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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↗

Female multiple mating behaviour in the common shrew as a strategy to reduce inbreeding.

Three hypotheses concerning potential genetic benefits of female multiple mating behaviour are evaluated for the common shrew. In a high-density population, many successful copulations took place between individuals estimated to be close relatives (e.g. full or half siblings). Juveniles resulting from such matings tended to be relatively small at weaning, and were generally less likely to survive to sexual maturity than more outbred individuals. Multiple paternity was discovered in eight of nine litters examined. The incidence of matings between close relatives, the cost of inbreeding, and the high incidence of multiple paternity presented are each consistent with the hypothesis that female multiple mating is a strategy to reduce inbreeding. That is, if females cannot always distinguish close kin, then they may copulate with several different males and so reduce the risk that all their offspring will be sired by a close relative.

Animals↗

Inbreeding avoidance and reproductive skew in a cooperative mammal.

In animal social groups, socially subordinate individuals frequently show low reproductive success or completely fail to breed. This suppression of subordinate reproduction is currently typically attributed to control by dominant individuals. However, subordinates in cooperative groups often lack access to unrelated mates, and an alternative possibility is that their reproduction is limited by inbreeding avoidance. Using the eusocial Damaraland mole-rat Cryptomys damarensis, this paper provides the first experimental evidence, to our knowledge, for this explanation. Subordinate, non-breeding female mole-rats were given access to unrelated mates while remaining in the presence of dominant females, and many became reproductively active soon after unrelated males were introduced. Inbreeding avoidance and the availability of unrelated mates provides a plausible and untested explanation for variation in reproductive skew across animal societies.

Aggression↗

Divorce in cooperatively breeding long-tailed tits: a consequence of inbreeding avoidance?

The decision of whether to divorce a breeding partner between reproductive attempts can significantly affect individual fitness. In this paper, we report that 63% of surviving pairs of long-tailed tits Aegithalos caudatus divorced between years. We examine three likely explanations for the high divorce rate in this cooperative breeder. The 'better option' hypothesis predicts that divorce and re-pairing increases an individual's reproductive success. However, divorcees did not secure better partners or more helpers and there was no improvement in their reproductive success following divorce. The 'inbreeding avoidance' hypothesis predicts that females should disperse from their family group to avoid breeding with philopatric sons. The observed pattern of divorce was consistent with this hypothesis because, in contrast to the usual avian pattern, divorce was typical for successful pairs (81%) and less frequent in unsuccessful pairs (36-43%). The 'forced divorce' hypothesis predicts that divorce increases as the number of competitors increases. The pattern of divorce among failed breeders was consistent with this hypothesis, but it fails to explain the overall occurrence of divorce because divorcees rarely re-paired with their partners' closest competitors. We discuss long-tailed tits' unique association between divorce and reproductive success in the context of dispersal strategies for inbreeding avoidance.

Animals↗

Habitat fragmentation causes bottlenecks and inbreeding in the European tree frog (Hyla arborea).

A genetic study of the European tree frog, Hyla arborea, in Denmark was undertaken to examine the population structure on mainland Jutland and the island of Lolland after a period of reduction in suitable habitat and population sizes. The two regions have experienced the same rate of habitat loss but fragmentation has been more severe on Lolland. Genetic variation based on 12 polymorphic DNA microsatellites was analysed in 494 tree frogs sampled from two ponds in Jutland and 10 ponds on Lolland. A significant overall deviation from Hardy-Weinberg expectations could be attributed to three ponds, all on Lolland. This was most probably caused by an inbreeding effect reducing fitness, which was supported by the observed significant negative correlation between larva survival and mean F(IS) value and mean individual inbreeding coefficient. A significant reduction in genetic variation (bottleneck) was detected in most of the ponds on Lolland. Population-structure analysis suggested the existence of at least 11 genetically different populations, corresponding to most of the sampled population units. The results indicated that the populations were unique genetic units and could be used to illustrate the migration pattern between newly established ponds arisen either by natural colonization of tree frogs or by artificial introduction. A high degree of pond fidelity in the tree frogs was suggested. A severe fragmentation process reducing population size and fitness within some of the populations probably caused the significant reduction in genetic variation of tree frog populations on Lolland.

