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Estimating long-term mating systems using DNA sequences.

Plant mating systems often involve a mixture of self fertilizations and outcross fertilizations. The degree of selfing has a large impact on the genetic composition of natural populations and on the evolution of the mating system itself in response to such factors as inbreeding depression. This paper describes a means of estimating the long-term rate of self-fertilization from samples of alleles taken from individuals in a population. Use is made of the genealogy of pairs of alleles at a locus within individuals and pairs between individuals. The degree of selfing is closely related to the extent to which the number of nucleotide sites differing within an individual is reduced relative to the number differing between individuals. Importantly, the estimate of long-term selfing is largely independent of population size and is not affected by historical fluctuations in population size; instead it responds directly to the mating system itself. The approach outlined here is most appropriate to evolutionary problems in which the long-term nature of the mating system is of interest, such as to determine the relationship between prior inbreeding and inbreeding depression.

Analysis of Variance↗

Heterosis studies for yield and its components in bread wheat over environments.

A set of diallel crosses involving 10 parents was made to have information on the extent of heterosis over mid-parent and better parent and inbreeding depression for yield and yield contributing characters under three different environments. Marked heterobeltiosis for grain yield and its important components were observed. For grain yield, 83 crosses showed significant positive heterobeltiosis in all the three sowing dates, however, twenty crosses showed significant consistent heterobeltiosis for grain yield per plant over all the three environments. The maximum heterobeltiosis for grain yield per plant observed was 50.94% (Raj 3765 x HD 2285), 121.08% (PBW 373 x HD 2329) and 93.96% (PBW 373 x HD 2329) under early, normal and late sowing conditions, respectively. Cross PBW 373 x HD 2329 in both early and normal plantings and cross Raj 3765 x HD 2285 under late planting were observed most heterotic for grain yield. The crosses showing heterosis for grain yield were not heterotic for all the characters. Heterosis for grain yield per spike followed by tillers per plant and 1000-grain weight was independently associated with heterosis for grain yield in early and normal plantings. However, heterosis for grain yield per spike, dwarf plant height and tillers per plant contributed maximum towards yield heterosis. Significant inbreeding depression was recorded frequently for yield and yield contributing traits, however, in a few traits it was observed significant negative indicated that F(2) was superior to F(1) considered desirable combination for trait(s). The study reveals good scope for commercial exploitation of heterosis as well as isolation of pure lines among the progenies of heterotic F(1) for improvement of yield levels in bread wheat.

Analysis of Variance↗

The effects of nectar addition on pollen removal and geitonogamy in the non-rewarding orchid Anacamptis morio.

It has been suggested that the absence of floral rewards in many orchid species causes pollinators to probe fewer flowers on a plant, and thus reduces geitonogamy, i.e. self-pollination between flowers, which may result in inbreeding depression and reduced pollen export. We examined the effects of nectar addition on pollinator visitation and pollen transfer by tracking the fate of colour-labelled pollen in Anacamptis morio, a non-rewarding orchid species pollinated primarily by queen bumble-bees. Addition of nectar to spurs of A. morio significantly increased the number of flowers probed by bumble-bees, the time spent on an inflorescence, pollinarium removal and the proportion of removed pollen involved in self-pollination through geitonogamy, but did not affect pollen carryover (the fraction of a pollinarium carried over from one flower to the next). Only visits that exceeded 18 s resulted in geitonogamy, as this is the time taken for removed pollinaria to bend into a position to strike the stigma. A mutation for nectar production in A. morio would result in an initial 3.8-fold increase in pollinarium removal per visit, but also increase geitonogamous self-pollination from less than 10% of pollen depositions to ca. 40%. Greater efficiency of pollen export will favour deceptive plants when pollinators are relatively common and most pollinaria are removed from flowers or when inbreeding depression is severe. These findings provide empirical support both for Darwin's contention that pollinarium bending is an anti-selfing mechanism in orchids and for the idea that floral deception serves to maximize the efficiency of pollen export.

Animals↗

Inbreeding, fluctuating asymmetry, and ejaculate quality in an endangered ungulate.

