RESEARCH PAPERS AND REVIEWS

• Popat AK, Simon RC, Aoyama BB, Wokhlu A, Ehrlich AT, Harwell CC, Margolis EB. Mu opioid receptor mRNA and protein localization across the rat and mouse habenula. J Comp Neurol 2026 June 534(6):e70175 Cited 2 times. PDF
• Margolis EB. A switch in kappa opioid receptor signaling from inhibitory to excitatory induced by stress in a subset of cortically-projecting dopamine neurons. bioRxiv. 2025 Sep 05 bioRxiv
• Liao SC, Kano K, Phanse S, Nguyen M, Margolis E, Fu Y, X Meng J, Moutaoufik MT, Chatterton Z, Saccon T, Broderick K, Aoki H, Simms J, Suteja FX, Sei Y, Huang EJ, McAvoy K, Manfredi G, Halliday G, Babu M, Nakamura K. CHCHD2 mutant mice link mitochondrial deficits to PD pathophysiology Sci Adv. 2025 Nov 14;11(46):eadu0726. Cited 4 times. PDF
• Rademacher K, Doric Z, Haddad D, Mamaligas A, Liao SC, Creed R, Kano K, Chatterton Z, Fu Y, Garcia JH, Vance VM, Sei YJ, Kreitzer A, Halliday G, Nelson AB, Margolis EB, Nakamura K Chronic hyperactivation of midbrain dopamine neurons causes preferential dopamine neuron degeneration. 2025. Cited 12 times. Elife. 2025 Aug 26; 13. PDF
• Varga BR, Bernhard SM, El Daibani A, Zaidi SA, Lam JH, Aguilar J, Appourchaux K, Nazarova AL, Kouvelis A, Shinouchi R, Hammond HR, Eans SO, Weinreb V, Margolis EB, Fay JF, Huang XP, Pradhan A, Katritch V, McLaughlin JP, Majumdar S, Che T. Structure-guided design of partial agonists at an opioid receptor. Nat Commun. 2025 Mar 13;16(1):2518. Cited 18 times. PDF
• Gomes I, Gupta A, Margolis EB, Fricker LD, Devi LA Ketamine and Major Ketamine Metabolites Function as Allosteric Modulators of Opioid Receptors 2024. Mol Pharmacol Oct 17; 106(5):240-252. Cited 23 times. PDF
• Margolis EB, Moulton MG, Lambeth PS, O'Meara MJ The life and times of endogenous opioid peptides: Updated understanding of synthesis, spatiotemporal dynamics, and the clinical impact in alcohol use disorder 2023. Neuropharmacology Mar 1; 225:109376. Cited 25 times. PDF
• Waung MW, Maanum KA, Driscoll JR, O'Brien C, Bryant S, Mansourian KA, Morales M, Barker DJ, Margolis EB, A diencephalic circuit for opioid analgesia but not positive reinforcement 2022. Nat Commun. Feb 9; 13(1):764. Cited 21 times. PDF
• Chakraborty S, DiBerto JF, Faouzi A, Bernhard SM, Gutridge AM, Ramsey S, Zhou Y, Provasi D, Nuthikattu N, Jilakara R, Nelson MNF, Asher WB, Eans SO, Wilson LL, Chintala SM, Filizola M, van Rijn RM, Margolis EB, Roth BL, McLaughlin JP, Che T, Sames D, Javitch JA, Majumdar S A novel mitragynine analog with low efficacy mu-opioid receptor agonism displays antinociception with attenuated adverse effects 2021. J Med Chem. Sep 10 64, 18, 13873-13892. Cited 86 times. PDF
• Miranda-Barrientos J, Chambers I, Mongia S, Liu B, Wang HL, Mateo-Semidey GE, Margolis EB, Zhang S, Morales M Ventral tegmental area GABA, glutamate, and glutamate-GABA neurons are heterogeneous in their electrophysiological and pharmacological properties 2021. European J Neurosci July 54(1)(12):4061-4084. Cited 56 times. PDF
• Margolis EB, Wallace TL, Van Orden LJ, Martin WJ Differential effects of novel kappa opioid receptor antagonists on dopamine neurons using acute brain slice electrophysiology 2020. PLoS One Dec 29;15(12):e0232864. Cited 17 times. PDF
• Driscoll JR, Wallace TL, Mansourian KA, Martin WJ, Margolis EB. Differential Modulation of Ventral Tegmental Area Circuits by the Nociceptin/Orphanin FQ System 2020. eNeuro Oct 19;7(5):ENEURO.0376-19.2020. Cited 21 times. PDF
