Height is a classic anthropometric quantitative trait in humans due to its ease of measurement, approximately normal distribution and relative stability in adulthood. Since the studies of height in the late 19^th and early 20^th centuries (Galton, 1886; Pearson and Lee, 1903; Fisher, 1919), twin, adoption and family studies have shown that height is one of the most heritable human quantitative phenotypes (Silventoinen, 2003). More recently, genetic linkage studies have helped to elucidate the location of genetic effects in the genome (Perola et al., 2007) and genome-wide association (GWA) studies allowed identification of loci consistently associated with height in populations of different ancestry (Cho et al., 2009; Hao et al., 2013; Lango Allen et al., 2010; N'Diaye et al., 2011; Wood et al., 2014). Besides the genetic factors, a multitude of environmental factors, such as nutrition and childhood diseases, operate during the growth period and can affect the final attained height. These and other proximate biological determinants of height are further associated with social and economic conditions, which in turn are associated with living standards (Bozzoli et al., 2009; Bogin, 2001; Eveleth and Tanner, 1990; Steckel, 2009). The secular trend of increasing height over the 20^th century observed in many parts of the world, which has slowed or stopped in most northern European countries, probably reflects the continuous improvement in the standard of living (Eveleth and Tanner, 1990; Cole, 2003; Stulp and Barrett, 2016). A recent study showed that the height difference between the tallest and shortest populations a century ago (19–20 cm) has remained the same for women and increased for men (NCD Risk Factor Collaboration (NCD-RisC), 2016) .

Twin and family studies have consistently estimated that the proportion of variation in adult height explained by genetic differences between individuals, or heritability, in general populations is approximately 0.80 (Fisher, 1919; Silventoinen et al., 2003; Stunkard et al., 1986). There is a hypothesis that heritability is not constant and can differ in environments having different amount of environmental variation. Accordingly, it has been suggested that heritability of height is lower in populations with low living standards compared with affluent populations since poverty can lead to a lack of basic necessities important for human growth in part of the population (Steckel, 2009). However, there is little direct evidence on this issue. A study in Finnish twins born between 1900 and 1957 showed that the heritability of height increased across birth cohorts born in the first half of the century when the standard of living increased and leveled off after World War II thus supporting this hypothesis (Silventoinen et al., 2000). Because this result needs to be replicated, we conducted an individual-based analysis of 40 twin cohorts from 20 countries. We aimed to analyze (i) the genetic and environmental contribution to the variation of adult height across nine birth-year cohorts covering more than 100 years and (ii) to assess whether the pattern varies by geographic-cultural region (Europe, North America and Australia, and East Asia).

In the pooled data (all twin cohorts together), mean height was greater in men than in women and increased over the birth-year cohorts in both sexes; the decrease ( > 1 cm) observed in the latest birth cohort mainly reflects differences in the distribution of different twin cohorts within each group (Table 1). Both means and variances were significantly different between twin cohorts in all birth-year and sex groups. Mean height was shorter in East Asia than in Europe and North America and Australia in all birth-year and sex groups. The increase in mean height over the birth cohorts (from 1940–1949 to 1980–1994) was substantially greater in East Asia than in the other two geographic-cultural regions. The variance of height was generally greater in men than in women, lowest in East Asia and greatest in North America and Australia, and showed a general trend to increase over the birth cohorts.

