Inbreeding Experiments Research Articles

Accelerated inbreeding depression suggests synergistic epistasis for deleterious mutations in Drosophila melanogaster

Epistasis may have important consequences for a number of issues in quantitative genetics and evolutionary biology. In particular, synergistic epistasis for deleterious alleles is relevant to the mutation load paradox and the evolution of sex and recombination. Some studies have shown evidence of synergistic epistasis for spontaneous or induced deleterious mutations appearing in mutation-accumulation experiments. However, many newly arising mutations may not actually be segregating in natural populations because of the erasing action of natural selection. A demonstration of synergistic epistasis for naturally segregating alleles can be achieved by means of inbreeding depression studies, as deleterious recessive allelic effects are exposed in inbred lines. Nevertheless, evidence of epistasis from these studies is scarce and controversial. In this paper, we report the results of two independent inbreeding experiments carried out with two different populations of Drosophila melanogaster. The results show a consistent accelerated inbreeding depression for fitness, suggesting synergistic epistasis among deleterious alleles. We also performed computer simulations assuming different possible models of epistasis and mutational parameters for fitness, finding some of them to be compatible with the results observed. Our results suggest that synergistic epistasis for deleterious mutations not only occurs among newly arisen spontaneous or induced mutations, but also among segregating alleles in natural populations.

Red flour beetle (Tribolium castaneum): From population genetics to functional genomics.

Tribolium castaneum is a small and low maintenance beetle that has emerged as a most suitable insect model for studying developmental biology and functional genetic analysis. Diverse population genetic studies have been conducted using Tribolium as the principal model to establish basic facts and principles of inbreeding experiments and response to the selection and other quantitative genetics fundamentals. The advanced molecular genetic studies presently focused on the use of Tribolium as a typical invertebrate model for higher diploid eukaryotes. After a whole genome sequencing of Tribolium, many areas of functional genomics were unraveled, which enabled the use of it in many technical approaches of genomics. The present text reviews the use of Tribolium in techniques such as RNAi, transgenic studies, immune priming, immunohistochemistry, in situ hybridization, gene sequencing for characterization of microRNAs, and gene editing using engineered endonuclease. In contrast to Drosophila, the T. castaneum holds a robust systemic RNAi response, which makes it an excellent model for comparative functional genetic studies.

Complementary sex determination, inbreeding depression and inbreeding avoidance in a gregarious sawfly.

Although most Hymenoptera reproduce via arrhenotokous haplodiploidy, the underlying genetic mechanisms vary. Of these, the most widespread mechanism appears to be single-locus complementary sex determination (sl-CSD), in which individuals that are diploid and heterozygous at a sex-determining locus are female, and individuals that are homozygous or hemizygous are male. Because inbreeding increases the probability of producing diploid males, which are often sterile or inviable, sl-CSD can generate substantial inbreeding depression. To counteract this, Hymenoptera with traits that promote inbreeding, such as gregariousness, may evolve one or more of the following: inbreeding avoidance, functional diploid males or alternative sex determination mechanisms. Here, we investigate sex determination, inbreeding depression and inbreeding avoidance in Neodiprion lecontei, a gregarious, pine-feeding sawfly in the family Diprionidae. First, via inbreeding experiments and flow cytometry, we demonstrate that this species has CSD. By modeling expected sex ratios under different conditions, we also show that our data are consistent with sl-CSD. Second, via tracking survival in inbred and outbred families, we demonstrate that inbred families have reduced larval survival and that this mortality is partly attributable to the death of diploid males. Third, using a no-choice mating assay, we demonstrate that females are less willing to mate with siblings than nonsiblings. Together, these results suggest that inbreeding depression stemming from CSD has shaped mating behavior in N. lecontei. These results also set the stage for future comparative work that will investigate the interplay between sex determination, ecology and behavior in additional diprionid species that vary in larval gregariousness.