• Fricker LD, Margolis EB, Gomes I, Devi LA. Five Decades of Research on Opioid Peptides: Current Knowledge and Unanswered Questions 2020. Molecular Pharmacology August 98(2), 96-108. Cited 187 times. PDF
• Gomes I, Sierra S, Lueptow L, Gupta A, Gouty S, Margolis EB, Cox BM, Devi LA. Biased Signaling by Endogenous Opioid Peptides 2020. PNAS May 26;117(21):11820-11828. Cited 151 times. PDF
• Waung MW, Margolis EB, Charbit AR, Fields HL. A Midbrain Circuit that Mediates Headache Aversiveness in Rats 2019. Cell Reports 28, 2739–2747. Cited 37 times. PDF
• Margolis EB, Karkhanis AN. Dopaminergic cellular and circuit contributions to kappa opioid receptor mediated aversion 2019. Neurochemistry International, 129 104504. Cited 71 times. PDF
• Breton JM, Charbit AR, Snyder BJ, Fong PTK, Dias EV, Himmels P, Lock H, Margolis EB. Relative contributions and mapping of ventral tegmental area dopamine and GABA neurons by projection target in rat. 2019. J Comp Neurol. 527(5) 916-941. Cited 154 times. PDF
• Saunders BT, Richard JM, Margolis EB, Janak PH. Dopamine neurons create Pavlovian conditioned stimuli with circuit-defined motivational properties. 2018 Nature Neurosci 21(8) 1072-83. Cited 462 times. PDF
• Margolis EB, Fujita W, Devi LA, Fields HL Two delta opioid receptor subtypes are functional in single ventral tegmental area neurons, and can interact with the mu opioid receptor. Neuropharmacology 2017 Sep 1;123:420-432. PMID: 28645621. Cited 49 times. PDF
• Morales M, Margolis EB. Ventral tegmental area: cellular heterogeneity, connectivity and behaviour. Nat Rev Neurosci. 2017 Jan 05. PMID: 28053327. Cited 1446 times. PDF
• Margolis EB, Fields HL. Mu Opioid Receptor Actions in the Lateral Habenula. PLoS One. 2016; 11(7):e0159097. PMID: 27427945. Cited 74 times. PDF
• Avegno EM, Salling MC, Borgkvist A, Mrejeru A, Whitebirch AC, Margolis EB, Sulzer D, Harrison NL. Voluntary adolescent drinking enhances excitation by low levels of alcohol in a subset of dopaminergic neurons in the ventral tegmental area. Neuropharmacology. 2016 Nov; 110(Pt A):386-95. PMID: 27475082. Cited 31 times. PDF
• Gomes I, Bobeck EN, Margolis EB, Gupta A, Sierra S, Fakira AK, Fujita W, Müller TD, Müller A, Tschöp MH, Kleinau G, Fricker LD, Devi LA. Identification of GPR83 as the receptor for the neuroendocrine peptide PEN. Sci Signal. 2016; 9(425):ra43. PMID: 27117253. PMCID: PMC5147544. Cited 98 times. PDF 
• Fields HL, Margolis EB. Understanding opioid reward. Trends in Neurosciences 2015 38(4):217-225. Cited 547 times. PDF
• Margolis EB, Hjelmstad GO, Fujita W, Fields HL. Direct Bidirectional µ-Opioid Control of Midbrain Dopamine Neurons. J Neurosci. 2014 Oct 29; 34(44):14707-16. PMID: 25355223. Cited 135 times. PDF
• Berthet A, Margolis EB, Zhang J, Hsieh I, Zhang J, Hnasko TS, Ahmad J, Edwards RH, Sesaki H, Hung EJ, Nakamura K. Loss of mitochondrial fission depletes axonal mitochondria in midbrain dopamine neurons. J Neurosci. 2014 Oct 22; 34(43):14304-17. PMID: 25339743. Cited 237 times. PDF
• Mitchell JM, Margolis EB, Coker AR, Allen DC, Fields HL. Intra-VTA Deltorphin, But Not DPDPE, Induces Place Preference in Ethanol-Drinking Rats: Distinct DOR-1 and DOR-2 Mechanisms Control Ethanol Consumption and Reward. Alcohol Clin Exp Res. 2014 Jan; 38(1):195-203. PMID: 24033469. Cited 27 times. PDF
• Hjelmstad GO, Xia Y, Margolis EB, Fields HL. Opioid modulation of ventral pallidal afferents to ventral tegmental area neurons. J Neurosci. 2013 Apr 10; 33(15):6454-9. PMID: 23575843. Cited 151 times. PDF
