Adam Boyko, PhD

Adam Boyko

Department of Biomedical Sciences

Associate Professor

Boyko Lab

Department of Biomedical Sciences
Cornell University College of Veterinary Medicine
T8 016A Veterinary Research Tower, Box 17
Ithaca, NY 14853-6401

Office: 607-253-3503
Fax: 607-253-4447

Research Interest

Canine genetics


  • PhD, Biology, Purdue University, 2005
  • MS, Computer Science, Purdue University, 2004
  • BS, Ecology, Ethology & Evolution, University of Illinois at Urbana-Champaign, 1999
  • BS, Computer Science, University of Illinois at Urbana-Champaign, 1999

Biography/Professional Experience

Adam Boyko is an assistant professor in Biomedical Sciences at the Cornell University College of Veterinary Medicine. His research focuses on genomic investigation of dogs as a model of genetic disease and evolutionary genetics. One aspect of this work is understanding the evolution and genetics of village dogs, the semi-feral pariah dogs found in much of the world today. Dr. Boyko is a graduate of the University of Illinois, Urbana-Champaign and received an MS in Computer Science and a PhD in Biology from Purdue University before his postdoctoral research in the Department of Biological Statistics and Computational Biology at Cornell University. He served as a Research Associate in the Genetics Department at the Stanford School of Medicine before beginning his faculty appointment at Cornell in 2011.

Dr. Boyko is a member of Cornell's Center for Vertebrate Genomics, the Center for Comparative and Population Genomics, and the fields of Comparative Biomedical Sciences; Computational Biology; Genetics, Genomics and Development; Molecular and Integrative Physiology and Zoology and Wildlife Conservation. He also serves as Founder and Chief Science Officer of Embark Veterinary, a canine genetic testing company.


