Benjamin Greenwood, Ph.D.

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, Stress Physiology

Ph.D., Neuroscience and Integrative Physiology, University of Colorado Boulder, 2005
M.S., Integrative Physiology, University of Colorado Boulder, 2002
B.A., Psychology and Integrative Physiology, University of Colorado Boulder, 1999

The Behavioral Exercise Neuroscience Lab uses rodent models to investigate the following:

  1. Mechanisms underlying stress resilience from exercise
  2. Neural circuits governing motivation for voluntary physical activity
  3. Exercise and dopaminergic modulation of fear extinction and relapse
  4. Sex differences therein

Out of 74 publications. *UCD graduate or undergraduate student trainees are marked with an asterisk.

Tanner MK*, Hohorst AA*, Mellert SM*, Loetz EC, Baratta MV, Greenwood BN. Female rats are more responsive than are males to the protective effects of voluntary physical activity against the behavioral consequences of inescapable stress. Stress. 2023 Nov;26(1):2245492. doi: 10.1080/10253890.2023.2245492.

Crombie KM, Adams TG, Dunsmoor JE, Greenwood BN, Smits JA, Nemeroff CB, Cisler JM. Aerobic exercise in the treatment of PTSD: An examination of preclinical and clinical laboratory findings, potential mechanisms, clinical implications, and future directions. J Anxiety Disord. 2023 Mar;94:102680. doi: 10.1016/j.janxdis.2023.102680.

Tanner MK*, Davis JKP*, Jaime J*, Moya NA*, Hohorst AA*, Bonar K*, Abrams KA*, Jamil N*, Han R*, Hubert TJ*, Brown N*, Loetz EC, Greenwood BN. Duration- and sex-dependent neural circuit control of voluntary physical activity. Psychopharmacology (Berl). 2022 Nov;239(11):3697-3709. doi: 10.1007/s00213-022-06243-0.

Moya NA*, Tanner MK*, Smith AM*, Balolia A*, Davis JKP*, Bonar K, Jaime J*, Hubert TJ*, Silva J*, Whitworth W*, Loetz EC, Bland ST, Greenwood BN. Acute exercise enhances fear extinction through a mechanism involving central mTOR signaling. Neurobiol Learn Mem. 2020 Dec;176:107328. doi: 10.1016/j.nlm.2020.107328.

Fallon IP, Tanner MK*, Greenwood BN, Baratta MV. Sex differences in resilience: Experiential factors and their mechanisms. Eur J Neurosci. 2020 Jul;52(1):2530-2547. doi: 10.1111/ejn.14639.

Arnold MR, Greenwood BN, McArthur JA, Clark PJ, Fleshner M, Lowry CA. Effects of repeated voluntary or forced exercise on brainstem serotonergic systems in rats. Behav Brain Res. 2020 Jan 27;378:112237. doi: 10.1016/j.bbr.2019.112237.

Greenwood BN, Fleshner M. Voluntary Wheel Running: A Useful Rodent Model for Investigating the Mechanisms of Stress Robustness and Neural Circuits of Exercise Motivation. Curr Opin Behav Sci. 2019 Aug;28:78-84. doi: 10.1016/j.cobeha.2019.02.001.

Greenwood BN. The role of dopamine in overcoming aversion with exercise. Brain Res. 2019 Jun 15;1713:102-108. doi: 10.1016/j.brainres.2018.08.030.

Hake HS*, Davis JKP*, Wood RR*, Tanner MK*, Loetz EC, Sanchez A*, Ostrovskyy M*, Oleson EB, Grigsby J, Doblin R, Greenwood BN. 3,4-methylenedioxymethamphetamine (MDMA) impairs the extinction and reconsolidation of fear memory in rats. Physiol Behav. 2019 Feb 1;199:343-350. doi: 10.1016/j.physbeh.2018.12.007.

Tanner MK*, Hake HS*, Bouchet CA*, Greenwood BN. Running from fear: Exercise modulation of fear extinction. Neurobiol Learn Mem. 2018 May;151:28-34. doi: 10.1016/j.nlm.2018.03.021.

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. 2018 Feb;43(3):665-672. doi: 10.1038/npp.2017.235.

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. Learn Mem. 2017 Aug;24(8):358-368. doi: 10.1101/lm.045195.117.

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 BN, Strong 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 D, Brooks 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