Boyer Winters

Neuroscience & Applied Cognitive Science
(519) 824-4120 x 52163
MacKinnon Extension

Accepting Graduate Students: 
Accepting New Experiential Learning Students: 

My group investigates the neurobiology of cognition, with an emphasis on learning and memory. Topics of interest include memory acquisition, consolidation, and reconsolidation in rats and mice, as well as cognitive testing in rodent models of human disorders such as Alzheimer's disease and schizophrenia.


Undergraduate (1994-1998): Experimental Psychology (with Dr. Richard Brown), Dalhousie University, Halifax, NS, Canada

Graduate (1998-2001): PhD in Experimental Psychology (with Dr. Barry Everitt and Dr. Trevor Robbins), University of Cambridge, Cambridge, UK

Post-doctoral (2001-2007): With Dr. Timothy Bussey and Dr. Lisa Saksida, University of Cambridge, Cambridge, UK


Mammalian memory is a complex cognitive process involving the integration of disparate types of information. Our research emphasizes the hierarchical and distributed nature of declarative memory (memory for facts and events) in the brain.

A primary interest of the lab is the neural basis of object recognition memory.  We use the spontaneous object recognition paradigm to study the anatomical, pharmacological, and molecular bases of memory acquisition, consolidation, and reconsolidation in rats and mice. Furthermore, we have developed various simple modifications to the basic object recognition task to enable the study of information integration and other cognitive functions in the service of higher-order memories. We use these tasks, in combination with numerous touchscreen-based operant tests, to study the neural bases of cognitive processes in animal models of the normal and diseased brain.

This approach has the potential to provide insight into not only the ‘building blocks' of declarative memory, but also the progressive decline of cognitive function in neurodegenerative diseases such as Alzheimer's disease, in which rather specific memory deficits (e.g., failing to recognise objects such as faces) expand into a marked global amnesia affecting all aspects of autobiographical and factual memory.

Selected Publications

Creighton, S.D., Collett, H.A., Zonneveld, P.M., Pandit, R.A., Huff, A.E., Jardine, K.H., McNaughton, B.L., and Winters, B.D. (2019). Development of an “object category recognition task” for mice: involvement of muscarinic acetylcholine receptors. Behavioral Neuroscience, 133, 527-536.

Mitchnick, K.A., Mendell, A.L., Wideman, C.E., Jardine, K.H., Creighton, S.D., Mueller, A., Choleris, E., MacLusky, N.J., & Winters, B.D. (2019). Dissociable involvement of estrogen receptors in perirhinal cortex-mediated object-place memory in male rats. Psychoneuroendocrinology, 107, 98-108.

Duong, A.T.H., Reitz, C.J., Louth, E.L., Creighton, S.D., Rasouli, M., Zwaiman, A., Kroetsch, J.T., Bolz, S., Winters, B.D., Bailey, C.D.C., & Martino, T.A. (2019). The clock mechanism influences neurobiology and adaptations to heart failure in clock∆19/∆19 mice with implications for circadian medicine. Scientific Reports, 9: 4994; DOI:10.1038/s41598-019-41469-7.

Creighton, S.D., Mendell, A.L., Palmer, D., Kalisch, B.E., MacLusky, N.J., Prado, V. F., Prado, M. A. M., & Winters, B.D. (2019). Dissociable cognitive impairments in two strains of transgenic Alzheimer’s disease mice revealed by a battery of object-based tests. Scientific Reports, 9: 57; DOI:10.1038/s41598-018-37312-0.

Wolter, M., Huff, E., Speigel, T., Winters, B.D., & Leri. F. (2019). Cocaine, nicotine, and their conditioned contexts enhance consolidation of object memory in rats. Learning & Memory, 26, 46-55.

Wideman, C.E., Jardine, K.H., & Winters, B.D. (2018). Involvement of classical neurotransmitter systems in memory reconsolidation: focus on destabilization. Neurobiology of Learning and Memory, 156, 68-79.

Creighton, S.D., Palmer, D., Mitchnick, K.A., & Winters, B.D. (2018). Exploiting novelty and oddity exploratory preferences in rodents to study multisensory object memory and perception. In A. Ennaceur & M. de Souza Silva (Eds), Handbook of Research on Object Novelty Recognition (pp. 103-123). London: Elsevier.

