Google scholar • ResearchGate • Open Science Framework • University of Minnesota

Education

Doctor of Philosophy (Psychology). University of Minnesota Twin Cities, January 2021.

Master of Arts (Psychology). University of Minnesota Twin Cities, September 2019.

Bachelor of Arts (Biology). Thomas Edison State University, December 2012.

Publications

Under review and in preparation

Lee, J. J., & Willoughby, E. A. (Revision invited, Cognition). Intelligence and reaction time in the two-digit number-comparison task. SSRN preprint: 6520324.

Brown, T. L., & Willoughby, E. A. (Revision invited, Attention, Perception, & Psychophysics). What can color-word Stroop effects tell us about attention?

Brodeur, A., ... Willoughby, E. A. ... & Institute for Replication (Under review). Mass reproducibility in psychological science.

Tan, T., Jayashankar, H., Guan, J., ... Willoughby, E. A. ... & Young, A. S. (Preprint). Family-GWAS reveals effects of environment and mating on genetic associations. medRxiv DOI: 10.1101/2024.10.01.24314703.

McGue, M., Willoughby, E. A., Krueger, R. F., & Roisman, G. I. (Forthcoming). Characterizing the nature of gene–environment interplay through analysis of twin, adoption and family data. In K. Marceau & C. R. Beam (Eds.), The Cambridge handbook of developmental gene-environment interplay: A lifespan approach. Cambridge, UK: Cambridge University Press.

2026

Schwaba, T., Clapp Sullivan, M. L., Akingbuwa, W. A., Ilves, K., Tanksley, P. T., Williams, C. M., ... Willoughby, E. A. ... & Tucker-Drob, E. M. (2026). Robust inference and correlates from genetic associations with personality. Nature. Online First. DOI: 10.1038/s41586-026-10992-9.

O’Donnell, K. P., Xie, X., Huizenga, B., Herman, A., Willoughby, E. A., Lee, J. J., Krueger, R. F., Wilson, S., Roisman, G. I., Luciana, M., Iacono, W. G., Malone, S., & Vrieze, S. (2026). Growing up with parents who smoke: Genetic and environmental effects on offspring substance use and externalizing psychopathology from a 20-year Minnesota study of adoptive families. Behavior Genetics. Online First. DOI: 10.1007/s10519-026-10278-w.

Dobson, R. T., Edwards, T., Giannelis, A., Miller, G. F., March, M. J., Willoughby, E. A., Krueger, R. F., Roisman, G. I., & McGue, M. (2026). Higher quality parent-offspring relationships predict lower-risk adolescent sexual behavior: A genetically informed sibling comparison. Journal of Adolescent Health, 79(4), 476–482. DOI: 10.1016/j.jadohealth.2026.05.007.

Willoughby, E. A., Edwards, T., & Lee, J. J. (2026). Durable lessons on intelligence from twin and adoption research. Behavior Genetics. Online First. DOI: 10.1007/s10519-026-10270-4.

Starr, A., Ruks, M., Giannelis, A., Willoughby, E., Pettersson, O., Pahnke, C. K. L., ... & von Stumm, S. (2026). Within- and between-family genetic effects on educational achievement vary across countries and ages. Molecular Psychiatry, 31(4), 2029–2037. DOI: 10.1038/s41380-025-03342-0.

2025

Giannelis, A., Willoughby, E. A., Edwards, T., McGue, M., & Lee, J. J. (2025). A genetic common factor underlying self-reported math ability and highest math class taken. Molecular Psychiatry, 30(12), 5701–5709. DOI: 10.1038/s41380-025-03237-0.

Willoughby, E. A., Love, A. C., McGue, M., Edwards, T., Giannelis, A., & Lee, J. J. (2025). The freedom to believe in free will: Evidence from an adoption study against the first law of behavioral genetics. Philosophical Psychology, 1–25. Online First. DOI: 10.1080/09515089.2025.2533519.

Kweon, H., Burik, C. A. P., ... Willoughby, E. A. ... Benjamin, D. J., & Koellinger, P. D. (2025). Associations between common genetic variants and income provide insights about the socio-economic health gradient. Nature Human Behaviour, 9(4), 794–805. DOI: 10.1038/s41562-024-02080-7.

