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Partial male sterility and the evolution of nuclear gynodioecy in plants.

Gynodioecy, a genetic dimorphism of females and hermaphrodites, is pertinent to an understanding of the evolution of plant gender, mating and genetic variability. Classical models of nuclear gynodioecy attribute the maintenance of the dimorphism to frequency-dependent selection in which the female phenotype has a fitness advantage at low frequency owing to a doubled ovule fertility. Here, I analyse explicit genetic models of nuclear gynodioecy that expand on previous work by allowing partial male sterility in combination with either fixed or dynamically evolving mutational inbreeding depression. These models demonstrate that partial male sterility causes fitness underdominance at the mating locus, which can prevent the spread of females. However, if partial male sterility is compensated by a change in selfing rate, overdominance at the mating locus can cause the spread of females. Overdominance at introduction of the male sterility allele can be caused by high inbreeding depression and a lower selfing rate in the heterozygote, by purging of mutations by a higher selfing rate in the heterozygote, and by low inbreeding depression and a higher selfing rate in the heterozygote. These processes might be of general importance in the maintenance of mating polymorphisms in plants.

Biological Evolution↗

Lethals in subdivided populations.

The fate of lethal alleles in populations is of interest in evolutionary and conservation biology for several reasons. For instance, lethals may contribute substantially to inbreeding depression. The frequency of lethal alleles depends on population size, but it is not clear how it is affected by population structure. By analysing the case of the infinite island model by numerical approaches and analytical approximations it is shown that, like population size, population structure affects the fate of lethal alleles if dominance levels are low. Inbreeding depression caused by such alleles is also affected by the population structure, whereas the mutation load is only weakly affected. Heterosis also depends on population structure, but it always remains low, of the order of the mutation rate or less. These patterns are compared with those caused by mildly deleterious mutations to give a general picture of the effect of population structure on inbreeding depression, heterosis, and the mutation load.

Alleles↗

Effects of inbreeding and heterozygosity on preweaning traits in a closed population of Herefords under selection.

Records of five inbred lines at the Livestock and Range Research Laboratory were used to evaluate effects of inbreeding and heterozygosity on preweaning traits. Members of each line were descendants of a single founder Hereford bull. A total of 8,065 records of birth weight and 7,380 records of preweaning daily gain and weaning weight were analyzed by derivative-free REML using a model that included fixed effects of sex, combination of year and month of birth and parity of dam, with covariates for direct and maternal genetic fractions of inheritance from the genetic groups, inbreeding, and heterozygosity fractions. Heterozygosity fractions were computed for crosses between lines. The random model effects were direct and maternal genetic and uncorrelated maternal permanent environmental and temporary environmental. Direct inbreeding and heterozygosity fractions averaged .098 and .343, and maternal inbreeding and heterozygosity fractions averaged .075 and .294. Regression coefficients of traits on direct and maternal inbreeding fractions were -5.8 +/- 1.1 and -4.7 +/- 1.3 for birth weight, -.189 +/- .031 and -.252 +/- .039 for preweaning daily gain, and -44.5 +/- 6.6 and -56.1 +/- 8.4 kg for weaning weight. Estimates for direct heritability, maternal heritability, and direct-maternal genetic correlations were .37, .12, and -.01 for birth weight; .16, .25, and -.27 for daily gain; and .17, .26, and -.21 for weaning weight. Results suggest that heterosis represents recovery of accumulated inbreeding depression. Results also indicate that selection can overcome inbreeding depression and antagonism exists between direct and maternal genetic effects for preweaning traits.

Animals↗

The evolution and maintenance of monoecy and dioecy in Sagittaria latifolia (Alismataceae).

Plant species rarely exhibit both monoecious and dioecious sexual systems. This limits opportunities to investigate the consequences of combined versus separate sex function on mating patterns and genetic variation and the analysis of factors responsible for the evolution and maintenance of the two sexual systems. Populations of the North American clonal aquatic Sagittaria latifolia are usually either monoecious or dioecious and often grow in close geographic proximity. We investigated mating patterns, genetic structure, and relationships between the two sexual systems using allozyme variation in populations from southern Ontario, Canada. As predicted, selfing rates in monoecious populations (n = 6, mean = 0.41) were significantly higher than in dioecious populations (n = 6, mean = 0.11). Moreover, marker-based estimates of inbreeding depression (delta) indicated strong selection against inbred offspring in both monoecious (mean delta = 0.83) and dioecious (mean delta = 0.84) populations. However, the difference in selfing rate between the sexual systems was not reflected in contrasting levels of genetic variation. Our surveys of 12 loci in 15 monoecious and 11 dioecious populations revealed no significant differences in the proportion of polymorphic loci (P), number of alleles per locus (A), or observed and expected heterozygosity (H(o) and H(e), respectively). Strong inbreeding depression favoring survival of outcrossed offspring may act to maintain similar levels of diversity between monoecious and dioecious populations. Despite geographical overlap between the two sexual systems in southern Ontario, a dendrogram of genetic relationships indicated two distinct clusters of populations largely corresponding to monoecious and dioecious populations. Reproductive isolation between monoecious and dioecious populations appears to be governed, in part, by observed differences in habitat and life history. We suggest that selfing and inbreeding depression in monoecious populations are important in the transition from monoecy to dioecy and that the maintenance of distinct sexual systems in S. latifolia is governed by interactions between ecology, life history, and mating.