Animal Migration↗

Inbreeding and sex-biased gene flow in the ant Formica exsecta.

The objective of this study was to assess breeding and dispersal patterns of both males and females in a monogyne (a single queen per colony) population of ants. Monogyny is commonly associated with extensive nuptial flights, presumably leading to considerable gene flow over large areas. Opposite to these expectations we found evidence of both inbreeding and sex-biased gene flow in a monogyne population of Formica exsecta. We found a significant degree of population subdivision at a local scale (within islands) for queens (females heading established colonies) and workers, but not for colony fathers (the males mated to the colony queens). However, we found little evidence of population subdivision at a larger scale (among islands). More conclusive support for sex-biased gene flow comes from the analysis of isolation by distance on the largest island, and from assignment tests revealing differences in female and male philopatry. The genetic similarity between pairs of queens decreased significantly when geographical distance increased, demonstrating limited dispersal and isolation by distance in queens. By contrast, we found no such pattern for colony fathers. Furthermore, a significantly greater fraction of colony queens were assigned as having originated from the population of residence, as compared to colony fathers. Inbreeding coefficients were significantly positive for workers, but not for mother queens. The queen-male relatedness coefficient of 0.23 (regression relatedness) indicates that mating occurs between fairly close relatives. These results suggest that some monogyne species of ants have complex dispersal and mating systems that can result in genetic isolation by distance over small geographical scales. More generally, this study also highlights the importance of identifying the relevant scale in analyses of population structure and dispersal.

Animals↗

Inbreeding depression and multiple regions showing heterozygote advantage in Drosophila melanogaster exposed to stress.

Recent studies that reveal a correlation between heterozygosity and fitness in natural populations have rekindled interest in whether balancing selection is widespread or an evolutionary oddity. We therefore quantified heterozygote advantage at 12 microsatellite markers in both inbred and outbred crosses of Drosophila grown under different forms of environmental stress. As expected, inbreeding depression reduces fitness relative to the outbred controls. In addition, many loci exhibit heterozygote advantage over and above any effect due to inbreeding, with approximately 30% of markers showing an effect in any given culture condition and approximately 75% of markers showing an effect in at least one of the four culture conditions. To explore the extent of linkage disequilibrium surrounding these loci we further typed four new markers close to each of the three strongest hits. We find a pattern where the extent of heterozygote excess tends to decline to nonsignificance within around 1.5 megabases (Mb) either side of the original hit. Crude extrapolation suggests 12 genes or regions experience detectable levels of heterozygote advantage in any one condition and as many as 25 overall. Thus, balancing selection is widespread and is likely to play an important role in maintaining genetic variability.

Animals↗

Size asymmetry in intraspecific competition and the density-dependence of inbreeding depression in a natural plant population: a case study in cassava (Manihot esculenta Crantz, Euphorbiaceae).

The effects of competition on the genetic composition of natural populations are not well understood. We combined demography and molecular genetics to study how intraspecific competition affects microevolution in cohorts of volunteer plants of cassava (Manihot esculenta) originating from seeds in slash-and-burn fields of Palikur Amerindians in French Guiana. In this clonally propagated crop, genotypic diversity is enhanced by the incorporation of volunteer plants into farmers' stocks of clonal propagules. Mortality of volunteer plants was density-dependent. Furthermore, the size asymmetry of intraspecific competition increased with local clustering of plants. Size of plants was correlated with their multilocus heterozygosity, and stronger size-dependence of survival in clusters of plants, compared with solitary plants, increased the magnitude of inbreeding depression when competition was severe. The density-dependence of inbreeding depression of volunteer plants helps explain the high heterozygosity of volunteers that survive to harvest time and thus become candidates for clonal propagation. This effect could help favour the maintenance of sex in this 'vegetatively' propagated crop plant.

