier locus and the viability loci promote evolutionary increases in the rate of self-fertilization.

Biometry↗
PubMed2005

Long-term responses, changes in genetic variances and inbreeding depression from 122 generations of selection on increased litter size in mice.

Data on mice selected for litter size over 122 generations have been analysed in order to reveal the effect of long-term selection on responses and changes in variances over a long selection period. Originally, three lines were established from the same base population, namely an H line selected for large litter size, an L line selected for small litter size and a K line without selection. In generation 122, the mean number of pups born alive (NBA) was 22 for the H line and 11 for the K line. Phenotypic response to selection is reduced over generations, but crossing of plateaued lines increased responses and realized heritabilities. Both realized heritabilities and heritabilities from residual maximal likelihood (REML) analyses were, in general, calculated from generation (-1)-44 (period 1), 45-70 (period 2) and 71-122 (period 3) separately. Realized heritabilities were in general smaller than heritabilities estimated from mixed model analysis. An overall estimate of heritability for NBA was found to be 0.19 (+/- 0.01) by REML analysis. Additive variance is constant over all periods in the high line and the control line, but is reduced over periods in the low line. The reduction of additive variance in the low line could probably be explained by changes in gene frequencies. In all lines, environmental variances increased over periods. Inbreeding reduced the mean litter size by 0.72 (+/- 0.10) pups per 10% increase in inbreeding, with substantial variance between periods and lines.

Animals↗
PubMed2006

Inbreeding depression, increased phenotypic variance, and a trade-off between gonads and appendages in selfed progeny of the aphid Prociphilus oriens.

Trade-offs are potentially common among two or more traits whose development is dependent on the same resources. To detect genetic trade-offs, the techniques of quantitative genetics, pedigree analyses, and selection experiments have been used. This study demonstrates genetically based trade-offs between gonads and appendages in hatched larvae of the aphid Prociphilus oriens by focusing on enlarged variance among the families of selfed progeny. The selfed and outbred families were compared in respect to the size of morphological traits, gonad volume, and hatch dates as well as egg volume. Selfing not only increased the among-family variance component in all larval traits examined, but it also increased the mean size of all the morphological traits significantly. In contrast, gonad volume, a fitness component, was reduced with selfing. Calculation of the allometry (log-transformed regression) of larval traits to egg volume indicated that in the outbred group, morphological traits grew slowly relative to egg volume with slopes below 0.25, whereas gonads exhibited isometric growth. With selfing, most morphological traits had significantly steeper slopes, whereas the slope for gonads was greatly decreased. When the effect of egg volume was statistically removed from the means of selfed families, significant negative correlation was detected between the adjusted means of gonad volume and those of tibia length. This result suggests genetic trade-offs between gonad volume and tibia length. Thus, the evidence implies that at the loci governing the development of appendages, the dominant alleles function to canalize the development of tibiae into an optimal size, irrespective of egg volume. It is hypothesized that increased homozygosity of the deleterious recessive alleles reduced gonad volume through increasing the resource allocation to tibiae. The hypothesis of the gonad-appendage trade-off could be applied to explain the phenotypic evolution in some aphid species.

Animals↗
PubMed2000

Use of computerized mate selection programs to control inbreeding of Holstein and Jersey cattle in the next generation.

The expected role of computerized mate selection programs with regard to inbreeding and lifetime profitability of Holstein and Jersey cattle was examined using data from 25 large registered herds of each breed. Sire selection and mate allocation were carried out using linear programming with the following objectives: 1) minimum inbreeding, 2) maximum net merit subject to a fixed inbreeding threshold, and 3) maximum expected lifetime profit after adjustment for inbreeding depression. Inbreeding of actual matings was similar to inbreeding from random matings, indicating that current inbreeding avoidance programs in these herds are ineffective. Inbreeding was reduced by 1.6 and 1.9% in Holsteins and Jerseys, respectively, when a mate allocation program was applied with service sires and usage levels fixed at the actual values. Benefits of mate selection programs increased when both sire selection and mate pair allocation were considered. Maximization of mean net merit with inbreeding restricted to a fixed level (5% in Holsteins and 8% in Jerseys) led to decreases in inbreeding of 0.9 and 1.4% and increases in lifetime profit of $16.66 and $26.86 in Holsteins and Jerseys, respectively, relative to programs that ignored inbreeding. Maximization of mean expected lifetime profit after adjustment for inbreeding depression decreased inbreeding by 1.8 and 2.8% and increased lifetime profit by $37.37 and $59.77 in Holsteins and Jerseys, respectively. Inbreeding coefficients estimated with pedigree traced to 1985 were inadequate predictors of inbreeding coefficients estimated with pedigrees traced to 1960. Mate selection programs cannot function optimally unless extensive historical pedigree data are available, particularly for service sires. Computerized mate selection programs can reduce inbreeding in the next generation, which will lead to an increase in farm profitability. However, if genetic diversity is to be maintained in the long term, procedures for selecting parents of AI sires must also be considered.

