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Benjamin N Greenwood

TitleAssociate Professor
InstitutionUniversity of Colorado Denver - Denver Campus
DepartmentCLAS-Psychology
Phone303/315-7038

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    Publications listed below are automatically derived from MEDLINE/PubMed and other sources, which might result in incorrect or missing publications. Faculty can login to make corrections and additions.
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    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 11; 26(1):2245492. PMID: 37549016.
      View in: PubMed
    2. 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 03; 94:102680. PMID: 36773486.
      View in: PubMed
    3. 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. PMID: 36195731.
      View in: PubMed
    4. 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. Neurobiol Learn Mem. 2020 12; 176:107328. PMID: 33075479.
      View in: PubMed
    5. 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. Am J Physiol Regul Integr Comp Physiol. 2020 08 01; 319(2):R171-R183. PMID: 32551825.
      View in: PubMed
    6. 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. Dev Psychobiol. 2021 01; 63(1):74-87. PMID: 32524583.
      View in: PubMed
    7. Fallon IP, Tanner MK, Greenwood BN, Baratta MV. Sex differences in resilience: Experiential factors and their mechanisms. Eur J Neurosci. 2020 07; 52(1):2530-2547. PMID: 31800125.
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    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. Behav Brain Res. 2020 01 27; 378:112237. PMID: 31525404.
      View in: PubMed
    9. Tanner MK, Fallon IP, Baratta MV, Greenwood BN. Voluntary exercise enables stress resistance in females. Behav Brain Res. 2019 09 02; 369:111923. PMID: 31028765.
      View in: PubMed
    10. 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. PMID: 32766411.
      View in: PubMed
    11. 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 02 01; 199:343-350. PMID: 30529341.
      View in: PubMed
    12. Greenwood BN. The role of dopamine in overcoming aversion with exercise. Brain Res. 2019 06 15; 1713:102-108. PMID: 30171838.
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    13. Tanner MK, Hake HS, Bouchet CA, Greenwood BN. Running from fear: Exercise modulation of fear extinction. Neurobiol Learn Mem. 2018 05; 151:28-34. PMID: 29614374.
      View in: PubMed
    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. J Comp Physiol B. 2018 05; 188(3):541-551. PMID: 29075844.
      View in: PubMed
    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 02; 43(3):665-672. PMID: 28976945.
      View in: PubMed
    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. Learn Mem. 2017 08; 24(8):358-368. PMID: 28716955.
      View in: PubMed
    17. Ishiwata T, Hasegawa H, Greenwood BN. Involvement of serotonin in the ventral tegmental area in thermoregulation of freely moving rats. Neurosci Lett. 2017 Jul 13; 653:71-77. PMID: 28527719.
      View in: PubMed
    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. Behav Brain Res. 2017 04 14; 323:56-67. PMID: 28130174.
      View in: PubMed
    19. Thompson RS, Roller R, Mika A, Greenwood BN, Knight R, Chichlowski M, Berg BM, 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. Front Behav Neurosci. 2016; 10:240. PMID: 28119579.
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    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 07; 27(17):1287-1292. PMID: 27741215.
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    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. Eur J Neurosci. 2017 02; 45(3):342-357. PMID: 27763700.
      View in: PubMed
    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 09; 157(9):2012-23. PMID: 27355182.
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    23. Reber SO, Siebler PH, Donner NC, Morton JT, Smith DG, Kopelman JM, Lowe KR, Wheeler KJ, Fox JH, Hassell JE, Greenwood BN, Jansch C, Lechner A, Schmidt D, Uschold-Schmidt N, F?chsl AM, Langgartner D, Walker FR, Hale MW, Lopez Perez G, Van Treuren W, Gonz?lez 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. Proc Natl Acad Sci U S A. 2016 May 31; 113(22):E3130-9. PMID: 27185913.
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    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 05; 19(3):312-24. PMID: 27124542.
      View in: PubMed
    25. Nicastro TM, 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. Curr Zool. 2016 Jun; 62(3):293-306. PMID: 29491917.
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    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. Eur J Neurosci. 2016 05; 43(9):1190-202. PMID: 26833814.
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    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; 10(11):e0141898. PMID: 26555633.
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    28. Mika A, Bouchet CA, Bunker P, Hellwinkel JE, Spence KG, Day HE, Campeau S, Fleshner M, Greenwood BN. Voluntary exercise during extinction of auditory fear conditioning reduces the relapse of fear associated with potentiated activity of striatal direct pathway neurons. Neurobiol Learn Mem. 2015 Nov; 125:224-35. PMID: 26454156.
