1 Evolutionary Genetics of Personality Revisited 1 The Evolutionary Genetics of Personality Revisited Lars Penke Georg Elias Müller Institute of Psychology, Georg August University Göttingen, Gosslerstr. 14, Göttingen, Germany Markus Jokela Institute of Behavioural Sciences, University of Helsinki, Siltavuorenpenger 1A, University of Helsinki, Finland Correponding author: Lars Penke Running Head: Evolutionary Genetics of Personality Revisited In press, Current Opinion in Psychology
2 Evolutionary Genetics of Personality Revisited 2 Abstract Like all human individual differences, personality traits and intelligence are substantially heritable. From an evolutionary perspective, this poses the question what evolutionary forces maintain their genetic variation. Information about the genetic architecture and associations with evolutionary fitness permit inferences about these evolutionary forces. As our understanding of the genomics of personality and its associations with reproductive success have grown considerably in recent years, it is time to revisit this question. While mutations clearly affect the very low end of the intelligence continuum, individual differences in the normal intelligence range seem to be surprisingly robust against mutations, suggesting that they might have been canalized to withstand such perturbations. Most personality traits, by contrast, seem to be neither neutral to selection nor under consistent directional or stabilizing selection. Instead evidence is in line with balancing selection acting on personality traits, likely supported by human tendencies to seek out, construct and adapt to fitting environments. Key words: Personality traits; intelligence; evolutionary genetics; molecular genetics; reproductive success Highlights: - Evolutionary forces that maintain genetic variance in traits can be inferred from their genetic architecture and fitness correlates - A substantial amount of new data on the genomics and reproductive success associated with personality traits and intelligence has recently become available - Intelligence differences seem to have been selected for robustness against mutations - Human tendencies to select, create and adapt to environments might support the maintenance of personality traits through balancing selection
3 Evolutionary Genetics of Personality Revisited 3 Personality traits are relatively stable dimensions of individual differences in cognition, affect and behavior. Human personality traits can be organized around five independent dimensions: Extraversion, Agreeableness, Neuroticism, Conscientiousness and Openness to Experience. These dimensions are of interest, as they are substantial predictors of important life outcomes, from educational and occupational attainment to lifetime reproductive success and longevity [1-3]. The strongest single predictor of any personality trait is the standing of one s biological parents on the same trait: About 50% of the variation in broad personality traits is genetically heritable . The finding is hardly surprising, as virtually all human traits ever studied with quantitative behavior genetic designs (e.g., twin and adoption studies) show substantial genetic components  a finding so robust that it has been enshrined as the first law of behavior genetics . This raises the question of how genetic variation in personality traits has been maintained in populations over evolutionary time. In 2007, Penke, Denissen and Miller [7,8] proposed that the maintaining evolutionary forces can be inferred from the genetic architecture of traits as well as their associations with reproductive fitness. After summarizing the evidence available at that time, they concluded that personality traits are unlikely to be neutral to selection. For general intelligence, a balance between steadily occurring deleterious mutations and directional selection towards higher intelligence appeared to be the best explanation. Personality traits, by contrast, seemed to be under balancing selection, where selection pressures in different directions affected the same traits at different times or in different places, in a way that no genetic variant underlying personality traits is consistently favored over others. Recent evidence on the genetic architecture of intelligence A lot of progress has been made since the publication of Penke and colleagues article [7,8], particularly in the field of genomics. We now have an ever more elaborate toolkit to infer selective regimes from genetically informative data, and the necessary data is increasingly becoming available . For general intelligence, we know that several hundreds of rare mutations with large effect
