Contact Information

  • Lab: +1 301 402 3519
  • Fax: +1 301 480 1952

Overview of the Lab

The overarching mission of the lab is to understand the neural basis of cognitive and emotional regulation and how these critical mental processes are mediated by discrete neural circuits and moderated in function by genetic variation and environmental insults, including stress and alcohol.

Selected Publications


Bjorni M, Rovero NG, Yang ER, Holmes A, Halladay LR. (2020) Phasic signaling in the bed nucleus of the stria terminalis during fear learning predicts within- and across-session cued fear expression. Learning and Memory. bioRxiv.


Halladay LR, Kocharian A, Piantadosi PT, Authement ME, Lieberman AG, Spitz NA, Coden K, Glover LR, Costa VD, Alvarez VA, Holmes A. (2019) Prefrontal regulation of punished ethanol self-administration. Biological Psychiatry. 19(31): 854-2 PubMed (Media: Why is drinking in moderation so difficult for some people? SceinceDaily)

Rovira-Esteban L, Gunduz-Cinar O, Bukalo O, Limoges A, Brockway E, Müller K, Fenno L, Kim SY, Ramakrishnan C, Andrási T, Deisseroth K, Homes A, Hájos N. (2019) Excitation of Diverse Classes of Cholecystokinin Interneurons in the Basal Amygdala Facilitates Fear Extinction. eNeuro.  6(6): 0220-19 PubMed

Sengupta A, Holmes A. (2019) A discrete dorsal raphe to basal amygdala 5-HT circuit calibrates aversive memory. Neuron. 103(3):489-505 PubMed (Previewed by Tiwari P, Fanibunda SE, Vaidya VA. (2019) The Ministry of Fear: 'The Conjuring' of Fright in the Amygdala by the Raphe. Neuron. 103(3):356-358 PubMed)

Zhang WH, Liu WZ, He Y, You WJ, Zhang JY, Xu H, Tian XL, Li BM, Mei L, Holmes A, Pan BX. (2019) Chronic stress causes projection-specific adaptation of amygdala neurons via small-conductance calcium-activated potassium channel downregulation. Biological Psychiatry. 85(10):812-28. PubMed (Previewed by Morel C, Montgomery S, Ming-Hu H. (2019) Small-Conductance, Calcium-Activated Potassium Channels: A Key Circuit Determinant for Stress-Induced Amygdala Dysfunction. Biological Psychiatry. 85(10):784-786. PubMed)

Ferrazzo S, Gunduz-Cinar O, Stefanova N, Pollack GA, Holmes A, Schmuckermair C, Ferraguti F. (2019) Increased anxiety-like behavior following circuit-specific catecholamine denervation in mice. Neurobiology of Disease. 125:55-66. PubMed

Hardaway JA, Halladay LR, Mazzone CM, Pati D, Bloodgood DW, Kim M, Jensen J, DiBerto JF, Boyt KM, Shiddapur A, Erfani A. (2019) Central amygdala prepronociceptin-expressing neurons mediate palatable food consumption and reward. Neuron. PubMed

Holmes A. (2019) Biological clues to an enigmatic treatment for traumatic stress. Nature. 566:335-336. PubMed

Marquardt K, Josey M, Kenton JA, Cavanagh JF, Holmes A, Brigman JL. (2019) Impaired cognitive flexibility following NMDAR-GluN2B deletion is associated with altered orbitofrontal-striatal function. Neuroscience. PubMed

Radke AK, Zweifel LS, Holmes A. (2019) NMDA receptor deletion on dopamine neurons disrupts visual discrimination and reversal learning. Neuroscience Letters. 699:109-114. PubMed

Zhang JY, Liu TH, He Y, Pan HQ, Zhang WH, Yin XP, Tian XL, Li BM, Wang XD, Holmes A, Yuan TF. (2019) Chronic stress remodels synapses in an amygdala circuit–specific manner. Biological Psychiatry. 85(3):189-201. PubMed

Milton AL, Holmes A. (2019) The psychopharmacology of extinction—from theory to therapy. Psychopharmacology. 236(1):1-6. PubMed

Holmes A. (2019) The neuroscience and genomics of social behavior. Genes, Brain and Behavior. 18(1):e12551. PubMed

Cover image

Piantadosi PT, Lieberman AG, Pickens CL, Bergstrom HC, Holmes, A. (2019) A novel multichoice touchscreen paradigm for assessing cognitive flexibility in mice. Learn. Mem. 26:24-30. PubMed

Jan 2019 Learning & Memory Cover

Singewald N, Holmes A. (2019) Rodent models of impaired fear extinction. Psychopharmacology. 236(1):21-32. PubMed

O’Connor RM, McCafferty CP, Bravo JA, Singewald N, Holmes A, Cryan JF. (2019) Increased amygdalar metabotropic glutamate receptor 7 mRNA in a genetic mouse model of impaired fear extinction. Psychopharmacology. 236(1):265-72. PubMed


Delgado-Acevedo C, Estay SF, Radke AK, Sengupta A, Escobar AP, Henríquez-Belmar F, Reyes CA, Haro-Acuña V, Utreras E, Sotomayor-Zárate R, Cho A. (2018) Behavioral and synaptic alterations relevant to obsessivecompulsive disorder in mice with increased EAAT3 expression. Neuropsychopharmacology. 0:1-11. PubMed

Kenton JA, Castillo R, Holmes A, Brigman JL. (2018) Cortico-hippocampal GluN2B is essential for efficient visual-spatial discrimination learning in a touchscreen paradigm. Neurobiology of Learning and Memory. 156:60-67. PubMed

Singewald N, Holmes A. (2018) Rodent models of impaired fear extinction. Psychopharmacology. PubMed

Jury NJ, Radke AK, Pati D, Kocharian A, Mishina M, Kash TL, Holmes A. (2018) NMDA receptor GluN2A subunit deletion protects against dependence-like ethanol drinking. Behav Brain Res.353:124-128. Pubmed

Radke AK, Kocharian A, Covey DP, Lovinger DM, Cheer JF, Mateo Y, Holmes A. (2018) Contributions of nucleus accumbens dopamine to cognitive flexibility. European Journal of Neuroscience. Pubmed

Zhang JY, Liu TH, He Y, Pan HQ, Zhang WH, Yin XP, Tian XL, Li BM, Wang XD, Holmes A, Yuan TF, Pan BX. (2018) Chronic stress remodels synapses in an amygdala circuit-specific manner. Biol Psychiatry. pii: S0006-3223(18)31633-0. doi: 10.1016/j.biopsych.2018.06.019.  Pubmed

O'Connor RM, McCafferty CP, Bravo JA, Singewald N, Holmes A, Cryan JF. (2018) Increased amygdalar metabotropic glutamate receptor 7 mRNA in a genetic mouse model of impaired fear extinction. Psychopharmacology. Pubmed

Lowery-Gionta EG, Crowley NA, Bukalo O, Silverstein S, Holmes A, Kash TL. (2018) Chronic stress dysregulates amygdalar output to the prefrontal cortex. Neuropharmacology.139:68-75. Pubmed