Natural resistance to experimental feline infectious peritonitis virus infection is decreased rather than increased by positive genetic selection

A previous study demonstrated the existence of a natural resistance to feline infectious peritonitis virus (FIPV) among 36% of randomly bred laboratory cats. A genome wide association study (GWAS) on this population suggested that resistance was polygenic but failed to identify any strong specific associations. In order to enhance the power of GWAS or whole genome sequencing to identify strong genetic associations, a decision was made to positively select for resistance over three generations. The inbreeding experiment began with a genetically related parental (P) population consisting of three toms and four queens identified from among the survivors of the earlier study and belonging to a closely related subgroup (B). The subsequent effects of inbreeding were measured using 42 genome-wide STR markers. P generation cats produced 57 first filial (F1) kittens, only five of which (9.0%) demonstrated a natural resistance to FIPV infection. One of these five F1 survivors was then used to produce six F1/P-backcrosses kittens, only one of which proved resistant to FIP. Six of eight of the F1 and F1/P survivors succumbed to a secondary exposure 4–12 months later. Therefore, survival after both primary and secondary infection was decreased rather than increased by positive selection for resistance. The common genetic factor associated with this diminished resistance was a loss of heterozygosity.

Pattern of inbreeding depression, condition dependence, and additive genetic variance in Trinidadian guppy ejaculate traits

In polyandrous species, a male's reproductive success depends on his fertilization capability and traits enhancing competitive fertilization success will be under strong, directional selection. This leads to the prediction that these traits should show stronger condition dependence and larger genetic variance than other traits subject to weaker or stabilizing selection. While empirical evidence of condition dependence in postcopulatory traits is increasing, the comparison between sexually selected and ‘control’ traits is often based on untested assumption concerning the different strength of selection acting on these traits. Furthermore, information on selection in the past is essential, as both condition dependence and genetic variance of a trait are likely to be influenced by the pattern of selection acting historically on it. Using the guppy (Poecilia reticulata), a livebearing fish with high levels of multiple paternity, we performed three independent experiments on three ejaculate quality traits, sperm number, velocity, and size, which have been previously shown to be subject to strong, intermediate, and weak directional postcopulatory selection, respectively. First, we conducted an inbreeding experiment to determine the pattern of selection in the past. Second, we used a diet restriction experiment to estimate their level of condition dependence. Third, we used a half-sib/full-sib mating design to estimate the coefficients of additive genetic variance (CVA) underlying these traits. Additionally, using a simulated predator evasion test, we showed that both inbreeding and diet restriction significantly reduced condition. According to predictions, sperm number showed higher inbreeding depression, stronger condition dependence, and larger CVA than sperm velocity and sperm size. The lack of significant genetic correlation between sperm number and velocity suggests that the former may respond to selection independently one from other ejaculate quality traits. Finally, the association between sperm number and condition suggests that this trait may mediate the genetic benefits of polyandry which have been shown in this species.

Loss of genetic variability in social spiders: genetic and phylogenetic consequences of population subdivision and inbreeding

The consequences of population subdivision and inbreeding have been studied in many organisms, particularly in plants. However, most studies focus on the short-term consequences, such as inbreeding depression. To investigate the consequences of both population fragmentation and inbreeding for genetic variability in the longer term, we here make use of a natural inbreeding experiment in spiders, where sociality and accompanying population subdivision and inbreeding have evolved repeatedly. We use mitochondrial and nuclear data to infer phylogenetic relationships among 170 individuals of Anelosimus spiders representing 23 species. We then compare relative mitochondrial and nuclear genetic variability of the inbred social species and their outbred relatives. We focus on four independently derived social species and four subsocial species, including two outbred–inbred sister species pairs. We find that social species have 50% reduced mitochondrial sequence divergence. As inbreeding is not expected to reduce genetic variability in the maternally inherited mitochondrial genome, this suggests the loss of variation due to strong population subdivision, founder effects, small effective population sizes (colonies as individuals) and lineage turnover. Social species have < 10% of the nuclear genetic variability of the outbred species, also suggesting the loss of genetic variability through founder effects and/or inbreeding. Inbred sociality hence may result in reduction in variability through various processes. Sociality in most Anelosimus species probably arose relatively recently (0.1–2 mya), with even the oldest social lineages having failed to diversify. This is consistent with the hypothesis that inbred spider sociality represents an evolutionary dead end. Heterosis underlies a species potential to respond to environmental change and/or disease. Inbreeding and loss of genetic variability may thus limit diversification in social Anelosimus lineages and similarly pose a threat to many wild populations subject to habitat fragmentation or reduced population sizes.