The variance of adult height explained by additive genetic, shared environmental and unique environmental factors by birth-year cohorts is presented in Figure 1 (estimates with 95% confidence intervals (CIs) are available in Supplementary file 1A). In men, there was a trend for an increasing total variance from birth cohort 1940–1949 onwards; genetic variance also increased during this period but especially in the two latest birth-year cohorts (1970–1979 and 1980–1994). Height variance due to the environment shared by co-twins was significant from birth cohorts 1920–1929 to 1970–1979, being greatest from 1950 to 1969. The effect of environmental factors unique to each twin individual including measurement error was more similar across birth-year cohorts. Heritability estimates ranged from 0.69 to 0.84 and were greatest in the two earliest and the two latest birth-year cohorts (Table 2). In women, although the total variance also started to increase from birth cohort 1940–1949, genetic variance showed an increasing trend from the earliest birth-year cohort. Both shared and unique environmental factors explained variation in height in all analyzed birth-year cohorts; whereas the shared environmental variance was somewhat greater in the latest cohorts (1970–1979 and 1980–1994) unique environmental variance was greatest in the earliest one. Although the variance components differed between sexes in all birth-year cohorts, the relative contribution of the genetic and environmental variance components did not differ by sex from 1930–1939 to 1960–1969 (Supplementary file1B). In contrast to the results in men, heritability estimates in women (0.53 to 0.78) were lowest in the earliest and latest cohorts, particularly in 1886–1909. When we studied the effect of birth year on the genetic variance by using gene-environment interaction models, modest but statistically significant increase was found. The interaction effect was 0.050 (95% CI 0.018–0.082) in men and 0.043 (95% CI 0.019–0.071) in women for the genetic path coefficient per 10 years. This turns to 1.37 (95% CI 0.50–2.27) increase of genetic variance in men and 1.07 (95% CI 0.46–1.79) increase of genetic variance in women per 25 years, i.e. approximately one human generation.

Figure 1

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Univariate quantitative genetic models for height were then conducted separately in the three geographic-cultural regions (Figure 1 and Supplementary file 1A). The pattern in Europe was practically the same as that observed for the pooled data because it represents a large proportion of the total sample. In North America and Australia, the total variance of height was greater than in Europe, but the pattern of genetic and environmental variances was less consistent across birth-year cohorts. In East Asia, because of the smaller sample size, the magnitude of the variance components between the birth-year cohorts fluctuated more than in the other two geographic-cultural regions. Genetic variance was generally greater in men than in women in the three geographic-cultural regions. Variance components of height (both raw and relative proportion) showed a similar pattern across birth-year cohorts when analyses were performed for men and women together (Supplementary file 1C).

This very large twin study showed no clear pattern in the heritability of height across birth-year cohorts and thus does not support the hypothesis that the heritability of height is lower in populations with low living standards compared with affluent populations, nor that the heritability of height will increase within a population as living standards improve. Since infant mortality rates are higher in men than in women, both in singletons (Drevenstedt et al., 2008) and twins (Pongou, 2013), the higher heritability observed for men in the earliest cohorts could be explained by selection effects since those who survived were the genetically more advantaged and thus less vulnerable to environmental conditions. The greater relative environmental effect on height variation in women than in men, although unexpected because women’s growth is considered to be more resistant to environmental influences, is in agreement with the findings in Finnish twins born prior to 1958 (Silventoinen et al., 2000). This might indicate differential access to food and medical care (Eveleth and Tanner, 1990). Women are also more likely to develop osteoporosis leading to shrinking in old age (National Institute of Arthritis and musculoskeletal and Skin Diseases, 2014), which may affect the greater influence of unique environmental factors in women born in 1886–1910. This idea is supported by results showing that although genetic factors play an important role in bone loss in early postmenopausal women, their effect weakens with age and completely disappears with advanced aging (Moayyeri et al., 2012).

Total and genetic variance of height generally increased across birth-year cohorts; gene-birth year interaction analysis showed that the genetic variance increase was only modest even when it was statistically significant in this very large twin cohort. However, part of the increase in total variance in some birth-year cohorts was also due to the increase in shared environmental variance. This suggests that both greater ethnic diversity and variation in living standards have contributed to the secular increase in height variation. The greatest total height variation in North America and Australia was due to both genetic and environmental factors and the pattern of variance components across the birth cohorts was less consistent than in Europe. A recent study across 14 European countries found that many independent loci contribute to population genetic differences in height and estimated that these differences account for 24% of the captured additive genetic variance (Robinson et al., 2015). Therefore, it may be that both allelic frequencies and the effects of genes affecting height vary between the geographic-cultural regions. It has been previously shown that even when the total variance of height was greater in Western populations than in East Asian populations, heritability estimates were largely similar in adolescence (Hur et al., 2008) and from 1 to 19 years of age (Jelenkovic et al., 2016); however, the limited statistical power in the data from East Asia does not allow for comparisons across birth cohorts.

