Tracy Bale, PhD
Visiting Professor, Psychiatry-Psychopharmacology

Graduate School :
  • PhD, University of Washington (1997)
  • Salk Institute, Neuroscience
Department: Psychiatry-Psychopharmacology

Professional Titles

  • Professor and Endowed Chair
  • The Anschutz Foundation Endowed Chair in Women's Integrated Mental and Physical Health Research
  • Director, InterGenerational Stress and Health
  • Director, Department of Psychiatry Sex Differences Research

Recognition & Awards

  • Maryland Top 100 Women, The Daily Record (2021)
  • Joseph Erlanger Award and Distinguished Lecture, American Physiological Society (2019)
  • NIH Award and Key Lecture, International Federation of Placenta Associations (2019)
  • Daniel H. Efron Research Award, American College of Neuropsychopharmacology (2016)
  • Medtronic Society for Women’s Health Research Prize, Society for Women's Health Research (2012)


  • Montgomery KR, Bridi MS, Folts LM, Marx-Rattner R, Zierden HC, Wulff AB, Kodjo EA, Thompson SM, Bale TL. Chemogenetic activation of CRF neurons as a model of chronic stress produces sex-specific physiological and behavioral effects. Neuropsychopharmacology. 2024 Jan;49(2):443-454. PubMed PMID: 37833589
  • Robertson CD, Davis P, Richardson RR, Iffland PH 2nd, Vieira DCO, Steyert M, McKeon PN, Romanowski AJ, Crutcher G, Jašarevic E, Wolff SBE, Mathur BN, Crino PB, Bale TL, Dick IE, Poulopoulos A. Rapid modeling of an ultra-rare epilepsy variant in wild-type mice by in utero prime editing. bioRxiv. 2023 Dec 19. PubMed PMID: 38106154
  • Rock KD, Folts LM, Zierden HC, Marx-Rattner R, Leu NA, Nugent BM, Bale TL. Developmental transcriptomic patterns can be altered by transgenic overexpression of Uty. Sci Rep. 2023 Nov 30;13(1):21082. PubMed PMID: 38030664
  • Cissé YM, Montgomery KR, Zierden HC, Hill EM, Kane PJ, Huang W, Kane MA, Bale TL. Maternal preconception stress produces sex-specific effects at the maternal:fetal interface to impact offspring development and phenotypic outcomes. Biol Reprod. 2023 Nov 16. [Epub ahead of print] PubMed PMID: 37971364
  • Herb BR, Glover HJ, Bhaduri A, Colantuoni C, Bale TL, Siletti K, Hodge R, Lein E, Kriegstein AR, Doege CA, Ament SA. Single-cell genomics reveals region-specific developmental trajectories underlying neuronal diversity in the human hypothalamus. Sci Adv. 2023 Nov 10;9(45):eadf6251. PubMed PMID: 37939194
  • Kramer AC, Jansson T, Bale TL, Powell TL. Maternal-fetal cross-talk via the placenta: influence on offspring development and metabolism. Development. 2023 Oct 15;150(20). PubMed PMID: 37831056
  • Zierden HC, Marx-Rattner R, Rock KD, Montgomery KR, Anastasiadis P, Folts L, Bale TL. Extracellular vesicles are dynamic regulators of maternal glucose homeostasis during pregnancy. Sci Rep. 2023 Mar 20;13(1):4568. PubMed PMID: 36941297
  • Duffy KA, Sammel MD, Johnson RL, Kim DR, Wang EY, Ewing G, Hantsoo L, Kornfield SL, Bale TL, Epperson CN. Maternal adverse childhood experiences impact fetal adrenal volume in a sex-specific manner. Biol Sex Differ. 2023 Feb 17;14(1):7. PubMed PMID: 36803442
