Kin detection cues and sibling relationship quality in adulthood: The role of childhood co-residence duration and maternal perinatal association

Cues facilitating kin detection among children have been suggested to have a profound impact on adult sibling relationships. Using a large and population-based data of younger and middle-aged Finns, we test how childhood co-residence duration and maternal perinatal association (MPA) correlate with contact frequency, emotional closeness and provision of help between adult siblings. Employing sibling fixed-effect regression models we show that duration of co-residence in childhood and MPA are indeed associated with better relationship quality in all three measures. Provided MPA, sibling relationship quality is high independently of co-residence length, but in the absence of MPA, increased co-residence duration is associated with better relationship quality. Co-residence duration is more strongly associated with provision of help between opposite-gender than same-gender sibling dyads. Full siblings report better relationship quality than half siblings do, although the co-residence duration mediates the effect of genetic relatedness in emotional closeness between full and maternal half siblings and in provision of help between full and paternal half siblings. Moreover, MPA serves as a mediator in the case of emotional closeness between full and maternal half siblings. Our results provide solid support for the importance of childhood kin detection cues for sibling relationship quality in adulthood, and how such cues interact with genetic relatedness and gender.     

Kin recognition in Rana cascadae tadpoles: maternal and paternal effects

Cascades frog (Rana cascadae) tadpoles preferentially associate with full siblings over half sibling and half siblings over non-siblings when reared with siblings or as isolates. These tadpoles can use cues of maternal or paternal origin in distinguishing siblings from non-siblings, but maternal cues are preferred over paternal cues. This suggests that a hierarchy of cue importance may exist. Our results are consistent with both a phenotypic matching and a genetic recognition system of kin recognition. Thus, both learned and innate components may play a role in R. cascadae sibling recognition. Kin recognition may facilitate preferential treatment of kin, such as cooperation in food finding or in warning against predators, and therefore those individuals behaving altruistically in kin groups can increase their inclusive fitness.     

Cannibalistic siblicide in praying mantis nymphs (Miomantis caffra)

Inclusive fitness theory predicts that cannibalism should be more likely to arise if close relatives can be avoided, suggesting that cannibalistic species will possess mechanisms for minimizing predation on kin. Juvenile Miomantis caffra are good candidates for the possession of such traits because; (1) groups of siblings hatch together into the same locale, (2) they are aggressive hunters, and (3) they are strongly cannibalistic. In this study, the possibility of kin recognition or avoidance in M. caffra is investigated by laboratory comparison of cannibalism rates between groups of differing relatedness. In order to examine the likelihood of encounters between early instar siblings, the extent of dispersal away from the ootheca in the days following hatching is also observed. Nymphs did not rapidly disperse after hatching, so the chances of full siblings encountering one another in the wild appear to be high. Despite this, cannibalism was equally high in groups of full siblings and groups of mixed parenthood. We suggest that for M. caffra, a generalist ambush predator, the benefits of indiscriminate aggression may outweigh any inclusive fitness benefits that would be gained from kin discrimination.     

Intralocus sexual conflict for fitness: sexually antagonistic alleles for testosterone

Intralocus sexual conflict occurs when a trait encoded by the same genetic locus in the two sexes has different optima in males and females. Such conflict is widespread across taxa, however, the shared phenotypic traits that mediate the conflict are largely unknown. We examined whether the sex hormone, testosterone (T), that controls sexual differentiation, contributes to sexually antagonistic fitness variation in the bank vole, Myodes glareolus. We compared (opposite-sex) sibling reproductive fitness in the bank vole after creating divergent selection lines for T. This study shows that selection for T was differentially associated with son versus daughter reproductive success, causing a negative correlation in fitness between full siblings. Our results demonstrate the presence of intralocus sexual conflict for fitness in this small mammal and that sexually antagonistic selection is acting on T. We also found a negative correlation in fitness between parents and their opposite-sex progeny (e.g. father-daughter), highlighting a dilemma for females, as the indirect genetic benefits of selecting reproductively successful males (high T) are lost with daughters. We discuss mechanisms that may mitigate this disparity between progeny quality.     

