Research Papers

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Chova PE, Ferrer E, Goetze LR, Dufek ME, Freeman SM, Bales KL. (2023) Neural and behavioral reactions to partners and strangers in monogamous female titi monkeys (Plecturocebus cupreus). Behav Brain Res. Feb 11:114334.


Berendzen KM, Sharma R, Mandujano MA, Wei Y, Rogers FD, Simmons TC, Seelke AMH, Bond JM, Larios R, Goodwin NL, Sherman M, Parthasarthy S, Espineda I, Knoedler JR, Beery A, Bales KL, Shah NM, Manoli DS. (2022) Oxytocin receptor is not required for social attachment in prairie voles. Neuron. Dec 28:S0896-6273(22)01084-4.

Rogers FD, Bales KL. (2022) Introduction to Special Issue on Affective Science in Animals: Toward a Greater Understanding of Affective Processes in Non-Human Animals. Affect Sci. Dec 3;3(4):697-702.

Zablocki-Thomas PB, Rogers FD, Bales KL. (2022) Neuroimaging of human and non-human animal emotion and affect in the context of social relationships. Front Behav Neurosci. Oct 21;16:994504.

Baxter A, Maxwell JA, Bales KL, Finkel EJ, Impett EA, Eastwick PW. (2022) Initial impressions of compatibility and mate value predict later dating and romantic interest. Proc Natl Acad Sci U S A. Nov 8;119(45):e2206925119.

Bales KL, Rogers FD. (2022) Interactions between the κ opioid system, corticotropin-releasing hormone and oxytocin in partner loss. Philos Trans R Soc Lond B Biol Sci. Aug 29;377(1858):20210061.

Frehner SS, Dooley KT, Palumbo MC, Smith AL, Goodman MM, Bales KL, Freeman SM. (2022) Effect of sex and autism spectrum disorder on oxytocin receptor binding and mRNA expression in the dopaminergic pars compacta of the human substantia nigra. Philos Trans R Soc Lond B Biol Sci. Aug 29;377(1858):20210118.

Arias Del Razo R, Velasco Vazquez ML, Turcanu P, Legrand M, Lau AR, Weinstein TAR, Goetze LR, Bales KL. (2022) Effects of Chronic and Acute Intranasal Oxytocin Treatments on Temporary Social Separation in Adult Titi Monkeys (Plecturocebus cupreus). Front Behav Neurosci. Jun 22;16:877631.

Mederos SL, Duarte RC, Mastoras M, Dennis MY, Settles ML, Lau AR, Scott A, Woodward K, Johnson C, Seelke AMH, Bales KL. (2022) Effects of pairing on color change and central gene expression in lined seahorses. Genes Brain Behav. Jun;21(5):e12812.

Asangba AE, Mugisha L, Rukundo J, Lewis RJ, Halajian A, Cortés-Ortiz L, Junge RE, Irwin MT, Karlson J, Perkin A, Watsa M, Erkenswick G, Bales KL, Patton DL, Jasinska AJ, Fernandez-Duque E, Leigh SR, Stumpf RM. (2022) Large Comparative Analyses of Primate Body Site Microbiomes Indicate that the Oral Microbiome Is Unique among All Body Sites and Conserved among Nonhuman Primates. Microbiol Spectr. Jun 29;10(3):e0164321.

Conley AJ, Berger T, Del Razo RA, Cotterman RF, Sahagún E, Goetze LR, Jacob S, Weinstein TAR, Dufek ME, Mendoza SP, Bales KL. (2022) The onset of puberty in colony-housed male and female titi monkeys (Plecturocebus cupreus): Possible effects of oxytocin treatment during peri-adolescent development. Horm Behav. Jun;142:105157.

Razo RA, Velasco Vazquez ML, Turcanu P, Legrand M, Floch M, Weinstein TAR, Goetze LR, Freeman SM, Baxter A, Witczak LR, Sahagún E, Berger T, Jacob S, Lawrence RH, Rothwell ES, Savidge LE, Solomon M, Mendoza SP, Bales KL. (2022) Long term effects of chronic intranasal oxytocin on adult pair bonding behavior and brain glucose uptake in titi monkeys (Plecturocebus cupreus). Horm Behav. Apr;140:105126.