The main strength of the present study is the very large sample size of our multinational database of twin cohorts, with adult height data from individuals born between year 1886 and 1994, allowing a more detailed investigation of the genetic and environmental contributions to individual differences in height across birth cohorts than in the previous studies. Important advantages of individual-based data are improved opportunities for statistical modeling and lack of publication bias. This type of analysis is difficult to perform by using literature-based meta-analyses because most of the published studies do not provide the needed statistics by birth-year cohorts. However, our study also has limitations. Countries and/or ethnic-cultural regions are not equally represented and the database is heavily weighted toward populations following Westernized lifestyles. In the classical twin design, parental phenotypic assortment increases dizygotic correlations and thus inflates the shared environmental component when not accounted for in the modeling. In our database, we do not have information on parental height and thus could not take into account assortative mating, which may thus explain part of the shared environmental variation. In addition, most of the height measures were self-reported (Silventoinen et al., 2015), which may bias our analyses toward higher estimates of unique environmental effects due to increased measurement error. However, these sources of bias are unlikely to explain our main result, i.e., relatively similar heritability estimates of adult height over birth cohorts. Finally, since we previously showed that there was no zygosity difference in height variance (Jelenkovic et al., 2015), variance components estimates should not be affected by changes in the proportion of MZ to DZ twins across birth-year cohorts.

In conclusion, although the genetic variance of height showed a slightly increasing trend with birth year, heritability estimates did not present any clear pattern of secular changes across birth-year cohorts from 1886 to 1994. Thus, our findings do not support the hypothesis that the heritability of height increases along with increasing living standards and diminishing rate of absolute poverty within populations.

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Author details

1. Aline Jelenkovic

   1. Department of Social Research, University of Helsinki, Helsinki, Finland
   2. Department of Genetics, Physical Anthropology and Animal Physiology, University of the Basque Country, Leioa, Spain

   Contribution

   AJ, In charge of data management, Conducted the analyses, Wrote the first draft of the manuscript and has primary responsibility of the final content

   For correspondence

   aline.jelenkovic@helsinki.fi

   Competing interests

   The authors declare that no competing interests exist.

2. Yoon-Mi Hur

   Department of Education, Mokpo National University, Jeonnam, South Korea

   Contribution

   Y-MH, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

3. Reijo Sund

   Department of Social Research, University of Helsinki, Helsinki, Finland

   Contribution

   RS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

4. Yoshie Yokoyama

   Department of Public Health Nursing, Osaka City University, Osaka, Japan

   Contribution

   YY, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

5. Sisira H Siribaddana

   1. Institute of Research & Development, Battaramulla, Sri Lanka
   2. Faculty of Medicine & Allied Sciences, Rajarata University of Sri Lanka, Saliyapura, Sri Lanka

   Contribution

   SHS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

6. Matthew Hotopf

   NIHR Mental Health Biomedical Research Centre, South London and Maudsley NHS Foundation Trust and, Institute of Psychiatry Psychology and Neuroscience, King's College London, London, United Kingdom

   Contribution

   MH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

7. Athula Sumathipala

   1. Institute of Research & Development, Battaramulla, Sri Lanka
   2. Research Institute for Primary Care and Health Sciences, School for Primary Care Research, Faculty of Health, Keele University, Staffordshire, United Kingdom

   Contribution

   ASu, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

8. Fruhling Rijsdijk

   MRC Social, Genetic & Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King's College London, London, United Kingdom

   Contribution

   FRi, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

9. Qihua Tan

   Epidemiology, Biostatistics and Biodemography, Institute of Public Health, University of Southern Denmark, Odense, Denmark

   Contribution

   QT, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

   Competing interests

   The authors declare that no competing interests exist.