An ever-increasing number of species are suffering marked reductions in population size as a consequence of human activities. To understand the impact of these changes it is essential to assess how small population size affects individual fitness and the viability of populations. This issue acquires special relevance among endangered species in which numbers have decreased to such an extent that captive breeding must be established with a few founders. A major risk associated with small population size is inbreeding depression. The effects of inbreeding upon male reproductive traits are the subject of an ongoing controversy, since the evidence linking lack of genetic variability and poor ejaculate quality at the population level has been criticized recently by several authors. We report that among Gazella cuvieri males, inbreeding coefficient shows a strong inverse relationship with ejaculate quality. Furthermore, the degree of fluctuating asymmetry is positively related to the coefficient of inbreeding and negatively related to the proportion of normal sperm, suggesting that it is a reliable indicator of genetic stress and of ejaculate quality.

Animals↗

Inbreeding and outbreeding depression in male courtship song characters in Drosophila montana.

In Drosophila montana, male courtship song frequency is closely associated with male courtship success and offspring survival. Other pulse characters (pulse length and cycle number) may also affect female mate choice, whereas pulse train characters (interpulse interval, pulse number and pulse train length) are not associated with these male fitness components. Inbreeding depression in these song characters was investigated by comparing the songs of inbred and outbred fly strains. The average change in most song characters as a result of inbreeding was only a few percent. However, in male song frequency the average inbreeding depression was about 14%, suggesting that this song character is associated with fitness. Outbreeding depression and the genetic architecture of song characters were investigated with interpopulation crosses and joint scaling tests. For pulse train characters the generation means show only evidence of additivity, and the existence of dominance or epistasis in these characters was strongly rejected in each case. In pulse characters the means of the F1 males were lower than the average of the parental generations. In pulse length and cycle number this difference was attributable to dominance alone. In frequency there was outbreeding depression also in the F2 generation, suggesting a break-up of favourable epistatic gene combinations. The outbreeding depression in this character in the F1 generation was caused by dominance, and in the F2 also by duplicate epistasis between dominant decreasers. The possible role of outbreeding depression and epistasis in speciation is discussed.

Animal Communication↗

Mating patterns and demography in the tristylous daffodil Narcissus triandrus.

Mating patterns in plant populations are influenced by interactions between reproductive traits and ecological conditions, both factors that are likely to vary geographically. Narcissus triandrus, a wide-ranging heterostylous herb, exhibits populations with either two (dimorphic) or three (trimorphic) style morphs and displays substantial geographical variation in demographic attributes and floral morphology. Here, we investigate this variation to determine if demography, morphology, and mating system differ between the two sexual systems. Our surveys in Portugal and NW Spain indicated that dimorphic populations were less dense, of smaller size, and had larger plants and flowers compared to trimorphic populations. Outcrossing rates estimated using allozyme markers revealed similar outcrossing rates in dimorphic and trimorphic populations (t(m) dimorphic=0.759; t(m) trimorphic=0.710). All populations experienced significant inbreeding in progeny (mean F=0.143). In contrast, parental estimates of inbreeding were not significantly different from zero (mean F=0.062), implying that few inbred offspring survive to reproductive maturity due to inbreeding depression. Although the majority of inbreeding results from selfing, significant levels of biparental inbreeding were also detected in eight of the nine populations (mean s(s)-s(m)=0.081). Density was negatively associated with levels of selfing but positively associated with biparental inbreeding. Population size was positively associated with outcrossing but not biparental inbreeding. There were no consistent differences among the style morphs in outcrossing or biparental inbreeding indicating that the maintenance of trimorphism vs dimorphism is unlikely to be associated with inbreeding of maternal parents.

Crosses, Genetic↗

A measure of the within-chromosome synergistic epistasis for Drosophila viability.

In order to detect possible synergistic epistasis for viability in Drosophila melanogaster we assayed the relative viability of chromosomes II in: (i) panmixia, (ii) forced total homozygosity, and (iii) homozygosity for, on the average, half of their loci. As these genotypes were constructed using exactly the same set of chromosomes in the three cases, the design allows us to estimate the inbreeding depression rate at two different inbreeding levels in the absence of purging natural selection. Overall, no consistent synergistic epistasis was found. However, there was a small fraction of chromosomes whose severely deleterious effect when homozygous was almost significantly larger than expected from their viability when homozygous for half of their loci. This suggests occasional but important synergistic epistasis, which might confer evolutionary advantage to recombination in tightly linked genomes. Nevertheless, such epistasis is unlikely to be an evolutionary advantage driving the evolution of sexual anisogamous reproduction, as its contribution to overall viability is small when compared with the two-fold cost of anisogamy.