INBREEDING INFLUENCES WITHIN‐BROOD HETEROZYGOSITY‐FITNESS CORRELATIONS (HFCS) IN AN ISOLATED PASSERINE POPULATION

Molecular estimates of inbreeding may be made using genetic markers such as microsatellites, however the interpretation of resulting heterozygosity‐fitness correlations (HFCs) with respect to inbreeding depression is not straightforward. We investigated the relationship between pedigree‐determined inbreeding coefficients (f) and HFCs in a closely monitored, reintroduced population of Stewart Island robins (Petroica australis rakiura) on Ulva Island, New Zealand. Using a full sibling design, we focused on differences in juvenile survival associated specifically with individual sibling variation in standardized multilocus heterozygosity (SH) when expected f was identical. We found that within broods, siblings with higher SH at microsatellite loci experienced a higher probability of juvenile survival. This effect, however, was detected primarily within broods that experienced inbreeding or when inbreeding had occurred in their pedigree histories (i.e., at the parents’ level). Thus we show, for the first time in a wild population, that the strength of an HFC is partially dependent on the presence of inbreeding events in the recent pedigree history. Our results illustrate the importance of realized effects of inbreeding on genetic variation and fitness and the value of full‐sibling designs for the study of HFCs in the context of small, inbred populations.     

Parental substance misuse and reproductive timing in offspring: A genetically informed study

Parents with substance misuse provide their children with a potentially risky rearing environment. According to the evolutionary life history theory, such environments steer individuals towards faster reproductive strategy. However, parents providing their children with hazardous environments may also pass on genes associated with early parenthood. In register data on individuals born in Sweden 1973–1993 (N = 2,176,128), the hazard ratio of entering parenthood by age 25 was 1.70 (95% CI 1.69–1.72) and 1.85 (1.82–1.89) among offspring of fathers and mothers with substance misuse, respectively, compared to others. In analyses using an offspring-of-siblings design in three types of parental sibling pairs (half siblings, full siblings and dizygotic twins, and monozygotic twins) increasingly controlling for genetic confounding, associations between parental substance misuse and offspring's early reproduction gradually attenuated. Our results suggest that the association between parental substance misuse and earlier parenthood in offspring is at least partly genetically confounded. Life history theory should be further tested with genetically informative research designs.     

Parentage and Sibship Inference From Multilocus Genotype Data Under Polygamy

Likelihood methods have been developed to partition individuals in a sample into sibling clusters using genetic marker data without parental information. Most of these methods assume either both sexes are monogamous to infer full sibships only or only one sex is polygamous to infer full sibships and paternal or maternal (but not both) half sibships. We extend our previous method to the more general case of both sexes being polygamous to infer full sibships, paternal half sibships, and maternal half sibships and to the case of a two-generation sample of individuals to infer parentage jointly with sibships. The extension not only expands enormously the scope of application of the method, but also increases its statistical power. The method is implemented for both diploid and haplodiploid species and for codominant and dominant markers, with mutations and genotyping errors accommodated. The performance and robustness of the method are evaluated by analyzing both simulated and empirical data sets. Our method is shown to be much more powerful than pairwise methods in both parentage and sibship assignments because of the more efficient use of marker information. It is little affected by inbreeding in parents and is moderately robust to nonrandom mating and linkage of markers. We also show that individually much less informative markers, such as SNPs or AFLPs, can reach the same power for parentage and sibship inferences as the highly informative marker simple sequence repeats (SSRs), as long as a sufficient number of loci are employed in the analysis.     

Sibling cannibalism in the ladybird beetleHarmonia axyridis: Fitness consequences for mother and offspring

The adaptive significance of sibling cannibalism was analyzed in the ladybird beetle Harmonia axyridis at two prey densities. Possible costs and benefits were considered from three points of view: the mother, the cannibal and the victim. Cannibals ate both infertile and fertile eggs, thereby increasing in body length and survival rate with the intensity of sibling cannibalism. The cannibalistic trait was clearly beneficial to the cannibal when aphid density was low. However, it was not always beneficial when aphid density was high and the victims were full siblings. The altruistic behavior of being a victim was beneficial only when the victim was cannibalized by full siblings at low aphid density. The mother attained almost equal fitness at low aphid density, regardless of the intensity of sibling cannibalism. This suggests that sibling cannibalism is not maladaptive for the mother. At high aphid density, however, mother's fitness decreased with the intensity of sibling cannibalism, indicating that sibling cannibalism is maladaptive for the mother when larval food availability is high.     