• Volman SF, Lammel S, Margolis EB, Kim Y, Richard JM, Roitman MF, Lobo MK. New Insights into the specificity and plasticity of reward and aversion encoding in the mesolimbic system. J Neurosci 2013 33(45):17569-76. Cited 236 times. PDF
• Margolis EB, Toy B, Himmels P, Morales M, Fields HL. Identification of rat ventral tegmental area GABAergic neurons. PLoS One. 2012; 7(7):e42365. PMID: 22860119. Cited 273 times. PDF
• Mitchell JM, Margolis EB, Coker AR, Fields HL. Alcohol self-administration, anxiety, and cortisol levels predict changes in delta opioid receptor function in the ventral tegmental area. Behav Neurosci. 2012 Aug; 126(4):515-22. PMID: 22708955. Cited 18 times. PDF
• Margolis EB, Mitchell JM, Hjelmstad GO, Fields HL. A novel opioid receptor-mediated enhancement of GABAA receptor function induced by stress in ventral tegmental area neurons. J Physiol. 2011 Sep 1; 589(Pt 17):4229-42. PMID: 21690191. Cited 33 times. PDF
• Xia Y, Driscoll JR, Wilbrecht L, Margolis EB, Fields HL, Hjelmstad GO. Nucleus accumbens medium spiny neurons target non-dopaminergic neurons in the ventral tegmental area. J Neurosci. 2011 May 25; 31(21):7811-6. PMID: 21613494. Cited 306 times. PDF
• Margolis EB, Coker AR, Driscoll JR, Lemaître AI, Fields HL. Reliability in the identification of midbrain dopamine neurons. PLoS One. 2010; 5(12):e15222. PMID: 21151605. Cited 102 times. PDF
• Xia YF, Margolis EB, Hjelmstad GO. Substance P inhibits GABAB receptor signalling in the ventral tegmental area. J Physiol. 2010 May 1; 588(Pt 9):1541-9. PMID: 20231139. Cited 19 times. PDF
• Dobi A, Margolis EB, Wang HL, Harvey BK, Morales M. Glutamatergic and nonglutamatergic neurons of the ventral tegmental area establish local synaptic contacts with dopaminergic and nondopaminergic neurons. J Neurosci. 2010 Jan 6; 30(1):218-29. PMID: 20053904. Cited 298 times. PDF
• Margolis EB, Fields HL, Hjelmstad GO, Mitchell JM. Delta-opioid receptor expression in the ventral tegmental area protects against elevated alcohol consumption. J Neurosci. 2008 Nov 26; 28(48):12672-81. PMID: 19036960. Cited 111 times. PDF
• Margolis EB, Mitchell JM, Ishikawa J, Hjelmstad GO, Fields HL. Midbrain dopamine neurons: projection target determines action potential duration and dopamine D(2) receptor inhibition. J Neurosci. 2008 Sep 3; 28(36):8908-13. PMID: 18768684. Cited 304 times. PDF
• Fields HL, Hjelmstad GO, Margolis EB, Nicola SM. Ventral tegmental area neurons in learned appetitive behavior and positive reinforcement Annu Rev Neurosci 2007 30:289-316. Cited 890 times. PDF
• Margolis EB, Lock H, Hjelmstad GO, Fields HL. The ventral tegmental area revisited: is there an electrophysiological marker for dopaminergic neurons? J Physiol. 2006 Dec 15; 577(Pt 3):907-24. PMID: 16959856. Cited 697 times. PDF
• Margolis EB, Lock H, Chefer VI, Shippenberg TS, Hjelmstad GO, Fields HL. Kappa opioids selectively control dopaminergic neurons projecting to the prefrontal cortex. Proc Natl Acad Sci U S A. 2006 Feb 21; 103(8):2938-42. PMID: 16477003. Cited 414 times. PDF
• Margolis EB, Hjelmstad GO, Bonci A, Fields HL. Both kappa and mu opioid agonists inhibit glutamatergic input to ventral tegmental area neurons. J Neurophysiol. 2005 Jun; 93(6):3086-93. PMID: 15615834 Topic of "Editorial Focus" in this issue. Cited 112 times. PDF
• Margolis EB, Hjelmstad GO, Bonci A, Fields HL. Kappa-opioid agonists directly inhibit midbrain dopaminergic neurons. J Neurosci. 2003 Nov 5; 23(31):9981-6. PMID: 14602811 "This week in the Journal" feature. Cited 361 times. PDF