Edwards, T., March, M. J., Willoughby, E. A., & Giannelis, A. (2025). Intelligence and individual differences in astrological belief. Journal of Individual Differences, 46(1), 50–57. DOI: 10.1027/1614-0001/a000434.

Edwards, T., Dawes, C. T., Willoughby, E. A., McGue, M., & Lee, J. J. (2025). More than g: Verbal and performance IQ as predictors of socio-political attitudes. Intelligence, 108, 101876. DOI: 10.1016/j.intell.2024.101876.

Johnson, E. C., Lai, D., Balbona, J. V., ... Willoughby, E. A. ... & Agrawal, A. (2025). Multi-ancestral genome-wide association study of clinically defined nicotine dependence reveals strong genetic correlations with other substance use disorders and health-related traits. Psychological Medicine, 55, e234. DOI: 10.1017/S0033291725100883.

2024

Edwards, T., Giannelis, A., Willoughby, E. A., & Lee, J. J. (2024). Predicting political beliefs with polygenic scores for cognitive performance and educational attainment. Intelligence, 104, 101831. DOI: 10.1016/j.intell.2024.101831.

Willoughby, E. A., Giannelis, A., Iacono, W. G., McGue, M., & Vrieze, S. I. (2024). A century of behavioral genetics at the University of Minnesota—Corrigendum. Twin Research and Human Genetics, 27(4–5), 262. DOI: 10.1017/thg.2023.5.

2023

Kim, Y., Saunders, G. R. B., Giannelis, A., Willoughby, E. A., DeYoung, C. G., & Lee, J. J. (2023). Genetic and neural bases of the neuroticism general factor. Biological Psychology, 184, 108692. DOI: 10.1016/j.biopsycho.2023.108692.

Howe, L. J., Rasheed, H., Jones, P. R., Boomsma, D. I., Evans, D. M., Giannelis, A., ... Willoughby, E. A. ... Brumpton, B., & Davies, N. M. (2023). Educational attainment, health outcomes and mortality: A within-sibship Mendelian randomization study. International Journal of Epidemiology, 52(5), 1579–1591. DOI: 10.1093/ije/dyad079.

Willoughby, E. A., Polderman, T. J. C., & Boutwell, B. B. (2023). Behavioural genetics methods. Nature Reviews Methods Primers, 3, Article 10. DOI: 10.1038/s43586-022-00191-x.

Willoughby, E. A., Kim, Y., Lee, J. J., & DeYoung, C. G. (2023). The construct validity of Intellect and Openness as distinct aspects of personality through differential associations with reaction time. Journal of Intelligence, 11(2), 30. DOI: 10.3390/jintelligence11020030.

Giannelis, A., Willoughby, E. A., Corley, R., Hopfer, C., Hewitt, J. K., Iacono, W. G., Anderson, J., Rustichini, A., Vrieze, S. I., McGue, M., & Lee, J. J. (2023). The association between saving disposition and financial distress: A genetically informed approach. Journal of Economic Psychology, 96, 102610. DOI: 10.1016/j.joep.2023.102610.

2022

Willoughby, E. A., Giannelis, A., Iacono, W. G., McGue, M., & Vrieze, S. I. (2022). A century of behavioral genetics at the University of Minnesota. Twin Research and Human Genetics, 25(6), 211–225. DOI: 10.1017/thg.2023.1.

McGue, M., Anderson, E. L., Willoughby, E. A., Giannelis, A., Iacono, W. G., & Lee, J. J. (2022). Not by g alone: The benefits of a college education among individuals with low levels of general cognitive ability. Intelligence, 92, 101642. DOI: 10.1016/j.intell.2022.101642.

Howe, L. J., Nivard, M. G., Morris, T. T., Hansen, A. F., Rasheed, H., Cho, Y., ... Willoughby, E. A. ... Brumpton, B., Hemani, G., & Davies, N. M. (2022). Within-sibship genome-wide association analyses decrease bias in estimates of direct genetic effects. Nature Genetics, 54(5), 581–592. DOI: 10.1038/s41588-022-01062-7.