  1. Adam H. Freedman, AH, RM Schweizer, DO Vecchyo, E Han, B Davis, I Gronau, PM Silva, M Galaverni, Z Fan, P Marx, B Lorente-Galdos, O Ramirez, F Hormozdiari, C Alkan, C Vilà, K Squire, E Geffen, J Kusak, AR Boyko, HG Parker, C Lee, V Tadigotla, A Siepel, CD Bustamante, TT Harkins, SF Nelson, T Marques-Bonet, EA Ostrander, RK Wayne and J Novembre. 2016 Demographicallybased evaluation of genomic regions under selection in domestic dogs. PLOS Genet, 12, e1005851.
  2. Hayward, JJ, MG Castelhano, KC Oliveira, E Corey, C Balkman, TL Baxter, ML Casal, SA Center, M Fang, SJ Garrison, SE Kalla, P Korniliev, MI Kotlikoff, NS Moise, LM Shannon, KW Simpson, NB Sutter, RJ Todhunter and AR Boyko. 2016. Complex disease and phenotype mapping in the domestic dog. Nat Comm, 7, 10460. Adam R. Boyko 2
  3. Schlamp, F, J van der Made, R Stambler, L Chesebrough, AR Boyko* and PW Messer*. 2016. Evaluating the performance of selection scans to detect selective sweeps in domestic dogs. Mol Ecol, 25, 342–356.
  4. White, ME, JJ Hayward, T Stokol, AR Boyko. 2015 Genetic mapping of novel loci affecting canine blood phenotypes. PLOS One, 10, e0145199.
  5. Shannon, LM, RH Boyko, MG Castelhano, E Corey, JJ Hayward, C McLean, ME White, M Abi Said, BA Anita, NB Ikombe, J Calero, A Galov, M Hedimbi, B Imam, R Khalap, D Lally, A Masta, KC Oliveira, J Randall, NM Tam, C Valeriano, FJT Cornejo, L Pérez, NB Sutter, RJ Todhunter, CD Bustamante and AR Boyko. 2015. Genetic structure in village dogs reveals a Central Asian domestication origin. Proc Natl Acad Sci U S A, 112, 13639–13644
  6. Li, Y, D Wu, AR Boyko, G Wang, S Wu, DM Irwin and Y-P Zhang. 2014. Population variation revealed high altitude adaptation of Tibetan Mastiffs. Mol Biol Evol, 31, 1200-1205.
  7. Boyko, AR, SA Brooks, AL Behan, E Corey, KC Oliveira, RJ Todhunter, Z Zhang, MG Castelhano, DM Ainsworth and NE Robinson. 2014. Genomic analysis establishes correlation between growth and laryngeal neuropathy in Thoroughbreds. BMC Genom, 15, 259.
  8. Freedman, HA, RM Schweizer, I Gronau, E Han, DO Vecchyo, PM Silva, M Galavemi, Z Fan, P Marx, B Lorente-Galdos, H Beale, O Ramierz, F Hormozdiari, C Alkan, C Vilà, K Squire, E Geffen, J Kusak, AR Boyko, H Parker, C Lee, V Tadigotla, A Siepel, CD Bustamante, TT Harkins, SF Nelson, EA Ostrander, T Marques-Bonet, RK Wayne and J Novembre. 2014. Genome sequencing highlights the dynamic early history of dogs. PLOS Genet, 10, e1004016.
  9. Tang, R, HJ Noh, D Wang, S Sigurdsson, R Swofford, M Perloski, M Duxbury, EE Pettersson, J Albright, M Castelhano, A Auton, AR Boyko, G Feng, K Lindblad-Toh and EK Karlsson. 2014. Candidate genes and functional noncoding variants identified in a canine model of obsessive-compulsive disorder. Genom Biol, 15, R25.
  10. Auton, A, Y Rui Li, J Kidd, K Oliveira, J Nadel, JK Holloway, JJ Hayward, PE Cohen, JM Greally, J Wang, CD Bustamante and AR Boyko. 2013. Genetic recombination is targeted towards gene promoter regions in dogs. PLOS Genet, 9, e1003984.
  11. Goldstein, O, J Mezey, P Schweitzer, AR Boyko, C Gao, C Bustamante, J Jordan, G Aguirre and G Acland. 2013. IQCB1 and PDE6B mutations cause similar early onset retinal degenerations in two closely related terrier dog breeds. Invest Ophthalmol Vis Sci, 54, 7005–7019.
  12. Li, Y, BM vonHoldt, G Wang, A Reynolds, AR Boyko, RK Wayne, D Wu and Y Zhang. 2013. Artificial selection on brain expressed genes during the domestication of dog. Mol Biol Evol, 30, 1867–1876.
  13. Schoenebeck, JJ, SA Hutchinson, A Byers, HC Beale, B Carrington, DL Faden, M Rimbault, B Decker, JM Kidd, R Sood, AR Boyko, JW Fondon III, RK Wayne, CD Bustamante, B Ciruna and EA Ostrander. 2012. Variation of BMP3 contributes to dog breed skull diversity. PLOS Genet, 8, e1002849.
  14. Xie, X, J Molina, R Hernandez, A Reynolds, AR Boyko, CD Bustamante and MD Purugganan. 2011. Levels and patterns of nucleotide variation in domestication QTL regions on rice chromosome 3 suggest lineage-specific selection. PLOS One, 6, e20670.
  15. Molina, J, M Sikora, N Garud, JM Flowers, S Rubinstein, A Reynolds, P Huang, SA Jackson, BA Schaal, CD Bustamante, AR Boyko and MD Purugganan. 2011. Reply to Ge and Sang: A single origin of domesticated rice. Proc Natl Acad Sci U S A, 108, e756.