Bergstrom HC, Lipkin AM, Lieberman AG, Pinard CR, Gunduz-Cinar O, Brockway ET, Taylor WW, Nonaka M, Bukalo O, Wills TA, Rubio FJ, Li X, Pickens CL, Winder DG, Holmes A. (2018) Dorsolateral striatum engagement interferes with early discrimination learning. Cell Rep. 23(8):2264-2272. Pubmed

Yuan JT, Gatti DM, Philip VM, Kasparek S, Kreuzman AM, Mansky B, Sharif K, Taterra D, Taylor WM, Thomas M, Ward JO, Holmes A, Chesler EJ, Parker CC (2018) Genome-wide association for testis weight in the diversity outbred mouse population Mamm Genome. 29(5-6):310-324. Pubmed

Bloodgood DW, Sugam JA, Holmes A, Kash TL. (2018) Fear extinction requires infralimbic cortex projections to the basolateral amygdala. Transl Psychiatry.8(1):60. Pubmed

Bukalo O, Holmes A. (2018) Everything in Its Right Place: A Prefrontal-Midbrain Circuit for Contextual Fear Discrimination. Neuron.97(4):732-733. Pubmed

Gunduz-Cinar O, Brockway E, Lederle L, Wilcox T, Halladay LR, Ding Y, Oh H, Busch EF, Kaugars K, Flynn S, Limoges A, Bukalo O, MacPherson KP, Masneuf S, Pinard C, Sibille E, Chesler EJ, Holmes A. (2018) Identification of a novel gene regulating amygdala-mediated fear extinction. Mol Psychiatry.  Pubmed


Jury NJ, Pollack GA, Ward MJ, Bezek JL, Ng AJ, Pinard CR, Bergstrom HC, Holmes A. (2017) Chronic Ethanol During Adolescence Impacts Corticolimbic Dendritic Spines and Behavior. Alcohol Clin Exp Res.41(7):1298-1308. Pubmed

 Alcohol cover

Craske MG, Stein MB, Eley TC, Milad MR, Holmes A, Rapee RM, Wittchen HU. (2017) Anxiety disorders. Nat Rev Dis Primers.3:17024Review. Erratum in: Nat Rev Dis Primers. 2017 Dec 14;3:17100.  Pubmed

Patel S, Hill MN, Cheer JF, Wotjak CT, Holmes A. (2017) The endocannabinoid system as a target for novel anxiolytic drugs. Neurosci Biobehav Rev. 76(Pt A):56-66.  Pubmed

Holmes A. (2017) Sex and Orexins: Uncovering a mechanism underlying sex differences in stress susceptibility. Biol Psychiatry.81(8):642-644. Pubmed

Bogdan R, Salmeron BJ, Carey CE, Agrawal A, Calhoun VD, Garavan H, Hariri AR, Heinz A, Hill MN, Holmes A, Kalin NH, Goldman D. (2017) Imaging genetics and genomics in psychiatry: A critical review of progress and potential. Biol Psychiatry. pii: S0006-3223(17)30034-3. Pubmed

Peterson VL, Jury NJ, Cabrera-Rubio R, Draper LA, Crispie F, Cotter PD, Dinan TG, Holmes A, Cryan JF. (2017) Drunk bugs: Chronic vapour alcohol exposure induces marked changes in the gut microbiome in mice Behav Brain Res.323:172-176. Pubmed

Radke AK, Jury NJ, Delpire E, Nakazawa K, Holmes A. (2017) Reduced ethanol drinking following selective cortical interneuron deletion of the GluN2B NMDA receptor subunit. Alcohol. 58:47-51. Pubmed

Holmes A.(2017) Drug addictions: new insight into causes, comorbidity and potential treatments. Genes Brain Behav.16(1):5-7. Pubmed

Williams RW, Holmes A. (2017) Preface to a special issue on genetic models of alcoholism and alcohol-stress interactions. Alcohol. 58:23-24. Pubmed

Halladay LR, Kocharian A, Holmes A (2017) Mouse strain differences in punished ethanol self-administration. Alcohol. 58:83-92. Pubmed

Lohoff FW, Sorcher JL, Rosen AD, Mauro KL, Fanelli RR, Momenan R, Hodgkinson CA, Vendruscolo LF, Koob GF, Schwandt M, George DT, Jones IS, Holmes A, Zhou Z, Xu MJ, Gao B, Sun H, Phillips MJ, Muench C, Kaminsky ZA. (2017) Methylomic profiling and replication implicates deregulation of PCSK9 in alcohol use disorder. Molecular Psychiatry. Pubmed


Jury NJ, DiBerto JF, Kash TL, Holmes A. (2016) Sex differences in the behavioral sequelae of chronic ethanol exposure. Alcohol. Aug 18. pii: S0741-8329(16)30032-5.  Pubmed

Marcinkiewcz CA, Mazzone CM, D'Agostino G, Halladay LR, Hardaway JA, DiBerto JF, Navarro M, Burnham N, Cristiano C, Dorrier CE, Tipton GJ, Ramakrishnan C, Kozicz T, Deisseroth K, Thiele TE, McElligott ZA, Holmes A, Heisler LK, Kash TL. (2016) Serotonin engages an anxiety and fear-promoting circuit in the extended amygdala. Nature. 537(7618):97-101 Pubmed

Knoll AT, Halladay LR, Holmes AJ, Levitt P (2016) Quantitative Trait Loci and a Novel Genetic Candidate for Fear Learning. J Neurosci.36(23):6258-68. Pubmed (covered by media, e.g. (link is external))

Gamble-George JC, Baldi R, Halladay L, Kocharian A, Hartley N, Silva CG, Roberts H, Haymer A, Marnett LJ, Holmes A, Patel S (2016) Cyclooxygenase-2 inhibition reduces stress-induced affective pathology. Elife. 2016 May 10;5. pii: e14137. doi: 10.7554/eLife.14137. Pubmed

Izquierdo A, Brigman JL, Radke AK, Rudebeck PH, Holmes A (2016) The neural basis of reversal learning: An updated perspective. Neuroscience. Pubmed

Didion JP, Morgan AP, Yadgary L, Bell TA, McMullan RC, Ortiz de Solorzano L, Britton-Davidian J, Bult CJ, Campbell KJ, Castiglia R, Ching YH, Chunco AJ, Crowley JJ, Chesler EJ, Förster DW, French JE, Gabriel SI, Gatti DM, Garland T Jr, Giagia-Athanasopoulou EB, Giménez MD, Grize SA, Gündüz İ, Holmes A, Hauffe HC, Herman JS, Holt JM, Hua K, Jolley WJ, Lindholm AK, López-Fuster MJ, Mitsainas G, da Luz Mathias M, McMillan L, Ramalhinho MD, Rehermann B, Rosshart SP, Searle JB, Shiao MS, Solano E, Svenson KL, Thomas-Laemont P, Threadgill DW, Ventura J, Weinstock GM, Pomp D, Churchill GA, Pardo-Manuel de Villena F (2016) Mol Biol Evol. R2d2 Drives Selfish Sweeps in the House Mouse. (Covered by media, e.g., is external) and is external) ). Pubmed

