Curriculum vitaeGoogle scholarResearchGateOpen Science FrameworkUniversity of Minnesota


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.


Under review and in preparation

Willoughby, E. A., Giannelis, A., McGue, M., Iacono, W. G., Lee, J. J., & Love, A. C. (In preparation). The etiology of beliefs about free will and determinism in adoptive and biological families.

Giannelis, A., Willoughby, E. A., Saunders, Wolfram, T., G., Iacono, W.G., McGue, M. & Lee, J. J. (In preparation). A genome-wide association study of specific mathematical ability.

Giannelis, A., Willoughby, E. A., Iacono, W.G., Anderson J., Rustichini, A., Vrieze S., McGue, M. & Lee, J. J. (In preparation). The intergenerational transmission of saving disposition and financial distress in adoptive and biological families.

Brown, T. L. & Willoughby, E. A. (Under review). The color-word Stroop task as a measure of selective attention and cognitive control in experimental and individual differences research.

Edwards, T., Giannelis, A., Willoughby, E. A., & Lee, James J. (In preparation). Predicting political beliefs with polygenic scores for cognitive performance and educational attainment.


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, dyad079. DOI: 10.1093/ije/dyad079.

Willoughby, E. A., Giannelis, A. and Iacono, W. G., McGue, M., & Vrieze, S. (2023). 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.

Willoughby, E. A., Polderman, T. J. C. & Boutwell, B. (2023). Behavior genetics methods. Nature Reviews Methods Primers, 3(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., 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.


McGue, M., Anderson, E. A., 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., Hansen, A., Rasheed, H., Cho, Y., Chittoor, G., Lind, P., Palviainen, T., ... Giannelis, A., Willoughby, E. A., Andreassen, O., Ask, H. Corfield, E., Hughes, A., Magnus, P., Njolstad, P. R., Stoltenberg, C. Lee, J. J., Oskarsson, S. (2022). Within- sibship GWAS of 25 phenotypes improve estimates of direct genetic effects. Nature Genetics, 54, 581–592. DOI: 10.1038/s41588-022-01062-7.


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 brain imaging and genetics data. In: The Cambridge handbook of intelligence and neuroscience. Cambridge, UK: Cambridge University Press. ISBN: 978-1108480543

Willoughby, E. A., McGue, M., Iacono, W. G., & Lee, James J. (2021). The heritability of intelligence estimated in biological and adoptive families with 30-year-old offspring. Intelligence, 88(101579). DOI: 10.1016/j.intell.2021.101579.

Willoughby, E. A., & Lee, James J. (2021). Parsing information flow in speeded cognitive tasks: The associations of g with 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 non-adoptive families. Psychological Science, 32(12), 2023–2034. DOI: 10.1177/09567976211021844.


Anderson, E. L., Saunders, G. R. B., Willoughby, E. A., Iacono, W. G., McGue, M. (2020). The role of the shared environment in college attainment: An adoption study. Journal of Personality, 89(3), 580–593. DOI: 10.1111/jopy.12600

McGue, M., Willoughby, E. A., Rustichini, A., Johnson, W., Iacono, W., & Lee, J. J. (2020). The contribution of cognitive and non-cognitive 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

Willoughby, E. A., McGue, M., Iacono, W., Rustichini, A., & Lee, J. J. (2020). The role of parental genotype in predicting offspring years of education: Evidence for genetic nurture. Molecular Psychiatry, 26, 3896–3904. DOI: 10.1038/s41380-019-0494-1

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


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


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

Lee, J. J., Wedow, R., Okbay, A., Kong, E., Maghzian, O., Zacher, M., Nguyen-Viet, T. A., Bowers, P., Sidorenko, J., Linner, R. K., Fontana, M. A., Kundu, T., Lee, C., Li, H., Li, R., Royer, R., Timshel, P. N., Walters, R. K., Willoughby, E. A. . . . Cesarini, D. (2018). Gene discovery and polygenic prediction from a 1.1-million-person GWAS of educational attainment. Nature Genetics, 50(8), 1112–1121. DOI: 10.1038/s41588-018-0147-3


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

Popular science articles

"The compassionate way to combat creationism" at Quillette Magazine, published Oct 2017

"Amber preserves detail of ancient bird wings" at, published Aug 2016

"Nests, neurons and the evolution of behavior" at, published Aug 2016

"Velociraptor needs your help" at, published July 2016

"Bird eyes: the greatest eyes on earth" at, published June 2016

"Big, beautiful bird brains" at, published March 2016

"Dakotaraptor: giant raptor straight out of Hell Creek" at, published Nov 2015