Variation In Inbreeding Depression Research Articles

Comparison among nine alfalfa breeding schemes based on actual biomass yield gains

AbstractThe definition of efficient selection methods for biomass yield of alfalfa and other open‐pollinated forage crops has lagged behind, despite its crucial importance for breeders. This study, performed in northern Italy, aimed to compare nine breeding schemes encompassing the evaluation of replicated clones and half‐sib (HS), first selfing generation (S1) or second selfing generation (S2) families without or with within‐family selection. The comparison was based on actual gains for dry‐matter yield over 43 mo and final plant survival across irrigated and rainfed cropping of advanced generation synthetics (Syn‐3) built from 12 selected parents. The study envisaged biomass yield selection over 31 mo under dense planting among 125 S0 plants and/or their progenies, after a stage of stratified mass selection which acted as a benchmark for genetic gains and whose genetic progress over the original genetic base was concurrently assessed. Half‐sib progeny‐based selection exhibited highest genetic gains with respect to unit time and return on yearly investment for direct costs for both biomass yield and crop persistence. Among‐and‐within‐HS family selection ranked second, and S1 progeny‐based selection ranked third, for yield gains, whereas three schemes implying two selfing generations were valuable for improving persistence but not yield. The initial stage of stratified mass selection was time‐ and cost‐efficient for improving yield but not persistence. We provided information also on self‐fertility and inbreeding depression variation across selfing generations, genetic variation of clonal, HS, S1 and S2 material, and consistency of S0 parent value across different types of material. Our results may have implications for field‐based and genomic selection.

Invasion success in polyploids: the role of inbreeding in the contrasting colonization abilities of diploid versus tetraploid populations of Centaurea stoebe s.l.

Summary As a consequence of founder effects, inbreeding can hamper colonization success: First, in species with self‐incompatibility controlled by an S‐locus, inbreeding may decrease cross‐compatibility, mainly due to the sharing of identical S‐alleles between closely related mating partners. Secondly, inbreeding can reduce fitness of inbred relative to outbred offspring (i.e. inbreeding depression). Polyploids often show reduced inbreeding depression compared to diploids, which may contribute to the overrepresentation of polyploids among invasive species. This is the first study that tests how the effects of inbreeding differ between geocytotypes (i.e. ploidy levels within a given range). Our model organism, Centaurea stoebe, is strictly self‐incompatible and comprises three geocytotypes: diploids are more frequent than tetraploids in the native range, while only tetraploids occur in the invasive range. We conducted a breeding experiment (sib‐mating vs. outcrossing) with 14 native diploid, 13 native tetraploid and 15 invasive tetraploid populations. We recorded cross‐compatibility and estimated a cumulative index for offspring fitness. Since frequent inbreeding can result in purging of genetic load responsible for inbreeding depression, our analyses included a metric for within‐population relatedness, based on eight microsatellite markers, to assess the effect of purging. Inbreeding was found to reduce cross‐compatibility, which was similarly pronounced in diploids and tetraploids. It also caused inbreeding depression in cumulative fitness, which was significant in diploids but not in tetraploids. No evidence of purging was observed as inbred fitness was not affected by within‐population relatedness. Synthesis. Our results provide new insights into the contrasting invasion success of the cytotypes of C. stoebe. As the effects of cross‐compatibility and purging were comparable between cytotypes, both processes can be ruled out to affect the colonization success of diploids versus tetraploids. Our findings are consistent with the hypothesis that polyploidy increases the masking of recessive mutations, which maintains high fitness in inbred tetraploids and may thus facilitate colonization of new ranges. We highlight that reduced inbreeding depression may add to previously acknowledged advantages of polyploids in range expansions, a mechanism that may hitherto have been underestimated due to a lack of data on variation in inbreeding depression across geocytotypes.

Maternal effects alter the severity of inbreeding depression in the offspring.

