Adjusting for Heritable Covariates Can Bias Effect Estimates in Genome-Wide Association Studies

Principal components analysis corrects for stratification in genome-wide association studies

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Genetic relationship between five psychiatric disorders estimated from genome-wide SNPs

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Overadjustment bias and unnecessary adjustment in epidemiologic studies.

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Genome-wide association yields new sequence variants at seven loci that associate with measures of obesity

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Understanding mechanisms underlying human gene expression variation with RNA sequencing

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Discerning asthma endotypes through comorbidity mapping

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Genetic architectures of cerebral ventricles and their overlap with neuropsychiatric traits

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Controlling population structure in the genomic prediction of tropical maize hybrids

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Lung function, COPD and cognitive function: a multivariable and two sample Mendelian randomization study

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Genomic analysis of intracranial and subcortical brain volumes yields polygenic scores accounting for variation across ancestries

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Clarifying the causes of consistent and inconsistent findings in genetics

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Modelling local gene networks increases power to detect trans-acting genetic effects on gene expression

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Rare variants of small effect size in neuronal excitability genes influence clinical outcome in Japanese cases of SCN1A truncation-positive Dravet syndrome

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Genetic variants affecting bone mineral density and bone mineral content at multiple skeletal sites in Hispanic children

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Re-evaluating the relationship between missing heritability and the microbiome

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Blood pressure and risk of cancer: a Mendelian randomization study

Chan, I. I., Kwok, M. K., & Schooling, C. M. (2021) · BMC Cancer · doi:10.1186/s12885-021-09067-x · Cited by 24

Exome-Derived Adiponectin-Associated Variants Implicate Obesity and Lipid Biology

Spracklen, C. N., Karaderi, T., Yaghootkar, H., Schurmann, C., Fine, R. S., Kutalik, Z., Preuss, M. H., Lu, Y., Wittemans, L. B. L., Adair, L. S., et al. (2019) · The American Journal of Human Genetics · doi:10.1016/j.ajhg.2019.05.002 · Cited by 25

Problems in interpreting and using GWAS of conditional phenotypes illustrated by 'alcohol GWAS'

Holmes, M. V., & Smith, G. D. (2018) · Molecular Psychiatry · doi:10.1038/s41380-018-0037-1 · Cited by 23

A rare variant analysis framework using public genotype summary counts to prioritize disease-predisposition genes

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Genetics-informed precision treatment formulation in schizophrenia and bipolar disorder

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Adjusting for Principal Components of Molecular Phenotypes Induces Replicating False Positives

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A versatile, fast and unbiased method for estimation of gene-by-environment interaction effects on biobank-scale datasets

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Genetic Variants in the Insulin-like Growth Factor Pathway and Colorectal Cancer Risk in the Netherlands Cohort Study

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Genetic variants associated with type 2 diabetes and adiposity and risk of intracranial and abdominal aortic aneurysms

Hof, F. N. G. V. ’., Vaucher, J., Holmes, M. V., de Wilde, A., Baas, A. F., Blankensteijn, J. D., Albert Hofman, D., Kiemeney, L. A. L. M., Rivadeneira, F., Uitterlinden, A. G., et al. (2017) · European Journal of Human Genetics · doi:10.1038/ejhg.2017.48 · Cited by 23

Genetic basis for plasma amino acid concentrations based on absolute quantification: a genome-wide association study in the Japanese population

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ALDH2 genotype modulates the association between alcohol consumption and AST/ALT ratio among middle-aged Japanese men: a genome-wide G × E interaction analysis

Sut­oh, Y., Hachiya, T., Suzuki, Y., Komaki, S., Ohmomo, H., Kakisaka, K., Wang, T., Takikawa, Y., & Shimizu, A. (2020) · Scientific Reports · doi:10.1038/s41598-020-73263-1 · Cited by 21

A Method to Exploit the Structure of Genetic Ancestry Space to Enhance Case-Control Studies

Bodea, C. A., Neale, B. M., Ripke, S., Daly, M. J., Devlin, B., Roeder, K., Barclay, M. L., Peyrin‐Biroulet, L., Chamaillard, M., Colombel, J.-F., et al. (2016) · The American Journal of Human Genetics · doi:10.1016/j.ajhg.2016.02.025 · Cited by 21

The influence of rare variants in circulating metabolic biomarkers

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Body mass index: Has epidemiology started to break down causal contributions to health and disease?

