Peer-reviewed publications since 2015
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*denotes CU Denver undergraduate or graduate student trainee

1. 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.
2. Kevin M Crombie, Tom G Adams, Joseph E Dunsmoor, Benjamin N Greenwood, Jasper A Smits, Charles B Nemeroff, Josh M Crisler. Aerobic exercise in the treatment of PTSD: An examination of preclinical and clinical laboratory findings, potential mechanisms, clinical implications, and future directions. Journal of Anxiety Disorders, 2023 Mar, 94:102680. doi: 10.1016/j.janxdis.2023.102680.
3. Margaret K Tanner*, Jazmyne K P Davis*, Jennifer Jaime*, Nicolette A Moya*, Alyssa A Hohorst*, Kelsey Bonar*, Kelsey A Abrams*, Nashra Jamil*, Rebecca Han*, Troy J Hubert*, Nadja Brown*, Esteban C Loetz, Benjamin N Greenwood. Duration- and sex-dependent neural circuit control of voluntary physical activity. Psychopharmacology, 2022 Oct 4, doi: 10.1007/s00213-022-06243-0.
4. Dawud LM, Loetz EC, Lloyd B*, Beam R, Tran S, Cowie K, Browne K, Khan T, Montoya R, Greenwood BN, Bland ST. A novel social fear conditioning procedure alters social behavior and mTOR signaling in differentially housed adolescent rats. Developmental Psychobiology, 2021 Jan;63(1):74-87. doi: 10.1002/dev.22001.
5. Moya NA*, Tanner MK*, Smith AM*, Balolia A*, Davis JKP*, Bonar K*, Jaime J*, Hubert T*, Silva J*, Whitworth W*, Loetz EC, Bland ST, Greenwood BN. Acute exercise enhances fear extinction through a mechanism involving central mTOR signaling. Neurobiology of Learning and Memory, 2020 Dec;176:107238. 
6. Foright RM, Johnson GC, Kahn D, Charleston CA, Presby DM, Bouchet CA*, Wellberg EA, Sherk VD, Jackman MR, Greenwood BN, MacLean PS. Compensatory eating behaviors in male and female rats in response to exercise training. American Journal of Physiology, Regul Integr Comp Physiol. 2020 Aug 1;319(2):R171-R183.
7. Fallon IP, Tanner MK*, Greenwood BN, Baratta MV. Sex differences in resilience: Experiential factors and their mechanisms. European Journal of Neuroscience, 2020 Jul;52(1):2530-2547.
8. 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. Behavioral Brain Research​, 2020 Jan 27;378:112237.
9. Tanner MK*, Fallon IP, Baratta MV, Greenwood BN. Voluntary exercise enables stress resistance in females. Behavioral Brain Research​, 2019 2019 Sep 2;369:111923.
10. Greenwood BN, Fleshner M. Voluntary wheel running: a useful rodent model for investigating mechanisms of stress robustness and exercise motivation. Current Opinion in Behavioral Sciences​, 2019 August; 28:78-84. Invited review
11. Greenwood BN. The Role of Dopamine in Overcoming Aversion with Exercise. Brain Research, 2019 Jun 15;1713:102-108. Invited review
12. 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. Physiology and Behavior, 2019 Feb; 199:343-350.
13. Tanner MK*, Hake HS*, Bouchet CA*, Greenwood BN. Running from Fear: Exercise Modulation of Fear Extinction. Neurobiology of Learning and Memory, 2018 May;151:28-34. Invited review
14. Ishiwata T, Greenwood BN. Changes in thermoregulation and monoamine release in freely moving rats during cold exposure and inhibition of the ventromedial, dorsomedial, or posterior hypothalamus. Journal of Comparative Physiology B, 2018 May;188(3):541-551.
15. 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. Figure 1 chosen for February cover.
16. 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 Memory​, 2017 Jul 17;24(8):358-368.
17. Ishiwata T, Hasegawa H, Greenwood BN. Involvement of serotonin in the ventral tegmental area in thermoregulation of freely moving rats. Neuroscience Letters​, 2017 May 17;653:71-77.
18. 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. Behavioral Brain Research 2017 Apr 14;323:56-67.
19. Thompson RS, Roller R, Mika A, Greenwood BN, Knight R, Chichlowski M, Berg B, Fleshner M. Dietary Prebiotics and Bioactive Milk Fractions Improve NREM Sleep, Enhance REM Sleep Rebound and Attenuate the Stress-Induced Decrease in Diurnal Temperature and Gut Microbial Alpha Diversity. Frontiers in Behavioral Neuroscience​, 2017 Jan 10;10:240.
20. Ishiwata T, Oshimoto A, Saito T, Kotani Y, Nomoto S, Aihara Y, Hasegawa H, Greenwood BN. Possible mechanisms of hypothermia after inhibition of the median or dorsal raphe nucleus of freely moving rats. Neuroreport​, 2016 Dec 7;27(17):1287-1292.
21. Mika A, Day HE, Martinez A, Rumian NL, Greenwood BN, Chichlowski M, Berg BM, Fleshner M. Early life diets with prebiotics and bioactive milk fractions attenuate the impact of stress on learned helplessness behaviours and alter gene expression within neural circuits important for stress resistance. European Journal of Neuroscience​, 2016 Feb;45(3):342-357.
22. Grace PM, Fabisiak TJ, Green-Fulgham SM, Anderson ND, Strand KA, Kwilasz AJ, Galer EL, Walker FR, Greenwood BN, Maier SF, Fleshner M, Watkins LR. Prior voluntary wheel running attenuates neuropathic pain. Pain​, 2016 Sep;157(9):2012-23
23. 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 016 May 31;113(22):E3130-9
24. Thompson RS, Roller R, Greenwood BN, Fleshner M. Wheel running improves REM sleep and attenuates stress-induced flattening of diurnal rhythms in F344 rats. Stress​, 2016 May;19(3):312-24.
25. Nicastro T*, Greenwood BN. Central monoaminergic 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. Invited review
26. Herrera JJ, Fedynska S*, Ghasem PR, Wieman T, Clark PJ, Gray N*, Loetz E, Campeau S, Fleshner M, Greenwood BN. Neurochemical and behavioural indices of exercise reward are independent of exercise controllability. European Journal of Neuroscience​, 2016 43(9):1190-202.
27. Clark PJ, Amat J, McConnell SO, Ghasem PR, Greenwood BN, Maier SF, Fleshner M. Running Reduces Uncontrollable Stress-Evoked Serotonin and Potentiates Stress-Evoked Dopamine Concentrations in the Rat Dorsal Striatum. PLoS One​, 2015 Nov 10;10(11).
28. 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.
29. Woodruff ER, Greenwood BN, Chun LE, Fardi S, Hinds LR, Spencer RL. Adrenal-dependent diurnal modulation of conditioned fear extinction learning. Behavioral Brain Research​, 2015 Jun 1;286:249-55.