Analysis of Variance↗

Inbreeding effects on reproductive traits in the ring-necked pheasant.

Ten inbred lines of Ring-necked pheasants were established in 1978 and mated for four generations using a system of repeated backcrossing of daughters to a common sire. In the event the old sire died, the surviving daughters were mated to a surviving brother or half-brother of the same generation. Only 4 of 10 original inbred lines survived four generations of backcrossing; two involved matings with the original sire and two with brothers or half-brothers of Generations 1 and 3, respectively. Egg production, hatchability, and viability were the three traits most affected by inbreeding depression. For 4 generations of inbreeding, the coefficients of regression for all inbred lines on a 10% increase in inbreeding were -5.89, -.42, -1.73, and -3.04 for egg production, egg weight, fertility, and hatchability, respectively. Inbreeding had less severe effects on reproductive traits in two of the four surviving lines. There is evidence that intense early selection among lines for high performance after one generation of inbreeding F = .250 will enhance the success of establishing highly viable inbred lines of pheasants.

Animals↗

Quantitative genetics in conservation biology.

Most of the major genetic concerns in conservation biology, including inbreeding depression, loss of evolutionary potential, genetic adaptation to captivity and outbreeding depression, involve quantitative genetics. Small population size leads to inbreeding and loss of genetic diversity and so increases extinction risk. Captive populations of endangered species are managed to maximize the retention of genetic diversity by minimizing kinship, with subsidiary efforts to minimize inbreeding. There is growing evidence that genetic adaptation to captivity is a major issue in the genetic management of captive populations of endangered species as it reduces reproductive fitness when captive populations are reintroduced into the wild. This problem is not currently addressed, but it can be alleviated by deliberately fragmenting captive populations, with occasional exchange of immigrants to avoid excessive inbreeding. The extent and importance of outbreeding depression is a matter of controversy. Currently, an extremely cautious approach is taken to mixing populations. However, this cannot continue if fragmented populations are to be adequately managed to minimize extinctions. Most genetic management recommendations for endangered species arise directly, or indirectly, from quantitative genetic considerations.

Animals↗

Selection for scrapie resistance and simultaneous restriction of inbreeding in the rare sheep breed "Mergellander".

Scrapie is a fatal infectious neurodegenerative disease for which susceptibility is associated with polymorphisms in the ovine prion protein (PrP) gene. Scrapie-eradication programmes are based on eliminating the susceptible VRQ allele and/or breeding for the resistant ARR allele. In rare breeds or breeds with a low frequency of the ARR allele this can lead to unacceptably high inbreeding rates with associated increased risk of genetic defects and inbreeding depression. The conservation status of populations with inbreeding rates (DeltaF) above 1% is considered critical. In the Dutch rare sheep breed the Mergellander animals carrying ARR alleles are closely related to one another, and could reach 1.53% when only ARR/ARR animals are used as parents. Inbreeding rates can be reduced by selecting the set of parents according to their average co-ancestry. We minimised inbreeding rates by calculating the optimal contribution of each ram and selection of ewes. This resulted in inbreeding rates of -0.17% with exclusive use of homozygous ARR rams, and -0.38% if use of heterozygous rams was allowed as well. Thus sophisticated breeding programs can prevent unacceptably high inbreeding rates when breeding for scrapie resistance.

Alleles↗

Stabilizing selection on genomic divergence in a wild fish population.

Conservation programs use breeding protocols to increase genomic divergence (by mating genetically dissimilar individuals) in an attempt to circumvent population declines resulting from inbreeding depression. However, disruption of either beneficial gene complexes or local genetic adaptations can lead to outbreeding depression, and thus, there should be a reduction in fitness of individuals at either end of the genomic divergence continuum. Although such simultaneous inbreeding and outbreeding depression has been observed in plant populations, it rarely has been demonstrated in animal populations. Here, I use both genetic and phenotypic measures to show that there is stabilizing selection on genomic divergence in a wild population of bluegill sunfish (Lepomis macrochirus). I also show that breeding individuals that exercise mate choice produce offspring that are closer to the optimal level of genomic divergence than random mating alone would predict.

Animals↗

Impact of nonadditive genetic effects in the estimation of breeding values for fertility and correlated traits.