4 Evolutionary Genetics of Personality Revisited 4 explain a substantial amount of the variation at the very low end of the distribution, that is, in cases of intellectual disability. However, other genetic variants in the same genetic regions that cause low intelligence when mutated do not contribute to individual differences in the normal range . Neither do overall burdens of rare copy number variants (variation in how often a genetic region is repeated in the DNA), exomic mutations (those that can alter the amino acids produced by genes) or de novo mutations (that occurred from one generation to the next) explain any substantial amount of variance of intelligence in the general population, though one recent study suggested lower burden of exomic mutations in extremely intelligent individuals (IQ > 170) compared to those with average intelligence . Thus, the genomic mutations studied so far have not been associated with intelligence in the normal range. Instead, genome-wide complex trait analyses (GCTA), which estimate the overall contribution of genetic variants with common frequencies in the population to traits based on around a million genetic markers across the genome, suggest that 28-51% of the variance in intelligence is due to the cumulative effect of thousands of common genetic variants, each with miniscule effect size. For a brief review, see . Common genetic variants are unlikely to be deleterious mutations . The GCTA estimates still leave a substantial proportion of heritable variance in intelligence unaccounted for, and it still remains possible that a burden of very specific rare mutations that are older and intronic (non-coding for amino acids) contribute to the unexplained fraction of the heritability . Alternative explanations for the unaccounted heritability in GCTA include interactions between variants at different genetic loci (epistasis), or between genes and environments. The overall picture suggests that only the low and perhaps the very high end of the intelligence spectrum can be directly explained by mutation-selection balance. The rest of the spectrum might be effectively more neutral to selection than generally appreciated, perhaps due to trade-offs between benefits of higher intelligence and energetic costs of developing and maintaining a highly intelligent brain. Alternatively, normal-range intelligence could have been actively selected for robustness against mutational insults, which can result in both highly redundant (and thus compensatory) common genetic variation and widespread epistasis [11,12,14].
5 Evolutionary Genetics of Personality Revisited 5 Recent evidence on the genetic architecture of personality Compared to intelligence, much less genomic data is available on personality traits. What can be said with high certainty is that none of the candidate genes for personality (including DRD4, 5HTT- LPR and COMT) have held up in meta-analyses. If these genes are associated with personality at all, their individual effects are tiny [15,16]. The lack of individual genetic variants with large effects is in line with genome-wide association studies (GWAS), which scan the genome for individual genetic variants linked to traits. So far GWAS of personality have not found a single replicable hit [17-19]. These results suggest that a large number of genetic variants with individually tiny effects explain a substantial part of the heritability of personality, which is similar to what has been found for intelligence and indeed any human behavioral, clinical, and physical traits. As this seems to be a general pattern, it has recently been proposed as the fourth law of behavioral genetics (; the first was discussed above, the second and third being that environmental influences do not contribute much to the similarity of family members, but substantially to their dissimilarity ). A puzzling finding that diverges from the patterns generally found for other human traits is that GCTA estimates of the overall contribution of common genetic variants to personality traits are low: Zero to 21% variance explained (highest for Openness to Experience and Neuroticism, zero for Agreeableness and Conscientiousness), with confidence intervals often touching or including zero [17-19,21]. These estimates are markedly lower than the heritabilities for personality traits found in quantitative genetic studies . Explanations for the surprisingly high proportion of missing heritability in personality traits remain unclear. Some of it might be explained by rare mutations, which are not captured by GCTA estimates and have hardly been studied directly for personality, or by widespread epistasis and gene-environment interactions. Notably, while most quantitative genetic designs are not well suited to isolate epistasis, those few studies that have used appropriate designs have consistently identified substantial non-additive genetic components (including epistasis) for most, if not all, broad personality traits .