Hsp90 is important for fecundity, longevity, and buffering of cryptic deleterious variation in wild fly populations

BackgroundIn the laboratory, the Drosophila melanogaster heat shock protein Hsp90 can buffer the phenotypic effects of genetic variation. Laboratory experiments either manipulate Hsp90 activity pharmacologically, or they induce mutations with strong effects in the gene Hsp83, the single-copy fly gene encoding Hsp90. It is unknown whether observations from such laboratory experiments are relevant in the wild.ResultsWe here study naturally occurring mutations in Hsp83, and their effects on fitness and phenotypic buffering in flies derived from wild populations. We examined more than 4500 flies from 42 Drosophila populations distributed world-wide for insertions or deletions of mobile DNA in or near the Hsp83 gene. The insertions we observed occur at low population frequencies, and reduce Hsp83 gene expression. In competition experiments, mutant flies performed much more poorly than wild-type flies. Mutant flies were also significantly less fecund and shorter-lived than wild-type flies, as well as less well buffered against cryptic deleterious variation, as we show through inbreeding experiments. Specifically, in Hsp83 mutant flies female fecundity dropped to much lower levels after inbreeding than in wild-type flies. At even slightly elevated temperatures, inbred mutant Hsp83 populations went extinct, whereas inbred wild-type populations persisted.ConclusionsOur work shows that Hsp90, a regulator of the stress response and of signaling, helps buffer deleterious variation in fruit flies derived from wild population, and that its buffering role becomes even more important under heat stress.

Hsp90 is important for fecundity, longevity, and buffering of cryptic deleterious variation in wild fly populations

BackgroundIn the laboratory, the Drosophila melanogaster heat shock protein Hsp90 can buffer the phenotypic effects of genetic variation. Laboratory experiments either manipulate Hsp90 activity pharmacologically, or they induce mutations with strong effects in the gene Hsp83, the single-copy fly gene encoding Hsp90. It is unknown whether observations from such laboratory experiments are relevant in the wild.ResultsWe here study naturally occurring mutations in Hsp83, and their effects on fitness and phenotypic buffering in flies derived from wild populations. We examined more than 4500 flies from 42 Drosophila populations distributed world-wide for insertions or deletions of mobile DNA in or near the Hsp83 gene. The insertions we observed occur at low population frequencies, and reduce Hsp83 gene expression. In competition experiments, mutant flies performed much more poorly than wild-type flies. Mutant flies were also significantly less fecund and shorter-lived than wild-type flies, as well as less well buffered against cryptic deleterious variation, as we show through inbreeding experiments. Specifically, in Hsp83 mutant flies female fecundity dropped to much lower levels after inbreeding than in wild-type flies. At even slightly elevated temperatures, inbred mutant Hsp83 populations went extinct, whereas inbred wild-type populations persisted.ConclusionsOur work shows that Hsp90, a regulator of the stress response and of signaling, helps buffer deleterious variation in fruit flies derived from wild population, and that its buffering role becomes even more important under heat stress.

