Multifactorial inheritance with cultural transmission and assortative mating. II. a general model of combined polygenic and cultural inheritance.

A general linear model of combined polygenic-cultural inheritance is described. The model allows for phenotypic assortative mating, common environment, maternal and paternal effects, and genic-cultural correlation. General formulae for phenotypic correlation between family members in extended pedigrees are given for both primary and secondary assortative mating. A FORTRAN program BETA, available upon request, is used to provide maximum likelihood estimates of the parameters from reported correlations. American data about IQ and Burks' culture index are analyzed. Both cultural and genetic components of phenotypic variance are observed to make significant and substantial contributions to familial resemblance in IQ. The correlation between the environments of DZ twins is found to equal that of singleton sibs, not that of MZ twins. Burks' culture index is found to be an imperfect measure of midparent IQ rather than an index of home environment as previously assumed. Conditions under which the parameters of the model may be uniquely and precisely estimated are discussed. Interpretation of variance components in the presence of assortative mating and genic-cultural covariance is reviewed. A conservative, but robust, approach to the use of environmental indices is described.     

Variation in human mate choice: simultaneously investigating heritability, parental influence, sexual imprinting, and assortative mating

Human mate choice is central to individuals' lives and to the evolution of the species, but the basis of variation in mate choice is not well understood. Here we looked at a large community-based sample of twins and their partners and parents ([Formula: see text] individuals) to test for genetic and family environmental influences on mate choice, while controlling for and not controlling for the effects of assortative mating. Key traits were analyzed, including height, body mass index, age, education, income, personality, social attitudes, and religiosity. This revealed near-zero genetic influences on male and female mate choice over all traits and no significant genetic influences on mate choice for any specific trait. A significant family environmental influence was found for the age and income of females' mate choices, possibly reflecting parental influence over mating decisions. We also tested for evidence of sexual imprinting, where individuals acquire mate-choice criteria during development by using their opposite-sex parent as the template of a desirable mate; there was no such effect for any trait. The main discernible pattern of mate choice was assortative mating; we found that partner similarity was due to initial choice rather than convergence and also at least in part to phenotypic matching.     

Sex differences and non-additivity in the effects of genes on personality.

New large-sample data show that non-additive genetic effects, probably epistatic interactions between loci, and sex-limited gene expression are significant features of the genetic architecture of human personality as measured by questionnaire scales of extraversion and neuroticism. Three large data sets--new data on large samples (n = 20,554) of US twins, their spouses, parents, siblings and children, correlations for Australian twins (n = 7,532), and previously published twin data from Finland (n = 14,288)--are subjected to an integrated analysis to test alternative hypotheses about the genetic causes of family resemblance in personality. When allowance is made for differences in reliability of the scales, the combined data are consistent with the same model for variation. There are significant amounts of genetic non-additivity for both dimensions of personality. The evidence favours additive x additive epistatic interactions rather than dominance. In the case of neuroticism, there is especially strong evidence of sex differences in genetic architecture favouring a greater relative contribution of non-additive genetic effects in males. The data confirm previous claims to find no major contribution of the shared environment of twins and siblings to these dimensions of personality. Correlations between spouses are zero, and the correlations for very large samples of siblings and non-identical twins do not differ significantly.     

Mate selection in man: Evidence,theory, and outcome

Abstract

This review presents a comprehensive survey of the literature on mate selection and non‐random mating in man. The topics discussed include: (1) genetic aspects of non‐random mating for complex traits; (2) evidence on resemblance between spouses on a large variety of traits such as intelligence, personality, physical characteristics, and sociocultural traits; (3) a critical review of sociological and psychological theories offered to account for assortative mating, and (4) implications of assortative mating for marital satisfaction. It is suggested that the factors leading to choice of marriage partners need to be studied from the point of view of multivariate profiles rather than single traits. Such studies will require sophisticated methodologies of research design and data analysis.     

The Genetic Correlation between Height and IQ: Shared Genes or Assortative Mating?

