Portrait photo of Ben Greenwood
Associate Professor
Behavioral/Cognitive Neuroscience

Mailing Address:
Department of Psychology 
Campus Box 173, PO Box 173364 
Denver, CO 80217-3364

Physical Location:
North Classroom Building
1200 Larimer Street
Room 5005F (5th floor)

Office Hours:
By appointment

Expertise Areas:
Behavioral Neuroscience Exercise Neuroscience

PhD, Neuroscience and Integrative Physiology, University of Colorado Boulder, 2005
MS, Integrative Physiology, University of Colorado Boulder, 2002
BA, Psychology and Integrative Physiology, University of Colorado Boulder, 1999

Research in the Greenwood lab investigates the neural mechanisms underlying effects of exercise on stress and learning and memory of traumatic events; neural circuits controlling initiation and maintenance of voluntary exercise behavior; signals by which the experience of exercise is communicated to the brain to result in exercise-induced changes in brain and behavior; and sex differences in the effects of exercise. We are also interested in the role of dopamine in fear extinction.

Out of 59 publications. *UCD graduate or undergraduate student trainees are marked with an asterisk.

Bouchet CA*, Miner MA*, Loetz EC, Rosberg AJ*, Hake HS*, Farmer CE*, Ostrovskyy M*, Gray N*, Greenwood BN. Activation of nigrostriatal dopamine neurons during fear extinction prevents the renewal of fear. Neuropsychopharmacology, in press.

Bouchet CA*, Lloyd BA*, Loetz EC, Farmer CE*, Ostrovskyy M*, Haddad N*, Foright RM*, Greenwood BN. Acute exercise enhances the consolidation of fear extinction memory and reduces conditioned fear relapse in a sex-dependent manner. Learning and Memory2017 Jul 17;24(8):358-368.

Lloyd BA*, Hake HS*, Ishiwata T, Farmer CE*, Loetz EC, Fleshner M. Bland ST, Greenwood BN. Exercise increases mTOR signaling in brain regions involved in cognition and emotional behavior. Behavioural Brain Research 2017 Apr 14;323:56-67.

Reber SO, Siebler PH, Donner NC, Morton JT, Smith DG, Kopelman JM, Lowe KR, Wheeler KJ, Fox JH, Hassell JE Jr, Greenwood BN, Jansch C, Lechner A, Schmidt D, Uschold-Schmidt N, Füchsl AM, Langgartner D, Walker FR, Hale MW, Lopez Perez G, Van Treuren W, González A, Halweg-Edwards AL, Fleshner M, Raison CL, Rook GA, Peddada SD, Knight R, Lowry CA. Immunization with a heat-killed preparation of the environmental bacterium Mycobacterium vaccae promotes stress resilience in mice. Proceedings of the National Academy of Sciences USA 2016 May 31;113(22):E3130-9

Nicastro T*, Greenwood BN. Noradrenergic and serotonergic systems are a site of convergence of signals conveying the experience of exercise to brain circuits involved in cognition and emotional behavior. Current Zoology 2016 62(3):293-306.

Herrera JJ, Fedynska S*, Ghasem PR, Wieman T, Clark PJ, Gray N*, Loetz E, Campeau S, Fleshner M, Greenwood BN. Neurochemical and behavioral indices of exercise reward are independent of exercise controllability. European Journal of Neuroscience 2016 43(9):1190-202.

Mika A, Bouchet CA*, Bunker P, Hellwinkel JE, Spence KG, Day HEW, Campeau S, Fleshner M, Greenwood BN. Voluntary exercise during extinction of auditory fear conditioning reduces the relapse of fear associated with potentiated activation of striatal direct pathway neurons. Neurobiology of Learning and Memory 2015 Nov 125:224-235.

Greenwood BN and Fleshner M. Exercise, stress resistance and central serotonin systems. Exercise and Sports Science Reviews 2011 Jul;39(3):140-9.

Rozeske RR, Greenwood BN, Fleshner M, Watkins LR, Maier SF. Voluntary exercise prevents stress-induced potentiation of morphine conditioned place preference. Behavioral Brain Research 2011 Jun 1;219(2):378-81.

Greenwood BN, Foley TE, Le TV, Strong PV, Loughridge AB, Day HEW, Fleshner M. Long term voluntary exercise is rewarding and produces plastic changes in the mesolimbic reward pathway. Behavioral Brain Research 2011 Mar 1;217(2):354-62.

Greenwood BN, Strong PV, Foley TE, Fleshner M. A behavioral analysis of the impact of voluntary physical activity on hippocampal-dependent contextual conditioning. Hippocampus2009 Oct; 19(10):988-1001.

Greenwood BNStrong PV, Foley TE, Thompson RS, Fleshner M. Learned helplessness is independent of levels of brain-derived neurotrophic factor in the hippocampus. Neuroscience2007 Feb 23; 144(4):1193-208.

Greenwood BN, Foley TE, Day HEW, Burhans DBrooks L, Campeau S, Fleshner M. Wheel running alters serotonin (5-HT) transporter, 5-HT1A, 5-HT1B, and alpha 1b-adrenergic receptor mRNA in the rat raphe nuclei. Biological Psychiatry 2005 Mar 1; 57(5):559-68.

Greenwood BN, Foley TE, Day HEW, /campisi J, Hammack SH, Campeau S, Maier SF, Fleshner M. Wheel running prevents learned helplessness / behavioral depression: role of dorsal raphe serotonergic neurons. Journal of Neuroscience 2003 Apr 1; 23(7):2889-98.

PSYC 2220 Biological Basis of Behavior
PSYC 3264 Exercise Brain and Behavior