The action of evolutionary forces on metric traits

Fisher's theorem of natural selection implies that the population genetic variance of quasi-neutral traits should be mostly additive. In the case of fitness component traits, however, that variance would be characterised by a substantial contribution from non-additive loci. In parallel, Robertson's theorem states that selection will change the population mean of a trait proportionally to the magnitude of the genetic correlation between that trait and fitness, which should be weak for quasi-neutral traits or strong for the mean fitness components. Drosophila data from inbreeding and artificial selection experiments are discussed within that theoretical framework. In addition, the process of regeneration by mutation of the genetic variance of a quasi-neutral trait (abdominal bristle number) in a Drosophila population initially homozygous at all loci has been analysed. After 485 generations of mutation accumulation, the levels of additive variance found in this population closely approached those commonly observed in laboratory populations. Furthermore, these values, together with previously reported estimates for natural populations, could be jointly explained by a model assuming weak causal stabilising selection.

The ultrastructure of a characteristic spermhead-defect in the boar: the SME-defect.

Semen samples from an eight-month-old landrace boar showing poor sperm motility were investigated at the boars' arrival to an A.I. centre. Sperm morphology was found abnormal in +/- 30% of the sperm heads, which contained a rounded, often circular hypochromatic body (eosin-nigrosin stain) about 2 mum in diameter. The body was located in the front half of the sperm head, i.e. within the region of the acrosomal cap. Later a similar defect (+/- 50%) was found in an imported 2 1/2-year-old Yorkshire boar with decreased fertility. Electron microscopy of ejaculates and testis tissue showed that the hypochromatic body was a cyst containing a body of variable size but of moderate electron density. Usually the cyst was located within the nucleus and sometimes with direct communication to the acrosonal system. During spermiogenesis the abnormal spermids may show a deep invagination into the nucleus of the acrosomal granule. Inbreeding experiments are planned in order to determine if a possible hereditary factor is involved in the defect.

Inbreeding, eugenics, and Helen Dean King (1869–1955)

Helen Dean King's scientific work focused on inbreeding using experimental data collected from standardized laboratory rats to elucidate problems in human heredity. The meticulous care with which she carried on her inbreeding experiments assured that her results were dependable and her theoretical explanations credible. By using her nearly homozygous rats as desired commodities, she also was granted access to venues and people otherwise unavailable to her as a woman. King's scientific career was made possible through her life experiences. She earned a doctorate from Bryn Mawr College under Thomas Hunt Morgan and spent a productive career at the Wistar Institute of Anatomy and Biology in Philadelphia where she had access to the experimental subjects which made her career possible. In this paper I examine King's work on inbreeding, her participation in the debates over eugenics, her position at the Wistar Institute, her status as a woman working with mostly male scientists, and her involvement with popular science.

Single-locus sex determination in the parasitoid wasp Cotesia glomerata (Hymenoptera: Braconidae)

The parasitoid Cotesia glomerata usually produces female-biased sex ratios in the field, which are presumably caused by inbreeding and local mate competition (LMC); yet, sibling mating increases the production of males, leading to the male-biased sex ratio of broods in the laboratory. Previous studies have suggested that the sex allocation strategy of C. glomerata is based on both partial LMC in males and inbreeding avoidance in females. The current study investigated the presence of single-locus complementary sex determination (sl-CSD) as a sex-determining mechanism in this species through inbreeding experiment, cytological examination and microsatellite analysis. Cytological examination detected diploid males in nine of 17 single pairs of sibling mating, thus in agreement with the proportion of matched matings predicted by the sl-CSD model. Sex ratio shifts in these matched sibling matings were consistent with the sl-CSD model with less viable diploid males. The haploid males have a single set of maternal chromosomes (n = 10), whereas diploid males possess a double set of chromosomes (2n = 20). Microsatellite analyses confirmed that diploid males produced from the matched matings inherited segregating genetic materials from both parents. Thus, this study provides the first solid evidence for the presence of sl-CSD as a sex-determining mechanism in the braconid genus Cotesia.