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The total and direct effects of systolic and diastolic blood pressure on cardiovascular disease and longevity using Mendelian randomisation

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Adiposity, metabolites, and colorectal cancer risk: Mendelian randomization studyPreprint

Bull, C. J., Bell, J. A., Murphy, N., Sanderson, E. C. M., Smith, G. D., Timpson, N. J., Banbury, B. L., Albanes, D., Berndt, S. I., Bezieau, S., et al. (2020) · medRxiv · doi:10.1101/2020.03.19.20031138 · Cited by 19

ARMC5 Variants and Risk of Hypertension in Blacks: MH‐GRID Study

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Discordant associations of educational attainment with ASD and ADHD implicate a polygenic form of pleiotropy

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Larger cerebral cortex is genetically correlated with greater frontal area and dorsal thickness

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150 risk variants for diverticular disease of intestine prioritize cell types and enable polygenic prediction of disease susceptibility

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The overlapping genetic architecture of psychiatric disorders and cortical brain structure

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The shared genetic architecture of modifiable risk for Alzheimer's disease: a genomic structural equation modelling study

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Bone mineral density loci specific to the skull portray potential pleiotropic effects on craniosynostosis

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Unraveling the association between depression and telomere length using genomics

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Case-control study on the association between the GATA2 gene and premature myocardial infarction in the Iranian population

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Genome-wide association for testis weight in the diversity outbred mouse population

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Bias in two-sample Mendelian randomization when using heritable covariable-adjusted summary associationsPreprint

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Genome-wide association studies of binge-eating behaviour and anorexia nervosa yield insights into the unique and shared biology of eating disorder phenotypesPreprint

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HostSeq: a Canadian whole genome sequencing and clinical data resource

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Beyond apples and pears: sex-specific genetics of body fat percentage

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Genetic Risk of Trigger Finger: Results of a Genomewide Association Study

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Assessing causal estimates of the association of obesity-related traits with coronary artery disease using a Mendelian randomization approach

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The impact of assortative mating, participation bias and socioeconomic status on the polygenic risk of behavioural and psychiatric traits

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Exploring the causal associations of micronutrients on urate levels and the risk of gout: A Mendelian randomization study

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Pitfalls in performing genome-wide association studies on ratio traits

McCaw, Z. R., Dey, R., Somineni, H., Amar, D., Mukherjee, S., Sandor, K. P., Karaletsos, T., Koller, D., Aschard, H., Smith, G. D., et al. (2025) · Human Genetics and Genomics Advances · doi:10.1016/j.xhgg.2025.100406 · Cited by 14

Genome-wide Association Studies Reveal Novel Locus With Sex-/Therapy-Specific Fracture Risk Effects in Childhood Cancer Survivors

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Contrasting regional architectures of schizophrenia and other complex diseases using fast variance components analysisPreprint

Loh, P., Bhatia, G., Gusev, A., Finucane, H. K., Bulik-Sullivan, B. K., Pollack, S. J., de Candia, T. R., Lee, S., Wray, N. R., Kendler, K. S., et al. (2015) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/016527 · Cited by 13

Letter in response to ‘Bias in two-sample Mendelian randomization when using heritable covariable-adjusted summary associations’—‘Interpreting Mendelian randomization studies pre-adjusted for the heritable covariable survival to recruitment’

Schooling, C. M., Zhao, J. V., Yeung, S. L. A., & Kwok, M. K. (2021) · International Journal of Epidemiology · doi:10.1093/ije/dyab126 · Cited by 13

Sources of bias in genomics research of oral and dental traits

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Impact of Liability to Periodontitis on Glycemic Control and Type II Diabetes Risk: A Mendelian Randomization Study

Shah, P. D., Schooling, C. M., & Borrell, L. N. (2021) · Frontiers in Genetics · doi:10.3389/fgene.2021.767577 · Cited by 14

Genetic Correlation and Bidirectional Causal Association Between Type 2 Diabetes and Pulmonary Function

Zhu, J., Zhao, H., Chen, D., Tse, L. A., Kinra, S., & Li, Y. (2021) · Frontiers in Endocrinology · doi:10.3389/fendo.2021.777487 · Cited by 12

Distinct impact modes of polygenic disposition to dyslexia in the adult brain

Soheili‐Nezhad, S., Schijven, D., Mars, R. B., Fisher, S. E., & Francks, C. (2024) · Science Advances · doi:10.1126/sciadv.adq2754 · Cited by 11

The future of epidemiology: methods or matter?