A maternal effect is a causal influence of the maternal phenotype on the offspring phenotype over and above any direct effects of genes. There is abundant evidence that maternal effects can have a major impact on offspring fitness. Yet, no previous study has investigated the potential role of maternal effects in influencing the severity of inbreeding depression in the offspring. Inbreeding depression is a reduction in the fitness of inbred offspring relative to outbred offspring. Here, we tested whether maternal effects due to body size alter the magnitude of inbreeding depression in the burying beetle Nicrophorus vespilloides We found that inbreeding depression in larval survival was more severe for offspring of large females than offspring of small females. This might be due to differences in how small and large females invest in an inbred brood because of their different prospects for future breeding opportunities. To our knowledge, this is the first evidence for a causal effect of the maternal phenotype on the severity of inbreeding depression in the offspring. In natural populations that are subject to inbreeding, maternal effects may drive variation in inbreeding depression and therefore contribute to variation in the strength and direction of selection for inbreeding avoidance.

The fitness consequences of inbreeding in natural populations and their implications for species conservation – a systematic map

Threatened species often have small and isolated populations where mating among relatives can result in inbreeding depression increasing extinction risk. Effective management is hampered by a lack of syntheses summarising the magnitude of, and variation in inbreeding depression. Here we describe the nature and scope of the literature examining phenotypic/fitness consequences of inbreeding, to provide a foundation for future syntheses and management. We searched the literature for articles documenting the impact of inbreeding in natural populations. Article titles, abstracts and full-texts were assessed against a priori defined criteria, and information relating to study design, quality and other factors that may influence inbreeding responses (e.g. population size) was extracted from relevant articles. The searches identified 11457 articles, of which 614 were assessed as relevant and included in the systematic map (corresponding to 703 distinct studies). Most studies (663) assessed within-population inbreeding resulting from self-fertilisation or consanguineous pairings, while 118 studies assessed among-population inbreeding due to drift load. Plants were the most studied taxon (469 studies) followed by insects (52 studies) and birds (43 studies). Most studies investigated the effects of inbreeding on components of fitness (e.g. survival or fecundity; 648 studies) but measurements were typically under laboratory/greenhouse conditions (486 studies). Observations were also often restricted to the first inbred generation (607 studies) and studies frequently lacked contextual information (e.g. population size). Our systematic map describes the scope and quality of the evidence describing the phenotypic consequences of inbreeding. The map reveals substantial evidence relating to inbreeding responses exists, but highlights information is still limited for some aspects, including the effects of multiple generations of inbreeding. The systematic map allowed us to define several conservation-relevant questions, where sufficient data exists to support systematic reviews, e.g. How do inbreeding responses vary with population size? However, we found that such syntheses are likely to be constrained by incomplete reporting of critical contextual information. Our systematic map employed the same rigorous literature assessment methods as systematic review, including a novel survey of study quality and thus provides a robust foundation to guide future research and syntheses seeking to inform conservation decision-making.

Within and between population variation in inbreeding depression in the locally threatened perennial Scabiosa columbaria

Inbreeding depression plays a central role within the conservation genetics paradigm. Until now inbreeding depression is incorporated into models of population viability as a mean value (e.g. number of lethal equivalents) for all traits in a population. In this study of the locally threatened perennial plant species Scabiosa columbaria we investigated both the mean and the variance among families of inbreeding depression in eight life history traits for five natural populations varying in size from 300 to more than 120,000 individuals. Significant inbreeding depression was found in all populations and all traits. The mean inbreeding depression value per trait was never correlated to population size. Within each population, highly significant variation in inbreeding depression between families (VIFLID) was found. Per trait, families with inbreeding depression next to families with outbreeding depression were often found within the same population. Inbreeding depression at the family level was in many cases not correlated among traits and independent of correlations between traits themselves. VIFLID was negatively correlated with population size: in two traits these correlations were significant. The results underline that inbreeding depression is a complex, highly dynamic phenomenon. Models of viability should incorporate inbreeding depression distributions, with a trait specific mean and variance. Moreover, models of metapopulation dynamics should incorporate genotype quality as factor in colonization success.