10. Dongfeng Zhang

    Department of Public Health, Qingdao University Medical College, Qingdao, China

    Contribution

    DZ, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

11. Zengchang Pang

    Department of Noncommunicable Diseases Prevention, Qingdao Centers for Disease Control and Prevention, Qingdao, China

    Contribution

    ZP, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

12. Sari Aaltonen

    1. Department of Social Research, University of Helsinki, Helsinki, Finland
    2. Department of Public Health, University of Helsinki, Helsinki, Finland

    Contribution

    SA, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

13. Kauko Heikkilä

    Department of Public Health, University of Helsinki, Helsinki, Finland

    Contribution

    KH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

14. Sevgi Y Öncel

    Department of Statistics, Faculty of Arts and Sciences, Kirikkale University, Kirikkale, Turkey

    Contribution

    SYÖ, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

15. Fazil Aliev

    1. Faculty of Business, Karabuk University, Karabuk, Turkey
    2. Department of Psychology, Virginia Commonwealth University, Richmond, United States
    3. Department of African American Studies, Virginia Commonwealth University, Richmond, United States

    Contribution

    FA, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

16. Esther Rebato

    Department of Genetics, Physical Anthropology and Animal Physiology, University of the Basque Country, Leioa, Spain

    Contribution

    ER, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

17. Adam D Tarnoki

    1. Department of Radiology and Oncotherapy, Semmelweis University, Budapest, Hungary
    2. Hungarian Twin Registry, Budapest, Hungary

    Contribution

    ADT, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

18. David L Tarnoki

    1. Department of Radiology and Oncotherapy, Semmelweis University, Budapest, Hungary
    2. Hungarian Twin Registry, Budapest, Hungary

    Contribution

    DLT, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

19. Kaare Christensen

    1. The Danish Twin Registry, University of Southern Denmark, Odense, Denmark
    2. Department of Public Health, Epidemiology, Biostatistics & Biodemography, University of Southern Denmark, Odense, Denmark
    3. Department of Clinical Biochemistry and Pharmacology, Odense University Hospital, Odense, Denmark
    4. Department of Clinical Genetics, Odense University Hospital, Odense, Denmark

    Contribution

    KC, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

20. Axel Skytthe

    1. The Danish Twin Registry, University of Southern Denmark, Odense, Denmark
    2. Department of Public Health, Epidemiology, Biostatistics & Biodemography, University of Southern Denmark, Odense, Denmark

    Contribution

    ASk, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

21. Kirsten O Kyvik

    1. Department of Clinical Research, University of Southern Denmark, Odense, Denmark
    2. Odense Patient data Explorative Network (OPEN), Odense University Hospital, Odense, Denmark

    Contribution

    KOK, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

22. Judy L Silberg

    Department of Human and Molecular Genetics, Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, Virginia, United States

    Contribution

    JLS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

23. Lindon J Eaves

    Department of Human and Molecular Genetics, Virginia Institute for Psychiatric and Behavioral Genetics, Virginia Commonwealth University, Richmond, Virginia, United States

    Contribution

    LJE, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

24. Hermine H Maes

    Department of Human and Molecular Genetics, Psychiatry & Massey Cancer Center, Virginia Commonwealth University, Richmond, Virginia, United States

    Contribution

    HHM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

25. Tessa L Cutler

    The Australian Twin Registry, Centre for Epidemiology and Biostatistics, The University of Melbourne, Melbourne, Australia

    Contribution

    TLC, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

26. John L Hopper

    1. The Australian Twin Registry, Centre for Epidemiology and Biostatistics, The University of Melbourne, Melbourne, Australia
    2. Department of Epidemiology, School of Public Health, Seoul National University, Seoul, Korea

    Contribution

    JLH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

27. Juan R Ordoñana

    1. Department of Human Anatomy and Psychobiology, University of Murcia, Murcia, Spain
    2. IMIB-Arrixaca, Murcia, Spain

    Contribution

    JRO, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

28. Juan F Sánchez-Romera

    1. IMIB-Arrixaca, Murcia, Spain
    2. Department of Developmental and Educational Psychology, University of Murcia, Murcia, Spain