Biological Evolution↗

Inbreeding effects on metrical phenotypes among North Indian Children.

This study aimed to examine the effects of inbreeding on 12 quantitative phenotypes like body weight, height, sitting height, head circumference, head length, head breadth, chest circumference, verbal, performance and full scale intelligence quotients, systolic and diastolic blood pressures among North Indian Children. The sample consisted of 3,253 subjects (1,683 males and 1,570 females) including offspring of first cousins, first cousins once removed, second cousins, (inbred part of samples) and unrelated spouses (non-inbred part of samples) ranging in age from 6 to 14 years from the Aligarh district, Uttar Pradesh in North India. Samples were selected purposively to estimate the average inbreeding coefficients based on 3 ancestral generations and using Wright's path method. The average inbreeding coefficient of the present inbred part of sample is 0.04609. The mean ages of males and females were almost equal for both inbred and non-inbred individuals. A significant reduction of means (p < 0.05) or inbreeding depression has been observed in inbred series for all anthropometric and psychometric traits for both the sexes. A significant elevation of mean with inbreeding is observed for systolic and diastolic blood pressure. The average inbreeding depression per 10% inbreeding appeared very high in verbal IQ (> or =23), performance IQ (> or =40), full scale IQ (> or =28), systolic blood pressure (> or =10) and diastolic blood pressure (> or =12) among both sexes. On the whole, relatively greater amount of inbreeding effects are apparent among all measures of intelligence quotient and blood pressures compared to the anthropometric traits. The results of the present work have thrown light on the nature and mechanism of genetic effects of inbreeding on certain quantitative traits in human.

Adolescent↗

Mating system, sex ratio, and persistence of females in the gynodioecious shrub Daphne laureola L. (Thymelaeaceae).

Although in gynodioecious populations male steriles require a fecundity advantage to compensate for their gametic disadvantage, southern Spanish populations of the long-lived shrub Daphne laureola do not show any fecundity advantage over hermaphrodites in terms of seed production and early seedling establishment. By using allozyme markers, we assess the mating system of this species in five populations differing in sex ratio, and infer levels of inbreeding depression over the whole life cycle by comparing the inbreeding coefficients at the seed and adult plant stages. Extremely low outcrossing rates (0.001<t<0.125) were consistently found for hermaphrodites in all populations, whereas, as expected, female progeny were entirely outcrossed. In most populations, offspring were much more inbred than their parents, and heterozygosity of adults was greater than expected from outcrossing rate estimates, with very few selfed progeny appearing to reproduce in the field. The combination of extensive selfing in hermaphrodites and a strong inbreeding depression expressed late in the life cycle (and thus, only estimable by indirect measures based on genetic markers) may explain the persistence and high frequency of D. laureola females in southern Spanish populations.

Biological Evolution↗

On the evolution of genetic incompatibility systems. VI. A three-locus modifier model for the origin of gametophytic self-incompatibility.

Recent genetic analyses have demonstrated that self-incompatibility in flowering plants derives from the coordinated expression of a system of loci. To address the selective mechanisms through which a genetic system of this kind evolves, I present a three-locus model for the origin of gametophytic self-incompatibility. Conventional models assume that a single locus encodes all physiological effects associated with self-incompatibility and that the viability of offspring depends only on whether they were derived by selfing or outcrossing. My model explicitly represents the genetic determination of offspring viability by a locus subject to symmetrically overdominant selection. Initially, the level of expression of the proto-S locus is insufficient to induce self-incompatibility. Weak gametophytic self-incompatibility arises upon the introduction of a rare allele at an unlinked modifier locus which enhances the expression of the proto-S locus. While conventional models predict that the origin of self-incompatibility requires at least two- to threefold levels of inbreeding depression, I find that the comparatively low levels of inbreeding depression generated by a single overdominant locus can ensure the invasion of an enhancer of self-incompatibility under sufficiently high rates of receipt of self-pollen. Associations among components of the incompatibility system promote the origin of self-incompatibility. Enhancement of heterozygosity at the initially neutral proto-S locus improves offspring viability through associative overdominance. Further, the modifier that enhances the expression of self-incompatibility develops a direct association with heterozygosity at the overdominant viability locus. These results suggest that the evolutionary processes by which incompatibility systems originate may differ significantly from those associated with their breakdown. The genetic mechanism explored here may apply to the evolution of other systems that restrict reproduction, including maternal-fetal incompatibility in mammals.