  • Morgan CP, Meadows VE, Marx-Rattner R, Cisse YM, Bale TL. HA-tag CD63 is a novel conditional transgenic approach to track extracellular vesicle interactions with sperm and their transfer at conception. Sci Rep. 2023 Jan 13;13(1):707. PubMed PMID: 36639735
  • Morrison KE, Stenson AF, Marx-Rattner R, Carter S, Michopoulos V, Gillespie CF, Powers A, Huang W, Kane MA, Jovanovic T, Bale TL. Developmental Timing of Trauma in Women Predicts Unique Extracellular Vesicle Proteome Signatures. Biol Psychiatry. 2022 Feb 1;91(3):273-282. PubMed PMID: 34715991
  • Cole AB, Montgomery K, Bale TL, Thompson SM. What the hippocampus tells the HPA axis: Hippocampal output attenuates acute stress responses via disynaptic inhibition of CRF PVN neurons. Neurobiol Stress. 2022 Sep;20:100473. PubMed PMID: 35982732
  • Moon N, Morgan C, Bale TL. Cross Talk proposal: The kids will be fine: parental stress rodent models are good for assessing influences on human neurobiology. J Physiol. 2022 Oct;600(20):4409-4411. PubMed PMID: 36184259
  • Moon N, Morgan C, Bale TL. Rebuttal from Nickole Moon, Christopher Morgan and Tracy L. Bale. J Physiol. 2022 Oct;600(20):4417-4418. PubMed PMID: 36190177
  • Morrison KE, Stenson AF, Marx-Rattner R, Carter S, Michopoulos V, Gillespie CF, Powers A, Huang W, Kane MA, Jovanovic T, Bale TL. Developmental Timing of Trauma in Women Predicts Unique Extracellular Vesicle Proteome Signatures. Biol Psychiatry. 2022 Feb 1;91(3):273-282. PubMed PMID: 34715991
  • Cole AB, Montgomery K, Bale TL, Thompson SM. What the hippocampus tells the HPA axis: Hippocampal output attenuates acute stress responses via disynaptic inhibition of CRF+ PVN neurons. Neurobiol Stress. 2022 Sep;20:100473. PubMed PMID: 35982732
  • Duffy KA, Bale TL, Epperson CN. Germ Cell Drivers: Transmission of Preconception Stress Across Generations. Front Hum Neurosci. 2021;15:642762. PubMed PMID: 34322003
  • Bale TL, Jovanovic T. The critical importance in identifying the biological mechanisms underlying the effects of racism on mental health. Neuropsychopharmacology. 2021 Jan;46(1):233. PubMed PMID: 32792682
  • Arumugasaamy N, Rock KD, Kuo CY, Bale TL, Fisher JP. Microphysiological systems of the placental barrier. Adv Drug Deliv Rev. 2020;161-162:161-175. PubMed PMID: 32858104
  • Morrison KE, Cole AB, Kane PJ, Meadows VE, Thompson SM, Bale TL. Pubertal adversity alters chromatin dynamics and stress circuitry in the pregnant brain. Neuropsychopharmacology. 2020 Jul;45(8):1263-1271. PubMed PMID: 32045935
  • Ramsteijn AS, Jašarevic E, Houwing DJ, Bale TL, Olivier JD. Antidepressant treatment with fluoxetine during pregnancy and lactation modulates the gut microbiome and metabolome in a rat model relevant to depression. Gut Microbes. 2020 Jul 3;11(4):735-753. PubMed PMID: 31971855
  • Cissé YM, Chan JC, Nugent BM, Banducci C, Bale TL. Brain and placental transcriptional responses as a readout of maternal and paternal preconception stress are fetal sex specific. Placenta. 2020 Oct;100:164-170. PubMed PMID: 32980048
  • Bale TL. Sex matters. Neuropsychopharmacology. 2019 Jan;44(1):1-3. PubMed PMID: 30361539
  • Epperson CN, Bale TL. Deciphering the Brain Before Birth. Biol Psychiatry. 2019 Jan 15;85(2):90. PubMed PMID: 30573050