Mitchnick, K.A., Wideman, C., Huff, A.E., Palmer, D., McNaughton, B., & Winters, B.D. (2018). Development of novel tasks for studying view-invariant object recognition in rodents: sensitivity to scopolamine. Behavioral Brain Research, 344, 48-56.

Stiver, M.L., Cloke, J.M., Nightingale, N., Rizos, J., & Winters, B.D. (2017). Linking muscarinic receptor activation to UPS-mediated object memory destabilization: implications for long-term memory modification and storage. Neurobiology of Learning and Memory, 145, 151-164.

Lymer, J., Robinson, A., Winters, B.D., Choleris, E. (2017). Rapid effects of dorsal hippocampal G-protein coupled estrogen receptor on learning in female mice. Psychoneuroendocrinology, 77, 131-140.

Cloke, J.M., Nguyen, R., Chung, B. Y. T., Wasserman, D.I., De Lisio, S., Kim, J.C., Bailey, C.D.C., & Winters, B.D. (2016). A novel multisensory integration task reveals robust deficits in rodent models of schizophrenia: converging evidence for remediation via nicotinic receptor stimulation of inhibitory transmission in the prefrontal cortex. Journal of Neuroscience, 36, 12570-12585. Featured Article.

Chung, B.Y.T., Bignell, W., Jacklin, D.L., Winters, B.D., & Bailey, C.D.C. (2016). Postsynaptic nicotinic acetylcholine receptors facilitate excitation of developing CA1 pyramidal neurons. Journal of Neurophysiology, 116, 2043-2055.

Mitchnick, K.A., Creighton, S.D., van Tiggelen, M., Zaika, O., O’Hara, M., Christen, B., Cloke, J.M., Kalisch, B.E., & Winters, B.D. (2016). Dissociable roles for histone acetyltransferases p300 and PCAF in hippocampus- and perirhinal cortex-mediated object memory. Genes, Brain, and Behavior, 15, 542-557.

Palmer, D., Creighton, S.D., Prado, V.F., Prado, M.A.M., Choleris, E., & Winters, B.D. (2016). Mice deficient for striatal Vesicular Acetylcholine Transporter (VAChT) display impaired short-term but normal long-term object recognition memory. Behavioural Brain Research, 311, 267-278.

Jacklin, D.L., Cloke, J.M., Potvin, A., Garrett, I., & Winters, B.D. (2016). The dynamic multisensory engram: neural circuitry underlying crossmodal object recognition in rats changes with the nature of object experience. Journal of Neuroscience, 36, 1273-1289.

Mitchnick, K.A., Creighton, S.D., O’Hara, M., Kalisch, B.E., & Winters, B.D. (2015). Dissociable contributions of de novo and maintenance DNA methyltransferases to object memory processing in the rat hippocampus and perirhinal cortex. European Journal of Neuroscience, 41, 773-786.

Stiver, M.L., Jacklin, D.L., Mitchnick, K.A, Vicic, N., Carlin, J., O’Hara, M., & Winters, B.D. (2015). Cholinergic manipulations bidirectionally regulate object memory destabilization. Learning & Memory, 22, 203-214.

Jacklin, D.L., Kelly, P., Bianchi, C., MacDonald, T., Traquair, H., & Winters, B.D. (2015). Evidence for a specific role for muscarinic receptors in crossmodal object recognition in rats. Neurobiology of Learning and Memory, 118, 125-132.

Cloke, J.M. & Winters, B.D. (2015). α4β2 nicotinic receptor stimulation of the GABAergic system within the orbitofrontal cortex ameliorates the severe crossmodal object recognition impairment in ketamine-treated rats: implications for cognitive dysfunction in schizophrenia. Neuropharmacology, 90, 42-52.

Sticht, M.A., Jacklin, D.L., Parker, L.A., & Winters, B.D. (2015). Intra-perirhinal cortex administration of the synthetic cannabinoid, HU210, disrupts object recognition memory in rats. NeuroReport, 26, 258-262.