The effects of inbreeding, heterosis, recombination loss, and percentage Holstein on the estimation of predicted transmitting abilities for fertility traits (calving interval, number of days from calving to first insemination, nonreturn rate, number of inseminations) and correlated traits (milk yield at test nearest d 110 and body condition score) were examined in a mixed population of Holstein and Friesian cattle. An unfavorable effect of percentage Holstein on calving interval was observed, resulting in a 12-d increase for pure Holsteins compared with pure Friesians. Insemination traits were less affected by percentage Holstein, with 3% more animals returning to first service within 56 d and 0.1 more inseminations required for Holstein animals. Heterosis and recombination loss affected some of the traits. Heterosis had a favorable effect on yield, with a 0.35-kg difference between a pure and cross-bred animal for test milk. There was a reduction of 1 d to first insemination between a pure and first-crossbred animal. Inbreeding had a significant and unfavorable effect on all traits. The difference between a noninbred animal and an animal with an inbreeding coefficient of 10% was a 2.8-d increase in calving interval, a 1.7-d increase in days to first insemination, a 1% increased probability to return to estrus at first service, 0.03 more inseminations, a 0.27-unit decrease in body condition, and a 0.54-kg decrease in milk on test nearest d 110. The effect of inbreeding depression was more pronounced at higher levels of inbreeding. The rank correlations between the predicted transmitting abilities for fertility and correlated traits, with and without the additional nonadditive effects in the model, were over 0.99. Steps should be taken to control the rise in inbreeding, or the effects on fertility and correlated traits such as milk production will begin to manifest themselves.

Animals↗

Inbreeding, outbreeding and environmental effects on genetic diversity in 46 walleye (Sander vitreus) populations.

Genetic diversity is recognized as an important population attribute for both conservation and evolutionary purposes; however, the functional relationships between the environment, genetic diversity, and fitness-related traits are poorly understood. We examined relationships between selected lake parameters and population genetic diversity measures in 46 walleye (Sander vitreus) populations across the province of Ontario, Canada, and then tested for relationships between six life history traits (in three categories: growth, reproductive investment, and mortality) that are closely related to fitness, and genetic diversity measures (heterozygosity, d2, and Wright's inbreeding coefficient). Positive relationships were observed between lake surface area, growing degree days, number of species, and hatchery supplementation versus genetic diversity. Walleye early growth rate was the only life history trait significantly correlated with population heterozygosity in both males and females. The relationship between FIS and male early growth rate was negative and significant (P < 0.01) and marginally nonsignificant for females (P = 0.06), consistent with inbreeding depression effects. Only one significant relationship was observed for d2: female early growth rate (P < 0.05). Stepwise regression models showed that surface area and heterozygosity had a significant effect on female early growth rate, while hatchery supplementation, surface area and heterozygosity had a significant effect on male early growth rate. The strong relationship between lake parameters, such as surface area, and hatchery supplementation, versus genetic diversity suggests inbreeding and outbreeding in some of the populations; however, the weak relationships between genetic diversity and life history traits indicate that inbreeding and outbreeding depression are not yet seriously impacting Ontario walleye populations.

Animals↗

Genome-wide analysis on inbreeding effects on gene expression in Drosophila melanogaster.

The deleterious consequences of inbreeding, especially in the form of inbreeding depression, are well known. However, little is known about how inbreeding affects genome-wide gene expression. Here, we show that inbreeding changes transcription levels for a number of genes. Gene expression profiles of Drosophila melanogaster lines inbred to F approximately = 0.67 at different rates changed relative to those of noninbred lines, but the rate of inbreeding did not significantly affect gene expression patterns. Genes being differentially expressed with inbreeding are disproportionately involved in metabolism and stress responses, suggesting that inbreeding acts like an environmental stress factor.

Animals↗

Epistasis and the conversion of non-additive to additive genetic variance at population bottlenecks.

The effect of population bottlenecks on the mean and the additive variance generated by two neutral independent epistatic loci has been studied theoretically. Six epistatic models, used in the analysis of binary disease traits, were considered. Ancestral values in an infinitely large panmictic population were compared with their expectations at equilibrium, after t consecutive bottlenecks of equal size N (derived values). An increase in the additive variance after bottlenecks (inversely related to N and t) will occur only if the frequencies of the negative allele at each locus are: (1) low, invariably associated to strong inbreeding depression; (2) high, always accompanied by an enhancement of the mean with inbreeding. The latter is an undesirable property, making the pertinent models unsuitable for the genetic analysis of disease. For the epistatic models considered, it is unlikely that the rate of evolution may be accelerated after population bottlenecks, in spite of occasional increments of the derived additive variance over its ancestral value.