Hartley ND, Gunduz-Cinar O, Halladay L, Bukalo O, Holmes A, Patel S. (2016)  2-arachidonoylglycerol signaling impairs short-term fear extinction.  Transl Psychiatry  doi: 10.1038/tp.2016.26. Pubmed


Radke AK, Jury NJ, Kocharian A, Marcinkiewcz CA, Lowery-Gionta EG, Pleil KE, McElligott ZA, McKlveen JM, Kash TL, Holmes A. (2015) Chronic EtOH effects on putative measures of compulsive behavior in mice. Addiction Biology  doi: 10.1111/adb.12342. Pubmed

Gunduz-Cinar O, Flynn S, Brockway E, Kaugars K, Baldi R, Ramikie TS, Cinar R, Kunos G, Patel S, Holmes A. (2015) Fluoxetine Facilitates Fear Extinction through Amygdala Endocannabinoids. Neuropsychopharmacology. doi: 10.1038/npp.2015.318. Pubmed

Holmes A, Chen A. (2015) GABA receptors in a state of fear. Nat Neurosci. Pubmed

Radke AK, Nakazawa K, Holmes A (2015) Cortical GluN2B deletion attenuates punished suppression of food reward-seeking. Psychopharmacology.  DOI: 10.1007/s00213-015-4033-8. is external)

Maren S, Holmes A (2015) Stress and Fear Extinction. Neuropsychopharmacology. Pubmed

Brigman JL, Daut RA, Saksida L, Bussey TJ, Nakazawa K, Holmes A  (2015) Impaired discrimination learning in interneuronal NMDA-GluN2B mutant mice. NeuroReport. Pubmed

Holmes A, Jury N, Halladay L, Nakazawa K, Mishina M, Sprengel R, Xu W, Grant S, Svenningsson P, Kiselycznyk C (2015)  NMDA receptor subunits and associated signaling molecules mediating the antidepressant-related effects of NMDA-GluN2B antagonism.  Behavioral Brain Research.  Pubmed

Thomson SM, Josey M, Holmes A, Brigman, J (2015) Conditional loss of GluNR2B in cortex and hippocampus impairs attentional set formation.  Behavioral Neuroscience.  Pubmed

Hariri AR, Holmes A. (2015) Finding translation in stress research. Nature Neuroscience 18(10):1347-1352. Pubmed

Bukalo O, Pinard CR, Silverstein S, Brehm  C, Hartley NH, Whittle N, Colacicco G, Busch E, Patel S, Singewald N, Holmes (2015). Prefrontal inputs to the amygdala instruct fear extinction memory formation. Science Advances 1(6). Pubmed

Fitzgerald PJ, Pinard CR, Camp MC, Feyder M, Sah A, Bergstrom HC, Graybeal C, Liu Y, Schlüter OM, Grant SG, Singewald N, Xu W, Holmes A (2015) Durable fear memories require PSD-95.  Molecular Psychiatry. 20(7):901-912. Pubmed



Singewald N, Schmuckermair C, Whittle N, Holmes A, Ressler KJ (2014) Pharmacology of cognitive enhancers for exposure-based therapy of fear, anxiety and trauma-related disorders. Pharmacology and Therapeutics.  Pubmed

Bukalo O, Pinard C, Holmes A (2014) Mechanisms to medicines: elucidating neural and molecular substrates of fear extinction to identify novel treatments for anxiety disorders.  British Journal of Pharmacology.  Pubmed

Masneuf S, Lowery-Gionta E, Colacicco G, Pleil KE, Li C, Crowley N, Flynn S, Holmes A, Kash T (2014)  Glutamatergic mechanisms associated with stress-induced amygdala excitability and anxiety-related behavior.  Neuropharmacology. Pubmed.

Graybeal C, Bachu M, Mozhui K, Saksida LM, Bussey TJ, Sagalyn E, Williams RW, Holmes A (2014) Strains and stressors: an analysis of touchscreen learning in genetically diverse mouse strains.  PLoS One.  9:e87745. Pubmed.

Radke AK, Pickens CL, Holmes A (2014) “Stress and Alcohol-Drinking Behavior.”  Chapter in Neurobiology of Alcohol Dependence.  6:97-105.

Daut RA, Busch EF, Ihne J, Fisher D, Mishina M, Grant S, Camp M, Holmes A (2014) Tolerance to ethanol intoxication after chronic ethanol: role of GluN2A and PSD-95.  Addiction Biology. Pubmed.

Fitzgerald PJ, Whittle N, Flynn SM, Graybeal C, Pinard C, Gunduz-Cinar O, Kravitz A, Singewald N, Holmes A (2014) Pre-frontal single unit firing associated with deficient extinction in mice.  Neurobiology of Learning and Memory.  Pubmed

Neurobiology of Learning and Memory-cover


Wellman CL, Camp M, Jones VM, Macpherson KP, Ihne J, Fitzgerald P, Maroun M, Drabant E, Bogdan R, Hariri AR, Holmes A (2013) Convergent effects of mouse Pet-1 deletion and human PET-1 variation on amygdala fear and threat processing.  Experimental Neurology.  Pubmed.

Horner AE, Heath CJ, Hvoslef-Eide M, Kent BA, Kim CH, Nilsson SR, Alsiö J, Oomen CA, Holmes A, Saksida LM, Bussey TJ (2013) The touchscreen operant platform for testing learning and memory in rats and mice.  Nature Protocols.  Pubmed

Mar AC, Horner AE, Nilsson S, Alsiö J, Kent BA, Kim CH, Holmes A, Saksida LM, Bussey TJ (2013) The touchscreen operant platform for assessing executive function in rats and mice.  Nature Protocols. Pubmed

Gunduz-Cinar O, Hill MN, McEwen BS, Holmes A (2013) Amygdala FAAH and anandamide: mediating protection and recovery from stress.  Trends in Pharmacological Sciences. Pubmed.  Covered by NPR News: "Could Pot Help Veterans With PTSD?  Brain Scientists Say Maybe” (link is external)


Holmes A, Spanagel R, Krystal JH (2013) Glutamatergic targets for new alcohol medications.  Psychopharmacology Pubmed


Brigman JL, Daut RA, Wright T, Gunduz-Cinar O, Graybeal C, Davis MI, Jiang Z, Saksida LM, Jinde S, Pease M, Bussey TJ, Lovinger DM, Nakazawa K, Holmes A (2013) GluN2B in corticostriatal circuits governs choice learning and choice shifting.  Nature Neuroscience 16:1101-1110.  Pubmed is external)

Maroun M, Ioannides PJ, Bergman KL, Kavushansky A, Holmes A, Wellman CL (2013)  Fear extinction deficits following acute stress associated with increased spine density and dendritic retraction in basolateral amygdala neurons.  European Journal of Neuroscience. 38(4):2611-20Pubmed

Macpherson K, Whittle N, Camp M, Gunduz-Cinar O, Singewald N, Holmes A (2013) Temporal factors in the extinction of fear in inbred mouse strains differing in extinction efficacy.  Biology of Mood & Anxiety Disorders 3:13. Pubmed

Holmes A, Singewald N (2013) Individual differences in recovery from traumatic fear. Trends in Neurosciences 36:23-31. Pubmed