Cloke, J.M., Jacklin, D.L., & Winters, B.D. (2014). The neural bases of crossmodal object recognition in non-human primates and rodents: a review. Behavioural Brain Research, 285, 118-130.

O’Brien, L., Limebeer, C.L., Parker, L.A., & Winters, B.D. (2014). CB1 receptor antagonism in the granular insular cortex or somatosensory area facilitates consolidation of object recognition memory. Neuroscience Letters, 578, 192-196.

Bartko, S.J., Winters, B.D., Saksida, L.M. & Bussey, T.J. (2014). Different roles for M1 and M2 receptors within perirhinal cortex in object recognition and discrimination. Neurobiology of Learning and Memory, 110, 16-26.

Reid, J.M., Jacklin, D.L., & Winters, B.D. (2014). Delineating prefrontal cortex region contributions to crossmodal object recognition in rats. Cerebral Cortex, 24, 2108-2119.

Rkieh, N., Cloke, J.M., Gallagher, N., Winters, B.D., & Leri, F. (2014). Drugs of abuse as memory modulators: a study of cocaine in rats. Psychopharmacology, 231, 2339-2348.

Flavell, C.R., Lambert, E., Winters, B.D., & Bredy, T.W. (2013). Mechanisms governing the reactivation-dependent destabilization of memories and their role in extinction. Frontiers in Behavioral Neuroscience, 7:214. doi: 10.3389/fnbeh.2013.00214

Reid, J.M., Jacklin, D.L., & Winters, B.D. (2012). Crossmodal object recognition in rats with and without multimodal object pre-exposure: no effect of hippocampal lesions. Neurobiology of Learning and Memory, 98, 311-319.

Jacklin, D.L., Goel, A., Clementino, K., Hall, A., Talpos, J.C., & Winters, B.D. (2012). Severe crossmodal object recognition deficits in rats treated sub-chronically with NMDA receptor antagonists are reversed by systemic nicotine: implications for abnormal multisensory integration in schizophrenia. Neuropsychopharmacology, 37, 2322-2331.

Melichercik, A., Elliott, K.S., Bianchi, C., Ernst, S.M., & Winters, B.D. (2012). Nicotinic receptor activation in perirhinal cortex and hippocampus enhances object memory in rats. Neuropharmacology, 62, 2096-2105

Tuerke, K.J., Winters, B.D., & Parker, L.A. (2012). Ondansetron interferes with unconditioned lying-on-belly and acquisition of conditioned gaping induced by LiCl as models of nausea-induced behaviors in rats. Physiology & Behavior, 105, 856-860.

Winters, B.D., Tucci, M.C., Jacklin, D.L., Reid, J.M., & Newsome, J. (2011). On the dynamic nature of the engram: evidence for circuit-level reorganization of object memory traces following reactivation. Journal of Neuroscience, 31, 17719-17728.

McTighe S.M., Cowell, R.A., Winters, B.D., Bussey, T.J., & Saksida, L.M. (2010). Paradoxical false memory for objects after brain damage. Science, 330, 1408-1410.

Bartko, S.J., Cowell, R.A., Winters, B.D., Saksida, L.M. & Bussey, T.J. (2010). Heightened susceptibility to interference in an animal model of amnesia: impairment in encoding, storage, retrieval - or all three? Neuropsychologia, 48, 2987-2997.

Winters, B.D., Bussey, T.J., & Saksida, L.M. (2010). Implications of animal object memory research for human amnesia. Neuropsychologia, 48, 2251-2261.

Winters, B.D. & Reid, J.M. (2010). A distributed cortical representation underlies crossmodal object recognition in rats. Journal of Neuroscience, 30, 6253-6261.

Winters, B.D., Bartko, S.J., Saksida, L.M., & Bussey, T.J. (2010). Muscimol, AP5, or scopolamine infused into perirhinal cortex impairs two-choice visual discrimination learning in rats. Neurobiology of Learning and Memory, 93, 221-228.

Winters, B.D., Tucci, M.C, & DaCosta-Furtado, M. (2009). Older and stronger object memories are selectively destabilized by reactivation in the presence of new information. Learning and Memory, 16, 545-553.