Ebrahim, S. B. J., Ferrie, J. E., & Smith, G. D. (2016) · International Journal of Epidemiology · doi:10.1093/ije/dyx032 · Cited by 13

Assessing the genetic relationship between gastro-esophageal reflux disease and risk of COVID-19 infection

Ong, J., Gharahkhani, P., Vaughan, T. L., Whiteman, D. C., Kendall, B. J., & MacGregor, S. (2021) · Human Molecular Genetics · doi:10.1093/hmg/ddab253 · Cited by 11

Kidney disease progression and collider bias in GWAS

Khan, A., & Kiryluk, K. (2022) · Kidney International · doi:10.1016/j.kint.2022.06.018 · Cited by 10

Multi-omic association study identifies DNA methylation-mediated genotype and smoking exposure effects on lung function in children living in urban settings

Dapas, M., Thompson, E. E., Wentworth-Sheilds, W., Clay, S. M., Visness, C. M., Calatroni, A., Sordillo, J. E., Gold, D. R., Wood, R. A., Makhija, M., et al. (2023) · PLoS Genetics · doi:10.1371/journal.pgen.1010594 · Cited by 10

Using Genetics to Inform Interventions Related to Sodium and Potassium in Hypertension

Reay, W. R., Clarke, E. D., Eslick, S., Riveros, C., Holliday, E. G., McEvoy, M., Peel, R., Hancock, S., Scott, R. J., Attia, J., et al. (2024) · Circulation · doi:10.1161/circulationaha.123.065394 · Cited by 12

Understanding and applying gene–environment interactions: a guide for nutrition professionals with an emphasis on integration in African research settings

Nienaber‐Rousseau, C. (2024) · Nutrition Reviews · doi:10.1093/nutrit/nuae015 · Cited by 13

Genetically Predicted Circulating C-Reactive Protein Concentration and Colorectal Cancer Survival: A Mendelian Randomization Consortium Study

Hua, X., Dai, J. Y., Lindström, S., Harrison, T. A., Lin, Y., Alberts, S. R., Alwers, E., Berndt, S. I., Brenner, H., Buchanan, D. D., et al. (2021) · Cancer Epidemiology Biomarkers & Prevention · doi:10.1158/1055-9965.epi-20-1848 · Cited by 10

Genetic prediction of micronutrient levels and the risk of colorectal polyps: A mendelian randomization study

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Distinct genetic subtypes of adiposity and glycemic changes in response to weight-loss diet intervention: the POUNDS Lost trial

Chen, Y., Zhou, T., Sun, D., Li, X., Ma, H., Liang, Z., Heianza, Y., Pei, X., Bray, G. A., Sacks, F. M., et al. (2020) · European Journal of Nutrition · doi:10.1007/s00394-020-02244-x · Cited by 10

Causal relationships between genetically determined metabolites and human intelligence: a Mendelian randomization study

Yang, J., Zhao, B., Li, Q., Gao, F., Fan, Y., He, X., Ma, Q., Yang, L., Yan, B., Wang, W., et al. (2021) · Molecular Brain · doi:10.1186/s13041-021-00743-4 · Cited by 11

The consequences of adjustment, correction and selection in genome-wide association studies used for two-sample Mendelian randomization

Walker, V. M., Harrison, S., Carter, A. R., Gill, D., Tzoulaki, I., & Davies, N. M. (2021) · Wellcome Open Research · doi:10.12688/wellcomeopenres.16752.1 · Cited by 9

Investigating the role of insulin in increased adiposity: Bi-directional Mendelian randomization studyPreprint

Richmond, R. C., Wade, K. H., Corbin, L. J., Bowden, J., Hemani, G., Timpson, N. J., & Smith, G. D. (2017) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/155739 · Cited by 11

Genome-wide association study of liver fat, iron, and extracellular fluid fraction in the UK BiobankPreprint