Crops, Agricultural↗

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↗

Inbreeding and relatedness in Scandinavian grey wolves Canis lupus.

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

Alleles↗

Inbreeding and the incidence of childhood genetic disorders in Karnataka, South India.

Consanguineous marriages are strongly favoured among the populations of South India. In a study conducted on 407 infants and children, a total of 35 genetic diseases was diagnosed in 63 persons: 44 with single gene defects, 12 with polygenic disorders, and seven with Down's syndrome. The coefficient of inbreeding of the total study group, F = 0.0414, was significantly higher than that previously calculated for the general population, F = 0.0271, and autosomal recessive disorders formed the largest single disease category diagnosed. The results suggest that long term inbreeding may not have resulted in appreciable elimination of recessive lethals and sub-lethals from the gene pool.

Genes, Dominant↗

The major histocompatibility complex and mate choice: inbreeding avoidance and selection of good genes.

It has been known for decades that MHC genes play a critical role in the cellular immune response, but only recent research has provided a better understanding of how these molecules might affect mate choice. Original studies in inbred mouse strains revealed that mate choice was influenced by MHC dissimilarity. Detection of MHC differences between individuals in these experiments was related to olfactory cues, primarily in urine. Recent studies in humans have shown an analogous picture of MHC-based mating. Taken together, these findings could support either the hypothesis of MHC-based inbreeding avoidance or the hypothesis of MHC-related avoidance of reproductive failure, since studies in mice, humans and pigtailed macaques have shown that parental sharing of certain MHC alleles correlates with frequent spontaneous abortion or prolonged intergestational intervals. Data from many mammalian species clearly demonstrate that reproductive failure occurs as a result of inbreeding. Therefore, MHC similarity might serve as an indicator of genome-wide relatedness. In contrast, increased fitness due to the presence of individual MHC alleles in a pathogenic environment could explain MHC-based selection of currently good genes. Specifically, the physical condition of long-living animals depends on the ability to respond to immunological challenge and an individual's MHC alleles determine the response, since, unlike the T cell receptors, MHC alleles are not somatically recombined. Therefore, sexual selection of condition-dependent traits during mate choice could be used to select successful MHC alleles, thereby providing offspring with a higher relative immunity in their pathogenic environment.

Animals↗

Non-random mating for selection with restricted rates of inbreeding and overlapping generations.

Minimum coancestry mating with a maximum of one offspring per mating pair (MC1) is compared with random mating schemes for populations with overlapping generations. Optimum contribution selection is used, whereby DeltaF is restricted. For schemes with DeltaF restricted to 0.25% per year, 256 animals born per year and heritability of 0.25, genetic gain increased with 18% compared with random mating. The effect of MC1 on genetic gain decreased for larger schemes and schemes with a less stringent restriction on inbreeding. Breeding schemes hardly changed when omitting the iteration on the generation interval to find an optimum distribution of parents over age-classes, which saves computer time, but inbreeding and genetic merit fluctuated more before the schemes had reached a steady-state. When bulls were progeny tested, these progeny tested bulls were selected instead of the young bulls, which led to increased generation intervals, increased selection intensity of bulls and increased genetic gain (35% compared to a scheme without progeny testing for random mating). The effect of MC1 decreased for schemes with progeny testing. MC1 mating increased genetic gain from 11-18% for overlapping and 1-4% for discrete generations, when comparing schemes with similar genetic gain and size.

Age Factors↗

Effect of inbreeding on juvenile mortality in some small mammal species.