Talpos, J.C., Winters, B.D., Dias, R., Saksida, L.M., & Bussey, T.J. (2009). A novel touchscreen-automated paired-associate learning (PAL) task sensitive to pharmacological manipulation of the hippocampus: a translational rodent model of cognitive impairments in neurodegenerative disease. Psychopharmacology, 205, 157-168.

Bussey, T.J., Padain, T.L., Skillings, E.A., Winters, B.D., Morton, A.J., & Saksida, L.M. (2008). The touchscreen cognitive testing method for rodents: how to get the best out of your rat. Learning and Memory, 15, 516-523.

Winters, B.D., Saksida, L.M. & Bussey, T.J. (2008). Object recognition memory: Neurobiological mechanisms of encoding, consolidation and retrieval. Neuroscience and Biobehavioral Reviews, 32, 1055-1070.

Bartko, S.J., Winters, B.D., Cowell, R.A., Saksida, L.M. & Bussey, T.J. (2007). Perirhinal cortex resolves feature ambiguity in configural object recognition and perceptual oddity tasks. Learning and Memory, 14, 821-832.

Winters, B.D., Bartko, S.J., Saksida, L.M. & Bussey, T.J. (2007). Scopolamine infused into perirhinal cortex improves object recognition memory by blocking the acquisition of interfering object information. Learning and Memory, 14, 590-596.

Bartko, S.J., Winters, B.D., Cowell, R.A., Saksida, L.M. & Bussey, T.J. (2007). Perceptual impairments following perirhinal cortex lesions in rats: zero-delay object recognition and simultaneous oddity discriminations. Journal of Neuroscience, 27, 2548-2559.

Winters, B.D., Saksida, L.M. & Bussey, T.J. (2006). Paradoxical facilitation of object recognition memory after infusion of scopolamine into perirhinal cortex: implications for cholinergic system function. Journal of Neuroscience, 26, 9520-9529.

Winters, B.D. & Bussey, T.J. (2005). Glutamate receptors in perirhinal cortex mediate encoding, retrieval, and consolidation of object recognition memory. Journal of Neuroscience, 25, 4243-4251. Recommended by Faculty of 1000; see

Winters, B.D. & Bussey, T.J. (2005). Removal of cholinergic input to perirhinal cortex disrupts object recognition but not spatial working memory in the rat. European Journal of Neuroscience, 21, 2263-2270.

Winters, B.D. & Bussey, T.J. (2005). Transient inactivation of perirhinal cortex disrupts encoding, retrieval, and consolidation of object recognition memory. Journal of Neuroscience, 25, 52-61.

Forwood, S.E., Winters, B.D., & Bussey, T.J. (2005). Hippocampal lesions that abolish spatial maze performance spare object recognition memory at delays of up to 48 hours. Hippocampus, 15, 347-355.

Winters, B.D., Forwood, S.E., Cowell, R., Saksida, L.M., & Bussey, T.J. (2004). Double dissociation between the effects of peri-postrhinal cortex and hippocampal lesions on tests of object recognition and spatial memory: heterogeneity of function within the temporal lobe. Journal of Neuroscience, 24, 5901-5908.

Winters, B.D. & Dunnett, S.B. (2004). Selective lesioning of the cholinergic septo-hippocampal pathway does not disrupt spatial short-term memory: a comparison with the effects of fimbria-fornix lesions. Behavioral Neuroscience, 118, 546-562.

Winters, B.D., Robbins, T.W., & Everitt, B.J. (2004). Selective cholinergic denervation of the cingulate cortex impairs the acquisition and performance of a conditional visual discrimination in rats. European Journal of Neuroscience, 19, 490-496.

Winters, B.D., Matheson, W.R., McGregor, I.S., & Brown, R.E. (2000). An automated two-choice test of olfactory working memory in the rat: effect of scopolamine. Psychobiology, 28(1), 21-31.

I regularly teach the following courses:

PSYC*2410 - Behavioural Neuroscience I

PSYC*3030 - Neurochemical Basis of Behaviour

PSYC*4470 - Advanced Topics in Behavioural and Cognitive Neuroscience

PSYC*6800 - Neurobiology of Learning (Graduate Seminar)