Traits that are attractive to the opposite sex are often positively correlated when scaled such that scores increase with attractiveness, and this correlation typically has a genetic component. Such traits can be genetically correlated due to genes that affect both traits (“pleiotropy”) and/or because assortative mating causes statistical correlations to develop between selected alleles across the traits (“gametic phase disequilibrium”). In this study, we modeled the covariation between monozygotic and dizygotic twins, their siblings, and their parents (total N = 7,905) to elucidate the nature of the correlation between two potentially sexually selected traits in humans: height and IQ. Unlike previous designs used to investigate the nature of the height–IQ correlation, the present design accounts for the effects of assortative mating and provides much less biased estimates of additive genetic, non-additive genetic, and shared environmental influences. Both traits were highly heritable, although there was greater evidence for non-additive genetic effects in males. After accounting for assortative mating, the correlation between height and IQ was found to be almost entirely genetic in nature. Model fits indicate that both pleiotropy and assortative mating contribute significantly and about equally to this genetic correlation.     

Assortive mating for personaltiy traits, educational level, religious affiliation, height, weight, adn body mass index in parents of Korean twin sample.

The degree of assortative mating for psychological and physical traits in Asian societies in relatively unknown. The present study examined assortative mating for educational level, personality traits, religious affiliation, height, weight, and body mass index in a korean sample. Age-adjusted spouse correlations were high for educational level (r = .63) and religious affiliation (r = .67), modest for most personality traits (rs = -.01 to .26), and trivial for height (r = .04), weight (r = .05)m and body mass index (r = .11). These results were remarkably similar to those found from the western samples. Implications of the present findings in behavior genetic studies and human mating patterns were briefly discussed.     

Behavioral,social, and physiological determinants of lipoprotein concentrations

Abstract

The Lipoprotein Family Study of Japanese‐Americans in Hawaii is described. Measured effects of the behavioral, social, and physiological milieu (apart from age and sex) account for little of the variance. Subsequent papers will consider the genetic and cultural causes of family resemblance in this material.     

Direct and indirect assortative mating: a multivariate approach to plant flowering schedules

This paper develops methods to partition the phenotypic correlation between mates for a focal trait--the standard measure for assortative mating--into a direct component and additional indirect components. Indirect assortative mating occurs when a nonassorting trait is correlated within individuals to a directly assorting trait. Direct and indirect assortative mating is assessed for flowering phenology in Brassica rapa. The flowering time of pollen recipients (mothers) was strongly correlated (rho=0.67) to that of potential pollen donors (fathers). Similarly, recipients and donors were correlated for duration of their flowering periods (rho=0.32) and stem diameters (rho=0.52). A partitioning of between-mate correlations revealed direct assortative mating for flowering time and period duration. However, assortment for stem diameter is explained solely through its correlation to flowering time. Examination of standard quantitative genetic theory shows that indirect assortative mating inflates genetic variance in a focal trait and the genetic covariance between focal and phenotypically correlated traits.     

Multifactorial inheritance with cultural transmission and assortative mating. III. Family structure and the analysis of separation experiments.

Demographic data about family composition or structure in the United States is reviewed. About 25% of white children and a majority of black children are reared in either broken or extended families, and this must be taken into consideration for valid studies of cultural inheritance. Atypical family structures are described including those in which parents include: biological parents, stepparents, grandparents, uncles, aunts, sibs, foster parents, and their spouses. General formulae for a wide variety of kinship correlations are derived using path analysis. The multifactorial model presented allows for cultural inheritance, polygenic inheritance, correlated sibling environments, and phenotypic assortative mating (as previously described for intact families) plus extensions necessary for the analysis of separation experiments. These extensions allow for variable family structure and differences in parental influence due to separation, age or stage of development of the child, birth order, or type of relationship. Family structure is observed to have a marked effect on familial resemblance. Computer simulation studies demonstrate marked heterogeneity among phenotypic correlations for kinships of the same degree of genetic relationship arising in different family structures. Analyses of multiple types of sibs and other relatives in variable family structures offer great promise for the study of cultural inheritance.     

Assortative mating in unwed birth parents,adoptive, and nonadoptive parents

Abstract

Correlation between parents for a given characteristic substantively affects estimates of both genetic and environmental parameters. Spouse similarity for biological, adoptive, and nonadoptive parents was examined in a study using a full‐adoption design. With regard to isophormic comparisons, moderate assortment was found for age, educational attainment, performance on tests of verbal ability, family background, and habits such as alcohol and current smoking behavior. The effects of cross‐assortative mating on population covariance and cross‐correlation between relatives are discussed. Although of considerable theoretical interest, little cross‐trait assortative mating for personality and cognition was found in the present study, suggesting that its effects will not be large, at least for these variables. Because “assortative mating” may differ from “assortative marriage,” comparisons among estimates of homogamy for birth, adoptive, and nonadoptive parents were made. Results indicated differential assortment among the three types of parents for some of the variables examined.     