Animals↗

Inbreeding and reproduction in mice divergently selected for response to a dietary toxin.

This study was conducted to determine whether inbreeding coefficients of selected parents or of progeny differed between lines of mice selected for increased or decreased responsiveness to a nutritional toxicosis. A second objective was to determine whether the influence of inbreeding of parents and/or progeny on reproductive traits differed between those lines. Mice were selected divergently for 8 generations for the effect on post-weaning growth of endophyte-infected fescue seed in their diet. Forty pairs (or in Generation 7, 20 pairs) were selected and mated per generation in each line. Inbreeding increased 0.5 to 0.6% per generation in both lines, a rate close to that predicted from genetic theory. Inbreeding coefficients of selected parents were not higher in the susceptible than in the resistant line. A difference would have been expected if the inbreeding coefficient had been correlated with susceptibility to toxicosis. The magnitudes of inbreeding depression for reproductive traits did not differ significantly between lines. The average inbreeding coefficient of the potential litter tended to be higher in nonfertile than fertile matings (P = 0.10), but inbreeding coefficients of sires and dams did not differ between successful and unsuccessful matings. Inbred litters tended to be born earlier than noninbred litters (P = 0.10). Inbred dams produced smaller litters than noninbred dams (main effect P < 0.05) but only when the litter also was inbred (interaction P < 0.01). Sex ratio was not influenced by inbreeding of sire, dam or litter, but there was a higher proportion of male progeny in the susceptible than in the resistant line (P = 0.01). To avoid reduced reproductive fitness, laboratory animal populations should be managed to minimize inbreeding of progeny and dam.

Animals↗

Effects of inbreeding in three life stages of Drosophila buzzatii after embryos were exposed to a high temperature stress.

The interaction between inbreeding and high-temperature stress was examined in the cactophilic fruit fly, Drosophila buzzatii. Embryos of four inbreeding levels (F = 0, F = 0.25, F = 0.375, F = 0.5) were either maintained at 25 degrees C throughout egg-to-adult development or were exposed to 41.5 degrees C for 110 min at an age of 20 h. Hatching, larva-to-pupa survival, pupa-to-adult survival, and egg-to-adult survival were estimated. Heat shock reduced hatching rates, but survival to adulthood for individuals that hatched was unaffected by the heat shock. Inbreeding reduced the proportion of eggs hatching in the 25 degrees C control group only. For larva-to-pupa and pupa-to-adult survival there was no interaction between inbreeding and stress. The effect of inbreeding on egg-to-adult survival was stronger in the 25 degrees C control group compared with the group exposed to heat shock. The results imply environmental dependency of inbreeding depression and suggest that stress tolerance may not always be reduced by inbreeding. The thermal microenvironment of cactus rots in the field was assessed by measuring temperatures inside 17 rots. Internal rot temperatures varied with a maximum temperature of 48 degrees C during the day. Selection for temperature tolerance in nature may have depleted genetic variation for this trait limiting the effect of inbreeding on thermal resistance.

Animals↗

The effects of inbreeding on testicular sperm concentration in Peromyscus polionotus.

Inbreeding effects on fitness have most often been quantified via juvenile mortality. However, inbred adults may suffer from inbreeding depression if their fertility or fecundity is compromised as a consequence of inbreeding. Here, the effects of inbreeding on male fertility in oldfield mice, Peromyscus polionotus, were examined. Testicular sperm concentration was assessed in 93 males, 68 of which were paired for breeding. Forty of the 68 paired males failed to produce offspring. Total testicular sperm count, sperm count (g testis)(-1), and testis mass all declined significantly with increasing inbreeding coefficient. Sperm concentration did not significantly impact reproductive success. Although sperm concentration in males of most species can decline to low levels before reproductive impairment is detectable, the declines in testicular sperm concentration found here suggest that inbreeding can affect fertility in adult males. Furthermore, monitoring testicular sperm concentration could provide a mechanism to monitor potential declines in reproductive performance before population-level reproductive success is irreparably impaired. The implications for the management of small, captive and wild populations may be substantial.

Animals↗