Human ability to detect kinship in strangers' faces: effects of the degree of relatedness

The resemblance between human faces has been shown to be a possible cue in recognizing the relatedness between parents and children, and more recently, between siblings. However, the general inclusive fitness theory proposes that kin-selective behaviours are also relevant to more distant relatives, which requires the detection of larger kinship bonds. We conducted an experiment to explore the use of facial clues by ‘strangers’, i.e. evaluators from a different family, to associate humans of varying degrees of relatedness. We hypothesized that the visual capacity to detect relatedness should be weaker with lower degrees of relatedness. We showed that human adults are capable of (although not very efficient at) assessing the relatedness of unrelated individuals from photographs and that visible facial cues vary according to the degree of relatedness. This sensitivity exists even for kin pair members that are more than a generation apart and have never lived together. Collectively, our findings are in agreement with emerging knowledge on the role played by facial resemblance as a kinship cue. But we have progressed further to show how the capacity to distinguish between related and non-related pairs applies to situations relevant to indirect fitness.     

Postcopulatory inbreeding avoidance in guppies

In many species, the negative fitness effects of inbreeding have facilitated the evolution of a wide range of inbreeding avoidance mechanisms. Although avoidance mechanisms operating prior to mating are well documented, evidence for postcopulatory mechanisms of inbreeding avoidance remain scarce. Here, we examine the potential for paternity biases to favour unrelated males when their sperm compete for fertilizations though postcopulatory inbreeding avoidance mechanisms in the guppy, Poecilia reticulata. To test this possibility, we used a series of artificial inseminations to deliver an equal number of sperm from a related (either full sibling or half sibling) and unrelated male to a female while statistically controlling for differences in sperm quality between rival ejaculates. In this way, we were able to focus exclusively on postcopulatory mechanisms of inbreeding avoidance and account for differences in sperm competitiveness between rival males. Under these carefully controlled conditions, we report a significant bias in paternity towards unrelated males, although this effect was only apparent when the related male was a full sibling. We also show that sperm competition generally favours males with highly viable sperm and thus that some variance in sperm competitiveness can be attributed to difference in sperm quality. Our findings for postcopulatory inbreeding avoidance are consistent with prior work on guppies, revealing that sperm competition success declines linearly with the level of relatedness, but also that such effects are only apparent at relatedness levels of full siblings or higher. These findings reveal that postcopulatory processes alone can facilitate inbreeding avoidance.     

Ontogenetic isolation favours sibling cannibalism in mites

Inclusive fitness theory suggests that cannibalistic individuals should preferentially eat unrelated prey when given a choice between related and unrelated individuals. Using the predatory mite Phytoseiulus persimilis, I investigated whether having contact with siblings during a sensitive ontogenetic phase or not determines whether cannibalistic protonymphs tend to eat more sibling or nonsibling larvae in choice situations. Cannibals reared with siblings preferentially ate nonsibling larvae when these had been reared in a sibling group but showed no preference when the larvae had been reared in isolation. Cannibals reared in isolation showed no preference when the larvae had been reared in a sibling group and preferentially ate sibling larvae when these had been reared alone. The occurrence of preferential sibling cannibalism when both cannibal and prey had been reared in isolation suggests the use of self-referent phenotype matching to distinguish between siblings and nonsiblings.     

Genetics of obesity in adult adoptees and their biological siblings.

An adoption study of genetic effects on obesity in adulthood was carried out in which adoptees separated from their natural parents very early in life were compared with their biological full and half siblings reared by their natural parents. The adoptees represented four groups who by sampling from a larger population were categorised as either thin, medium weight, overweight, or obese. Weight and height were obtained for 115 full siblings of 57 adoptees and for 850 half siblings of 341 adoptees. In full siblings body mass index (kg/m2) significantly increased with weight of the adoptees. Body mass index of the half siblings showed a steady but weaker increase across the four weight groups of adoptees. There were no significant interactions with sex of the adoptees, sex of the siblings, or (for the half siblings) sex of the common parent. In contrast with the findings in half siblings and (previously) the natural parents there was a striking, significant increase in body mass index between full siblings of overweight and obese adoptees. The degree of fatness in adults living in the same environment appears to be influenced by genetic factors independent of sex, which may include polygenic as well as major gene effects on obesity.     