O’Dushlaine, C. T., Germino, M., Verweij, N., Nielsen, J. B., Yadav, A. K., Benner, C., Backman, J., Lin, N., Abecasis, G. R., Baras, A., et al. (2021) · medRxiv · doi:10.1101/2021.10.25.21265127 · Cited by 10

Impact of Alcohol Consumption on Lifespan: a Mendelian randomization study in Europeans

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Multi-ancestry meta-analysis of genome-wide association studies discovers 67 new loci associated with chronic back pain

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Examining the Association between Mitochondrial Genome Variation and Coronary Artery Disease

Vilne, B., Sawant, A. A., & Rudaka, I. (2022) · medRxiv · doi:10.3390/genes13030516 · Cited by 8

Limited Effect of Y Chromosome Variation on Coronary Artery Disease and Mortality in UK Biobank—Brief Report

Timmers, P. R. H. J., & Wilson, J. F. (2022) · Arteriosclerosis Thrombosis and Vascular Biology · doi:10.1161/atvbaha.122.317664 · Cited by 9

Genome-wide association study of the human brain functional connectome reveals strong vascular component underlying global network efficiency

Bell, S., Tozer, D. J., & Markus, H. S. (2022) · Scientific Reports · doi:10.1038/s41598-022-19106-7 · Cited by 8

KLKB1 and CLSTN2 are associated with HDL-mediated cholesterol efflux capacity in a genome-wide association study

Schachtl‐Riess, J. F., Schönherr, S., Lamina, C., Forer, L., Coassin, S., Streiter, G., Kheirkhah, A., Li, Y., Meiselbach, H., Maio, S. D., et al. (2023) · Atherosclerosis · doi:10.1016/j.atherosclerosis.2023.01.022 · Cited by 10

Genetic determinants of activity and antigen levels of contact system factors

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Quantification of genetic components of population differentiation in UK Biobank traits reveals signals of polygenic selectionPreprint

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Genome-Wide Association Study for eGFR in a Taiwanese Population

Chen, Y., Wong, H. S.-C., Wu, M., Chou, W., Kao, C., Chao, C.-H., Chang, W., & Wu, K. (2022) · Clinical Journal of the American Society of Nephrology · doi:10.2215/cjn.02180222 · Cited by 8

Twelve Loci Associated With Bone Density in Middle-aged and Elderly Chinese: The Shanghai Changfeng Study

Zeng, H., Ge, J., Xu, W., Ma, H., Chen, L., Xia, M., Pan, B., Lin, H., Wang, S., & Gao, X. (2022) · The Journal of Clinical Endocrinology & Metabolism · doi:10.1210/clinem/dgac597 · Cited by 8

Risk factors affecting polygenic score performance across diverse cohortsPreprint

Hui, D., DUDEK, S. M., Kiryluk, K., Walunas, T. L., Kullo, I. J., Wei, W., Tiwari, H. K., Peterson, J. F., Chung, W. K., Davis, B. H., et al. (2023) · eLife · doi:10.7554/elife.88149 · Cited by 8

The Genetic Architecture of the Human Corpus Callosum and its Subregions

Bhatt, R. R., Gadewar, S. P., Shetty, A., Gari, I. B., Haddad, E., Javid, S., Ramesh, A., Nourollahimoghadam, E., Zhu, A. H., de Leeuw, C., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-64791-3 · Cited by 7

Is Genetic Risk for Sleep Apnea Causally Linked With Glaucoma Susceptibility?

Ingold, N., Campos, A. I., Han, X., Ong, J., Gharahkhani, P., Mackey, D. A., Rentería, M. E., Law, M. H., & MacGregor, S. (2022) · Investigative Ophthalmology & Visual Science · doi:10.1167/iovs.63.1.25 · Cited by 7

Circulating micronutrient levels and their association with sepsis susceptibility and severity: a Mendelian randomization study

Wei, Z., Liu, Y., Mei, X., Zhong, J., & Huang, F. (2024) · Frontiers in Genetics · doi:10.3389/fgene.2024.1353118 · Cited by 7

Causal associations of antioxidants with Alzheimer’s disease and cognitive function: a Mendelian randomisation study