    Contribution

    JFS-R, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

29. Lucia Colodro-Conde

    1. Department of Human Anatomy and Psychobiology, University of Murcia, Murcia, Spain
    2. QIMR Berghofer Medical Research Institute, Brisbane, Australia

    Contribution

    LC-C, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

30. Wendy Cozen

    1. Department of Preventive Medicine, Keck School of Medicine of USC, University of Southern California, Los Angeles, United States
    2. USC Norris Comprehensive Cancer Center, Los Angeles, United States

    Contribution

    WC, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

31. Amie E Hwang

    Department of Preventive Medicine, Keck School of Medicine of USC, University of Southern California, Los Angeles, United States

    Contribution

    AEH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

32. Thomas M Mack

    1. Department of Preventive Medicine, Keck School of Medicine of USC, University of Southern California, Los Angeles, United States
    2. USC Norris Comprehensive Cancer Center, Los Angeles, United States

    Contribution

    TMM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

33. Joohon Sung

    1. Department of Epidemiology, School of Public Health, Seoul National University, Seoul, Korea
    2. Institute of Health and Environment, Seoul National University, Seoul, South-Korea

    Contribution

    JS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

34. Yun-Mi Song

    Department of Family Medicine, Samsung Medical Center, Sungkyunkwan University School of Medicine, Seoul, South-Korea

    Contribution

    Y-MS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

35. Sarah Yang

    1. Department of Epidemiology, School of Public Health, Seoul National University, Seoul, Korea
    2. Institute of Health and Environment, Seoul National University, Seoul, South-Korea

    Contribution

    SY, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

36. Kayoung Lee

    Department of Family Medicine, Busan Paik Hospital, Inje University College of Medicine, Busan, Korea

    Contribution

    KL, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

37. Carol E Franz

    Department of Psychiatry, University of California, San Diego, San Diego, United States

    Contribution

    CEF, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

38. William S Kremen

    1. Department of Psychiatry, University of California, San Diego, San Diego, United States
    2. VA San Diego Center of Excellence for Stress and Mental Health, La Jolla, CA, United States

    Contribution

    WSK, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

39. Michael J Lyons

    Department of Psychology, Boston University, Boston, United States

    Contribution

    MJL, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

40. Andreas Busjahn

    HealthTwiSt GmbH, Berlin, Germany

    Contribution

    AB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

41. Tracy L Nelson

    Department of Health and Exercise Sciences and Colorado School of Public Health, Colorado State University, Colorado, United States

    Contribution

    TLN, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

42. Keith E Whitfield

    Psychology and Neuroscience, Duke University, Durham, United States

    Contribution

    KEW, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

43. Christian Kandler

    Department of Psychology, Bielefeld University, Bielefeld, Germany

    Contribution

    CK, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

44. Kerry L Jang

    Department of Psychiatry, University of British Columbia, Vancouver, Canada

    Contribution

    KLJ, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

45. Margaret Gatz

    1. Department of Psychology, University of Southern California, Los Angeles, United States
    2. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    Contribution

    MG, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

46. David A Butler

    Health and Medicine Division, The National Academies of Sciences, Engineering, and Medicine, Washington, United States

    Contribution

    DAB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

47. Maria A Stazi

    Istituto Superiore di Sanità - National Center for Epidemiology, Surveillance and Health Promotion, Rome, Italy

    Contribution

    MAS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

48. Corrado Fagnani

    Istituto Superiore di Sanità - National Center for Epidemiology, Surveillance and Health Promotion, Rome, Italy

    Contribution

    CF, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

49. Cristina D'Ippolito

    Istituto Superiore di Sanità - National Center for Epidemiology, Surveillance and Health Promotion, Rome, Italy

    Contribution

    CD, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

50. Glen E Duncan

    Washington State Twin Registry, Washington State University - Health Sciences Spokane, Spokane, United States

    Contribution

    GED, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

51. Dedra Buchwald

    Washington State Twin Registry, Washington State University, Seattle, United States

    Contribution

    DB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

52. Catherine A Derom

    1. Centre of Human Genetics, University Hospitals Leuven, Leuven, Belgium
    2. Department of Obstetrics and Gynaecology, Ghent University Hospitals, Ghent, Belgium