Social runaway: Fisherian elaboration (or reduction) of socially selected traits via indirect genetic effects

Our understanding of the evolutionary stability of socially selected traits is dominated by sexual selection models originating with R. A. Fisher, in which genetic covariance arising through assortative mating can trigger exponential, runaway trait evolution. To examine whether nonreproductive, socially selected traits experience similar dynamics—social runaway—when assortative mating does not automatically generate a covariance, we modeled the evolution of socially selected badge and donation phenotypes incorporating indirect genetic effects (IGEs) arising from the social environment. We establish a social runaway criterion based on the interaction coefficient, ψ , which describes social effects on badge and donation traits. Our models make several predictions. (1) IGEs can drive the original evolution of altruistic interactions that depend on receiver badges. (2) Donation traits are more likely to be susceptible to IGEs than badge traits. (3) Runaway dynamics in nonsexual, social contexts can occur in the absence of a genetic covariance. (4) Traits elaborated by social runaway are more likely to involve reciprocal, but nonsymmetrical, social plasticity. Models incorporating plasticity to the social environment via IGEs illustrate conditions favoring social runaway, describe a mechanism underlying the origins of costly traits, such as altruism, and support a fundamental role for phenotypic plasticity in rapid social evolution.     

Effect of genetic heterogeneity and assortative mating on linkage analysis: a simulation study.

Linkage studies of complex genetic traits raise questions about the effects of genetic heterogeneity and assortative mating on linkage analysis. To further understand these problems, I have simulated and analyzed family data for a complex genetic disease in which disease phenotype is determined by two unlinked disease loci. Two models were studied, a two-locus threshold model and a two-locus heterogeneity model. Information was generated for a marker locus linked to one of the disease-defining loci. Random-mating and assortative-mating samples were generated. Linkage analysis was then carried out by use of standard methods, under the assumptions of a single-locus disease trait and a random-mating population. Results were compared with those from analysis of a single-locus homogeneous trait in samples with the same levels of assortative mating as those considered for the two-locus traits. The results show that (1) introduction of assortative mating does not, in itself, markedly affect the estimate of the recombination fraction; (2) the power of the analysis, reflected in the LOD scores, is somewhat lower with assortative rather than random mating. Loss of power is greater with increasing levels of assortative mating; and (3) for a heterogeneous genetic disease, regardless of mating type, heterogeneity analysis permits more accurate estimate of the recombination fraction but may be of limited use in distinguishing which families belong to each homogeneous subset. These simulations also confirmed earlier observations that linkage to a disease "locus" can be detected even if the disease is incorrectly defined as a single-locus (homogeneous) trait, although the estimated recombination fraction will be significantly greater than the true recombination fraction between the linked disease-defining locus and the marker locus.     

A family study of anthropometric traits in a Punjabi community: II. An investigation of familial transmission

Path analysis is used to characterize family resemblance for anthropometrics in twins and nuclear families from the Punjabi population of India. Significant positive assortative mating exists with respect to many body measurements, but not for cranial or facial variables. Evidence of a maternal effect for five measurements of bone diameter is reported. Twin resemblance is increased by a component not found in other pairs of relatives for all variables except nasal height, facial length, ear length, and head breadth. Although all variables have significant transmissible components, many have parameter estimates which are not consistent with strictly polygenic inheritance. Some form of cultural transmission is implicated for such variables, especially for those related to fatness levels.     

Consanguineous marriages in Italy: A Note

Abstract

A general multifactorial model is given for the inheritance of traits that exhibit a sexual dimorphism. The model allows for polygenic inheritance, cultural transmission, phenotypic assortative mating, and a common environment of rearing. Several cultural mechanisms are described for which transmission from parent to offspring is sex‐dependent and for which many different patterns of sex‐specific correlations can result. A special case of the general model is described in which phenotypic differences between males and females are due only to differences in nontransmissible environmental factors and/or genetic factors that do not contribute to variability within a sex. Application of these models to human spatial visualizing ability, using data reported by others, gives an estimate of 45 per cent for the proportion of the variance that is accounted for by transmissible factors. Neither an X‐linked hypothesis nor a sex‐specific cultural mechanism is required to explain the transmission of spatial ability.     