Animals↗

Estimation of additive and nonadditive genetic variances in Hereford, Gelbvieh, and Charolais by Method R.

Parameters for direct and maternal dominance were estimated in models that included non-additive genetic effects. The analyses used weaning weight records adjusted for age of dam from populations of Canadian Hereford (n = 467,814), American Gelbvieh (n = 501,552), and American Charolais (n = 314,552). Method R estimates of direct additive genetic, maternal additive genetic, permanent maternal environment, direct dominance, and maternal dominance variances as a proportion of the total variance were 23, 12, 13, 19, and 14% in Hereford; 27, 7, 10, 18, and 2% in Gelbvieh; and 34, 15, 15, 23, and 2% in Charolais. The correlations between direct and maternal additive genetic effects were -0.30, -0.23, and -0.47 in Hereford, Gelbvieh, and Charolais, respectively. The correlations between direct and maternal dominance were -0.38, -0.02, and -0.04 in Hereford, Gelbvieh, and Charolais, respectively. Estimates of inbreeding depression were -0.20, -0.18, and -0.13 kg per 1% of inbreeding for Hereford, Gelbvieh, and Charolais, respectively. Estimates of the maternal inbreeding depression were -0.01, -0.02, and -0.02 kg, respectively. The high ratio of direct dominance to additive genetic variances provided some evidence that direct dominance effects should be considered in beef cattle evaluation. However, maternal dominance effects seemed to be important only for Hereford cattle.

Animals↗

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<r/(1+r) where r is relatedness and delta is the fecundity loss relative to an outbred mating. This amounts to fitness losses up to one-fifth for a half-sib mating and one-third for a full-sib mating, which lie in the upper range of inbreeding depression values currently reported in natural populations. The observation of active inbreeding avoidance in a polygynous species thus suggests that inbreeding depression exceeds this threshold in the species under scrutiny or that inbred matings at least partly forfeit other mating opportunities for males. Our model also shows that female choosiness should decline rapidly with search costs, stemming from, for example, reproductive delays. Species under strong time constraints on reproduction should thus be tolerant of inbreeding.

Animals↗

Inbreeding and heterosis effects on quantitative traits in a White Leghorn population under long-term reciprocal recurrent selection.

1. Inbreeding and heterosis effects for quantitative traits were estimated in two White Leghorn lines selected with reciprocal recurrent selection (RRS) since 1950. 2. Pedigreed progeny were produced from full-sib, half-sib and unrelated pure line as well as reciprocal cross line matings to estimate inbreeding and heterosis effects "within sires". 3. Inbreeding effects estimated as linear regressions agreed well with estimates considering inbreeding as fixed effect. The inbreeding depression per 10% inbreeding was highest for hatchability (6-7%), intermediate for production-related traits (3%), and about 1% for shell quality, body weight and egg weight. 4. Heterosis effects were estimated by comparing pure line and cross line progeny of the same sires. 5. Results indicate considerably less heterosis for egg production than found earlier, which is interpreted as a consequence of pure line selection.

Animals↗

Does heterozygosity estimate inbreeding in real populations?

Many recent studies report that individual heterozygosity at a handful of apparently neutral microsatellite markers is correlated with key components of fitness, with most studies invoking inbreeding depression as the likely underlying mechanism. The implicit assumption is that an individual's inbreeding coefficient can be estimated reliably using only 10 or so markers, but the validity of this assumption is unclear. Consequently, we have used individual-based simulations to examine the conditions under which heterozygosity and inbreeding are likely to be correlated. Our results indicate that the parameter space in which this occurs is surprisingly narrow, requiring that inbreeding events are both frequent and severe, for example, through selfing, strong population structure and/or high levels of polygyny. Even then, the correlations are strong only when large numbers of loci (~200) can be deployed to estimate heterozygosity. With the handful of markers used in most studies, correlations only become likely under the most extreme scenario we looked at, namely 20 demes of 20 individuals coupled with strong polygyny. This finding is supported by the observation that heterozygosity is only weakly correlated among markers within an individual, even in a dataset comprising 400 markers typed in diverse human populations, some of which favour consanguineous marriages. If heterozygosity and inbreeding coefficient are generally uncorrelated, then heterozygosity-fitness correlations probably have little to do with inbreeding depression. Instead, one would need to invoke chance linkage between the markers used and one or more gene(s) experiencing balancing selection. Unfortunately, both explanations sit somewhat uncomfortably with current understanding. If inbreeding is the dominant mechanism, then our simulations indicate that consanguineous mating would have to be vastly more common than is predicted for most realistic populations. Conversely, if heterosis provides the answer, there need to be many more polymorphisms with major fitness effects and higher levels of linkage disequilibrium than are generally assumed.