      View in: PubMed
    29. Woodruff ER, Greenwood BN, Chun LE, Fardi S, Hinds LR, Spencer RL. Adrenal-dependent diurnal modulation of conditioned fear extinction learning. Behav Brain Res. 2015 Jun 01; 286:249-55. PMID: 25746455.
      View in: PubMed
    30. Greenwood BN, Thompson RS, Opp MR, Fleshner M. Repeated exposure to conditioned fear stress increases anxiety and delays sleep recovery following exposure to an acute traumatic stressor. Front Psychiatry. 2014; 5:146. PMID: 25368585.
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    31. Clark PJ, Ghasem PR, Mika A, Day HE, Herrera JJ, Greenwood BN, Fleshner M. Wheel running alters patterns of uncontrollable stress-induced cfos mRNA expression in rat dorsal striatum direct and indirect pathways: A possible role for plasticity in adenosine receptors. Behav Brain Res. 2014 Oct 01; 272:252-63. PMID: 25017571.
      View in: PubMed
    32. Thompson RS, Strong PV, Clark PJ, Maslanik TM, Wright KP, Greenwood BN, Fleshner M. Repeated fear-induced diurnal rhythm disruptions predict PTSD-like sensitized physiological acute stress responses in F344 rats. Acta Physiol (Oxf). 2014 Jun; 211(2):447-65. PMID: 24447583.
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    33. Fleshner M, Greenwood BN, Yirmiya R. Neuronal-glial mechanisms of exercise-evoked stress robustness. Curr Top Behav Neurosci. 2014; 18:1-12. PMID: 24481547.
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    34. Speaker KJ, Cox SS, Paton MM, Serebrakian A, Maslanik T, Greenwood BN, Fleshner M. Six weeks of voluntary wheel running modulates inflammatory protein (MCP-1, IL-6, and IL-10) and DAMP (Hsp72) responses to acute stress in white adipose tissue of lean rats. Brain Behav Immun. 2014 Jul; 39:87-98. PMID: 24246250.
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    35. Christianson JP, Greenwood BN. Stress-protective neural circuits: not all roads lead through the prefrontal cortex. Stress. 2014 Jan; 17(1):1-12. PMID: 23574145.
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    36. Loughridge AB, Greenwood BN, Day HE, McQueen MB, Fleshner M. Microarray analyses reveal novel targets of exercise-induced stress resistance in the dorsal raphe nucleus. Front Behav Neurosci. 2013; 7:37. PMID: 23717271.
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    37. Thompson RS, Christianson JP, Maslanik TM, Maier SF, Greenwood BN, Fleshner M. Effects of stressor controllability on diurnal physiological rhythms. Physiol Behav. 2013 Mar 15; 112-113:32-9. PMID: 23454291.
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    38. Maslanik T, Tannura K, Mahaffey L, Loughridge AB, Beninson L, Benninson L, Ursell L, Greenwood BN, Knight R, Fleshner M. Commensal bacteria and MAMPs are necessary for stress-induced increases in IL-1? and IL-18 but not IL-6, IL-10 or MCP-1. PLoS One. 2012; 7(12):e50636. PMID: 23236381.
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    39. Greenwood BN, Spence KG, Crevling DM, Clark PJ, Craig WC, Fleshner M. Exercise-induced stress resistance is independent of exercise controllability and the medial prefrontal cortex. Eur J Neurosci. 2013 Feb; 37(3):469-78. PMID: 23121339.
      View in: PubMed
    40. Maslanik T, Mahaffey L, Tannura K, Beninson L, Greenwood BN, Fleshner M. The inflammasome and danger associated molecular patterns (DAMPs) are implicated in cytokine and chemokine responses following stressor exposure. Brain Behav Immun. 2013 Feb; 28:54-62. PMID: 23103443.
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    41. Greenwood BN, Strong PV, Loughridge AB, Day HE, Clark PJ, Mika A, Hellwinkel JE, Spence KG, Fleshner M. 5-HT2C receptors in the basolateral amygdala and dorsal striatum are a novel target for the anxiolytic and antidepressant effects of exercise. PLoS One. 2012; 7(9):e46118. PMID: 23049953.
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    42. Greenwood BN, Loughridge AB, Sadaoui N, Christianson JP, Fleshner M. The protective effects of voluntary exercise against the behavioral consequences of uncontrollable stress persist despite an increase in anxiety following forced cessation of exercise. Behav Brain Res. 2012 Aug 01; 233(2):314-21. PMID: 22610051.