  16. vonHoldt, BM, JP Pollinger, DA Earl, JC Knowles, AR Boyko, H Parker, E Geffen, M Pilot, W Jedrzejewski, B Jedrzejewska, V Sidorovich, C Greco, E Randi, M Musiani, R Kays, CD Bustamante, EA Ostrander, J Novembre and RK Wayne. 2011. A genome-wide perspective on the evolutionary history of enigmatic wolf-like canids. Genom Res, 21, 1294–1305.
  17. Molina, J, M Sikora, N Garud, JM Flowers, S Rubinstein, A Reynolds, P Huang, SA Jackson, BA Schaal, CD Bustamante, AR Boyko and MD Purugganan. 2011. Molecular evidence for a single origin of rice. Proc Natl Acad Sci U S A, 108, 8351–8356.
  18. Boyko, AR. 2011. The domestic dog: man’s best friend in the genomic era. Genom Biol, 12, 216.
  19. Myers, RA, F Casals, J Gauthier, FF Hamdan, J Keebler, AR Boyko, CD Bustamante, AM Piton, D Spiegelman, E Henrion, M Zilversmit, J Hussin, J Quinlan, Y Yang, RG Lafrenière, AR Griffing, EA Stone, GA Rouleau and P Awadalla. 2011. A population genetic approach to mapping neurological disorder genes using deep resequencing. PLOS Genetics, 7, e1001318.
  20. Myles, S, AR Boyko, C Owens, PJ Brown, F Grassi, MK Aradhya, B Prins, A Reynolds, J Chia, D Ware, CD Bustamante and ES Buckler. 2011. Genetic structure and domestication history of the grape. Proc Natl Acad Sci U S A, 108, 3530–3535.
  21. Boyko, AR*, P Quignon*, L Li*, JJ Schoenebeck, JD Degenhardt, KE Lohmueller, K Zhao, A Brisbin, HG Parker, BM vonHoldt, M Cargill, A Auton, A Reynolds, AG Elkahloun, M Castelhano, DS Mosher, NB Sutter, GS Johnson, J Novembre, MJ Hubisz, A Siepel, RK Wayne, CD Bustamante and EA Ostrander. 2010. A simple genetic architecture underlies morphological variation in dogs. PLOS Biol, 8, e1000451.
  22. Awadalla, P, J Gauthier, RA Myers, F Casals, FF Hamdan, AR Griffing, M Côté, E Henrion, D Spiegelman, J Tarabeux, A Piton, Y Yang, AR Boyko, CD Bustamante, L Xiong, É Fombonne, L Mottron, M Zilversmit, J Keebler, H Daoud, C Marineau, MH Roy-Gagnon, MP Dubé, A EyreWalker, P Drapeau, EA Stone, RG Lafrenière, GA Rouleau. 2010. Direct measure of the de novo mutation rate in autism and schizophrenia cohorts. Amer J Hum Genet, 87, 316–324.
  23. Goldstein, O, JG Mezey, AR Boyko, C Gao, W Wang, CD Bustamante, LJ Anguish, JA Jordan, SE Pearce-Kelling, GD Aguirre and GM Acland. 2010. An ADAM9 mutation in canine cone-rod dystrophy 3 establishes homology with human cone-rod dystrophy 9. Mol Vision, 16, 1549–1569.
  24. Williams, LM, X Ma, AR Boyko, CD Bustamante and MF Oleksiak. 2010. SNP identification, verifi- cation, and utility for population genetics in a non-model genus. BMC Genet, 11, 32.
  25. vonHoldt, BM, JP Pollinger, KE Lohmueller, E Han, HG Parker, P Quignon, JD Degenhardt, AR Boyko, DA Earl, A Auton, A Reynolds, K Bryc, A Brisbin, JC Knowles, DS Mosher, TC Spady, A Elkahloun, E Geffen, M Pilot, W Jedrzejewski, C Greco, E Randi, D Bannasch, A Wilton, J Shearman, M Musiani, M Cargill, PG PG Jones, Z Qian, W Huang, Z Ding, Y Zhang, CD Bustamante, EA Ostrander, J Novembre and RK Wayne. 2010. Genome-wide SNP and haplotype analyses reveal a rich history underlying dog domestication. Nature, 464, 898–902.
  26. Andrés, AM, MJ Hubisz, A Indap, DG Torgerson, JD Degenhardt, AR Boyko, RN Gutenkunst, TJ White, ED Green, CD Bustamante, AG Clark and R Nielsen. 2009. Targets of balancing selection in the human genome. Mol Biol Evol, 26, 2755–2764.
  27. Cadieu, E, M Neff, P Quignon, K Walsh, K Chase, HG Parker, BM vonHoldt, A Rhue, AR Boyko, A Byers, A Wong, DS Mosher, A Elkahoun, TC Spady, C André, KG Lark, M Cargill, CD Bustamante, RK Wayne and EA Ostrander. 2009. Coat variation in the domestic dog is governed by variants in three genes. Science, 326, 150–153.
  28. Torgerson, DG, AR Boyko, RD Hernandez, A Indap, X Hu, TJ White, JJ Sninsky, M Cargill, CD Bustamante and AG Clark. 2009. Evolutionary processes acting on candidate cis-regulatory regions in humans inferred from patterns of polymorphism and divergence. PLOS Genet, 5, e1000592.
  29. Boyko, AR, RH Boyko, CM Boyko, P Jones, HG Parker, M Castelhano, L Corey, J Degenhardt, A Auton, M Hedimbi, R Kityo, EA Ostrander, J Schoenebeck, RJ Todhunter and CD Bustamante. 2009. Complex population structure in African village dogs and its implications for inferring dog domestication history. Proc Natl Acad Sci U S A, 106, 13903–13908.