Wang, J., Huang, Y., Bei, C., Yang, H., Lin, Z., & Xu, L. (2024) · Journal of Epidemiology & Community Health · doi:10.1136/jech-2023-221184 · Cited by 7

Defining a Contemporary Ischemic Heart Disease Genetic Risk Profile Using Historical Data

Mosley, J. D., Driest, S. L. V., Wells, Q. S., Shaffer, C. M., Edwards, T. L., Bastarache, L. A., McCarty, C. A., Thompson, W. K., Chute, C. G., Jarvik, G. P., et al. (2016) · Circulation Cardiovascular Genetics · doi:10.1161/circgenetics.116.001530 · Cited by 7

Letter regarding article, “Associations of obesity and circulating insulin and glucose with breast cancer risk: a Mendelian randomization analysis”

Tan, V. Y., Yarmolinsky, J., Lawlor, D. A., & Timpson, N. J. (2019) · International Journal of Epidemiology · doi:10.1093/ije/dyz013 · Cited by 7

The Trans-Ancestral Genomic Architecture of Glycaemic TraitsPreprint

Chen, J., Spracklen, C. N., Marenne, G., Varshney, A., Corbin, L. J., Luan, J., Willems, S. M., Wu, Y., Zhang, X., Horikoshi, M., et al. (2020) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2020.07.23.217646 · Cited by 7

The genetics of cortical organisation and development: a study of 2,347 neuroimaging phenotypesPreprint

Warrier, V., Stauffer, E.-M., Huang, Q.-Q., Wigdor, E., Slob, E., Seidlitz, J., Ronan, L., Valk, S., Mallard, T., Grotzinger, A., et al. (2022) · bioRxiv · doi:10.1101/2022.09.08.507084 · Cited by 7

Selecting Covariates for Genome-Wide Association StudiesPreprint

Dor, E., Margaliot, I., Brandes, N., Zuk, O., Michal Linial, B., & Rappoport, N. (2023) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2023.02.07.527425 · Cited by 7

Adiposity distribution and risks of 12 obesity-related cancers: a Mendelian randomization analysis

Hazelwood, E., Goudswaard, L. J., Lee, M. A., Vabistsevits, M., Pournaras, D. J., Brenner, H., Buchanan, D. D., Gruber, S. B., Gsur, A., Li, L., et al. (2025) · JNCI Journal of the National Cancer Institute · doi:10.1093/jnci/djaf201 · Cited by 7

Imaging genetics of language network functional connectivity reveals links with language-related abilities, dyslexia and handedness

Amelink, J. S., Postema, M. C., Kong, X., Schijven, D., Carrión-Castillo, A., Soheili‐Nezhad, S., Sha, Z., Molz, B., Joliot, M., Fisher, S. E., et al. (2024) · Communications Biology · doi:10.1038/s42003-024-06890-3 · Cited by 6

Germline mechanisms of immunotherapy toxicities in the era of genome‐wide association studies

Gusev, A. (2023) · Immunological Reviews · doi:10.1111/imr.13253 · Cited by 7

Reproduction and In-Depth Evaluation of Genome-Wide Association Studies and Genome-Wide Meta-analyses Using Summary Statistics

Niu, Y., Ye, C., He, J., Han, F., Guo, L., Zheng, H., & Chen, G. (2017) · G3 Genes Genomes Genetics · doi:10.1534/g3.116.038877 · Cited by 6

Adjustment for covariates using summary statistics of genome‐wide association studies

Wang, T., Xue, X., Xie, X., Ye, K. Q., Zhu, X., & Elston, R. C. (2018) · Genetic Epidemiology · doi:10.1002/gepi.22148 · Cited by 6

Germline variants are associated with increased primary melanoma tumor thickness at diagnosis

Mangantig, E., MacGregor, S., Iles, M. M., Scolyer, R. A., Cust, A. E., Hayward, N. K., Montgomery, G. W., Duffy, D. L., Thompson, J. F., Henders, A. K., et al. (2020) · Human Molecular Genetics · doi:10.1093/hmg/ddaa222 · Cited by 6

Genome-wide association study of habitual physical activity in over 277,000 UK Biobank participants identifies multiple variants including CADM2 and APOEPreprint

