David J. Mokler, Ph.D.

David J. Mokler, Ph.D.

Professor Emeritus of Pharmacology


Biddeford Campus


Dr. Mokler joined the faculty at UNE in 1986.  He received his Bachelor of Science degree in Psychology from Michigan State University in 1974 and his dual Doctorate in Pharmacology/Toxicology and Neurosciences from Michigan State University in 1984.  

His research focuses on the limbic system of the brain.  He has published widely on the serotonergic system of the brain.  He has worked in the area of prenatal protein malnutrition effects on the brain since 1995.  Recent work has focused on interactions of dopamine, norepinephrine and serotonin in the prefrontal cortex on attention in rats. 



Michigan State University
Michigan State University

Post-Doctoral Training

Post-Doctoral Training, Department of Pharmacology
Medical College of Virginia, Virginia Commonwealth University (Richmond, Virginia)


Current research

How does the adolescent prefrontal cortex differ from the adult prefrontal cortex?; Dietary influences on behavior and brain chemistry in adolescent animals; The effects of prenatal protein malnutrition on the adolescent and adult brain.

Selected publications

Peter, C., Fischer, L.K., Kundakovic, M., Garg, P., Jakovcevski, M., Dincer, A., Amaral, A., Ratner, C., Galdzicka, M., Medford, G., Waber, D., Bryce, C.P., Ginns, E.I., Mokler, D., Champagne, F.A., Rosene, D.L., McGaughy, J.A., Sharp, A.J., Galler, J., Akbarian,S. DNA methylation signatures of early childhood malnutrition associated with impairments in attention and cognition. Biological Psychiatry. 2016 Nov 15;80(10):765-774. doi: 10.1016/j.biopsych.2016.03.2100. Epub 2016 Mar 26. PMID: 27184921; PMCID: PMC5036982.

Fischer, L.K., Bradshaw, S.E., McGaughy, J. A., Fitzmaurice, G., Amaral, A.C., Rosene, D.L., Weissner, W., Mokler, D.J., and Galler, J.R.  Prenatal Protein Level Impacts Homing Behavior in Long-Evans Rat Pups., Nutritional Neuroscience 19(5), 187-195, 2016. PMID: 25603489

Amaral, A.C., Jakovcevski, M., McGaughy, J.A., Calderwood, S.K., Mokler, D.J., Rushmore, R.J., Galler, J.R., Akbarian, S.A., Rosene, D.L. Prenatal protein malnutrition decreases kcnj3 and 2dg activity in rat prefrontal cortex. Neuroscience 286, 79-86, 2015. PMID: 25446346

Duncan, J.R., Garland, M., Stark, R.I., Myers, M.M., Fifer, W.P., Mokler, D.J. and Kinney, H.C. Prenatal nicotine exposure selectively affects nicotinic receptor expression in primary and associative visual cortices of the fetal baboon. Brain Pathology 25(2), 171-81, 2015. PMID: 24903536

McGaughy, J.A., Amaral, A.C., Rushmore, R.J., Mokler, D. J.Morgane, P.J., Rosene, D.L., and Galler, J.R. Prenatal malnutrition leads to deficits in attentional set shifting and decreases metabolic activity in prefrontal subregions that control executive function, Developmental Neuroscience 36(6), 532-41, 2014 PMID: 25342495

Rech, R.H., Mokler, D.J. and Briggs, S.L. Effects of Combined Opioids on Pain and Mood in Mammals. Pain Research and Treatment, in press, online, 2012.

Rech, R.H., Briggs, S.L. and Mokler, D.J. Fentanyl and spiradoline interactions in a place-conditioning black-white shuttle-box. Pharmaceuticals 4: 101-116, 2011.

Staiti, A.M., Morgane, P.J., Galler, J.R., Grivetti, J.Y., Bass, D.C., and Mokler, D.J.  A microdialysis study of the medial prefrontal cortex of adolescent and adult rats.  Neuropharmacology 61(3), 544-549, 2011.

Duncan, J.R., Paterson, D.S., Hoffman, J.M., Mokler, D.J., Borestein, N.S., Belliveau, R.A., Krous, H.F., Haas, E.A., Stanley, C., Nattie, E.E., Trachtenberg, F.L. and Kinney, H.C. Brainstem serotonergic deficiency in the sudden infant death syndrome.  Journal of the American Medical Association, 303 (5): 430-437, 2010.

Mokler, D.J., Dugal, J.R., Hoffman, J.M. and Morgane, P.J. Functional interrelations between nucleus raphé dorsalis and nucleus raphé medianus: A dual probe microdialysis study of glutamate-stimulated serotonin release. Brain Research Bulletin, 78 (4-5): 132-138, 2009.

Morgane, P.J. and Mokler, D.J. (eds.) Limbic Brain: Structure and Function. Special Issue, Neuroscience and Biobehavioral Reviews, 30(2), 2006.

Full NLM bibliography

Mokler DJ, Miller CE, McGaughy JA. Evidence for a role of corticopetal, noradrenergic systems in the development of executive function. Neurobiol Learn Mem. 2017 Sep;143:94-100. doi: 10.1016/j.nlm.2017.02.011. Epub 2017 Feb 17. Review. PubMed PMID: 28219751.

Bove GM, Chapelle SL, Hanlon KE, Diamond MP, Mokler DJ. Attenuation of postoperative adhesions using a modeled manual therapy. PLoS One. 2017;12(6):e0178407. doi: 10.1371/journal.pone.0178407. eCollection 2017. PMID: 28574997; PMCID: PMC5456066.

Bove GM, Chapelle SL, Boyle E, Mokler DJ, Hartvigsen J. A Novel Method for Evaluating Postoperative Adhesions in Rats. J Invest Surg. 2017 Apr;30(2):88-94. doi: 10.1080/08941939.2016.1229367. Epub 2016 Oct 3. PMID: 27690703; PMCID: PMC6347019.

Mokler DJ, McGaughy JA, Bass D, Morgane PJ, Rosene DL, Amaral AC, Rushmore RJ, Galler JR. Prenatal Protein Malnutrition Leads to Hemispheric Differences in the Extracellular Concentrations of Norepinephrine, Dopamine and Serotonin in the Medial Prefrontal Cortex of Adult Rats. Front Neurosci. 2019;13:136. doi: 10.3389/fnins.2019.00136. eCollection 2019. PMID: 30890908; PMCID: PMC6411819.

Newman LA, Baraiolo J, Mokler DJ, Rabinowitz AG, Galler JR, McGaughy JA. Prenatal Protein Malnutrition Produces Resistance to Distraction Similar to Noradrenergic Deafferentation of the Prelimbic Cortex in a Sustained Attention Task. Front Neurosci. 2019;13:123. doi: 10.3389/fnins.2019.00123. eCollection 2019. PubMed PMID: 30853881; PubMed Central PMCID: PMC6396814.

Other scholarly activity


Talylor, L., Gurley, C., Pratt, S., Mokler, D., The Dolphin Brain - A Digital Project, 2020.

Mokler, D.J., Rigdon, A., and Schildroth, S. The Herbal Cabinet. Biomedical Sciences Faculty Projects, 2016. Book 1.

Research interests

The effects of prenatal protein malnutrition on brain development

Research topics

Behavioral Neuroscience
COM Faculty Research
COM Neuroscience and Pain