Clink DJ, Lau AR, Kanthaswamy S, Johnson LM, Bales KL. (2022) Moderate evidence for heritability in the duet contributions of a South American primate. J Evol Biol. Jan;35(1):51-63.


Witczak LR, Arias Del Razo R, Baxter A, Conley AJ, Cotterman R, Dufek M, Goetze LR, Lau AR, Mendoza SP, Savidge LE, Bales KL. (2021) Relationships between cortisol and urinary androgens in female titi monkeys (Plecturocebus cupreus). Gen Comp Endocrinol. 314:113927.

Karaskiewicz CL, Witczak LR, Lau AR, Dufek ME, Bales KL. (2021) Parenting costs time: Changes in pair bond maintenance across pregnancy and infant rearing in a monogamous primate (Plecturocebus cupreus). New Dir Child Adolesc Dev. Online ahead of print.

Bales KL, Ardekani CS, Baxter A, Karaskiewicz CL, Kuske JX, Lau AR, Savidge LE, Sayler KR, Witczak LR. (2021) What is a pair bond? Horm Behav. 136:105062.

Rogers FD, Freeman SM, Anderson M, Palumbo MC, Bales KL. (2021) Compositional variation in early-life parenting structures alters oxytocin and vasopressin 1a receptor development in prairie voles (Microtus ochrogaster). J Neuroendocrinol. 33(8).

Baxter A, Capitanio JP, Bales KL, Kinnally EL. (2021) Biobehavioral organization shapes the immune epigenome in infant rhesus Macaques (Macaca mulatta). Brain Behav Immun. 96:256-270.

Simmons TC, Freeman SM, Lackey NS, Dreyer BK, Manoli DS, Bales KL. (2021) Cannabinoid receptor Type 1 densities reflect social organization in Microtus. J Comp Neurol. 529(5):1004-1017.

Safa A, Lau AR, Aten S, Schilling K, Bales KL, Miller VA, Fitzgerald J, Chen M, Hill K, Dzwigalski K, Obrietan K, Phelps MA, Sadee W, Oberdick J. (2021) Pharmacological Prevention of Neonatal Opioid Withdrawal in a Pregnant Guinea Pig Model. Front Pharmacol. 11: 613328.

Grebe NM, Sharma A, Freeman SM, Palumbo MC, Patisaul HB, Bales KL, Drea CM. (2021) Neural correlates of mating system diversity: oxytocin and vasopressin receptor distributions in monogamous and non-monogamous Eulemur. Sci Rep. 11(1):3746.

Danoff JS, Wroblewski KL, Graves AJ, Quinn GC, Perkeybile AM, Kenkel WM, Lillard TS, Parikh HI, Golino HF, Gregory SG, Carter CS, Bales KL, Connelly JJ. (2021) Genetic, epigenetic, and environmental factors controlling oxytocin receptor gene expression. Clin Epigenetics. 13(1):23.

Lau AR, Grote MN, Dufek ME, Franzetti TJ, Bales KL, Isbell LA. (2021) Titi monkey neophobia and visual abilities allow for fast responses to novel stimuli. Sci Rep. 11(1):2578.

Simmons TC, Singh ALK, Bales KL. (2021) Effects of systemic endocannabinoid manipulation on social and exploratory behavior in prairie voles (Microtus ochrogaster). Psychopharmacology (Berl). 238(1):293-304.


Lawrence RH, Palumbo MC, Freeman SM, Guoynes CD, Bales KL. (2020) Developmental Fluoxetine Exposure Alters Behavior and Neuropeptide Receptors in the Prairie Vole. Front Behav Neurosci. 14:584731.

Bales KL. (2020) Psychobiological development in primates: Introduction to joint special issue with Developmental Psychobiology. Am J Primatol. 82(11):e23207.