6 Evolutionary Genetics of Personality Revisited 6 Personality and reproductive success Evidence from a dozen studies suggests that personality is related to the most direct indicator of reproductive success: how many children people have. Many associations, however, have not been consistent across studies (Table 1), and the effect sizes tend to be small. In the largest study to date from the contemporary United States , higher offspring number was associated with higher extraversion (+0.12 more offspring per one standard deviation change in the personality trait), lower neuroticism ( 0.05), higher agreeableness (+0.07), lower conscientiousness ( 0.06), and lower openness to experience ( 0.19). Some of the associations between personality and offspring number may have emerged only recently, and thereby represent evolutionarily novel selection pressures. For example, in the United States higher openness to experience and women s higher conscientiousness were not associated with fertility among individuals born in the 1920s, but the associations strengthened in more recent birth cohorts . These time-varying associations probably reflect societal changes, such as women s broader participation in the workforce and the adoption of less traditional lifestyles where people prefer to have smaller families. Other associations may represent more universal effects of personality on mating success and family formation. For example, extraversion and neuroticism have been associated with offspring number in various samples, extraversion particularly in men and most consistently in traditional small-scale societies (Table 1). Except for one early study , no studies have found evidence for curvilinear associations where the highest number of children would be observed for intermediate levels of parent s personality trait. The lack of curvilinear associations speaks against stabilizing selection. Furthermore, there appears to be no intergenerational trade-offs between the numbers of offspring and grandoffspring that would negate the associations across more than one generation. A study in the United States demonstrated that higher extraversion, higher agreeableness, lower conscientiousness, and lower openness to experience were associated not only with the number of children but also with the number of grandchildren . These findings line up well with a recent
7 Evolutionary Genetics of Personality Revisited 7 estimate of the genetic correlation between the number of offspring and number of grandoffspring in modern humans as perfect , which implies that the number of offspring is a good indicator for long-term reproductive success. The situation is different for many other animals, in which an increasing number of offspring often leads to poorer average survival and mating success of these offspring because of limited parental resources, and it might have been different in ancestral humans as well, even though we have little knowledge if this was the case and when it might have changed. Despite the lack of evidence for trade-offs between the numbers of offspring and grandoffspring, there may still be important personality-dependent trade-offs between the number of offspring and parental investments in the offspring. For example, higher neuroticism is associated with lower offspring number but this association may reflect an adaptive pattern where smaller families are optimal for parents with high neuroticism, who might compensate with higher parental investment. Preliminary evidence for such trade-offs in modern humans was provided by a study showing that high parental neuroticism was associated with higher offspring education (a proxy measure for parental investment) in small families but lower offspring education in large families . What can we infer about evolutionary forces affecting personality traits? All the new data on the genomic and reproductive fitness correlates of personality traits should have helped to make better inferences about the evolutionary forces that affect personality traits. But in fact they have made the picture more complex. It is now clear that the genetic architecture of personality traits is not based on a limited number of genetic variants with substantial effect sizes and common frequency in the population, as assumed by the traditional candidate gene literature and expected by Penke and colleagues  based on simple models of balancing selection. At the same time, personality traits are unlikely to be neutral to selection, as that would predict underlying genetic variants with mostly common frequencies in the population and little epistasis. GCTA results and quantitative genetic analyses with appropriate designs suggest the exact opposite, and links with