  • Morgan CP, Chan JC, Bale TL. Driving the Next Generation: Paternal Lifetime Experiences Transmitted via Extracellular Vesicles and Their Small RNA Cargo. Biol Psychiatry. 2019 Jan 15;85(2):164-171. PubMed PMID: 30580777
  • Bale TL, Abel T, Akil H, Carlezon WA Jr, Moghaddam B, Nestler EJ, Ressler KJ, Thompson SM. The critical importance of basic animal research for neuropsychiatric disorders. Neuropsychopharmacology. 2019 Jul;44(8):1349-1353. PubMed PMID: 31048727
  • Jašarevic E, Bale TL. Prenatal and postnatal contributions of the maternal microbiome on offspring programming. Front Neuroendocrinol. 2019 Oct;55:100797. PubMed PMID: 31574280
  • Morrison KE, Cole AB, Thompson SM, Bale TL. Brexanolone for the treatment of patients with postpartum depression. Drugs Today (Barc). 2019 Sep;55(9):537-544. PubMed PMID: 31584571
  • Hudecova AM, Hansen KEA, Mandal S, Berntsen HF, Khezri A, Bale TL, Fraser TWK, Zimmer KE, Ropstad E. A human exposure based mixture of persistent organic pollutants affects the stress response in female mice and their offspring. Chemosphere. 2018 Apr;197:585-593. PubMed PMID: 29407821
  • Jašarevic E, Howard CD, Morrison K, Misic A, Weinkopff T, Scott P, Hunter C, Beiting D, Bale TL. The maternal vaginal microbiome partially mediates the effects of prenatal stress on offspring gut and hypothalamus. Nat Neurosci. 2018 Aug;21(8):1061-1071. PubMed PMID: 29988069
  • Chan JC, Nugent BM, Bale TL. Parental Advisory: Maternal and Paternal Stress Can Impact Offspring Neurodevelopment. Biol Psychiatry. 2018 May 15;83(10):886-894. PubMed PMID: 29198470
  • Morrison KE, Epperson CN, Sammel MD, Ewing G, Podcasy JS, Hantsoo L, Kim DR, Bale TL. Preadolescent Adversity Programs a Disrupted Maternal Stress Reactivity in Humans and Mice. Biol Psychiatry. 2017 Apr 15;81(8):693-701. PubMed PMID: 27776734
  • Chan JC, Houghton AB, Bale TL. Strained in Planning Your Mouse Background? Using the HPA Stress Axis as a Biological Readout for Backcrossing Strategies. Neuropsychopharmacology. 2017 Aug;42(9):1749-1751. PubMed PMID: 28361869
  • Bale TL, Epperson CN. Sex as a Biological Variable: Who, What, When, Why, and How. Neuropsychopharmacology. 2017 Jan;42(2):386-396. PubMed PMID: 27658485
  • Bronson SL, Chan JC, Bale TL. Sex-Specific Neurodevelopmental Programming by Placental Insulin Receptors on Stress Reactivity and Sensorimotor Gating. Biol Psychiatry. 2017 Jul 15;82(2):127-138. PubMed PMID: 28168960
  • Epperson CN, Sammel MD, Bale TL, Kim DR, Conlin S, Scalice S, Freeman K, Freeman EW. Adverse Childhood Experiences and Risk for First-Episode Major Depression During the Menopause Transition. J Clin Psychiatry. 2017 Mar;78(3):e298-e307. PubMed PMID: 28394509
  • Bale TL. The placenta and neurodevelopment: sex differences in prenatal vulnerability. Dialogues Clin Neurosci. 2016 Dec;18(4):459-464. PubMed PMID: 28179817
  • Jašarevic E, Morrison KE, Bale TL. Sex differences in the gut microbiome-brain axis across the lifespan. Philos Trans R Soc Lond B Biol Sci. 2016 Feb 19;371(1688):20150122. PubMed PMID: 26833840
  • Kim DR, Bale TL, Epperson CN. Prenatal programming of mental illness: current understanding of relationship and mechanisms. Curr Psychiatry Rep. 2015 Feb;17(2):5. PubMed PMID: 25617041