The influence of mating system on the intensity of parent-offspring conflict in primates

An evolutionary conflict of interest exists between parents and their offspring over the partitioning of parental investment (PI) among siblings. When the direct fitness benefits to offspring of increased PI, outweigh the inclusive fitness costs from lost future sibling fitness, selection should favour the evolution of offspring selfishness over altruism. In theory, this conflict is heightened when females are not strictly monogamous, as current offspring should be less altruistic towards future half-siblings than they would be towards full-siblings. Using data collected on foetal growth rate (representing prenatal PI) in primates, I test the prediction from theory that the resolution of the parent-offspring conflict will be closer to the offspring's evolutionary optima in polyandrous species than in more monandrous species. Using phylogenetic comparative analysis, and controlling for allometry, I show that offspring are able to obtain more PI when the probability of future full-siblings decreases, and that this is most pronounced in taxa where there is the opportunity for direct foetal access to the maternal bloodstream. These results support the hypothesis that the resolution of prenatal PI conflict is influenced by both a species' mating system and by its placental structure.     

Delayed Phenotypic Expression of Growth Hormone Transgenesis during Early Ontogeny in Atlantic Salmon (Salmo salar)?

Should growth hormone (GH) transgenic Atlantic salmon escape, there may be the potential for ecological and genetic impacts on wild populations. This study compared the developmental rate and respiratory metabolism of GH transgenic and non-transgenic full sibling Atlantic salmon during early ontogeny; a life history period of intense selection that may provide critical insight into the fitness consequences of escaped transgenics. Transgenesis did not affect the routine oxygen consumption of eyed embryos, newly hatched larvae or first-feeding juveniles. Moreover, the timing of early life history events was similar, with transgenic fish hatching less than one day earlier, on average, than their non-transgenic siblings. As the start of exogenous feeding neared, however, transgenic fish were somewhat developmentally behind, having more unused yolk and being slightly smaller than their non-transgenic siblings. Although such differences were found between transgenic and non-transgenic siblings, family differences were more important in explaining phenotypic variation. These findings suggest that biologically significant differences in fitness-related traits between GH transgenic and non-transgenic Atlantic salmon were less than family differences during the earliest life stages. The implications of these results are discussed in light of the ecological risk assessment of genetically modified animals.     

Accurate inference of relationships in sib-pair linkage studies.

Relative-pair designs are routinely employed in linkage studies of complex genetic diseases and quantitative traits. Valid application of these methods requires correct specification of the relationships of the pairs. For example, within a sibship, presumed full sibs actually might be MZ twins, half sibs, or unrelated. Misclassification of half-sib pairs or unrelated individuals as full sibs can result in reduced power to detect linkage. When other family members, such as parents or additional siblings, are available, incorrectly specified relationships usually will be detected through apparent incompatibilities with Mendelian inheritance. Without other family members, sibling relationships cannot be determined absolutely, but they still can be inferred probabilistically if sufficient genetic marker data are available. In this paper, we describe a simple likelihood ratio method to infer the true relationship of a putative sibling pair. We explore the number of markers required to accurately infer relationships typically encountered in a sib-pair study, as a function of marker allele frequencies, marker spacing, and genotyping error rate, and we conclude that very accurate inference of relationships can be achieved, given the marker data from even part of a genome scan. We compare our method to related methods of relationship inference that have been suggested. Finally, we demonstrate the value of excluding non-full sibs in a genetic linkage study of non-insulin-dependent diabetes mellitus.     