Epigenetics as a new avenue for the role of inbreeding depression in evolutionary ecology

Epigenetics as a new avenue for the role of inbreeding depression in evolutionary ecology

Variation in inbreeding depression and plasticity across native and non-native field environments

Since the early 1990s, research on genetic variation of phenotypic plasticity has expanded and empirical research has emphasized the role of the environment on the expression of inbreeding depression. An emerging question is how these two evolutionary ecology mechanisms interact in novel environments. Interest in this area has grown with the need to understand the establishment of populations in response to climate change, and to human-assisted transport to novel environments. We compare performance in the field of outcrossed (O) and inbred lines (S1, S2) from 20 maternal families from each of two native populations of Mimulus guttatus. The experiment was planted in California in each population's home site, in the other populations's home site, in a novel site within the native range of M. guttatus, and in a novel site within the non-native range in North America. The experiment included nearly 6500 individuals. Survival, sexual reproduction and above-ground biomass were examined in order to evaluate inbreeding depression, and stem diameter and plant height were examined in order to evaluate phenotypic plasticity. Across all field sites, approx. 36 % of plants survived to flowering. Inbreeding depression differed among sites and outcrossed offspring generally outperformed selfed offspring. However, in the native-novel site, self-progeny performed better or equally well as outcross progeny. Significant phenotypic plasticity and genetic variation in plasticity was detected in the two architectural traits measured. The absolute value of plasticity showed the most marked difference between home and non-native novel site or non-native-novel site. Evidence was detected for an interaction between inbreeding and plasticity for stem diameter. The results demonstrate that during initial population establishment, both inbreeding depression and phenotypic plasticity vary among field sites, and may be an important response to environments outside a species' currently occupied range. However, the interaction between inbreeding and plasticity may be limited and environment-dependent.

Among-individual variation in pollen limitation and inbreeding depression in a mixed-mating shrub

Variation in inbreeding depression (δ) among individual plants is considered to play a central role in mating system evolution and population genetics. Moreover, such variation could be linked to individual susceptibility to pollen limitation (PL) because those individuals strongly affected by δ for seed production will require more outcross pollen for setting a given number of fruits or seeds. However, no study has tested explicitly for associations between PL and δ at the individual plant level. This study assesses the extent of among-individual variation in PL and δ, the consistency of δ across life stages, and the relationships between individual PL and δ in the mixed-mating shrub Myrtus communis. Controlled hand-pollinations were performed in a natural M. communis population. Marked flowers were monitored until fruit production and a greenhouse experiment was conducted with the seeds produced. Compared with selfing, outcross-pollination enhanced seed number per fruit, germination rate and seedling growth, but did not enhance fruit-set. Only seed number per fruit was pollen limited and, thus, cumulative pollen limitation depended more on pollen quality (outcross pollen) than on quantity. The effects of δ varied considerably across life stages and individual plants. Cumulative δ was high across individuals (mean δ = 0·65), although there were no positive correlations between δ values at different life stages. Interestingly, maternal plants showing stronger δ for seed production were more pollen limited, but they were also less affected by δ for seedling growth because of a seed size/number trade-off. Results show a general inconsistency in δ across life stages and individuals, suggesting that different deleterious loci are acting at different stages. The association between δ and PL at the individual level corroborates the idea that pollen limitation may be 'genotype-dependent' regardless of other factors.

INBREEDING DEPRESSION INCREASES WITH ENVIRONMENTAL STRESS: AN EXPERIMENTAL STUDY AND META-ANALYSIS

Inbreeding-environment interactions occur when inbreeding leads to differential fitness loss in different environments. Inbred individuals are often more sensitive to environmental stress than are outbred individuals, presumably because stress increases the expression of deleterious recessive alleles or cellular safeguards against stress are pushed beyond the organism's physiological limits. We examined inbreeding-environment interactions, along two environmental axes (temperature and rearing host) that differ in the amount of developmental stress they impose, in the seed-feeding beetle Callosobruchus maculatus. We found that inbreeding depression (inbreeding load, L) increased with the stressfulness of the environment, with the magnitude of stress explaining as much as 66% of the variation in inbreeding depression. This relationship between L and developmental stress was not explainable by an increase in phenotypic variation in more stressful environments. To examine the generality of this experimental result, we conducted a meta-analysis of the available data from published studies looking at stress and inbreeding depression. The meta-analysis confirmed that the effect of the environment on inbreeding depression scales linearly with the magnitude of stress; a population suffers one additional lethal equivalent, on average, for each 30% reduction in fitness induced by the stressful environment. Studies using less-stressful environments may lack statistical power to detect the small changes in inbreeding depression. That the magnitude of inbreeding depression scales with the magnitude of the stress applied has numerous repercussions for evolutionary and conservation genetics and may invigorate research aimed at finding the causal mechanism involved in such a relationship.