In breeding records for 12 species of small mammals from 9 families in 3 orders, individuals with an inbreeding coefficient of 0 were classified as non-inbred, those with inbreeding coefficients greater than 0 as inbred. Juvenile mortality was defined as all deaths prior to 1/2 of the age of sexual maturity for each species. It was significantly higher for inbred than for noninbred animals in 11 of the 12 species using a 1-tailed sign test, and by Fisher's exact test in 3 species. Monodelphis domestica, Wiedomys pyrrhorhinos and and Dolichotis salinicola. The higher mortality of inbred young of the species with the largest sample size, Elephantulus rufescens, was irrespective of litter size and of the parity of the dam.

Age Factors↗

Questioning inbreeding: Could outbreeding affect productivity in the North African catfish in Thailand?

The North African catfish (Clarias gariepinus) is a significant species in aquaculture, which is crucial for ensuring food and nutrition security. Their high adaptability to diverse environments has led to an increase in the number of farms that are available for their production. However, long-term closed breeding adversely affects their reproductive performance, leading to a decrease in production efficiency. This is possibly caused by inbreeding depression. To investigate the root cause of this issue, the genetic diversity of captive North African catfish populations was assessed in this study. Microsatellite genotyping and mitochondrial DNA D-loop sequencing were applied to 136 catfish specimens, collected from three populations captured for breeding in Thailand. Interestingly, extremely low inbreeding coefficients were obtained within each population, and distinct genetic diversity was observed among the three populations, indicating that their genetic origins are markedly different. This suggests that outbreeding depression by genetic admixture among currently captured populations of different origins may account for the low productivity of the North African catfish in Thailand. Genetic improvement of the North African catfish populations is required by introducing new populations whose origins are clearly known. This strategy should be systematically integrated into breeding programs to establish an ideal founder stock for selective breeding.

Animals↗

Genetic basis of heterosis and inbreeding depression in rice (Oryza sativa L.).

The genetic basis of heterosis was studied through mid-parent, standard variety and better parent for 11 quantitative traits in 17 parental lines and their 10 selected hybrids in rice (Oryza sativa L.). The characters were plant height, days to flag leaf initiation, days to first panicle initiation, days to 100% flowering, panicle length, flag leaf length, days to maturity, number of fertile spikelet/panicle, number of effective tillers/hill, grain yield/10-hill, and 1000-grain weight. In general the hybrids performed significantly better than the respective parents. Significant heterosis was observed for most of the studied characters. Among the 10 hybrids, four hybrids viz., 17Ax45R, 25Ax37R, 27Ax39R, 31Ax47R, and 35Ax47R showed highest heterosis in 10-hill grain yield/10-hill. Inbreeding depression of F2 progeny was also studied for 11 characters of 10 hybrids. Both positive and negative inbreeding depression were found in many crosses for the studied characters, but none was found significant. Selection of good parents was found to be the most important for developing high yielding hybrid rice varieties.

Hybrid Vigor↗

Maximizing genetic gain over multiple generations with quantitative trait locus selection and control of inbreeding.

Stochastic computer simulation was used to investigate the potential extra genetic gains obtained from gene-assisted selection (GAS) by combining 1) optimization of genetic contributions for maximizing gain, while restricting the rate of inbreeding with 2) optimization of the relative emphasis given to the QTL over generations. The genetic model assumed implied a mixed inheritance model in which a single quantitative trait locus (i.e., QTL) is segregating together with polygenes. When compared with standard GAS (i.e., fixed contributions and equal emphasis on the QTL and polygenic EBV), combined optimization of contributions of selection candidates and weights on the QTL across generations allowed substantial increases in gain at a fixed rate of inbreeding and avoided the conflict between short- and long-term responses in GAS schemes. Most of the increase of gain was produced by optimization of selection candidates' contributions. Optimization of the relative emphasis given to the QTL over generations had, however, a greater effect on avoiding the long-term loss usually observed in GAS schemes. Optimized contribution schemes led to lower gametic phase disequilibrium between the QTL and polygenes and to higher selection intensities both on the QTL and polygenes than with standard truncation selection with fixed contributions of selection candidates.

Alleles↗