Evaluating indirect genetic effects of siblings using singletons

Estimating effects of parental and sibling genotypes (indirect genetic effects) can provide insight into how the family environment influences phenotypic variation. There is growing molecular genetic evidence for effects of parental phenotypes on their offspring (e.g. parental educational attainment), but the extent to which siblings affect each other is currently unclear. Here we used data from samples of unrelated individuals, without (singletons) and with biological full-siblings (non-singletons), to investigate and estimate sibling effects. Indirect genetic effects of siblings increase (or decrease) the covariance between genetic variation and a phenotype. It follows that differences in genetic association estimates between singletons and non-singletons could indicate indirect genetic effects of siblings if there is no heterogeneity in other sources of genetic association between singletons and non-singletons. We used UK Biobank data to estimate polygenic score (PGS) associations for height, BMI and educational attainment in self-reported singletons (N = 50,143) and non-singletons (N = 328,549). The educational attainment PGS association estimate was 12% larger (95% C.I. 3%, 21%) in the non-singleton sample than in the singleton sample, but the height and BMI PGS associations were consistent. Birth order data suggested that the difference in educational attainment PGS associations was driven by individuals with older siblings rather than firstborns. The relationship between number of siblings and educational attainment PGS associations was non-linear; PGS associations were 24% smaller in individuals with 6 or more siblings compared to the rest of the sample (95% C.I. 11%, 38%). We estimate that a 1 SD increase in sibling educational attainment PGS corresponds to a 0.025 year increase in the index individual’s years in schooling (95% C.I. 0.013, 0.036). Our results suggest that older siblings may influence the educational attainment of younger siblings, adding to the growing evidence that effects of the environment on phenotypic variation partially reflect social effects of germline genetic variation in relatives.     

Do sibling tits (<Emphasis Type="Italic">Parus major,P. caeruleus</Emphasis>) disperse over similar distances and in similar directions?

We studied dispersal movements by sibling pairs of great tits, Parus major, and blue tits, P. caeruleus, in a patchy environment, in order to test whether siblings are more similar in dispersal than expected by chance. Because of possible common environmental effects due to the heterogeneity and finiteness of the study area, we compared the similarity among siblings with the similarity between each sibling and an unrelated bird that fledged in the same patch and year, as close to the siblings nest as possible. Siblings of both species were not more similar in dispersal distance than they were to control birds. However, great tit siblings dispersed in similar directions compared to control birds, and this result was not affected by the degree of matching between sibling and control birds. As a consequence, siblings ended up breeding at closer distances from one another than control birds. Heritability values calculated from parent-offspring regressions were close to zero, suggesting that there is no additive genetic variance for dispersal distance or dispersal direction. We propose that similarity in dispersal direction originates from association of siblings during dispersal or during activities that influence the choice of direction, such as postfledging family movements. Our results show that non-independence in the choice of dispersal direction by siblings may influence small-scale kin structure in this population with high local recruitment.     

Consequences of mating with siblings and nonsiblings on the reproductive success in a leaf beetle

Choosing a suitable mating partner is crucial for the fitness of an individual, whereby mating with siblings often results in inbreeding depression. We studied consequences of mating with siblings versus nonsiblings in the mustard leaf beetle, Phaedon cochleariae (Coleoptera: Chrysomelidae), on lifetime reproductive traits. Furthermore, we analyzed whether cuticular hydrocarbon (CHC) profiles are family specific and could potentially influence the mating behavior of young adults. We hypothesized a reduced reproductive success of females mated with siblings and a more rapid mating of males with nonsiblings. The hatching rate from eggs of sibling pairs was lower compared to that of nonsibling pairs, pointing to inbreeding depression. Furthermore, the number of eggs laid by females decreased over time in both sibling and nonsibling pairs. Interestingly, the CHC profiles and the body mass differed between families. However, the beetles did not avoid siblings and accepted them as readily as nonsiblings for mating in no‐choice tests. In summary, although it had negative consequences to mate a sibling and although siblings could potentially be recognized by their CHC profiles, the beetles did not show a delayed mating with siblings. Our results indicate that P. cochleariae beetles have not developed a precopulatory mechanism to avoid inbreeding, at least under the test conditions applied here. We predict that instead a polyandrous mating system and/or postcopulatory mechanisms might have evolved in this species by which inbreeding costs can be reduced.     

Within-family variation in obesity

We use data from the Children of the National Longitudinal Survey of Youth 1979 to document the degree to which childhood obesity varies among siblings. We find considerable differences in body weight between siblings with over half of the siblings differing by more than 20 age-specific percentiles in terms of the body mass index. Even among identical twins, there is an average BMI difference of 12 percentiles. This variation is important for the use of econometric approaches that involve sibling comparisons.