Kiselycznyk C, Zhang X, Huganir R, Holmes A, Svenningsson, P. (2013) Reduced phosphorylation of GluA1 subunits relates to anxiety-like behaviours in mice. International Journal of Neuropsychopharmacology Pubmed


DePoy CL, Daut R, Brigman J, MacPherson KP, Crowley N, Gunduz-Cinar O, Pickens C, Cinar R, Saksida L, Kunos G, Lovinger D, Bussey TJ, Camp MC, Holmes A  (2013) Chronic alcohol produces neuroadaptations to prime learning. Proceedings of the National Academy of Sciences, USA 110:14738-14788 (Covered by various media, e.g., is external), Highlighted ‘In This Issue’ is external)Pubmed

Griebel G, Holmes A (2013) 50 years of hurdles and hope in anxiolytic drug discovery.  Nature Reviews Drug Discovery 12:667-687 Pubmed

Nat rev drug disc-cover

Gunduz-Cinar O, MacPherson KP, Cinar R, Gamble-George J, Sugden K, Williams B, Ramikie TS, Gorka AX, Makriyannis A, Poulton R, Patel S, Hariri AR, Caspi A, Moffitt TE, Kunos G, Holmes A (2013) Convergent translational evidence of a role for anandamide in amygdala-mediated fear extinction, threat processing and stress-reactivity.  Molecular Psychiatry 18:813-823  (Covered by MSNBC, Yahoo News and other media, is external)).  (Faculty 1000 2x ‘must read’ is external))  Pubmed


Debrouse L, Hurd B, Kiselycznyk C, Plitt A, Todaro A, Mishina M, Grant SG, Camp M, Gunduz-Cinar O, Holmes A (2012-Epub ahead of print) Probing the modulation of acute ethanol intoxication by pharmacological manipulation of the NMDAR glycine co-agonist site. Alcoholism: Clinical and Experimental Research. 37(2):223-33. Pubmed

Whittle N, Schmuckermair C, Gunduz Cinar O, Hauschild M, Ferraguti F, Holmes A, Singewald N (2013) Deep brain stimulation, histone deacetylase inhibitors and glutamatergic drugs rescue resistance to fear extinction in a genetic mouse model. Neuropharmacology. 64:414-423. Pubmed

Bussey TJ, Holmes A, Lyon L, Mar AC, McAllister K, Nithianantharajah J, Oomen CA, Saksida LM (2012) New translational assays for preclinical modelling of cognition in schizophrenia: the touchscreen testing method for mice and rats. Neuropharmacology 62:1191-1203. Pubmed

Graybeal C, Kiselycznyk C, Holmes A (2012) Stress-induced impairments in prefrontal-mediated behaviors and the role of the N-methyl-D-aspartate receptor.  Neuroscience 211:28-38. Pubmed

Karlsson RM, Adermark L, Molander A, Pereau-Lenz S, Singley E, Solomon M, Holmes A, Tanaka K, Lovinger DM, Spanagel R, Heilig M (2012) Reduced alcohol intake, lack of alcohol reward and impaired endocannabinoid signaling in mice with a deletion of the glutamate transporter GLAST. Neuropharmacology 63:181-189. Pubmed

Chesler EJ, Plitt A, Fisher D, Hurd B, Lederle L, Kiselycznyk C, Holmes A (2012) Quantitative trait loci for sensitivity to ethanol intoxication in a C57BL/6Jx129S1/SvImJ inbred mouse cross.  Mammalian Genome 23:305-321. Pubmed

Maroun M, Kavushansky A, Holmes A, Wellman C, Motanis H (2012) Enhanced extinction of aversive memories by high-frequency stimulation of the rat infralimbic cortex. PLoS One. 7:e35853. Pubmed

Hartley CA, McKenna MC, Salman R, Holmes A, Casey BJ, Phelps EA, Glatt CE (2012) Serotonin transporter polyadenylation polymorphism modulates the retention of fear extinction memory. Proceedings of the National Academy of Sciences, USA 109:5493-5498. Pubmed

Kiselycznyk C, Hoffman D, Holmes A (2012) Effects of genetic deletion of the Kv4.2 voltage-gated potassium channel on murine anxiety-, fear- and stress-related behaviors. Biology of Mood and Anxiety Disorders 2:5. Pubmed

Ihne J, Fitzgerald PF, Hefner K, Holmes A (2012) Pharmacological manipulation of stress-induced C57BL/6J anxiety-related behavior. Neuropharmacology 62:464-473 Pubmed

Camp MC, Macpherson KP, Lederle L, Graybeal C, Gaburro S, Debrouse LM, Ihne JL, Bravo JA, O'Connor RM, Ciocchi S, Wellman CL, Lüthi A, Cryan JF, Singewald N, Holmes A (2012) Genetic strain differences in learned fear inhibition associated with variation in neuroendocrine, autonomic, and amygdala dendritic phenotypes. Neuropsychopharmacology 37:1534-1547. Pubmed (Faculty 1000 ‘exceptional’ paper is external)).

Barkus C, Feyder M, Graybeal C, Wright T, Wiedholz L, Izquierdo A, Kiselycznyk C, Schmitt W, Sanderson DJ, Rawlins JN, Saksida LM, Bussey TJ, Sprengel R, Bannerman D, Holmes A (2012) Do GluA1 knockout mice exhibit behavioral abnormalities relevant to the negative or cognitive symptoms of schizophrenia and schizoaffective disorder? Neuropharmacology 62:1263-1272. Pubmed

Holmes A, Fitzgerald PJ, MacPherson K, DeBrouse L, Colacicco G, Flynn S, Masneuf S, Pleil KE, Li C, Marcinkiewcz CA, Kash TL, Gunduz-Cinar O, Camp M (2012) Chronic alcohol remodels prefrontal neurons and disrupts NMDA receptor-mediated fear extinction encoding. Nature Neuroscience 15:1359-1361 (Covered by media, e.g., is external)). (Faculty 1000 ‘must read’ is external)).  Pubmed


Kiselycznyk C, Svenningsson P, Delpire E, Holmes A (2011) Genetic, pharmacological and lesion analyses reveal a selective role for corticohippocampal GLUN2B in a novel repeated swim stress paradigm. Neuroscience 193:259-268. Pubmed

Camp M, Feyder M, Ihne J, Palachick B, Hurd B, Chen Y-C, Coba MP, Grant SGN, Holmes A (2011) A novel role for PSD-95 in mediating ethanol intoxication, drinking and place preference.  Addiction Biology 16:428-439. Pubmed

Lederle L, Weber S, Wright T, Feyder M, Brigman JL, Crombag H, Saksida LM, Bussey TJ, Holmes A (2011) Reward-related behavioral paradigms for addiction research in the mouse: performance of common inbred strains. PLoS One 6: e15536. Pubmed

Graybeal C, Schulman E, Saksida LM, Bussey TJ, Feyder M, Holmes A (2011) Paradoxical reversal learning enhancement by stress or prefrontal cortical damage: rescue with BDNF.  Nature Neuroscience 14:1507-1509. (Faculty 1000 ‘recommended’ paper is external))  Pubmed