Baxter A, Anderson M, Seelke AM, Kinnally EL, Freeman SM, Bales KL. (2020) Oxytocin receptor binding in the titi monkey hippocampal formation is associated with parental status and partner affiliation. Sci Rep. 10(1):17301.

Rothwell ES, Carp SB, Savidge LE, Mendoza SP, Bales KL. (2020) Relationship tenure differentially influences pair-bond behavior in male and female socially monogamous titi monkeys (Callicebus cupreus). Am J Primatol. 82(10).

Bales KL. (2020) Introduction to special section on COVID-19 in primatology. Am J Primatol. 82(8).

Simmons TC, Freeman SM, Lackey NS, Dreyer BK, Manoli DS, Bales KL. (2020) Cannabinoid receptor Type 1 densities reflect social organization in Microtus. Journal of Comparative Neurology.

Rogers FD, Bales KL. (2020) Revisiting paternal absence: Female alloparental replacement of fathers recovers partner preference formation in female, but not male prairie voles (Microtus ochrogaster). Dev Psychobiol. 62(5):573-590.

Mercier F, Witczak LR, Bales KL. (2020) Coppery titi monkey (Plecturocebus cupreus) pairs display coordinated behaviors in response to a simulated intruder. Am J Primatol. 82(7).

Lau AR, Clink DJ, Bales KL. (2020) Individuality in the vocalizations of infant and adult coppery titi monkeys (Plecturocebus cupreus). Am J Primatol. 82(6).

Arias Del Razo R, Berger T, Conley AJ, Freeman SM, Goetze LR, Jacob S, Lawrence RH, Mendoza SP, Rothwell ES, Savidge LE, Solomon M, Weinstein TAR, Witczak LR, Bales KL. (2020) Effects of chronic intranasal oxytocin on behavior and cerebral glucose uptake in juvenile titi monkeys. Psychoneuroendocrinology.113:104494.

Savidge LE, Bales KL. (2020) An Animal Model for Mammalian Attachment: Infant Titi Monkey (Plecturocebus cupreus) Attachment Behavior Is Associated With Their Social Behavior as Adults. Front Psychol.11:25.


Baxter A, Wood EK, Witczak LR, Bales KL, Higley JD. (2019) Sexual Dimorphism in Titi Monkeys’ Digit (2D:4D) Ratio is Associated with Maternal Urinary Sex Hormones During Pregnancy. Dev Psychobiol.

Rogers FD, Bales KL. (2019) Mothers, Fathers, and Others: Neural Substrates of Parental Care. Trends Neurosci. 42(8):552-562.

Rothwell ES, Mendoza SP, Ragen BJ, Bales KL. (2019) Dopamine D1-like receptors regulate agonistic components of pair bond maintenance behaviors in male titi monkeys (Callicebus cupreus). Psychoneuroendocrinology. 106:259-267.

Ryan AM, Freeman SM, Murai T, Lau AR, Palumbo MC, Hogrefe CE, Bales KL, Bauman MD. (2019) Non-invasive Eye Tracking Methods for New World and Old World Monkeys. Frontiers in Behavioral Neuroscience.

Perkeybile AM, Carter CS, Wroblewski KL, Puglia MH, Kenkel WM, Lillard TS, Karaoli T, Gregory SG, Mohammadi N, Epstein L, Bales KL, Connelly JJ. (2019) Early nurture epigenetically tunes the oxytocin receptor. Psychoneuroendocrinology. 99:128-136.

Clink DJ, Lau AR, Bales KL. (2019) Age-related changes and vocal convergence in titi monkey duet pulses. Behaviour 156(15):1471-1494.


Freeman, S.M., Palumbo, M., Larke, R.H., Smith, A., Goodman, M., Bales, K.L. (2018) Effect of age and autism spectrum disorder on oxytocin receptor density in human basal forebrain and midbrain. Translational Psychiatry.

Seelke, A.M.H., Bond, J., Simmons, T.C., Joshi, N., Settles, M., Stolzenberg, D., Rhemtulla, M., Bales, K.L. (2018) Fatherhood alters gene expression within the MPOA. Environmental Epigenetics.