Inbreeding-environment interactions for fitness: complex relationships between inbreeding depression and temperature stress in a seed-feeding beetle

It is commonly argued that inbred individuals should be more sensitive to environmental stress than are outbred individuals, presumably because stress increases the expression of deleterious recessive alleles. However, the degree to which inbreeding depression is dependent on environmental conditions is not clear. We use two populations of the seed-feeding beetle, Callosobruchus maculatus, to test the hypotheses that (a) inbreeding depression varies among rearing temperatures, (b) inbreeding depression is greatest at the more stressful rearing temperatures, (c) the degree to which high or low temperature is stressful for larval development varies with inbreeding level, and (d) inbreeding depression is positively correlated between different environments. Inbreeding depression (δ) on larval development varied among temperatures (i.e., there was a significant inbreeding-environment interaction). Positive correlations for degree of inbreeding depression were consistently found between all pairs of temperatures, suggesting that at least some loci affected inbreeding depression across all temperatures examined. Despite variation in inbreeding depression among temperatures, inbreeding depression did not increase consistently with our proxy for developmental stress. However, inbreeding changed which environments are benign versus stressful for beetles; although 20°C was not a stressful rearing temperature for outbred beetles, it became the most stressful environment for inbred larvae. The finding that inbreeding-environment interactions can cause normally benign environments to become stressful for inbred populations has important consequences for many areas of evolutionary genetics, artificial breeding (for conservation or food production), and conservation of natural populations.

Genetic variation and parental performance under inbreeding for growth in Eucalyptus globulus

• We aimed to better understand the genetic architecture of growth in E. globulus undergoing inbreeding by comparing families from selfing (SELF), open pollination (OP) and unrelated polymix crossing (POL) of common parents. Stem diameter at breast height (DBH) was assessed at 4, 6 and 10 years after planting in a field trial.• The OP heritability was overestimated at an early age relative to the POL heritability.• No significant correlations were found between the SELF and POL parental effects, indicating substantial non-additive genetic variation under inbreeding. The OP family effects were better correlated with the SELF than the POL population, and only at age 10 years, after substantial mortality of inbred progeny has occurred, was the positive correlation between OP and POL families significantly different from zero.• The estimated dominance variance arising from inbreeding was nearly 10-fold greater than the dominance variance associated with random mating and the additive variance, and appeared to be a major contributor to the variation in inbreeding depression amongst selfed families.

A Genomewide Assessment of Inbreeding Depression: Gene Number, Function, and Mode of Action

Although the genetic basis of inbreeding depression is still being debated, most fitness effects are thought to be the result of increased homozygosity for recessive or partially recessive deleterious alleles rather than the loss of overdominant genes. It is unknown how many loci are associated with inbreeding depression, the genes or gene pathways involved, or their mode of action. To uncover genes associated with variation in fitness following inbreeding, we generated a set of inbred lines of Drosophila melanogaster for which only the third chromosome varied among lines and measured male competitive reproductive success among these lines to estimate inbreeding depression. Male competitive reproductive success for different lines validated our prediction that equally inbred lines show variation in inbreeding depression. To begin to assess the molecular basis of inbreeding depression for male competitive reproductive success, we detected variation in whole-genome gene expression across these inbred lines with commercially available high-density oligonucleotide microarrays. A total of 567 genes were differentially expressed among these inbred lines, indicating that inbreeding directly or indirectly affects a large number of genes: genes that are disproportionately involved in metabolism, stress and defense responses. Subsequently, we generated a set of outbred lines by crossing the highest inbreeding depression lines to each other and contrasted gene expression between parental inbred lines and F(1) hybrids with transcript abundance as a quantitative phenotype to determine the mode of action of the genes associated with inbreeding depression. Although our results indicated that approximately 75% of all genes involved in inbreeding depression were additive, partially additive, or dominant, about 25% of all genes expressed patterns of overdominance. These results should be viewed with caution given that they may be confounded by issues of statistical inference or associative overdominance.