Computer Simulation↗

Genetic components of variation in Nemophila menziesii undergoing inbreeding: morphology and flowering time.

The standard approaches to estimation of quantitative genetic parameters and prediction of response to selection on quantitative traits are based on theory derived for populations undergoing random mating. Many studies demonstrate, however, that mating systems in natural populations often involve inbreeding in various degrees (i.e. , self matings and matings between relatives). Here we apply theory developed for estimating quantitative genetic parameters for partially inbreeding populations to a population of Nemophila menziesii recently obtained from nature and experimentally inbred. Two measures of overall plant size and two of floral size expressed highly significant inbreeding depression. Of three dominance components of phenotypic variance that are defined under partial inbreeding, one was found to contribute significantly to phenotypic variance in flower size and flowering time, while the remaining two components contributed only negligibly to variation in each of the five traits considered. Computer simulations investigating selection response under the more complete genetic model for populations undergoing mixed mating indicate that, for parameter values estimated in this study, selection response can be substantially slowed relative to predictions for a random mating population. Moreover, inbreeding depression alone does not generally account for the reduction in selection response.

Asteraceae↗

Curvilinear inbreeding effects on some performance traits in laying hens.

Quadratic partial regression coefficients were estimated for the inbreeding level on five performance traits (body weight, average egg weight, age at first egg, percentage of fertilized eggs, and hatchability of set eggs) of two strains of laying hens. Data on 5631 of H77 layers and 3563 of N88 layers from nine consecutive generations were analysed. Only dams were accounted for. Partial regression coefficients were estimated by REML with a single-trait animal model, which included fixed effects (generation and hatching period) and random effects (additive genetic and error effects). The mean inbreeding level was 0.87% in strain H77 and 1.08% in strain N88. The inbreeding effects were analysed based on the quadratic partial regression equations. A slight inbreeding depression was found for all the traits analysed in N88. In strain H77, negative effects of inbreeding were only noted for body weight and average egg weight. The small inbreeding effects shown here resulted from a relatively low level of homozygosity in the populations studied. The strains were found to differ in the effects of inbreeding. It is worth pointing out that differences were noted both between the inbreeding depression estimated from the partial linear regression equation and the quadratic partial regression equation, as well as different inbreeding levels.

Animals↗

Selection response in finite populations.

Formulae were derived to predict genetic response under various selection schemes assuming an infinitesimal model. Account was taken of genetic drift, gametic (linkage) disequilibrium (Bulmer effect), inbreeding depression, common environmental variance, and both initial segregating variance within families (sigma AW02) and mutational (sigma M2) variance. The cumulative response to selection until generation t(CRt) can be approximated as [equation: see text] where Ne is the effective population size, sigma AW infinity 2 = Ne sigma M2 is the genetic variance within families at the steady state (or one-half the genic variance, which is unaffected by selection), and D is the inbreeding depression per unit of inbreeding. R0 is the selection response at generation 0 assuming preselection so that the linkage disequilibrium effect has stabilized. beta is the derivative of the logarithm of the asymptotic response with respect to the logarithm of the within-family genetic variance, i.e., their relative rate of change. R0 is the major determinant of the short term selection response, but sigma Me2 Ne and beta are also important for the long term. A selection method of high accuracy using family information gives a small Ne and will lead to a larger response in the short term and a smaller response in the long term, utilizing mutation less efficiently.

Animals↗

Inbreeding effects on the parameters of the growth function in three strains of Iberian pigs.