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

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

Single-locus complementary sex determination in the inbreeding wasp Euodynerus foraminatus Saussure (Hymenoptera: Vespidae)

The Hymenoptera have arrhenotokous haplodiploidy in which males normally develop from unfertilized eggs and are haploid, while females develop from fertilized eggs and are diploid. Multiple sex determination systems are known to underlie haplodiploidy, and the best understood is single-locus complementary sex determination (sl-CSD) in which sex is determined at a single polymorphic locus. Individuals heterozygous at the sex locus develop as females; individuals that are hemizygous (haploid) or homozygous (diploid) at the sex locus develop as males. sl-CSD can be detected with inbreeding experiments that produce diploid males in predictable proportions as well as sex ratio shifts due to diploid male production. This sex determination system is considered incompatible with inbreeding because the ensuing increase in homozygosity increases the production of diploid males that are inviable or infertile, imposing a high cost on matings between close relatives. However, in the solitary hunting wasp Euodynerus foraminatus, a species suspected of having sl-CSD, inbreeding may be common due to a high incidence of sibling matings at natal nests. In laboratory crosses with E. foraminatus, we find that sex ratios and diploid male production (detected as microsatellite heterozygosity) are consistent with sl-CSD, but not with other sex determination systems. This is the first documented example of sl-CSD in a hymenopteran with an apparent natural history of inbreeding, and thus presents a paradox for our understanding of hymenopteran genetics.

INBREEDING, DEVELOPMENTAL STABILITY, AND CANALIZATION IN THE SAND CRICKET GRYLLUS FIRMUS

Inbreeding, the mating of close relatives, is known to have deleterious effects on fitness traits in organisms. Developmental stability (DS) and canalization may represent two processes that allow an organism to maintain a stable development that will produce the fittest phenotype. Inbreeding is thus expected to affect either DS or canalization. We tested if inbreeding affects DS and canalization using an inbreeding experiment on the cricket Gryllus firmus. We compared mean length, fluctuating asymmetry (as an index of DS), and morphological variation (as an index of canalization) of four limb traits between seven highly inbred lines, their F1 crosses, and outbred lines originated from the same stock population and maintained in the same environmental conditions. We show evidence for moderate inbreeding depression on the four measures of leg length. The nonsystematic difference in fluctuating asymmetry indices between breed types indicates that inbreeding or heterozygosity did not affect DS, or that fluctuating asymmetry is not a reliable index of DS. In contrast, inbreeding appears to affect canalization, as shown by the significantly higher variation in inbred lines compared to other lines. Identical low variation values in the crossbred and outbred lines indicate that heterozygosity could affect canalization. High variation in morphological variation and fluctuating asymmetry within crossbred or inbred lines, however, suggest the effect of recessive deleterious alleles on both canalization and DS. Although the strong correlation in morphological variation among traits suggests that identical genetic mechanisms govern canalization for all the limb traits, the absence of significant correlation in fluctuating asymmetry among traits causes us to reject this hypothesis for DS. For most of the traits, morphological variation and fluctuating asymmetry were not significantly correlated, which support the hypothesis that canalization and DS consist in two distinct mechanisms.

INBREEDING, DEVELOPMENTAL STABILITY, AND CANALIZATION IN THE SAND CRICKET GRYLLUS FIRMUS

Inbreeding, the mating of close relatives, is known to have deleterious effects on fitness traits in organisms. Developmental stability (DS) and canalization may represent two processes that allow an organism to maintain a stable development that will produce the fittest phenotype. Inbreeding is thus expected to affect either DS or canalization. We tested if inbreeding affects DS and canalization using an inbreeding experiment on the cricket Gryllus firmus. We compared mean length, fluctuating asymmetry (as an index of DS), and morphological variation (as an index of canalization) of four limb traits between seven highly inbred lines, their F1 crosses, and outbred lines originated from the same stock population and maintained in the same environmental conditions. We show evidence for moderate inbreeding depression on the four measures of leg length. The nonsystematic difference in fluctuating asymmetry indices between breed types indicates that inbreeding or heterozygosity did not affect DS, or that fluctuating asymmetry is not a reliable index of DS. In contrast, inbreeding appears to affect canalization, as shown by the significantly higher variation in inbred lines compared to other lines. Identical low variation values in the crossbred and outbred lines indicate that heterozygosity could affect canalization. High variation in morphological variation and fluctuating asymmetry within crossbred or inbred lines, however, suggest the effect of recessive deleterious alleles on both canalization and DS. Although the strong correlation in morphological variation among traits suggests that identical genetic mechanisms govern canalization for all the limb traits, the absence of significant correlation in fluctuating asymmetry among traits causes us to reject this hypothesis for DS. For most of the traits, morphological variation and fluctuating asymmetry were not significantly correlated, which support the hypothesis that canalization and DS consist in two distinct mechanisms.