Brigman JL, Wright T, Talani G, Prasad-Mulcare S, Jinde S, Seabold GK, Mathur P, Davis MI, Bock R,. Alvarez VA, Nakazawa K, Delpire E, Lovinger DM, Holmes A. (2010) Deletion of NR2B-containing NMDA receptors in corticohippocampal principle neurons impairs long-term depression and learning. The Journal of Neuroscience 30:4590-4600. Pubmed (Press release:

Holmes A, Quirk GJ (2010) Pharmacological facilitation of fear extinction and the search for adjunct treatments for anxiety disorders - the case of yohimbine. Trends in Pharmacological Sciences 31:2-7. Pubmed

Holmes A, Wellman CL (2010) Stress-induced prefrontal reorganization and executive dysfunction. Neuroscience & Biobehavioral Reviews 33:773-783. Pubmed

Feyder M, Karlsson RM, Mathur P, Lyman M, Bock R, Momenan R, Munasinghe J, Scattoni ML, Ihne J, Camp M, Graybeal C, Strathdee D, Begg A, Alvarez VA, Kirsch P, Rietschel M, Cichon S, Walter H, Meyer-Lindenberg A, Grant SG, Holmes A (2010) Association of mouse Dlg4 (PSD-95) gene deletion and human DLG4 gene variation with phenotypes relevant to autism spectrum disorders and Williams' syndrome. American Journal of Psychiatry 167:1508-1517. Pubmed

Baganz N, Horton R, Martin K, Holmes A, Daws LC (2010) Repeated swim impairs serotonin clearance via a corticosterone-sensitive mechanism: organic cation transporter 3, the smoking gun. Journal of Neuroscience 30:15185-15195. Pubmed

Mozhui K, Karlsson RM, Kash TL, Ihne J, Norcross M, Fitzgerald PF , Patel S, Camp M, Cameron HA, Ciobanu DC, Winder DG, Williams RW, Holmes A (2010) Neuronal and molecular correlates of trait and stress-induced variation in anxiety-like behavior between mouse strains. The Journal of Neuroscience 30:5357-5367. Pubmed

Fitzgerald PJ, Barkus C, Feyder M, Wiedholz LM, Chen YC, Karlsson RM, Machado-Vieira R, Graybeal C, Sharp T, Zarate C, Harvey-White J, Du J, Sprengel R, Gass P, Bannerman D, Holmes A. (2010) Does gene deletion of AMPA GluA1 phenocopy features of schizoaffective disorder? Neurobiology of Disease 40:608-21. Pubmed

Whittle N, Hauschild M, Lubec G, Holmes A, Singewald N (2010) Rescue of impaired fear extinction and normalization of cortico-amygdala circuit dysfunction in a genetic mouse model by dietary zinc restriction. Journal of Neuroscience 30:13586-13596. Pubmed

Brigman J, Harvey-White J, Izquierdo A, Saksida LM, Bussey TJ, Murphy DL, Deneris E, Holmes A (2010) Pharmacological or genetic inactivation of the serotonin transporter improves reward-based cognitive flexibility on a mouse touchscreen-based assay. Cerebral Cortex 20:1955-1963. Pubmed

Fraser L, Brown RE, Hussin A, Fontana M, Whittaker A, O’Leary T, Lederle L, Holmes A, Ramos A (2010) Measuring locomotion and anxiety-related behaviours in mice: a new way of using old models. Psychopharmacology 211:99-112. Pubmed

Feyder M, Camp MC, Holmes A, Chen Y-C (2010) Ethanol and NMDA receptor interactions: Implications for pharmacotherapeutic treatments. Journal of Medical Sciences 30:3-10. Journal Website(link is external)

Caspi A, Moffit TE, Holmes A, Hariri AR (2010) Gene x environment studies in psychiatry. American Journal of Psychiatry 167:509-527. Pubmed


Camp M, Norcross M, Whittle N, Feyder M, D’Hanis W, Yilmazer-Hanke D, Singewald N, Holmes A (2009) Impaired Pavlovian fear extinction is common across genetic lineages of the 129 inbred mouse strain. Genes, Brain & Behavior 8:744-752. Pubmed

Karlsson RM, Tanaka K, Saksida LM, Bussey TJ, Heilig M, Holmes A (2009) Assessment of glutamate transporter GLAST (EAAT1)-deficient mice for phenotypes relevant to the negative and executive/cognitive symptoms of schizophrenia. Neuropsychopharmacology 34:1578-1589. Pubmed

Zhou R, Holmes A, Du J, Malkesman O, Yuan P, Wang Y, Damschroder-Williams P, Chen G, Guitart X, Manji HK (2009) Genome-wide gene expression profiling in GluR1 knockout mice: Key role of the calcium signaling pathway in glutamatergically-mediated hippocampal transmission. European Journal of Neuroscience 30:2318-2326. Pubmed

Witkiewitz K, Holmes A, Ray LA, Murphy LG, Richardson HN, Chen YC, McDevitt-Murphy ME, Cruz MT, Roberto M (2009) Young investigator award symposium. Alcohol 43:499-508 Pubmed


Holmes A (2008) Genetic variation in cortico-amygdala serotonin function and risk for stress-related disease. Neuroscience & Biobehavioral Reviews 32:1293-1314. Pubmed

Karlsson RM, Tanaka K, Heilig M, Holmes A (2008) Loss of glial glutamate and aspartate transporter (excitatory amino acid transporter 1) causes locomotor hyperactivity and exaggerated responses to psychotomimetics: rescue by haloperidol and metabotropic glutamate 2/3 agonist. Biological  Psychiatry. 64:810-814. Pubmed (For example of media coverage of this article, see (link is external)).

Hefner K, Whittle N, Juhasz J, Norcross M, Karlsson RM, Saksida LM, Bussey TJ, Singewald N, Holmes A (2008) Impaired fear extinction learning and cortico-amygdala circuit abnormalities in a common genetic mouse strain. Journal of Neuroscience. 28:8074-8085. Pubmed  (Journal Club Article : Sotres-Bayon F, Corcoran KA, Peters J, Sierra-Mercado D (2008) Neural correlates of individual variability in fear extinction. The Journal of Neuroscience 28:1-3. Pubmed).

Wiedholz LM, Owens WA, Horton RE, Feyder M, Hefner KR, Sprengel R, Celikel T, Daws LC, Holmes A (2008) Mice lacking the AMPA GluR1 receptor exhibit hyperdopaminergia and schizophrenia-related behaviors. Molecular Psychiatry 13:631-640. Pubmed . (Commentary: Fisher Y, Andre VM, Cepeda C, Levine MS (2008) Dopamine-glutamate interactions at the forefront of schizophrenia research. Cell Science Reviews 5:7-16 Journal website (link is external)).