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    43. Strong PV, Christianson JP, Loughridge AB, Amat J, Maier SF, Fleshner M, Greenwood BN. 5-hydroxytryptamine 2C receptors in the dorsal striatum mediate stress-induced interference with negatively reinforced instrumental escape behavior. Neuroscience. 2011 Dec 01; 197:132-44. PMID: 21958863.
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    44. Greenwood BN, Fleshner M. Exercise, stress resistance, and central serotonergic systems. Exerc Sport Sci Rev. 2011 Jul; 39(3):140-9. PMID: 21508844.
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    45. Rozeske RR, Greenwood BN, Fleshner M, Watkins LR, Maier SF. Voluntary wheel running produces resistance to inescapable stress-induced potentiation of morphine conditioned place preference. Behav Brain Res. 2011 Jun 01; 219(2):378-81. PMID: 21262267.
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    46. Greenwood BN, Foley TE, Le TV, Strong PV, Loughridge AB, Day HE, Fleshner M. Long-term voluntary wheel running is rewarding and produces plasticity in the mesolimbic reward pathway. Behav Brain Res. 2011 Mar 01; 217(2):354-62. PMID: 21070820.
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    47. Campeau S, Nyhuis TJ, Sasse SK, Kryskow EM, Herlihy L, Masini CV, Babb JA, Greenwood BN, Fleshner M, Day HE. Hypothalamic pituitary adrenal axis responses to low-intensity stressors are reduced after voluntary wheel running in rats. J Neuroendocrinol. 2010 Aug; 22(8):872-88. PMID: 20406350.
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    48. Greenwood BN, Strong PV, Fleshner M. Lesions of the basolateral amygdala reverse the long-lasting interference with shuttle box escape produced by uncontrollable stress. Behav Brain Res. 2010 Jul 29; 211(1):71-6. PMID: 20226213.
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    49. Campeau S, Nyhuis TJ, Kryskow EM, Masini CV, Babb JA, Sasse SK, Greenwood BN, Fleshner M, Day HE. Stress rapidly increases alpha 1d adrenergic receptor mRNA in the rat dentate gyrus. Brain Res. 2010 Apr 06; 1323:109-18. PMID: 20138850.
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    50. Christianson JP, Ragole T, Amat J, Greenwood BN, Strong PV, Paul ED, Fleshner M, Watkins LR, Maier SF. 5-hydroxytryptamine 2C receptors in the basolateral amygdala are involved in the expression of anxiety after uncontrollable traumatic stress. Biol Psychiatry. 2010 Feb 15; 67(4):339-45. PMID: 19914601.
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    51. Greenwood BN, Strong PV, Foley TE, Fleshner M. A behavioral analysis of the impact of voluntary physical activity on hippocampus-dependent contextual conditioning. Hippocampus. 2009 Oct; 19(10):988-1001. PMID: 19115374.
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    52. Strong PV, Greenwood BN, Fleshner M. The effects of the selective 5-HT(2C) receptor antagonist SB 242084 on learned helplessness in male Fischer 344 rats. Psychopharmacology (Berl). 2009 May; 203(4):665-75. PMID: 19037632.
      View in: PubMed
    53. Sasse SK, Greenwood BN, Masini CV, Nyhuis TJ, Fleshner M, Day HE, Campeau S. Chronic voluntary wheel running facilitates corticosterone response habituation to repeated audiogenic stress exposure in male rats. Stress. 2008 Nov; 11(6):425-37. PMID: 19065456.
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    54. Greenwood BN, Strong PV, Brooks L, Fleshner M. Anxiety-like behaviors produced by acute fluoxetine administration in male Fischer 344 rats are prevented by prior exercise. Psychopharmacology (Berl). 2008 Aug; 199(2):209-22. PMID: 18454279.
      View in: PubMed
    55. Greenwood BN, Fleshner M. Exercise, learned helplessness, and the stress-resistant brain. Neuromolecular Med. 2008; 10(2):81-98. PMID: 18300002.
      View in: PubMed
    56. Greenwood BN, Strong PV, Dorey AA, Fleshner M. Therapeutic effects of exercise: wheel running reverses stress-induced interference with shuttle box escape. Behav Neurosci. 2007 Oct; 121(5):992-1000. PMID: 17907830.
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    57. Bland ST, Tamlyn JP, Barrientos RM, Greenwood BN, Watkins LR, Campeau S, Day HE, Maier SF. Expression of fibroblast growth factor-2 and brain-derived neurotrophic factor mRNA in the medial prefrontal cortex and hippocampus after uncontrollable or controllable stress. Neuroscience. 2007 Feb 23; 144(4):1219-28. PMID: 17197100.