    Contribution

    CAD, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

53. Robert F Vlietinck

    Centre of Human Genetics, University Hospitals Leuven, Leuven, Belgium

    Contribution

    RFV, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

54. Ruth JF Loos

    1. The Charles Bronfman Institute for Personalized Medicine, Icahn School of Medicine at Mount Sinai, New York, United States
    2. The Mindich Child Health and Development Institute, Icahn School of Medicine at Mount Sinai, New York, United States

    Contribution

    RJFL, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

55. Nicholas G Martin

    Genetic Epidemiology Department, QIMR Berghofer Medical Research Institute, Brisbane, Australia

    Contribution

    NGM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

56. Sarah E Medland

    Genetic Epidemiology Department, QIMR Berghofer Medical Research Institute, Brisbane, Australia

    Contribution

    SEM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

57. Grant W Montgomery

    Molecular Epidemiology Department, QIMR Berghofer Medical Research Institute, Brisbane, Australia

    Contribution

    GWM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

58. Hoe-Uk Jeong

    Department of Education, Mokpo National University, Jeonnam, South Korea

    Contribution

    H-UJ, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

59. Gary E Swan

    Stanford Prevention Research Center, Department of Medicine, Stanford University School of Medicine, Stanford, United States

    Contribution

    GES, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

60. Ruth Krasnow

    Center for Health Sciences, SRI International, Menlo Park, United States

    Contribution

    RK, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

61. Patrik KE Magnusson

    Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    Contribution

    PKEM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

62. Nancy L Pedersen

    Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    Contribution

    NLP, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

63. Anna K Dahl-Aslan

    1. Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden
    2. Institute of Gerontology and Aging Research Network – Jönköping (ARN-J), School of Health and Welfare, Jönköping University, Jönköping, Sweden

    Contribution

    AKD-A, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

64. Tom A McAdams

    MRC Social, Genetic & Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King's College London, London, United Kingdom

    Contribution

    TAM, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

65. Thalia C Eley

    MRC Social, Genetic & Developmental Psychiatry Centre, Institute of Psychiatry, Psychology & Neuroscience, King's College London, London, United Kingdom

    Contribution

    TCE, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

66. Alice M Gregory

    Department of Psychology, Goldsmiths, University of London, London, United Kingdom

    Contribution

    AMG, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

67. Per Tynelius

    Department of Public Health Sciences, Karolinska Institutet, Stockholm, Sweden

    Contribution

    PT, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

68. Laura A Baker

    Department of Psychology, University of Southern California, Los Angeles, United States

    Contribution

    LAB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

69. Catherine Tuvblad

    1. Department of Psychology, University of Southern California, Los Angeles, United States
    2. School of Law, Psychology and Social Work, Örebro University, Örebro, Sweden

    Contribution

    CT, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

70. Gombojav Bayasgalan

    Healthy Twin Association of Mongolia, Ulaanbaatar, Mongolia

    Contribution

    GB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

71. Danshiitsoodol Narandalai

    1. Healthy Twin Association of Mongolia, Ulaanbaatar, Mongolia
    2. Graduate School of Biomedical and Health Sciences, Hiroshima University, Hiroshima, Japan

    Contribution

    DN, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

72. Paul Lichtenstein

    Department of Medical Epidemiology and Biostatistics, Karolinska Institutet, Stockholm, Sweden

    Contribution

    PL, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

73. Timothy D Spector

    Department of Twin Research and Genetic Epidemiology, King's College, London, United Kingdom

    Contribution

    TDS, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

74. Massimo Mangino

    Department of Twin Research and Genetic Epidemiology, King's College, London, United Kingdom

    Contribution

    MMa, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

75. Genevieve Lachance

    Department of Twin Research and Genetic Epidemiology, King's College, London, United Kingdom

    Contribution

    GL, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

76. Meike Bartels

    Department of Biological Psychology, VU University Amsterdam, Amsterdam, Netherlands