Chen YC, Holmes A (2008) Effects of topiramate and other anti-glutamatergic drugs on the negative intoxicating actions of ethanol in mice: modulation by genetic strain and stress. Neuropsychopharmacology 34:1454-1466. Pubmed

Crusio WE, Goldowitz D, Holmes A, Wolfer D (2008) Standards for the publication of mouse mutant studies. Genes, Brain & Behavior 8:1-4. Pubmed

Murphy, DL, Fox MA, Timpano KR, Moya P, Ren-Patterson R, Andrews A, Holmes, A, Watts SW, Wendland J, Lesch KP (2008) How the serotonin story is being rewritten by new gene-based discoveries principally related to SLC6A4, the serotonin transporter gene, whose functions influence the entire serotonin system. Neuropharmacology 55:932-960 . Pubmed

Mathur P, Graybeal C, Feyder M, Holmes A (2008) Effects of systemic treatment with the NMDA NR2B subunit antagonist, Ro 25-6981, on mouse fear memory: attenuation with ageing. Pharmacology, Biochemistry & Behavior 91:453-460 . Pubmed

Brigman J, Feyder M, Saksida LM, Bussey TJ, Mishina M, Holmes A (2008) Impaired discrimination learning in mice lacking the NMDA receptor NR2A subunit. Learning and Memory 15:50-54. Pubmed

Davis AR, Shields AD, Brigman J, Norcross M, McElligott ZA, Holmes A, Winder DG (2008) Alpha2-adrenergic receptor regulation of extinction of cocaine conditioned place-preference. Learning and Memory 15:667-676. Pubmed

Fegley DB, Holmes A, Riordan T, Faber CA, Weiss JR, Ma S, Sleeman M, Usdin TB (2008) Regulation of anxiety and fear by tuberoinfundibular peptide of 39 residues. Genes, Brain & Behavior 7:933-942. Pubmed

Brigman J, Mathur P, Lu L, Williams RW, Holmes A (2008) Genetic relationship between fear- and anxiety-related behaviors in BXD recombinant inbred mice. Behavioural Pharmacology 20:204-209. Pubmed

Yang RJ, Mozhui K, Karlsson RM, Cameron HA, Williams RW, Holmes A (2008) Variation in mouse basolateral amygdala volume is associated with differences in stress reactivity and fear learning. Neuropsychopharmacology. 33:2595-2604. Pubmed

Boyce-Rustay JM, Palachick B, Hefner KM, Chen YC, Karlsson RM, Millstein RA, Harvey-White J, Holmes A (2008) Desipramine potentiation of the acute depressant effects of ethanol: modulation by alpha2-adrenoreceptors and stress. Neuropharmacology 55:803-811. Pubmed

Karlsson RM, Hefner KM, Sibley DR, Holmes A (2008) Constitutive loss of dopamine D1, but not D5, receptor prevents locomotor sensitization to cocaine in mice. Psychopharmacology 200:117-127. Pubmed

Norcross M, Mathur P, Enoch AJ, Karlsson RM, Brigman JL, Cameron HA, Harvey-White J, Holmes A (2008) Effects of adolescent fluoxetine treatment on fear-, anxiety- or stress-related behaviors in C57BL/6J or BALB/cJ mice. Psychopharmacology 200:413-424. Pubmed

Parker CC, Ponicsan H, Spencer RL, Holmes A, Johnson TE (2008) Restraint stress and exogenous corticosterone differentially alter sensitivity to the sedative-hypnotic effects of ethanol in ILS and ISS mice. Alcohol 42:477-485. Pubmed

Adamec RA, Holmes A, Blundell J (2008) Vulnerability to lasting anxiogenic effects of brief exposure to predator stimuli: sex, serotonin and other factors - relevance to PTSD. Neuroscience & Biobehavioral Reviews 32:1287-1292. Pubmed

Palachick B, Chen Y-C, Enoch AJ, Karlsson RM, Sprengel R, Mishina M, Holmes A (2008) MK-801 potentiation of ethanol intoxication is not dependent upon major NMDAR or AMPA receptor subunits. Alcoholism: Clinical and Experimental Research 32:1479-1492. Pubmed

Feyder M, Wiedholz LM, Sprengel R, Holmes A (2008) Impaired associative fear learning following complete or partial loss of AMPA GluR1 receptors in mice. Frontiers in Behavioral Neuroscience 1:4. Journal website(link is external)

Tonelli LH, Holmes A, Postlache TT (2008) Intranasal immune challenge induces sex-dependent depressive-like behavior and cytokine expression in the brain. Neuropsychopharmacology 33:1038-1048. Pubmed

Karlsson RM, Holmes A, Crawley, JN, Heilig, MA (2008) The neuropeptide Y Y1 receptor subtype is necessary for the anxiolytic-like but not antidepressant-like effects of neuropeptide Y. Psychopharmacology 195:547-557. Pubmed

Boyce-Rustay JM, Janos A, Holmes A (2008) Effects of chronic swim stress on EtOH-related behaviors in C57BL/6J, DBA/2J and BALB/cByJ mice. Behavioural Brain Research 188:133-137. Pubmed

Holmes A, Stein, MB, Steckler T (2008) Modeling anxiety in mice. In Animal and translational models of psychiatric disorders. RA McArthur and F Borsini (Eds). Elsevier, London.


Fox MA, Andrews AM, Wendland JR, Lesch KP, Holmes A, Murphy DL (2007) A pharmacological analysis of mice with a targeted disruption of the serotonin transporter. Psychopharmacology 195:147-166. Pubmed

Hefner KM, Holmes A (2007) Short-term and long-term effects of exposure to an unfamiliar male in C57BL/6J mice. Behavioural Brain Research 182:344-348. Pubmed

Hefner KM, Holmes A (2007) an investigation of the behavioral effects of ethanol across adolescence in mice. Psychopharmacology 191:311-322. Pubmed

Hasnie FS, Wallace VC, Hefner K, Holmes A, Rice AS (2007). Mechanical and cold hypersensitivity in nerve-injured C57BL/6J mice is not associated with fear-avoidance- and depression-related behaviour. British Journal of Anaesthesia 98:816-822. Pubmed

Boyce-Rustay JM, Cameron HA, Holmes A (2007) Chronic swim stress alters sensitivity to acute behavioral effects of ethanol in mice. Physiology & Behavior, 91:77-86. Pubmed

Fisch GS, Holmes A (2007) Recent developments in the use of animal models of psychiatric disease. Behavior Genetics 37:259-263. Pubmed

Holmes A (2007) The pharmacology of anxiolysis. In Handbook of Anxiety and Fear. R Blanchard, DC Blanchard, DJ Nutt (Eds). Elsevier, London.

Hefner KM, Holmes A (2007) Short-term and long-term effects of exposure to an unfamiliar male in C57BL/6J mice. Behavioural Brain Research, 191:311-322. Pubmed

Holmes A (2007) Mouse models of anxiety. In Handbook of Behavioral Genetics of the Mouse. WE Crusio, F Sluyter, RT Gerlai (Eds). Cambridge University Press, Cambridge.