CORRELATED EVOLUTION OF MATING STRATEGY AND INBREEDING DEPRESSION WITHIN AND AMONG POPULATIONS OF THE HERMAPHRODITIC SNAILPHYSA ACUTA

Inbreeding depression is one of the main forces opposing the evolution of self-fertilization. Of central importance is the hypothesis that inbreeding depression and selfing coevolve antagonistically, generating either low selfing rate and high inbreeding depression or vice versa. However, there is limited evidence for this coevolution within species. We investigated this topic in the hermaphroditic snail Physa acuta. In this species, isolated individuals delay the onset of egg laying compared to individuals having access to mates. Longer delays ("waiting times") indicate more intense selfing avoidance. We measured inbreeding depression and waiting time in a large quantitative-genetic experiment (281 outbred families derived from 26 natural populations). We observed large genetic variance for both traits and a strong positive genetic covariance between them, most of which resided within rather than among populations. It means that, within populations, individuals with higher mutation load avoided selfing more strongly on average. This genetic covariance may result from pleiotropy and/or linkage disequilibrium. Whatever its genetic architecture, the fact it emerges specifically when individuals are deprived of mates suggests it is not fortuitous and rather reflects the action of natural selection. We conclude that a diversity of mating strategies can arise within populations subjected to variation in inbreeding depression.

The genetic consequences of fluctuating inbreeding depression and the evolution of plant selfing rates

The magnitude of inbreeding depression, a central parameter in the evolution of plant mating systems, can vary depending on environmental conditions. However, the underlying genetic mechanisms causing environmental fluctuations in inbreeding depression, and the consequences of this variation for the evolution of self-fertilization, have been little studied. Here, we consider temporal fluctuations of the selection coefficient in an explicit genetic model of inbreeding depression. We show that substantial variance in inbreeding depression can be generated at equilibrium by fluctuating selection, although the simulated variance tends to be lower than has been measured in experimental studies. Our simulations also reveal that purging of deleterious mutations does not depend on the variance in their selection coefficient. Finally, an evolutionary analysis shows that, in contrast to previous theoretical approaches, intermediate selfing rates are never evolutionarily stable when the variation in inbreeding depression is due to fluctuations in the selection coefficient on deleterious mutations.

Plant–Mycorrhizal Fungus Interactions Affect the Expression of Inbreeding Depression in Wild Strawberry

The influence of biotic interactions on the expression of inbreeding depression has received only modest investigation; however, these interactions may be important in determining the magnitude and/or variation of inbreeding depression seen in the wild and invoked in models of mating and sexual system evolution. We present the first experimental test of the effects of plant–mycorrhizal fungus interactions on the expression of inbreeding depression. We inoculated selfed and outcrossed seedlings from eight hermaphrodite genotypes of wild strawberry (Fragaria virginiana) with mycorrhizal fungal spores and determined the effect on vegetative growth and sexual and asexual reproduction. We found that inoculated plants grew at rates similar to those of control plants but produced fewer flowers and more plantlets than controls. However, some of these effects varied with maternal genotype and cross type. As a consequence, mycorrhizal fungal inoculation had variable and trait‐dependent effects on the expression of ...

The Different Sources of Variation in Inbreeding Depression, Heterosis and Outbreeding Depression in a Metapopulation ofPhysa acuta

Understanding how parental distance affects offspring fitness, i.e., the effects of inbreeding and outbreeding in natural populations, is a major goal in evolutionary biology. While inbreeding is often associated with fitness reduction (inbreeding depression), interpopulation outcrossing may have either positive (heterosis) or negative (outbreeding depression) effects. Within a metapopulation, all phenomena may occur with various intensities depending on the focal population (especially its effective size) and the trait studied. However, little is known about interpopulation variation at this scale. We here examine variation in inbreeding depression, heterosis, and outbreeding depression on life-history traits across a full-life cycle, within a metapopulation of the hermaphroditic snail Physa acuta. We show that all three phenomena can co-occur at this scale, although they are not always expressed on the same traits. A large variation in inbreeding depression, heterosis, and outbreeding depression is observed among local populations. We provide evidence that, as expected from theory, small and isolated populations enjoy higher heterosis upon outcrossing than do large, open populations. These results emphasize the need for an integrated theory accounting for the effects of both deleterious mutations and genetic incompatibilities within metapopulations and to take into account the variability of the focal population to understand the genetic consequences of inbreeding and outbreeding at this scale.