8 Evolutionary Genetics of Personality Revisited 8 reproductive fitness, even if somewhat inconsistent in direction across studies, contradict selective neutrality as well. The lack of evidence for clear linear or curvilinear relationships with reproductive success also provides little support for directional or stabilizing mutation-selection balance, respectively, as the main evolutionary forces maintaining genetic variance in personality traits. An exception might be male extraversion, where a consistent positive association with reproductive success particularly in traditional societies is in line with directional mutation-selection balance. While classical models of balancing selection based on a limited number of maintained genetic variants can be rejected for personality traits, balancing selection can maintain highly polygenetic variation as well. The problem is that evidence for highly polygenic balancing selection is difficult to establish based on population genetic models  or genetic signatures in currently available genomic data, especially if the balanced selective forces are not long-standing but transient over evolutionary time [28,29]. The difficulties of testing balancing selection have sometimes led to a dismissal of this evolutionary force. We would argue that this conclusion is particularly premature in the case of humans, as individuals of our species are especially good at seeking out and creating environments that suite their individual preferences, needs and capabilities. This tendency, variably called active gene-environment correlation, experience-producing drives, or niche picking and construction, can systematically lead individuals to place themselves into environments where their specific personality profiles yield the best-possible fitness pay-off. As long as environments are diverse enough for everybody to find a suitable niche (which might be much more the case in modern complex than in traditional small-scale societies), this process can stabilize balancing selection [9,30]. Evidence that individuals choose their environments to correspond with their personality traits comes from studies of migration. For example, extraverted individuals tend to migrate away from secluded locations, such as islands and rural areas, to more populated and urban areas [31,32], and similar migration patterns have been observed for intelligence . Moreover, individuals with high extraversion and high openness to experience appear to enjoy higher life satisfaction when living in
9 Evolutionary Genetics of Personality Revisited 9 more populated locations compared to remote rural areas . Similarly, a study in the London metropolitan area showed that individuals with high openness to experience had the highest life satisfaction in areas where the average openness level of the residents was also high . With the important caveat that life satisfaction is a very indirect indicator of evolutionarily meaningful personality environment fit, these findings provide support for the idea that people seek environments that match their personality traits, potentially sustaining balancing selection on personality that way. Consistent with the niche picking perspective, a recent review on the developmental quantitative genetics of personality  found that the rank-order stability of personality traits increases markedly up to the thirties, mediated by environmental effects acting on several distinct sources of genetic variance per personality trait. This pattern differs markedly from the development of intelligence, where rank-order stability is already reached by late childhood and mediated by genetic factors acting on mostly a single general pool of genetic variance. These authors concur with our conclusion that intelligence is likely selected to be strongly canalized, whereas the pattern found for personality traits may have been selected to support extended adaptive behavioral plasticity while individuals continue to explore and conform to environmental niches with unclear selective pressures up into adulthood. This is in line with the conceptualization of personality traits as behavioral reaction norms [7,37], as well as recent functionalist models of personality development that see personality traits as emergent from self-regulatory dynamics in response to environmental affordances [38-41]. These perspectives do not view personality traits as unitary causal generators of behavior, but as complexes of behavioral tendencies that result from individuals seeking out and adapting to their environment. While explicit modelling is still pending, these perspectives seem in line with widespread gene-environment interactions as an explanation for low GCTA heritabilities of personality traits. To conclude, while normal-range intelligence differences seem to be selected for robustness against mutational disturbances, most personality differences seem to be selected for plasticity that
10 Evolutionary Genetics of Personality Revisited 10 allows individuals to seek out and adapt to their environmental niches, and balancing selection to maintain its genetic variance.