Rogers, F.D., Rhemtulla, M., Ferrer, E., Bales, K.L. (2018) Longitudinal trajectories and interparental dynamics of prairie vole biparental care. Frontiers in Ecology and Evolution.

Ondrasek, N.R., Freeman, S.M., Bales, K.L., Calisi, R.M. (2018) Nonapeptide receptor distributions in promising avian models for the neuroecology of flocking. Frontiers in Neuroscience. 12, 713.

Seelke AM, Rhine MA, Khun K, Shweyk AN, Scott AM, Bond JM, Graham JL, Havel PJ, Wolden-Hanson T, Bales KL, Blevins JE. (2018) Intranasal oxytocin reduces weight gain in diet-induced obese prairie voles. Physiol Behav 196:67-77.

Freeman SM, Ngo J, Singh B, Masnaghetti M, Bales KL, Blevins JE. (2018) Effects of Chronic Oxytocin Administration and Diet Composition on Oxytocin and Vasopressin 1a Receptor Binding in the Rat Brain. Neuroscience 392:241-251.

Freeman SM, Bales KL. (2018) Oxytocin, vasopressin, and primate behavior: Diversity and insight. Am J Primotology 80(10).

Witczak LR, Ferrer E, Bales KL. (2018) Effects of aggressive temperament on endogenous oxytocin levels in adult titi monkeys. Am J Primotology 80(10).

Bales KL, Witczak LR, Simmons TC, Savidge LE, Rothwell ES, Rogers FD, Manning RA, Heise MJ, Englund M, Arias Del Razo R. (2018) Social touch during development: Long-term effects on brain and behavior. Neurosci Biobehav Rev. 95:202-219.

Freeman SM, Rebout N, Bales KL. (2018) Effect of reward type on object discrimination learning in socially monogamous coppery titi monkeys (Callicebus cupreus). Am J Primotology 80.

Hartman S, Freeman SM, Bales KL, Belsky J. (2018) Prenatal Stress as a Risk-and an Opportunity-Factor. Psychological Science (Epub ahead of print).

Kanthaswamy S, Bales KL. (2018) Evaluating the genetic status of a closed colony of titi monkeys (Callicebus cupreus) using multigenerational pedigrees. J Med Primatol (Epub ahead of print).


Maninger N, Mendoza SP, Williams DR, Mason WA, Cherry SR, Rowland DJ, Schaefer T, Bales KL. (2017) Imaging, Behavior and Endocrine Analysis of “Jealousy” in a Monogamous Primate. Frontiers in Ecology and Evolution.

Bales, KL. (2017) American Journal of Primatology: Goals and priorities of a new editor. Am J Primatol 80.

Guoynes CD, Simmons TC, Downing GM, Jacob S, Solomon M, Bales KL. (2017) Chronic Intranasal Oxytocin has Dose-dependent Effects on Central Oxytocin and Vasopressin Systems in Prairie Voles (Microtus ochrogaster). Neuroscience 369:292-302.

Duque-Wilckens N, Steinman MQ, Busnelli M, Chini B, Yokoyama S, Pham M, Laredo SA, Hao R, Perkeybile AM, Minie VA, Tan PB, Bales KL, Trainor BC. (2017) Oxytocin Receptors in the Anteromedial Bed Nucleus of the Stria Terminalis Promote Stress-Induced Social Avoidance in Female California Mice. Biol Psychiatry 83(3):203-213.

Bales KL, Arias Del Razo R, Conklin QA, Hartman S, Mayer HS, Rogers FD, Simmons TC, Smith LK, Williams A, Williams DR, Witczak LR, Wright EC. (2017) Titi Monkeys as a Novel Non-Human Primate Model for the Neurobiology of Pair Bonding
. Yale J Biol Med 90(3):373-387.

Simmons TC, Balland JF, Dhauna J, Yang SY, Traina JL, Vazquez J, Bales KL. (2017) Early Intranasal Vasopressin Administration Impairs Partner Preference in Adult Male Prairie Voles (Microtus ochrogaster). Frontiers in Endocrinology 8:145. pdf

Larke RH, Toubiana A, Lindsay KA, Mendoza SP, Bales KL. (2017) Infant titi monkey behavior in the open field test and the effect of early adversity. American Journal of Primatology 79(9).