Inbreeding depression in an autotetraploid herb: a three cohort field study

Autotetraploids are predicted to have reduced inbreeding depression relative to diploids. However, recent theory and information on genomic changes following autopolyploidy suggest that inbreeding depression may be closer to diploids. In three consecutive years, self and outcross pollinations were conducted on autotetraploid Campanulastrum americanum, seeds were planted into native sites, and biennial offspring were followed through seed production. Inbred individuals had lower germination rates, reduced survival, were smaller, and flowered later, producing fewer fruits with fewer seeds. Inbred offspring had 6% of the cumulative fitness of outcross offspring. Although performance varied substantially among cohorts, inbreeding depression for cumulative fitness was relatively constant, with delta ranging only from 0.92 to 0.95. C. americanum, like many outcrossing species, expressed very high amounts of inbreeding depression. This supports the hypothesis that inbreeding depression of some autotetraploids may be similar to that of diploids. Furthermore, few studies have measured temporal variation in inbreeding depression. Constant inbreeding depression given a sixfold range in cohort performance suggests that inbreeding depression may be relatively robust to environmental variation experienced by natural populations.

Variation in inbreeding depression among populations of the seed beetle, Stator limbatus

AbstractInbreeding depression has been documented in many insect species, but the degree to which it varies among traits within populations and among populations within species is poorly understood. We used a single‐generation factorial breeding design to examine variation in inbreeding depression among three populations of the seed‐feeding beetle, Stator limbatus Horn (Coleoptera: Chrysomelidae: Bruchinae), from the southwestern USA. Eggs from sib matings were less likely to develop and hatch, and larval hatch‐to‐adult mortality was higher for offspring of sib matings. Overall, inbreeding resulted in a reduction in the proportion of eggs that produced an adult from >80% for outbred matings in all three populations to an average of only 54% for inbred matings. Of those larvae that survived to adult, inbred beetles took ∼1.5 days (>5%) longer to reach adult. The only measured trait not affected by inbreeding was adult body mass. The degree to which inbreeding increased mortality varied among the populations – inbreeding depression was lowest in the population that is most isolated. Although populations of S. limbatus are generally large in nature our results suggest that increased inbreeding associated with population fragmentation can have substantial effects on fitness of S. limbatus.

Inbreeding depression in the partially self-incompatible endemic plant species Scalesia affinis (Asteraceae) from Galápagos islands

A previous study showed that some individuals of the tetraploid Galapagos endemic Scalesia affinis were able to produce offspring after selfing. The present study compares the fitness of self-pollinated offspring with the fitness of cross-pollinated offspring. Germination success, seedling survival, and four different growth parameters was measured. In most of the studied characters selfed offspring were significantly inferior to outcrossed progeny. The effect was very clear in germination and survival. Outcrossed embryos were 3.4 times more likely to germinate than those that were selfed-fertilized, and the mortality was 84% higher among selfed individuals. Also, there was no genetic variation in inbreeding depression. The present study is based on material from a large population on Isabela Island, Galapagos. At other localities in the archipelago, populations have been through recent dramatic bottlenecks due to the grazing of introduced mammals. Considering the significant inbreeding depression found in the large population and the presence of a partial self-incompatibility system, these small populations are likely to be highly vulnerable and their future survival critically threatened.

Problems in measuring among‐family variation in inbreeding depression

Understanding the sources of variation in inbreeding depression within populations is important for understanding the evolution of selfing rates. At the population level, inbreeding depression is due to decreased heterozygosity caused by inbreeding, which decreases overdominance and increases the frequency of expression of recessive deleterious alleles. However, within individual families inbreeding has two distinct consequences: it reduces heterozygosity and it restricts the alleles present in offspring to those present in the parent. Outcrossing both increases heterozygosity and brings new alleles into a family (compared to the alleles present if the plant is self-pollinated). Both consequences of inbreeding affect offspring fitness, but the most common experimental design used to measure among-family variation in inbreeding depression cannot distinguish them. The result is that variance in inbreeding depression among families is confounded by genetic variation in the traits being measured. Also, correlations (among families) between measures of inbreeding depression or between inbreeding depression and mean trait values are confounded by genetic variation in the traits being measured. I conclude that more complex crossing designs that allow estimation of breeding values for individual families are required to accurately detect and measure among-family variation in inbreeding depression.