Matrilineal genetic structure within and among populations of the cooperatively breeding common marmoset,Callithrix jacchus

Common marmosets are members of the family Callitrichidae, South American primates characterized by highly social group living and cooperative breeding. In this study we analysed 1112 base pairs (bp) of the mitochondrial control region in 59 Callithrix jacchus individuals, sampled mainly from two geographically distinct field sites in N.E. Brazil. Analysis of molecular variation revealed a highly significant genetic structuring of haplotypes between social groups and between populations. Examination of matrilineal genetic structure within social groups revealed that seven of nine recorded breeding pairs were from different maternal lineages, indicating assortative mating and outbreeding. In addition to the breeders, at least six of 10 groups contained adult individuals from different matrilines, with five haplotypes present in one social group of nine animals. Groups of mixed lineages raise questions about potential reproductive conflicts of interest, and the extent of kin-selected altruism in the evolution and maintenance of cooperative breeding in this species.     

Sex ratio dynamics and fluctuating selection on personality

Fluctuating selection has often been proposed as an explanation for the maintenance of genetic variation in personality. Here I argue that the temporal dynamics of the sex ratio can be a powerful source of fluctuating selection on personality traits, and develop this hypothesis with respect to humans. First, I review evidence that sex ratios modulate a wide range of social processes related to mating and parenting. Since most personality traits affect mating and parenting behavior, changes in the sex ratio can be expected to result in variable selection on personality. I then show that the temporal dynamics of the sex ratio are intrinsically characterized by fluctuations at various timescales. Finally, I address a number of evolutionary genetic challenges to the hypothesis. I conclude that the sex ratio hypothesis is a plausible explanation of genetic variation in human personality, and may be fruitfully applied to other species as well.     

Genetic effects on mating success and partner choice in a social mammal

Mating behavior has profound consequences for two phenomena--individual reproductive success and the maintenance of species boundaries--that contribute to evolutionary processes. Studies of mating behavior in relation to individual reproductive success are common in many species, but studies of mating behavior in relation to genetic variation and species boundaries are less commonly conducted in socially complex species. Here we leveraged extensive observations of a wild yellow baboon (Papio cynocephalus) population that has experienced recent gene flow from a close sister taxon, the anubis baboon (Papio anubis), to examine how admixture-related genetic background affects mating behavior. We identified novel effects of genetic background on mating patterns, including an advantage accruing to anubis-like males and assortative mating among both yellow-like and anubis-like pairs. These genetic effects acted alongside social dominance rank, inbreeding avoidance, and age to produce highly nonrandom mating patterns. Our results suggest that this population may be undergoing admixture-related evolutionary change, driven in part by nonrandom mating. However, the strength of the genetic effect is mediated by behavioral plasticity and social interactions, emphasizing the strong influence of social context on mating behavior in socially complex species.     

Effect of assortative mating on genetic change due to selection

Summary Two mathematical models (A and B) were used to study joint effects of selection and assortative mating on genetic change. Computer simulation was used to verify and extend the results. In each model, the genotype was additive with equal effects at each of N loci and the environmental distribution was N(0, ²). In Model A, each locus had two alleles; in Model B, allelic effects at each locus followed a normal distribution. Using Model A, genetic change with assortative or random mating of selected parents was evaluated for combinations of number of loci (N = 1, 2, 3), heritability in base population (H^[0] = 0.2, 0.5, 0.8), allelic frequency in base population (p = 0.1, 0.5), and proportion selected ( = 0.20, 0.85). Using Model B, genetic change with or without assortative mating was calculated for combinations of N (1, 2, 3, 5, 10, 100, H^[0] (0.2, 0.5, 0.8) and (0.20, 0.85). Response to selection under both mating systems in a finite population was estimated using Model A from 200 replications of a computer simulation; this was done for all combinations of N (1,2, 3, 5, 10) and (0.20, 0.85), with H^[0] = 0.5 and p = 0.1. Results obtained with both models indicate that the effect of assortative mating on genetic change increases with H^[0] and , and decreases with p. With Model A, the relationship between N and the effect of assortative mating on genetic change was not clear; with Model B, however, the advantage of assortative over random mating increased with N, as expected. Simulation results were in agreement with theory of Model A. This study indicates that selection with assortative mating can have a sizable (10 to 20%) long-term advantage over selection with random mating of parents when H^[0] is high, p is low and is large.