  • Bale TL. Epigenetic and transgenerational reprogramming of brain development. Nat Rev Neurosci. 2015 Jun;16(6):332-44. PubMed PMID: 25921815
  • Rodgers AB, Morgan CP, Leu NA, Bale TL. Transgenerational epigenetic programming via sperm microRNA recapitulates effects of paternal stress. Proc Natl Acad Sci U S A. 2015 Nov 3;112(44):13699-704. PubMed PMID: 26483456
  • Bale TL, Epperson CN. Sex differences and stress across the lifespan. Nat Neurosci. 2015 Oct;18(10):1413-20. PubMed PMID: 26404716
  • Rodgers AB, Bale TL. Germ Cell Origins of Posttraumatic Stress Disorder Risk: The Transgenerational Impact of Parental Stress Experience. Biol Psychiatry. 2015 Sep 1;78(5):307-14. PubMed PMID: 25895429
  • Bale TL. CRF as the key component of stress response systems. Front Neuroendocrinol. 2014 Apr;35(2):159-60. PubMed PMID: 24656169
  • Morrison KE, Rodgers AB, Morgan CP, Bale TL. Epigenetic mechanisms in pubertal brain maturation. Neuroscience. 2014 Apr 4;264:17-24. PubMed PMID: 24239720
  • Howerton CL, Bale TL. Targeted placental deletion of OGT recapitulates the prenatal stress phenotype including hypothalamic mitochondrial dysfunction. Proc Natl Acad Sci U S A. 2014 Jul 1;111(26):9639-44. PubMed PMID: 24979775
  • Bale TL. Lifetime stress experience: transgenerational epigenetics and germ cell programming. Dialogues Clin Neurosci. 2014 Sep;16(3):297-305. PubMed PMID: 25364281
  • Rodgers AB, Morgan CP, Bronson SL, Revello S, Bale TL. Paternal stress exposure alters sperm microRNA content and reprograms offspring HPA stress axis regulation. J Neurosci. 2013 May 22;33(21):9003-12. PubMed PMID: 23699511
  • Bale TL, Chen A. Minireview: CRF and Wylie Vale: a story of 41 amino acids and a Texan with grit. Endocrinology. 2012 Jun;153(6):2556-61. PubMed PMID: 22492308
  • Morgan CP, Bale TL. Early prenatal stress epigenetically programs dysmasculinization in second-generation offspring via the paternal lineage. J Neurosci. 2011 Aug 17;31(33):11748-55. PubMed PMID: 21849535
  • Bale TL. Sex differences in prenatal epigenetic programming of stress pathways. Stress. 2011 Jul;14(4):348-56. PubMed PMID: 21663536
  • Bale TL, Baram TZ, Brown AS, Goldstein JM, Insel TR, McCarthy MM, Nemeroff CB, Reyes TM, Simerly RB, Susser ES, Nestler EJ. Early life programming and neurodevelopmental disorders. Biol Psychiatry. 2010 Aug 15;68(4):314-9. PubMed PMID: 20674602
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Professional Memberships

  • International Brain Research Organization, President
  • American College of Neuropsychopharmacology, Fellow
  • Society for Neuroscience, Member

Practice Locations

CU Medicine Psychiatry - Outpatient Clinic
1890 N Revere Ct
Anschutz Health Sciences Bldg, Suite 4020
Aurora, CO 80045

Specialty Information

  • Psychiatry
Public Speaking

General Information

Graduate Schools:
  • PhD, University of Washington (1997)
  • Salk Institute, Neuroscience (0001)
Department: Psychiatry-Psychopharmacology
Contact Us
CU Anschutz
Fitzsimons Building

13001 East 17th Place
Campus Box C290
Aurora, CO 80045

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