      View in: PubMed
    58. Greenwood BN, Strong PV, Foley TE, Thompson RS, Fleshner M. Learned helplessness is independent of levels of brain-derived neurotrophic factor in the hippocampus. Neuroscience. 2007 Feb 23; 144(4):1193-208. PMID: 17161541.
      View in: PubMed
    59. Foley TE, Greenwood BN, Day HE, Koch LG, Britton SL, Fleshner M. Elevated central monoamine receptor mRNA in rats bred for high endurance capacity: implications for central fatigue. Behav Brain Res. 2006 Nov 01; 174(1):132-42. PMID: 16934883.
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    60. Bland ST, Schmid MJ, Greenwood BN, Watkins LR, Maier SF. Behavioral control of the stressor modulates stress-induced changes in neurogenesis and fibroblast growth factor-2. Neuroreport. 2006 Apr 24; 17(6):593-7. PMID: 16603918.
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    61. Dishman RK, Berthoud HR, Booth FW, Cotman CW, Edgerton VR, Fleshner MR, Gandevia SC, Gomez-Pinilla F, Greenwood BN, Hillman CH, Kramer AF, Levin BE, Moran TH, Russo-Neustadt AA, Salamone JD, Van Hoomissen JD, Wade CE, York DA, Zigmond MJ. Neurobiology of exercise. Obesity (Silver Spring). 2006 Mar; 14(3):345-56. PMID: 16648603.
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    62. Johnson JD, Campisi J, Sharkey CM, Kennedy SL, Nickerson M, Greenwood BN, Fleshner M. Catecholamines mediate stress-induced increases in peripheral and central inflammatory cytokines. Neuroscience. 2005; 135(4):1295-307. PMID: 16165282.
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    63. Nickerson M, Elphick GF, Campisi J, Greenwood BN, Fleshner M. Physical activity alters the brain Hsp72 and IL-1beta responses to peripheral E. coli challenge. Am J Physiol Regul Integr Comp Physiol. 2005 Dec; 289(6):R1665-74. PMID: 16081876.
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    64. Greenwood BN, Foley TE, Day HE, 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. Biol Psychiatry. 2005 Mar 01; 57(5):559-68. PMID: 15737672.
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    65. Greenwood BN, Foley TE, Burhans D, Maier SF, Fleshner M. The consequences of uncontrollable stress are sensitive to duration of prior wheel running. Brain Res. 2005 Feb 08; 1033(2):164-78. PMID: 15694921.
      View in: PubMed
    66. Day HE, Greenwood BN, Hammack SE, Watkins LR, Fleshner M, Maier SF, Campeau S. Differential expression of 5HT-1A, alpha 1b adrenergic, CRF-R1, and CRF-R2 receptor mRNA in serotonergic, gamma-aminobutyric acidergic, and catecholaminergic cells of the rat dorsal raphe nucleus. J Comp Neurol. 2004 Jun 28; 474(3):364-78. PMID: 15174080.
      View in: PubMed
    67. Greenwood BN, Foley TE, Day HE, Campisi J, Hammack SH, Campeau S, Maier SF, Fleshner M. Freewheel running prevents learned helplessness/behavioral depression: role of dorsal raphe serotonergic neurons. J Neurosci. 2003 Apr 01; 23(7):2889-98. PMID: 12684476.
      View in: PubMed
    68. Elphick GF, Wieseler-Frank J, Greenwood BN, Campisi J, Fleshner M. B-1 cell (CD5+/CD11b+) numbers and nIgM levels are elevated in physically active vs. sedentary rats. J Appl Physiol (1985). 2003 Jul; 95(1):199-206. PMID: 12651863.
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    69. Greenwood BN, Kennedy S, Smith TP, Campeau S, Day HE, Fleshner M. Voluntary freewheel running selectively modulates catecholamine content in peripheral tissue and c-Fos expression in the central sympathetic circuit following exposure to uncontrollable stress in rats. Neuroscience. 2003; 120(1):269-81. PMID: 12849759.
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    70. Fleshner M, Campisi J, Deak T, Greenwood BN, Kintzel JA, Leem TH, Smith TP, Sorensen B. Acute stressor exposure facilitates innate immunity more in physically active than in sedentary rats. Am J Physiol Regul Integr Comp Physiol. 2002 Jun; 282(6):R1680-6. PMID: 12010750.
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