Klimentidis, Y. C., Raichlen, D. A., Bea, J. W., Garcia, D. O., Mandarino, L. J., Alexander, G. E., Chen, Z., & Going, S. B. (2017) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/179317 · Cited by 6

The genetic case for cardiorespiratory fitness as a clinical vital sign and the routine prescription of physical activity in healthcarePreprint

Hanscombe, K. B., Persyn, E., Traylor, M., Glanville, K. P., Hamer, M., Coleman, J. R. I., & Lewis, C. M. (2020) · medRxiv · doi:10.1101/2020.12.08.20243337 · Cited by 6

A genome‐wide association study of hand eczema identifies locus 20q13.33 and reveals genetic overlap with atopic dermatitis

Rosenberg, F. M., van der Most, P. J., Loman, L., Kamali, Z., Dittmar, D., Snieder, H., & Schuttelaar, M. L. A. (2024) · Contact Dermatitis · doi:10.1111/cod.14619 · Cited by 6

Variants in FtsJ RNA 2′-O-Methyltransferase 3 and Growth Hormone 1 are associated with small body size and a dental anomaly in dogs

Abrams, S. R., Hawks, A. L., Evans, J. M., Famula, T. R., Mahaffey, M. B., Johnson, G. S., Mason, J. M., & Clark, L. A. (2020) · Proceedings of the National Academy of Sciences · doi:10.1073/pnas.2009500117 · Cited by 5

Leveraging omics data to boost the power of genome-wide association studies

Lin, Z., Knutson, K. A., & Pan, W. (2022) · Human Genetics and Genomics Advances · doi:10.1016/j.xhgg.2022.100144 · Cited by 6

Phenotype integration improves power and preserves specificity in biobank-based genetic studies of MDDPreprint

Dahl, A. W., Thompson, M. J., An, U., Krebs, M., Appadurai, V., Border, R., Bacanu, S., Werge, T., Flint, J., Schork, A., et al. (2023) · bioRxiv · doi:10.1101/2022.08.15.503980 · Cited by 6

Modeling the genomic architecture of adiposity and anthropometrics across the lifespan

Arehart, C. H., Lin, M. H., Gibson, R. A., Anderson, H. D., Aquilante, C. L., Arbogast, K. L., Arehart, C. H., Brooks, I. M., Brunetti, T. M., Brutus-Lestin, J., et al. (2025) · Nature Communications · doi:10.1038/s41467-025-62730-w · Cited by 7

Genome-wide association study of delay discounting identifies 11 loci and reveals transdiagnostic associations across mental and physical health

Thorpe, H. H. A., Cupertino, R. B., Pakala, S. R., Fontanillas, P., Jennings, M. V., Yang, J., Meredith, J. J., Greenwood, T. A., Bianchi, S. B., Vilar‐Ribó, L., et al. (2025) · Molecular Psychiatry · doi:10.1038/s41380-025-03356-8 · Cited by 11

Interpreting disease genome‐wide association studies and polygenetic risk scores given eligibility and study design considerations

Schooling, C. M., & Terry, M. B. (2024) · Genetic Epidemiology · doi:10.1002/gepi.22567 · Cited by 4

Perspective The Covariate’s Dilemma

Mefford, J., & Witte, J. S. (2016)

Consequence of adjustments for demographic or clinical covariates and a recommended solution in genome-wide association studiesPreprint

Sun, X., Li, H., Yu, Y., Yuan, Z., Jin, C., Hou, L., Liu, X., Wáng, Q., & Xue, F. (2021) · bioRxiv (Cold Spring Harbor Laboratory) · doi:10.1101/2021.12.07.471675

Response to Day et al.

Aschard, H., Vilhjálmsson, B. J., Joshi, A. D., Price, A. L., & Kraft, P. (2016) · The American Journal of Human Genetics · doi:10.1016/j.ajhg.2015.12.020 · Cited by 1

Adjustment for covariates using summary statistics of genome‐wide association studies

Wang, T., Xue, X., Xie, X., Ye, K. Q., Zhu, X., & Elston, R. C. (2018) · Genetic Epidemiology · doi:10.1002/gepi.22148 · Cited by 6

The Covariate's Dilemma

Mefford, J., & Witte, J. S. (2012) · PLoS Genetics · doi:10.1371/journal.pgen.1003096 · Cited by 60