Algorithms↗
PubMed2005

Effects of inbreeding and rate of inbreeding in Drosophila melanogaster- Hsp70 expression and fitness.

Induction of heat shock proteins (Hsp) is a well-known mechanism through which cells cope with stressful conditions. Hsp are induced by a variety of extrinsic stressors. However, recently intrinsic stressors (aging and inbreeding) have been shown to affect expression of Hsp. Increased homozygosity due to inbreeding may disrupt cellular homeostasis by causing increased expression of recessive deleterious mutations and breakdown of epistatic interactions. We investigated the effect of inbreeding and the rate of inbreeding on the expression of Hsp70, larval heat resistance and fecundity. In Drosophila melanogaster we found that inbred lines (F approximately 0.67) had significantly up-regulated expression of Hsp70, and reduced heat resistance and fecundity as compared with outbred control lines. A significant negative correlation was observed between Hsp70 expression and resistance to an extreme heat stress in inbred lines. We interpreted this as an increased requirement for Hsp70 in the lines suffering most from inbreeding depression. Inbreeding depression for fecundity was reduced with a slower rate of inbreeding compared with a fast rate of inbreeding. Thus, the effectiveness of purging seems to be improved with a slower rate of inbreeding.

Adaptation, Physiological↗
PubMed1999

Demographic consequences of incest avoidance in the cooperatively breeding acorn woodpecker.

The avoidance of breeding with close relatives is an adaptation to inbreeding depression. Unfortunately, inbreeding depression has proved difficult to document or measure in the wild, despite being frequently observed among animals in captivity. We address this problem indirectly by determining the demographic cost of incest avoidance in the cooperatively breeding, polygynandrous acorn woodpecker, Melanerpes formicivorus, following the death or disappearance of all breeders of one sex within a group (a reproductive vacancy). Groups undergoing female vacancies that also contained female nonbreeding helpers experienced significantly lower reproductive success in each of the subsequent 3 years than those in which either no nonbreeding helpers or only male nonbreeding helpers were present, a decrease attributable to incest avoidance between the helper females and the related breeder males in the group. Using a computer simulation combined with a life-table analysis, we estimated that incest avoidance costs the population 9.2-12.1% in overall reproductive potential (measured in fledglings/female) and decreases the population rate of increase by 1.78-2.33%/year. These results suggest the presence, on average, of at least 1.2-1.8 lethal equivalents per individual, a value of the same magnitude as estimated for several other taxa, including humans. Incest avoidance may compound random demographic and environmental events and significantly facilitate the decline of threatened populations even prior to any detrimental effects of inbreeding depression per se. Copyright 1999 The Association for the Study of Animal Behaviour.

Journal Article↗
PubMed2005

Multigenerational effects of inbreeding in Cucurbita pepo ssp. texana (Cucurbitaceae).

The shape of the fitness function relating the decline in fitness with coefficient of inbreeding (f) can provide evidence concerning the genetic basis of inbreeding depression, but few studies have examined inbreeding depression across a range of f using noncultivated species. Futhermore, studies have rarely examined the effects of inbreeding depression in the maternal parent on offspring fitness. To estimate the shape of the fitness function, we examined the relationship between f and fitness across a range off from 0.000 to 0.875 for components of both male and female fitness in Cucurbita pepo ssp. texana. Each measure of female fitness declined with f, including pistillate flower number, fruit number, seed number per fruit, seed mass per fruit, and percentage seed germination. Several aspects of male fitness also declined with f, including staminate flower number, pollen number per flower, and the number of days of flowering, although cumulative inbreeding depression was less severe for male (0.34) than for female function (0.39). Fitness tended to decline linearly with f between f = 0.00 and f = 0.75 for most traits and across cumulative lifetime fitness (mean = 0.66), suggesting that individual genes causing inbreeding depression are additive and the result of many alleles of small effect. However, most traits also showed a small reduction in inbreeding depression between f = 0.75 and f = 0.875, and evidence of purging or diminishing epistasis was found for in vitro pollen-tube growth rate. To examine inbreeding depression as a maternal effect, we performed outcross pollinations on f = 0.0 and f = 0.5 mothers and found that depression due to maternal inbreeding was 0.07, compared to 0.10 for offspring produced through one generation of selfing. In at least some families, maternal inbreeding reduced fruit number, seed number and mass, staminate flower number, pollen diameter, and pollen-tube growth rate. Collectively these results suggest that, while the fitness function appears to be largely linear for most traits, maternal effects may compound the effects of inbreeding depression in multigenerational studies, though this may be partially offset by purging or diminishing epistasis.

Analysis of Variance↗