2021

Willoughby, E. A. (2021). Drawing and painting dinosaurs. Marlborough, UK: The Crowood Press. ISBN: 978-1785009556.

Willoughby, E. A., & Lee, J. J. (2021). Predicting cognitive-ability differences from genetic and brain-imaging data. In A. K. Barbey, S. Karama, & R. J. Haier (Eds.), The Cambridge handbook of intelligence and cognitive neuroscience (pp. 349–364). Cambridge University Press. DOI: 10.1017/9781108635462.022.

Willoughby, E. A., McGue, M., Iacono, W. G., & Lee, J. J. (2021). Genetic and environmental contributions to IQ in adoptive and biological families with 30-year-old offspring. Intelligence, 88, 101579. DOI: 10.1016/j.intell.2021.101579.

Willoughby, E. A., & Lee, J. J. (2021). Parsing information flow in speeded cognitive tasks: The role of g in perception and decision time. Journal of Experimental Psychology: Learning, Memory, and Cognition, 47(11), 1792–1809. DOI: 10.1037/xlm0001026.

Willoughby, E. A., Giannelis, A., Ludeke, S., Klemmensen, R., Nørgaard, A. S., Iacono, W. G., Lee, J. J., & McGue, M. (2021). Parent contributions to the development of political attitudes in adoptive and biological families. Psychological Science, 32(12), 2023–2034. DOI: 10.1177/09567976211021844.

Willoughby, E. A., McGue, M., Iacono, W. G., Rustichini, A., & Lee, J. J. (2021). The role of parental genotype in predicting offspring years of education: Evidence for genetic nurture. Molecular Psychiatry, 26(8), 3896–3904. DOI: 10.1038/s41380-019-0494-1.

Anderson, E. L., Saunders, G. R. B., Willoughby, E. A., Iacono, W. G., & McGue, M. (2021). The role of the shared environment in college attainment: An adoption study. Journal of Personality, 89(3), 580–593. DOI: 10.1111/jopy.12600.

2020

McGue, M., Willoughby, E. A., Rustichini, A., Johnson, W., Iacono, W. G., & Lee, J. J. (2020). The contribution of cognitive and noncognitive skills to intergenerational social mobility. Psychological Science, 31(7), 835–847. DOI: 10.1177/0956797620924677.

Gulsuner, S., Stein, D. J., Susser, E. S., Sibeko, G., Pretorius, A., Walsh, T., Majara, L., Mndini, M. M., Mqulwana, S. G., Ntola, O. A., Casadei, S., Ngqengelele, L. L., Korchina, V., van der Merwe, C., Malan, M., Fader, K. M., Feng, M., Willoughby, E. A. . . . & McClellan, J. M. (2020). Genetics of schizophrenia in the South African Xhosa. Science, 367(6477), 569–573. DOI: 10.1126/science.aay8833.

Kane, J., & Willoughby, E. (2020). God’s word or human reason? An inside perspective on creationism (Second edition). Portland, Oregon: Inkwater Press. ISBN: 978-1629016382. (Author & lead illustrator)

2019

Willoughby, E. A., Love, A. C., McGue, M., Iacono, W. G., Quigley, J., & Lee, J. J. (2019). Free will, determinism, and intuitive judgments about the heritability of behavior. Behavior Genetics, 49(2), 136–153. DOI: 10.1007/s10519-018-9931-1.

2018

Lee, J. J., Wedow, R., Okbay, A., Kong, E., Maghzian, O., Zacher, M., ... Willoughby, E. A. ... & Cesarini, D. (2018). Gene discovery and polygenic prediction from a genome-wide association study of educational attainment in 1.1 million individuals. Nature Genetics, 50(8), 1112–1121. DOI: 10.1038/s41588-018-0147-3.

Willoughby, E. A., & Boutwell, B. B. (2018). Importance of intelligence and emotional intelligence for physicians. Journal of the American Medical Association, 320(2), 205. DOI: 10.1001/jama.2018.6278.

2016

Kane, J., Willoughby, E., & Keesey, T. M. (2016). God’s word or human reason? An inside perspective on creationism (First edition). Portland, Oregon: Inkwater Press. ISBN: 1629013722. (Author & lead illustrator)