Mozhui K, Hamre K, Holmes A, Lu L, Williams RW (2007) Genetic and structural analysis of the mouse basolateral amygdala complex. Behavior Genetics, 37:223-243. Pubmed

Carroll J, Boyce-Rustay JM, Izquierdo A, Millstein RA, Yang RJ, Wiedholz LM, Murphy DL, Holmes A (2007) Effects of mild early life stress on abnormal emotional behaviors in 5-HTT knockout mice. Behavior Genetics, 37:214-222. Pubmed

Hefner KM, Holmes A (2007) Ontogeny of fear-, anxiety- and depression-related behavior across mouse adolescence. Behavioral Brain Research 176:210-215. Pubmed

Millstein RA, Holmes A (2007) Effects of repeated maternal separation on anxiety- and depression-related phenotypes in different mouse strains. Neuroscience & Biobehavioral Reviews 31:3-17. Pubmed

Caldwell HK, Stewart S, Wiedholz LM, Millstein RA, Holmes A, Young SW, Wersinger S (2007) The acute intoxicating effects of ethanol are not dependent on the vasopressin 1a or 1b receptors. Neuropeptides 40:325-337. Pubmed

Wellman CL, Izquierdo A, Garrett JE, Martin KP, Carroll J, Millstein RA, Lesch K-P, Murphy DL, Holmes A (2007) Impaired stress-coping and fear extinction and abnormal corticolimbic morphology in serotonin transporter knockout mice. The Journal of Neuroscience 27:684-691. Pubmed


Holmes A, Cryan JF (2006) Measuring anxiety- and depression-related behaviors in the mouse. In Transgenic and Knockout Models of Neuropsychiatric Disorders. GS Fisch and J Flint (Eds). Humana Press, New Jersey.

Boyce-Rustay JM, Holmes A (2006) Ethanol-related behaviors in mice lacking the NMDA receptor NR2A subunit. Psychopharmacology 187:455-466. Pubmed

Izquierdo A, Wellman CL, Holmes A (2006) Stress causes rapid dendritic retraction in infralimbic cortex and impairs fear extinction in mice. The Journal of Neuroscience 26:5733-5738. Pubmed

Hariri AR, Holmes A (2006) Genetics of emotional regulation: the role of the serotonin transporter in neural function. Trends in Cognitive Sciences 10:182-191. Pubmed

Boyce-Rustay JM, Wiedholz LM, Millstein RA, Carroll J, Murphy DL, Daws LC, Holmes A (2006) Behavioral responses to ethanol in serotonin transporter knockout mice. Alcohol: Clinical and Experimental Research 30:1957-1965. Pubmed

Wrenn CC, Holmes A (2006) Galanin; a novel target for stress-related disorders. Drug News and Perspectives, 26:753-778. Pubmed

Holmes A, Picciotto M (2006) Galanin; a novel target system for stress-related disorders and addiction? Current Drug Targets – CNS & Neurological Disorders 5:225-232. Pubmed

Boyce-Rustay JM, Holmes A (2006) Genetic inactivation of the NMDA NR2A subunit has anxiolytic- and antidepressant-like effects in mice. Neuropsychopharmacology, 31:2405-1414. Pubmed

Ren-Patterson R, Cochran LW, Holmes A, Lesch K-P, Lu B, Murphy DL (2006) Gender-dependent modulation of brain monoamines and anxiety-like behaviors in mice with genetic serotonin transporter and BDNF deficiencies. Molecular and Cellular Biology 26:753-778. Pubmed

Karlsson R, Holmes A (2006) Galanin; a modulator of stress and emotion. Amino Acids, 31:231-239. Pubmed

Millstein RA, Ralph RJ, Yang RJ, Holmes A (2006) Effects of repeated maternal separation on prepulse inhibition of startle across inbred mouse strains. Genes, Brain & Behavior 5:346-354. Pubmed

Izquierdo A, Wiedholz LM, Millstein, RA, Yang, RJ, Bussey TJ, Saksida LM, Holmes A (2006) Genetic and dopaminergic modulation of reversal learning in a touchscreen-based operant procedure for mice. Behavioral Brain Research, 171:181-188. Pubmed

Daws LC, Montañez S, Munn JL, Owens WA, Baganz NL, Boyce-Rustay JM, Millstein RA, Wiedholz LM, Murphy DL, Holmes A (2006) Ethanol inhibits clearance of brain serotonin by a serotonin transporter-independent mechanism. The Journal of Neuroscience 26:6431-6438. Pubmed . (News Highlight: McGown D (2006) Neurophysiology: Under the influence. Nature Reviews Neuroscience 7:593 Journal website (link is external)).

Adamec, RA, Burton, P, Blundell, J, Murphy, DL, Holmes A (2006) Vulnerability to mild predator stress in serotonin transporter knockout mice. Behavioral Brain Research 170:126-140. Pubmed(link is external)

Cryan, JF, Holmes A (2005) The ascent of mouse: advances in modeling human depression and anxiety. Nature Reviews Drug Discovery 4:775-790. Pubmed


Zhao S, Edwards J, Carroll J, Wiedholz L, Millstein RA, Murphy DL, Lanthorn TH, Holmes A (2005) Mutant mice reveal an essential functional role for the C-terminus of the 5-HT transporter. Neuroscience 140:321-334. Pubmed

Kim D-K, Tolliver TJ, Huang SJ, Martin BJ, Andrews AM, Wichems C, Holmes A, Lesch K-P, Murphy DL (2005) Altered serotonin synthesis, turnover and dynamic regulation regions of mice lacking the serotonin transporter. Neuropharmacology 49:798-810. Pubmed

Esaki T, Cook M, Shimoji K, Murphy DL, Sokoloff L, Holmes A (2005) Developmental disruption of serotonin transporter function impairs cerebral responses to whisker stimulation in mice. Proceedings of the National Academy of Sciences USA 102:5582-5587. Pubmed

Holmes A, le Guisquet AM, Vogel E, Millstein RA, Leman S, Belzung C (2005) Early life genetic, epigenetic and environmental factors shaping emotionality in rodents. Neuroscience & Biobehavioral Reviews 29:1335-1346. Pubmed

Rustay NR, Wrenn CC, Kinney JW, Holmes A, Bailey KR, Sullivan T, Harris AP, Long KC, Saavedra MC, Starosta G, Innerfield CE, Yang RJ, Dreiling JL, Crawley JN (2005) Galanin impairs performance on learning and memory tasks: findings from galanin transgenic and GAL-R1 knockout mice. Neuropeptides 39:237-241. Pubmed

Boyce-Rustay JM, Holmes A (2005) Functional roles of NMDA receptor NR2A and NR2B subunits in the acute intoxicating effects of ethanol in mice. Synapse 56:222-225. Pubmed

Ren-Patterson RF, Cochran LW, Holmes A, Sherrill S, Huang S-J, Tolliver T, Lesch, K-P, Lu B, Murphy DL (2005) Loss of BDNF gene allele exacerbates brain monoamine deficiencies and increases stress abnormalities of serotonin transporter knockout mice. Journal of Neuroscience Research 79:756-771. Pubmed

Karlsson RM, Holmes A, Heilig M, Crawley JN (2005) Comparative profiles of centrally-administered neuropeptide Y and galanin in mouse tasks for fear and anxiety-related behaviors. Pharmacology, Biochemistry and Behavior 80:427-436. Pubmed

Holmes A, Li Q, Koenig EA, Gold E, Stephenson D, Yang RY, Crawley JN (2005) Phenotypic assessment of galanin modulation of the acute behavioral effects of antidepressants in mice. Psychopharmacology 178:276-285. Pubmed