Inbreeding and building up small populations of stingless bees (Hymenoptera, Apidae)

A study of the viability of small populations of Hymenoptera is a matter of importance to gain a better zoological, ethological, genetical and ecological knowledge of these insects, and for conservation purposes, mainly because of the consequences to the survival of colonies of many species of bees, wasps, and ants. Based on the Whiting (1943) principle, Kerr & Vencovski (1982) presented a hypothesis that states that viable populations of stingless bees (Meliponini) should have at least 40 colonies to survive. This number was later extended to 44 colonies by Kerr (1985). This would be necessary to avoid any substantial amount of homozygosis in the pair of chromosomic sexual loci, by keeping at least six different sexual gene alleles in a reproductive population. In most cases this would prevent the production of useless diploid males. However, several facts weigh against considering this as a general rule. From 1990 to 2001, 287 colony divisions were made, starting with 28 foundation colonies, in the inbreeding and population experiments with the Meliponini reported here. These experiments constitute the most extensive and longest scientific research ever made with Meliponini bees. In ten different experiments presented here, seven species (one with two subspecies) of Meliponini bees were inbred in five localities inside their wide-reaching native habitats, and in two localities far away from these habitats. This was done for several years. On the whole, the number of colonies increased and the loss of colonies over the years was small. In two of these experiments, although these populations were far (1,000 km and 1,200 km) from their native habitat, their foundation colonies were multiplied successfuly. It was possible to build up seven strong and three expanding medium populations, starting with one, two, three or even five colonies. However, in six other cases examined here, the Whiting (1943) principle and the hypothesis of Kerr & Vencovski (1982) and Kerr (1985), possibly hold up. In two other cases, the results are still unclear. Outside native habitats, most inbreeding experiments failed, possibly because of conditions that cause ecological stress. Although much more data are still needed, a new working hypothesis on the molecular level was presented to explain the results of the experiments described here. In the absence of any considerable stress, and in the eventuality of a good nutritive situation, even individual bees that are homozygous in the pair of chromosomic sexual locus would produce a sufficient amount of a sex determining substance. Therefore, the female genes of all the diploid individuals of a colony, both homozygous and heterozygous, would be activated. However, situations of considerable stress would cause a poor physiological and nutritive condition. This, together with homozygosis in the pair of chromosomic sexual locus, would lead to a smaller production of the sex determining substance. When this happens in the diploid homozygous individuals of a colony, in relation to sex, only male genes are activated. As a result, all such homozygous diploid bees of the colony become useless males. However, when there is a heterozygous situation in the chromosomic sexual locus of all bees of a colony, all diploid individuals would produce a high amount of the sex determining substance. Consequently, all diploid individuals of such a colony would become females (queens and workers). Stresses, including ecological stress, as well as the nutritive condition and the genetic situation in the chromosomic sexual loci, will have a key influence in the life and behavior of the Meliponini, including sex determination. In relation to genetic factors, hybrid vigour may often cause a greater production of biological substances. This may be due to the presence of a greater number of copies of allelic genes when there is heterozygosis. This is a hypothesis requiring further research. However, in the experiments presented here, this hypothesis seems to apply well to the production of a sex determining substance in bees (Apoidea) and other Hymenoptera.

