John K. Belknap, Ph.D. (deceased) (Oregon Health Sciences University)

Postdoctoral Advisor


Everything Jeff knows about genetics he learned from John Belknap.  Although primarily an alcohol researcher, John performed the first real pain-relevant QTL mapping study, of morphine analgesia, in 1992. John Belknap died in 2017.  An obituary Jeff was invited to write for Genes, Brain and Behavior can be found here:


  • Marek, P., Mogil, J.S., Belknap, J.K., Sadowski, B., and Liebeskind, J.C. Levorphanol and swim stress-induced analgesia in selectively bred mice: evidence for genetic commonalties. Brain Res., 608:353-357, 1993.
  • Mogil, J.S., Marek, P., O'Toole, L.A., Helms, M.L., Sadowski, B., Liebeskind, J.C., and Belknap, J.K. Mu-opiate receptor binding is up-regulated in mice selectively bred for high stress-induced analgesia. Brain Res., 653:16-22, 1994.
  • Belknap, J.K., Mogil, J.S., Helms, M.L., O'Toole, L.A., Richards, S.P., and Bergeson, S.E. Genetic mapping to proximal Chromosome 10 of a locus influencing morphine- induced analgesia in C57BL/6 and DBA/2 mouse strains.Life Sci., 57:PL117-124, 1995.
  • Mogil, J.S., Kest, B., Sadowski, B., and Belknap, J.K. Differential genetic mediation of sensitivity to morphine in genetic models of opiate sensitivity: influence of nociceptive assay. J. Pharmacol. Exp. Ther., 276:532-544, 1996.
  • Mogil, J.S., Sternberg, W.F., Marek, P., Sadowski, B., Belknap, J.K. and Liebeskind, J.C. The genetics of pain and pain inhibition. Proc. Natl. Acad. Sci., USA, 93:3048-3055, 1996.
  • Mogil, J.S., Grisel, J.E., Reinscheid, R.K., Civelli, O., Belknap, J.K., and Grandy, D.K. Orphanin FQ is a functional anti-opioid peptide. Neuroscience, 75:333-337, 1996.
  • Mogil, J.S., Grisel, J.E., Zhang, G., Belknap, J.K., and Grandy, D.K. Functional antagonism of mu-, delta- and kappa-opioid antinociception by orphanin FQ.Neurosci. Lett., 214:131-134, 1996.
  • Grisel, J.E., Mogil, J.S., Belknap, J.K., and Grandy, D.K. Orphanin FQ acts as a supraspinal, but not a spinal, anti-opioid peptide. Neuroreport, 7:2125-2129, 1996.
  • Mogil, J.S. and Belknap, J.K. Sex and genotype determine the selective activation of neurochemically-distinct forms of swim stress-induced analgesia.Pharmacol. Biochem. Behav., 56:61-66, 1997.
  • Mogil, J.S., Richards, S.P., O'Toole, L.A., Helms, M.L., Mitchell, S.R., andBelknap, J.K. Genetic sensitivity to hot-plate nociception in DBA/2J and C57BL/6J inbred mouse strains: possible sex-specific mediation by delta2-opioid receptors. Pain, 70:267-277, 1997.
  • Mogil, J.S., Richards, S.P., O'Toole, L.A., Mitchell, S.R., Helms, M.L., Kest, B., and Belknap, J.K. Identification of sex-specific quantitative trait loci mediating non-opioid swim stress-induced analgesia in female mice. J. Neurosci., 17:7995-8002, 1997.
  • Kest, B., McLemore, G.L., Sadowski, B., Mogil, J.S., Belknap, J.K., and Inturrisi, C.E. Acute morphine dependence in mice selectively-bred for high and low analgesia. Neurosci. Lett., 256:120-122, 1998.
  • Kest, B., Jenab, S., Brodsky, M., Sadowski, B., Belknap, J.K., Mogil, J.S., and Inturrisi, C.E. Mu and delta opioid receptor analgesia, binding density, and mRNA levels in mice selectively bred for high and low analgesia. Brain Res., 816:381-389, 1999.
  • Mogil, J.S., Wilson, S.G., Bon, K., Lee, S.E., Chung, K., Raber, P., Pieper, J.O., Hain, H.S., Belknap, J.K., Hubert, L., Elmer, G.I., Chung, J.M., and Devor, M. Heritability of nociception. I. Responses of eleven inbred mouse strains on twelve measures of nociception. Pain, 80:67-82, 1999.
  • Mogil, J.S., Wilson, S.G., Bon, K., Lee, S.E., Chung, K., Raber, P., Pieper, J.O., Hain, H.S., Belknap, J.K., Hubert, L., Elmer, G.I., Chung, J.M., and Devor, M. Heritability of nociception. II. 'Types' of nociception revealed by genetic correlation analysis. Pain, 80:83-93, 1999.
  • Grisel, J.E., Mogil, J.S., Grahame, N.J., Rubinstein, M., Belknap, J.K., Crabbe, J.C., and Low, M. Ethanol oral self-administration is increased in mutant mice with decreased beta-endorphin expression. Brain Res., 835:62-67, 1999.
  • Hain, H., Belknap, J.K., and Mogil, J.S. Pharmacogenetic evidence for the involvement of 5? hydroxytryptamine (serotonin)-1B receptors in the mediation of morphine antinociceptive sensitivity. J. Pharmacol. Exp. Ther., 291:444-449, 1999.
  • Mogil, J.S., Grisel, J.E., Bales, J.R., Hayward, M.D., Rubinstein, M., Belknap, J.K., and Low, M. Disparate spinal and supraspinal opioid antinociceptive responses in beta-endorphin-deficient mutant mice. Neuroscience, 101:709-717, 2000.
  • Bergeson, S.E., Helms, M.L., O'Toole, L.A., Jarvis, M.W., Hain, H.S., Mogil, J.S., and Belknap, J.K. Quantitative trait loci influencing morphine antinociception in four mapping populations. Mamm. Genome, 12:546-553, 2001.
  • Wilson, S.G., Chesler, E.J., Hain, H., Rankin, A.J., Schwarz, J.Z., Call, S.B., Murray, M.R., West, E.E., Teuscher, C., Rodriguez-Zas, S., Belknap, J.K., and Mogil, J.S. Identification of quantitative trait loci for chemical/inflammatory nociception in mice. Pain, 96:385-391, 2002.
  • The Complex Trait Consortium. The nature and identification of quantitative trait loci: a community's view. Nature Genet. Rev., 4:911-916, 2003.

Please provide your name, e-mail address as well as the name and e-mail address of the intended recipient in the spaces provided below:

Your Full Name:
Your Email:
Recipient's Full Name:
Recipient's Email:
Department of Psychology 
McGill University 
1205 Dr. Penfield Avenue, Rm. N7/42
Montreal, QC  H3A 1B1

Tel.  514.398.6085
Fax.  514.398.4896
Lab.  514.398.2742
The Pain Genetics Lab is committed to maintaining our users' privacy on the internet. Any information gathered on this website will be used only for the purposes for which it was collected. In the case of any information gathered without the direct knowledge of the user, such information will not be used to identify the user. The Pain Genetics Lab will never share any information gathered with third parties except in cases where it is reasonable to assume implicit agreement or explicit consent has been given.