11 Evolutionary Genetics of Personality Revisited 11 References ** Berg V, Lummaa V, Lahdenperä M, Rotkirch A, Jokela M: Personality and long-term reproductive success measured by the number of grandchildren. Evol Hum Behav 2014, 35: This study is the first to show associations between personality traits and reproductive success across two generations. It finds that personality affects number of children and grandchildren in very similar ways, contradicting reproductive quantity-quality trade-offs in humans.  Ozer DJ, Benet-Martinez V: Personality and the prediction of consequential outcomes. Annu Rev Psychol 2006, 57:  Roberts BW, Kuncel NR, Shiner R, Caspi A, Goldberg LR: The power of personality: The comparative validity of personality traits, socio-economic status, and cognitive ability for predicting important life outcomes. Perspect Psychol Sci 2007, 2:  Johnson AM, Vernon PA, Feiler AR: Behavioral genetic studies of personality: An introduction and review of the results of 50+ years of research. In The Sage Handbook of Personality Theory and Assessment. Edited by Boyle GJ, Matthews G, Saklofske DH. Sage; 2008: * Polderman TJC, Benyamin B, de Leeuw CA, Sullivan PF, van Bochoven A, Visscher PM, Posthuma D: Meta-analysis of the heritability of human traits based on fifty years of twin studies. Nat Gen 2015, 47: This enormous meta-analysis of all classical twin studies on human traits consistently shows that all traits studied so far are substantially heritable, providing strong empirical support for Turkheimer s first law of behavior genetics.  Turkheimer E: Three laws of behavior genetics and what they mean. Cur Dir Psychol Sci 2000, 9:  Penke L, Denissen JJA, Miller GF: The evolutionary genetics of personality. Eur J Personal 2007, 21:  Penke L, Denissen JJA, Miller GF: Evolution, genes, and inter-disciplinary personality research. Eur J Personal 2007, 21:
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13 Evolutionary Genetics of Personality Revisited 13  Verweij KJH, Yang J, Lahti J, Veijola J, Hintsanen M, Pulkki-Råback L, Heinonen K, Pouta A, Pesonen AK, Widen E et al.: Maintenance of genetic variation in human personality: Testing evolutionary models by estimating heritability due to common causal variants and investigating the effect of distant inbreeding. Evolution 2012, 66:  Vinkhuyzen AA, Pedersen NL, Yang J, Lee SH, Magnusson PK, Iacono WG, McGue M, Madden PA, Heath AC, Luciano M et al.: Common SNPs explain some of the variation in the personality dimensions of neuroticism and extraversion. Transl Psychiatry 2012, 2:e102.  Chabris CF, Lee JJ, Cesarini D, Benjamin DJ, Laibson DI: The fourth law of behavior genetics. Cur Dir Psychol Sci 2015, 24: * Power RA, Pluess M: Heritability estimates of the Big Five personality traits based on common genetic variants. Transl Psychiatry 2015, 5:e604. First study estimating the heritability of all dimensions of the Five Factor Model of personality from common genetic variants (GCTA), confirming low estimates for extraversion and neuroticism and adding the surprising result that estimates center around zero for agreeableness and conscientiousness.  Jokela M, Alvergne A, Pollet TV, Lummaa V: Reproductive behavior and personality traits of the Five Factor Model. Eur J Personal 2011, 25:  Jokela M: Birth-cohort effects in the association between personality and fertility. Psychol Sci 2012, 23:  Eaves LJ, Martin NG, Heath AC, Hewitt JK, Neale MC: Personality and reproductive fitness. Behav Gen 1990, 20: * Zietsch BP, Kuja-Halkola R, Walum H, Verweij KJ: Perfect genetic correlation between number of offspring and grandoffspring in an industrialized human population. Proc Natl Acad Sci U S A 2014, 111: A twin study that show that reproductive success across two generations is perfectly correlated in modern humans, indicating that number of children is as good a measure of reproductive fitness as number of grandchildren.
14 Evolutionary Genetics of Personality Revisited 14  Jokela M, Alvergne A, Rotkirch A, Rickard IJ, Lummaa V: Associations between family size and offspring education depend on aspects of parental personality. Personal Individ Dif 2014, 58:  Gillespie JH: Sampling theory for alleles in a random environment. Nature 1977, 266:  Andrés AM, Hubisz MJ, Indap A, Torgerson DG, Degenhardt JD, Boyko AR, Gutenkunst RN, White TJ, Green ED, Bustamante CD et al.: Targets of balancing selection in the human genome. Mol Biol Evol 2009, 26: * Fijarczyk A, Babik W: Detecting balancing selection in genomes: Limits and prospects. Mol Ecol 2015, 14: Review of current methods to detect balancing selection from genomic data and their limitations. This is contrasted with theoretical work predicting that balancing selection is actually ubiquitous in nature.  Penke L: Bridging the gap between modern evolutionary psychology and the study of individual differences. In The Evolution of Personality and Individual Differences. Edited by Buss DM, Hawley P. Oxford University Press; 2010:  Camperio Ciani AC, Capiluppi C: Gene flow by selective emigration as a possible cause for personality differences between small islands and mainland populations. Eur J Personal 2011, 25:  Jokela M, Elovainio M, Kivimäki M, Keltikangas-Järvinen L. Temperament and migration patterns in Finland. Psychol Sci 2008, 19:  Jokela M: Flow of cognitive capital across urban and rural United States. Intelligence 2014, 46:  Murray G, Judd F, Jackson H, Fraser C, Komiti A, Hodgins G, Pattison P, Humphreys J, Robins G: The five factor model and accessibility/remoteness: Novel evidence for person environment interaction. Personal Individ Dif 2005, 39:  Jokela M, Bleidorn W, Lamb ME, Gosling SD, Rentfrow PJ: Geographically varying associations between personality and life satisfaction in the London metropolitan area. Proc Natl Acad Sci U S A 2015, 112: ** Briley DA, Tucker Drob EM: Comparing the developmental genetics of cognition and personality over the lifespan. J Personal, 2015.