  30. Gray, MM, JM Granka, CD Bustamante, NB Sutter, AR Boyko, L Zhu, EA Ostrander and RK Wayne. 2009. Linkage disequilibrium and demographic history of wild and domestic canids. Genetics, 181, 1493–1505.
  31. Auton, A, K Bryc, AR Boyko, KE Lohmueller, J Novembre, A Reynolds, A Indap, MH Wright, J Degenhardt, RN Gutenkunst, KS King, MR Nelson and CD Bustamante. 2009. Global distribution of genomic diversity underscores rich complex history of continental human populations. Genom Res, 19, 795–803.
  32. Nielsen, R, MJ Hubisz, I Hellmann, D Torgerson, AM Andrés, A Albrechtsen, R Gutenkunst, MD Adams, M Cargill, AR Boyko, A Indap, CD Bustamante and AG Clark. 2009. Darwinian and demographic forces affecting human protein coding genes. Genom Res, 19, 838–849.
  33. Novembre, J, T Johnson, K Bryc, Z Kutalik, AR Boyko, A Auton, A Indap, KS King, S Bergmann, MR Nelson, M Stephens and CD Bustamante. 2008. Genes mirror geography within Europe. Nature, 456, 98–101.
  34. Nelson, MR, K Bryc, KS King, A Indap, AR Boyko, J Novembre, LP Briley, Y Maruyama, DM Waterworth, G Waeber, P Vollenweider, JR Oksenberg, SL Hauser, HA Stirnadel, JS Kooner, JC Chambers, B Jones, V Mooser, CD Bustamante, AD Roses, DK Burns, MG Ehm and EH Lai. 2008. The population reference sample (POPRES): a resource for population, disease, and pharmacological genetics research. Amer J Hum Genet, 83, 347–358.
  35. Blekhman, R, M Oma, L Hermann, AR Boyko, A Indap, C Koisol, CD Bustamante, KM Teshima and M Przeworski. 2008. Natural selection on genes that underlie human disease susceptibility. Curr Biol, 18, 883–889.
  36. Boyko, AR, SH Williamson, AR Indap, JD Degenhardt, RD Hernandez, KE Lohmueller, MD Adams, S Schmidt, JJ Sninsky, SR Sunyaev, TJ White, R Nielsen, AG Clark and CD Bustamante. 2008. Assessing the evolutionary impact of amino acid mutations in the human genome. PLOS Genet, 4, e1000083.
  37. Lohmueller, KE, AR Indap, S Schmidt, AR Boyko, RD Hernandez, MJ Hubisz, JJ Sninsky, TJ White, SR Sunyaev, R Nielsen, AG Clark and CD Bustamante. 2008. Proportionally more deleterious genetic variation in European than in African populations. Nature, 451, 994–998.
  38. Boyko, AR. 2008. Optimal age difference cannot differ between monogamous males and females: a comment on Fieder and Huber. Biol Let, 4, 82.
  39. Caicedo, AL, SH Williamson, RD Hernandez, AR Boyko, A Fledel-Anon, TL York, NR Polato, KM Olsen, R Nielsen, SR McCouch, CD Bustamante and MD Purugganan. 2007. Genome-wide patterns of nucleotide polymorphism in domesticated rice. PLOS Genet, 3, 1745–1756.
  40. Boyko, RH, AR Boyko and MG Boyko. 2007. Referee bias contributes to home-field advantage in English premiership football. J Sports Sci, 25, 1185–1194.
  41. Boyko, AR, RM Gibson and JR Lucas. 2004. How predation risk affects the temporal dynamics of avian leks: sage grouse vs. golden eagles. Am Nat, 163, 154–165.

Browse PubMed Browse PubMed for a complete listing of Dr. Boyko's publications

Awards and Honors

  • Zoetis Award for Veterinary Research Excellence (2015)
  • Genome Technology magazine Young Investigator award (2010)
  • Walter M. Fitch finalist, Society for Molecular Biology and Evolution (2006)
  • H. E. Umbarger Outstanding Graduate Student in Research, Purdue University (2005)
  • Graduate Student Award for Outstanding Teaching, Purdue University (2005)
  • Andrews Fellowship, Purdue University (1999)
  • Chancellor’s Scholar, Campus Honor Program, University of Illinois (1995–1999)
  • James Scholar, College of Engineering, University of Illinois (1995–1999)

Professional/Academic Affiliations

  • 2011-present, Assistant Professor, Biomedical Sciences, Cornell University College of Veterinary Medicine, Ithaca, NY
  • 2015-present, Co-founder and Chief Science Officer, Embark Veterinary, Inc., Ithaca, NY
  • 2009-2011, Research Associate, Genetics, Stanford University School of Medicine, Stanford, CA
  • 2008-2009, Research Associate, Biological Statistics & Computational Biology, Cornell University, Ithaca, NY
  • 2005-2008, Postdoctoral Researcher, Biological Statistics & Computational Biology