THE EVOLUTION OF TRADE-OFFS: EFFECTS OF INBREEDING ON FECUNDITY RELATIONSHIPS IN THE CRICKET GRYLLUS FIRMUS

The evolution of traits is modulated by their interrelationships with each other, particularly when those relationships result in a fitness trade-off. In this paper we explore the consequences of genetic architecture on functional relationships between traits. Specifically, we address the consequences of inbreeding on these relationships. We show that the linear regression between two traits will not be affected if there is no dominance genetic variance in either trait, whereas the intercept but not the slope of the regression will change if there is dominance genetic variance in one trait only. We test the latter hypothesis using fecundity relationships in the cricket Gryllus firmus. Data from pedigree analysis and an inbreeding experiment show that there is significant dominance genetic variance in fecundity, but not head width (an index of body size) or dorsal longitudinal muscle (DLM) mass. Fecundity increases with head width, but decreases with DLM mass. As predicted, the intercepts of the regressions of fecundity on these two morphological traits decrease with inbreeding, but there is little or no change in slope. Gryllus firmus is wing dimorphic, with the macropterous (LW) morph having a lower fecundity than the micropterous (SW) morph. We hypothesize that the difference in fecundity arises primarily because of a competition for resources in the LW females between DLM maintenance (i.e., mass) and egg production. As a consequence, we predict that the fecundity within each morph should decline linearly with the inbreeding coefficient at the same rate in both morphs. The result of this will be a change in the relative fitness of the two morphs, that of the SW morph increasing with inbreeding. This prediction is supported. These results indicate that trade-offs will evolve and such changes will affect evolutionary trajectories by altering the pattern of relationships among fitness components.Corresponding Editor: D. Wheeler

The evolution of trade-offs: effects of inbreeding on fecundity relationships in the cricket Gryllus firmus.

The evolution of traits is modulated by their interrelationships with each other, particularly when those relationships result in a fitness trade-off. In this paper we explore the consequences of genetic architecture on functional relationships between traits. Specifically, we address the consequences of inbreeding on these relationships. We show that the linear regression between two traits will not be affected if there is no dominance genetic variance in either trait, whereas the intercept but not the slope of the regression will change if there is dominance genetic variance in one trait only. We test the latter hypothesis using fecundity relationships in the cricket Gryllus firmus. Data from pedigree analysis and an inbreeding experiment show that there is significant dominance genetic variance in fecundity, but not head width (an index of body size) or dorsal longitudinal muscle (DLM) mass. Fecundity increases with head width, but decreases with DLM mass. As predicted, the intercepts of the regressions of fecundity on these two morphological traits decrease with inbreeding, but there is little or no change in slope. Gryllus firmus is wing dimorphic, with the macropterous (LW) morph having a lower fecundity than the micropterous (SW) morph. We hypothesize that the difference in fecundity arises primarily because of a competition for resources in the LW females between DLM maintenance (i.e., mass) and egg production. As a consequence, we predict that the fecundity within each morph should decline linearly with the inbreeding coefficient at the same rate in both morphs. The result of this will be a change in the relative fitness of the two morphs, that of the SW morph increasing with inbreeding. This prediction is supported. These results indicate that trade-offs will evolve and such changes will affect evolutionary trajectories by altering the pattern of relationships among fitness components.

Inbreeding depression by recessive deleterious genes affecting female fecundity of a haplo-diploid mite

Abstract The effect of inbreeding on haplo-diploid organisms has been regarded as very low, because deleterious recessive genes on hemizygous (haploid) males were immediately purged generation by generation. However, we determined such recessive genes to decrease female fecundity in a population of Schizotetranychus miscanthi Saito which is known in the Acari as a subsocial species with haplo-diploidy. In mother–son inbreeding experiments, there was no depression in egg hatchability nor in the larval survival of progeny over four generations. There was, on the other hand, significant inbreeding depression in the fecundity with increasing f-value. Crosses between two lineages, one having deleterious effects on the fecundity and the other having no such effects, established during the inbreeding, revealed heterosis, and backcrosses showed that the depression was caused by deleterious recessive(s). These results strongly suggest the existence of some deleterious genes governing only the traits of adult females in wild populations of haplo-diploid organisms.