15 Evolutionary Genetics of Personality Revisited 15 Review of developmental behavior genetic studies on personality traits and intelligence, drawing on recent meta-analyses by the same authors. This article nicely illustrates the markedly different developmental patterns of personality traits and intelligence and puts them in an evolutionary perspective.  Dingemanse NJ, Kazem AJ, Réale D, Wright J: Behavioural reaction norms: animal personality meets individual plasticity. Trends Ecol Evol 2010, 25:  Denissen JJA, Wood D, Penke L: Passing to the functionalists instead of passing them by. Eur J Personal 2012, 26:  Denissen JJA, Wood D, Penke L, van Aken MAG: Self-regulation underlies temperament and personality: An integrative developmental framework. Child Dev Persp 2013, 7:  von Rueden CR, Lukaszewski AW, Gurven M: Adaptive personality calibration in a human society: effects of embodied capital on prosocial traits. Behav Ecol 2015, 26: * Wood D, Gardner MH, Harms PD: How functionalist and process approaches to behavior can explain trait covariation. Psychol Rev 2015, 122: An elaborate model how broad personality dimensions can emerge from similar environmental affordances acting on narrow personality traits, causing their covariation. This functionalist, process-oriented perspective contrasts with older perspectives that broad personality traits are coherent latent causal entities.  Nettle D: An evolutionary approach to the extraversion continuum. Evol Hum Behav 2005, 26:  Jokela M, Kivimäki M, Elovainio M, Keltikangas-Järvinen, L: Personality and having children: A twoway relationship. J Personal Soc Psychol 2009, 96:  Alvergne A, Jokela M, Lummaa V: Personality and reproductive success in a high-fertility human population. Proc Natl Acad Sci U S A 2010, 107:  Jokela M, Hintsa T, Hintsanen M, Keltikangas-Järvinen L: Adult temperament and childbearing over the life course. Eur J Personal 2010, 24:  Reis O, Dörnte M, von der Lippe H: Neuroticism, social support, and the timing of first parenthood: A prospective study. Personal Individ Dif 2011, 50:
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17 Evolutionary Genetics of Personality Revisited 17 Tables Table 1. Selected associations between personality traits and number of children (or probability of having children) Study Personality trait Country E N A C O Sample size  no no... Australia 1101  no.... UK 545 b +... Finland 1839  +m w... Senegal (rural) 136 a +. Finland 1535  + + USA .... Germany 244  +m no no no no Paraguay (Ache) 110  +m m +w w w UK 8336  +m m no w m Norway 7017  + no no USA  +m m no +m +m Bolivia (Tsimane) = positive association, = negative association, no = no association,. = personality trait not assessed in the study, m = association observed in men, w = association observed in women Unless otherwise noted, E = Extraversion, N = Neuroticism, A = Agreeableness, C = Conscientiousness, O = Openness to experience a E = Novelty seeking, N = Harm avoidance, A = Reward dependence, C = Persistence measured by the Temperament and Character Inventory (TCI) b E = Sociability, N = Emotionality assessed by the Emotionality, Activity, and Sociability (EAS) personality model
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