The growth performance of inbred pigs coming from full-sib matings was compared to that of contemporary animals coming from the same sires mated to control dams. The animals belong to three strains of Iberian pigs with different histories of previous inbreeding. The data consisted of 1,615 weight records between 50 and 250 d of age from 229 pigs. The statistical model assumed the two parameters that describe the linear growth function (a = intercept, b = slope) as different traits (weight at 120 d and daily gain) and the analysis was carried out in a Bayesian framework via Gibbs sampling. The means of the posterior distribution of heritabilities, common litter environmental coefficients, and genetic correlation were 0.328 (h(2)a), 0.332 (h2b), 0.084 (c(2)a), 0.060 (c(2)b) and 0.973 (rhoG). Inbreeding depression, expressed as the performance decrease relative to the mean, per 10% increase of the inbreeding coefficient, was -3.18, -2.31 and -5.37% for weight at 120 d (a), and -3.16, -2.08 and -6.49% for daily gain (b) in each of the three strains. The results indicate that the inbreeding effects are dependent on the level of previous inbreeding, the more previous inbreeding, the less inbreeding depression.

Animals↗

Heterozygosity, inbreeding and neonatal traits in Soay sheep on St Kilda.

We investigated whether birth weight and neonatal survival, a period within which 24% of all mortalities occur, were correlated with levels of inbreeding in St Kilda Soay sheep, using pedigree inbreeding coefficients and four marker-based estimators of inbreeding. None of the inbreeding estimators, either of the offspring, or of their mothers, explained significant variation in a lamb's birth weight or probability of surviving the neonatal period, suggesting low inbreeding depression for these traits. We evaluated the correlation between the marker-based measures of inbreeding and inbreeding coefficients obtained from the Soay pedigree, where paternal links were inferred using the same panel of microsatellite markers. Even when using a relatively complete portion of the pedigree, in which all individuals had known maternal and paternal grandparents, the correlation was found to be weak (r = -0.207, where mean f = 0.0168). These results add support to the recent prediction that when the mean and variance in inbreeding are low in a population, heterozygosity-fitness correlations can be very weak or even undetectable. The pursuit of more detailed pedigrees offers the best prospect for identifying inbreeding depression within this study population.

Animals↗

Experimental analysis of biparental inbreeding in a self-fertilizing plant.

Localized dispersal and mating may genetically structure plant populations, resulting in matings among related individuals. This biparental inbreeding has significant consequences for the evolution of mating systems, yet is difficult to estimate in natural populations. We estimated biparental inbreeding in two populations of the largely self-fertilizing plant Aquilegia canadensis using standard inference as well as a novel experiment comparing apparent selfing between plants that were randomly relocated within populations to experimental control plants. Using two allozyme markers, biparental inbreeding (b) inferred from the difference between single-locus and multilocus estimates of selfing (b = s(s) - s(m)) was low. Less than 3% of matings involved close relatives (mean b = 0.029). In contrast, randomly relocating plants greatly reduced apparent selfing (mean s(s) = 0.674) compared to control plants that had been dug up and replanted in their original locations (s(s) = 0.953, P = 0.002). Based on this difference in s(s), we estimated that approximately 30% of all matings involved close relatives (mean b = 0.279, 95% CL = 0.072-0.428). Inference from s(s) - s(m) underestimated b in these populations by more than an order of magnitude. Biparental inbreeding is thought to influence the evolution of self-fertilization primarily through reducing the genetic cost of outcrossing. This is unlikely to be of much significance in A. canadensis because inbreeding depression (a major cost of selfing) is much stronger than the cost of outcrossing. However, biparental inbreeding combined with strong inbreeding depression may influence selection on dispersal.

Aquilegia↗

Inbreeding studies in Brasilian schoolchildren.

We present results of comparisons between consanguineous and control couples and their children in Southern Brasil. Multiple regression, t-test and chi 2 analyses were applied to the data. The following variables were investigated; Parents: structure of first cousin marriages, latitude and longitude of birthplaces, cohabitation time, rurality, marriage age, occupation, number of liveborn children, and frequency of twinning. Schoolchildren: clinical data as classified into 24 discontinuous traits, each one subdivided into two categories (number and severity), and seven continuous traits (weight, height, pulse rate, respiratory rate, systolic pressure, diastolic pressure, and temperature); nutrition indices and biotype; family names; intelligence tests and school performance; birth order, age, hair color, hair type, eye color, and skin color; twinning in the sibship, and infant mortality among sibs of the propositi. Our study showed a) nonrandomness of consanguineous marriages as detected on structural, racial, geographical, and temporal levels; b) a heavy mutational load of about 1.5 lethons acting on infant mortality; c) no inbreeding effect on morbidity in general (with a possible exception of "ear morbidity", including hearing deficit); d) a modest inbreeding depression on height (a decrease of 2 cm with an increase of 10% of inbreeding); 3) a suggestion of inbreeding depression on "intelligence".

Brazil↗