    Contribution

    MB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

77. Toos CEM van Beijsterveldt

    Department of Biological Psychology, VU University Amsterdam, Amsterdam, Netherlands

    Contribution

    TCEMvB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

78. Gonneke Willemsen

    Department of Biological Psychology, VU University Amsterdam, Amsterdam, Netherlands

    Contribution

    GW, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

79. S Alexandra Burt

    Michigan State University, East Lansing, Michigan, United States

    Contribution

    SAB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

80. Kelly L Klump

    Michigan State University, East Lansing, Michigan, United States

    Contribution

    KLK, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

81. Jennifer R Harris

    Norwegian Institute of Public Health, Oslo, Norway

    Contribution

    JRH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

82. Ingunn Brandt

    Norwegian Institute of Public Health, Oslo, Norway

    Contribution

    IB, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

83. Thomas Sevenius Nilsen

    Norwegian Institute of Public Health, Oslo, Norway

    Contribution

    TSN, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

84. Robert F Krueger

    Department of Psychology, University of Minnesota, Minneapolis, United States

    Contribution

    RFK, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

85. Matt McGue

    Department of Psychology, University of Minnesota, Minneapolis, United States

    Contribution

    MMcGu, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

86. Shandell Pahlen

    Department of Psychology, University of Minnesota, Minneapolis, United States

    Contribution

    SP, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

87. Robin P Corley

    Institute for Behavioral Genetics, University of Colorado, Boulder, United States

    Contribution

    RPC, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

88. Jacob v B Hjelmborg

    1. The Danish Twin Registry, University of Southern Denmark, Odense, Denmark
    2. Department of Public Health, Epidemiology, Biostatistics & Biodemography, University of Southern Denmark, Odense, Denmark

    Contribution

    JvBH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

89. Jack H Goldberg

    Department of Epidemiology, School of Public Health, University of Washington, Seattle, United States

    Contribution

    JHG, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

90. Yoshinori Iwatani

    Osaka University Graduate School of Medicine, Osaka University, Osaka, Japan

    Contribution

    YI, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

91. Mikio Watanabe

    Osaka University Graduate School of Medicine, Osaka University, Osaka, Japan

    Contribution

    MW, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

92. Chika Honda

    Osaka University Graduate School of Medicine, Osaka University, Osaka, Japan

    Contribution

    CH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

93. Fujio Inui

    1. Osaka University Graduate School of Medicine, Osaka University, Osaka, Japan
    2. Faculty of Health Science, Kio University, Nara, Japan

    Contribution

    FI, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

94. Finn Rasmussen

    Department of Public Health Sciences, Karolinska Institutet, Stockholm, Sweden

    Contribution

    FRa, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

95. Brooke M Huibregtse

    Institute for Behavioral Genetics, University of Colorado, Boulder, United States

    Contribution

    BMH, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

96. Dorret I Boomsma

    Department of Biological Psychology, VU University Amsterdam, Amsterdam, Netherlands

    Contribution

    DIB, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

97. Thorkild I A Sørensen

    1. Novo Nordisk Foundation Centre for Basic Metabolic Research (Section on Metabolic Genetics), University of Copenhagen, Copenhagen, Denmark
    2. Department of Public Health, Faculty of Health and Medical Sciences, University of Copenhagen, Copenhagen, Denmark
    3. Institute of Preventive Medicine, Bispebjerg and Frederiksberg Hospitals, Copenhagen, Denmark

    Contribution

    TIAS, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

98. Jaakko Kaprio

    1. Department of Public Health, University of Helsinki, Helsinki, Finland
    2. Institute for Molecular Medicine FIMM, Helsinki, Finland

    Contribution

    JK, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

99. Karri Silventoinen

    1. Department of Social Research, University of Helsinki, Helsinki, Finland
    2. Osaka University Graduate School of Medicine, Osaka University, Osaka, Japan

    Contribution

    KS, Planned the study design of the CODATwins project, Collected the data used in this study, Commented the manuscript, Read and approved the final version of the manuscript

    Competing interests

    The authors declare that no competing interests exist.

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