Li Q, Holmes A, Ma L, Murphy DL (2004) Medial hypothalamic 5-HT1A receptors may be involved in the regulation of defensive behaviors, but not exploratory anxiety-like behaviors in mice. The Journal of Neuroscience 24:10868-10877. Pubmed

Holmes A, Lachowicz JE, Sibley DR (2004) Functional specificity of dopamine receptor subtypes; recent evidence from knockout mice. Neuropharmacology 47:1117-1134. Pubmed

Wrenn CC, Kinney JW, Marriott LK, Holmes A, Harris AP, Saavedra MC, Innerfield CE, Jacoby AS, Shine J, Wenk G, Iismaa TP, Crawley JN (2004) Learning and memory performance in GAL-R1 null mutant mice. European Journal of Neuroscience 19:1384-1396. Pubmed


Murphy DL, Ren-Patterson R, Holmes A, Hall FS, Uhl G (2003) Experimental gene interaction studies with mutant mice as models for human polygenic traits and disorders. Genes, Brain and Behavior 2:350-364. Pubmed

Holmes A, Yang RJ, Lesch KP, Crawley JN, Murphy DL (2003) Mice lacking the serotonin transporter exhibit 5-HT1A receptor-mediated abnormalities in anxiety-like and exploratory behavior. Neuropsychopharmacology 28:2077-2088. Pubmed

Holmes A, Rodgers RJ (2003) Prior exposure to the elevated plus-maze sensitizes mice to the acute behavioral effects of phenelzine and fluoxetine. European Journal of Pharmacology 459:221-230. Pubmed

Holmes A, Hariri AR (2003) Genetic variation in serotonin transporter function and negative emotionality; placing single gene effects in the context of genetic background and environment. Genes, Brain and Behavior 2:332-335. Pubmed

Holmes A, Kinney JW, Wrenn CC, Li Q, Yang RJ, Ma L, Vishwanath J, Saveedra MC, Innerfield CE, Jacoby AS, Shine J, Iismaa TP, Crawley JN (2003) Galanin GAL-R1 receptor null mutant mice display increased anxiety-like behavior specific to the elevated plus-maze test. Neuropsychopharmacology 28:1031-1044. Pubmed

Holmes A, Murphy DL, Crawley JN (2003) Abnormal behavioral phenotypes of serotonin transporter knockout mice; parallels with human anxiety and depression. Biological Psychiatry 54:953-959. Pubmed

Holmes A, Li Q, Murphy DL, Gold E, Crawley JN (2003) Abnormal anxiety-related behavior in serotonin transporter null mutant mice; the influence of genetic background. Genes, Brain and Behavior 2:365-380. Pubmed

Holmes A, Heilig M, Rupniak, NMJ, Steckler T, Griebel G (2003) Neuropeptides as novel therapeutic targets for stress-related disorders; recent developments. Trends in Pharmacological Sciences 24:580-558. Pubmed


Holmes A, Yang RJ, Crawley JN (2002) Evaluation of an anxiety-related phenotype in galanin overexpressing transgenic mice. Journal of Molecular Neuroscience 18:151-165. Pubmed

Wrenn CC, Marriott L, Kinney JW, Holmes A, Wenk G, Crawley JN (2002) Galanin peptide levels in hippocampus and cortex of galanin overexpressing transgenic mice evaluated for cognitive performance. Neuropeptides 36:413-426. Pubmed

Kinney JW, Starosta G, Holmes A, Wrenn CC, Yang RJ, Harris AP, Long KC, Crawley JN (2002) Deficits in trace cued fear conditioning in galanin-treated rats and galanin-overexpressing transgenic mice. Learning and Memory 9:178-190. Pubmed

Holmes A, Yang RJ, Crawley JN, Murphy DL (2002) Evaluation of antidepressant-related behavioral responses in mice lacking the serotonin transporter. Neuropsychopharmacology 27:914-923. Pubmed

Holmes A, Wrenn CC, Harris AP, Thayer K, Crawley JN (2002) Behavioral profiles of inbred strains on novel, olfactory, spatial and emotional tests for reference memory in mice. Genes, Brain and Behavior 1:55-69. Pubmed

Holmes A, Murphy DL, Crawley JN (2002) Reduced aggression in mice lacking the 5-HT transporter. Psychopharmacology 161:160-167. Pubmed

Williams RW, Dubnau J, Enoch MA, Flaherty L, Sluyter F, Gannon KS, Maxson SC, Riedl CAL, Williams KD, Holmes A, Bolivar VJ, Crusio WE (2002) Hot topics in behavioral and neural genetics. Genes, Brain and Behavior 2:117-130. Pubmed


Steiner RA, Hohmann JG, Holmes A, Wrenn CC, Cadd G, Jureus A, Clifton DK, Luo M, Gutshall M, Ma SY, Mufson EJ, Crawley JN (2001) Galanin overexpression provokes selective cognitive deficits and reduces cholinergic survival in a transgenic mouse model of Alzheimer's disease. Proceedings of the National Academy of Sciences USA 98:4184-4189. Pubmed

Holmes A, Iles JP, Mayell SJ, Rodgers RJ (2001) Prior test experience compromises the anxiolytic efficacy of chlordiazepoxide in the mouse light/dark exploration test. Behavioral Brain Research 122:159-167. Pubmed

Holmes A, Hollon TR, Liu Z, Sibley DR, Dreiling J, Gleason TC, Crawley JN (2001) Behavioral characterization of dopamine D5 receptor null mutant mice. Behavioral Neuroscience 115:1129-1144. Pubmed

Holmes A (2001) Transgenic and gene knockout approaches to the study of anxiety-like behavior in mice. Neuroscience & Biobehavioral Reviews 25:261-273. Pubmed


Holmes A, Crawley JN (2000) Promises and limitations of transgenic and knockout mice in modeling psychiatric symptoms. In Contemporary Issues in Modeling Psychopathology. M Myslobodsky and I Weiner (Eds). Kluwer, Boston.

Holmes A, Parmigiani S, Ferrari PF, Palanza PJ, Rodgers RJ (2000) Behavioral profile of wild mice in the elevated plus-maze test for anxiety. Physiology & Behavior 71:509-516. Pubmed


Rodgers RJ, Haller J, Holmes A, Halasz J, Walton T, Brain PF (1999) Corticosterone response to the plus-maze: high correlation with risk assessment in rats and mice. Physiology & Behavior 68:47-53. Pubmed

Holmes A, Rodgers RJ (1999) Influence of spatial and temporal manipulations on the anxiolytic efficacy of chlordiazepoxide in mice previously exposed to the elevated plus-maze. Neuroscience & Biobehavioral Reviews 23:971-980. Pubmed


Holmes A, Rodgers RJ (1998) Responses of Swiss-Webster mice to repeated plus-maze experience: further evidence for a qualitative shift in emotional state? Pharmacology, Biochemistry & Behavior 60:473-488. Pubmed


Rodgers RJ, Cao BJ, Dalvi A, Holmes A (1997) Animal models of anxiety: an ethological perspective. Brazilian Journal of Medical and Biological Research 30:289-304. Pubmed

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