Maninger, N., Hinde, K., Mendoza, S.P., Mason, W.A., Larke, R.H., Ragen, B.J., Jarcho, M.R., Cherry, S.R., Rowland, D.J., Ferrer, E., Bales, K.L. (2017) Pair bond formation leads to a sustained increase in global cerebral glucose metabolism in monogamous male titi monkeys (Callicebus cupreus). Neuroscience 348: 302-312. NIHMS: 855416

Bales, K.L. (2017) Parenting in Animals.  Current Opinion in Psychology 15:93-98. NIHMSID: 856537.

Duchemin, A., Seelke, A.M., Simmons, T.C., Freeman, S.M., Bales, K.L. (2017) Localization of oxytocin receptors in the prairie vole (Microtus ochrogaster) neocortex.  Neuroscience 348: 201-211.  PMCID: PMC5368034.

Freeman SM, Smith AL, Goodman MM, Bales KL. (2017) Selective localization of oxytocin receptors and vasopressin 1a receptors in the human brainstem. Social Neuroscience 12:113-123.

Hostetler, C.M., Hinde, K., Maninger, N., Mendoza, S.P., Mason, W.A., Rowland, D.J., Wang, G.B., Kukis, D., Cherry, S.R., Bales, K.L. (2017) Effects of pair bonding on dopamine D1 receptors in monogamous male titi monkeys (Callicebus cupreus). American Journal of Primatology 79: 1-9.

Perkeybile AM, Bales KL. (2017) Intergenerational transmission of sociality: the role of parents in shaping social behavior in monogamous and non-monogamous species. Journal of Experimental Biology 220:114-123. pdf


Muth C, Bales KL, Hinde K, Maninger N, Mendoza SP, Ferrer E. (2016).  Alternative Models for Small Samples in Psychological Research: Applying Linear Mixed Effects Models and Generalized Estimating Equations to Repeated Measures Data.  Education and Psychological Measurement, 76(1):64-87.

Hinde K, Muth C, Maninger N, Ragen BJ, Larke RH, Jarcho MR, Mendoza SP, Mason WA, Ferrer E, Cherry SR, Fisher-Phelps ML, Bales KL. (2016) Challenges to the Pair Bond: Neural and Hormonal Effects of Separation and Reunion in a Monogamous Primate. Frontiers in Behavioral Neuroscience. 10:221. pdf

Seelke A.M.H., Yuan S-M, Perkeybile AM, Krubitzer LA, Bales KL. (2016) Early experiences can alter the size of cortical fields in prairie voles (Microtus ochrogaster). Environmental Epigenetics. 1-9. pdf

Arias Del Razo R, Bales KL. (2016) Exploration in a dispersal task: Effects of early experience and correlation with other behaviors in prairie voles (Microtus ochrogaster). Behav Processes. 66-75. (Epub ahead of print). pdf

Larke RH, Maninger N, Ragen BJ, Mendoza SP, Bales KL. (2016) Serotonin 1A agonism decreases affiliative behavior in pair-bonded titi monkeys. Horm Behav. 71-77. (Epub ahead of print). pdf

Hostetler CM, Hinde K, Maninger N, Mendoza SP, Mason WA, Rowland DJ, Wang GB, Kukis D, Cherry SR, Bales KL. (2016) Effects of pair bonding on dopamine D1 receptors in monogamous male titi monkeys (Callicebus cupreus). American Journal of Primatology (online before print). pdf

Corbett BA, Bales KL, Swain D, Sanders K, Weinstein TA, Muglia LJ. (2016) Comparing oxytocin and cortisol regulation in a double-blind, placebo-controlled, hydrocortisone challenge pilot study in children with autism and typical development. J Neurodev Disord. 8-32. pdf

Duque-Wilckens N, Steinman MQ, Laredo SA, Hao R, Perkeybile AM, Bales KL, Trainor BC. (2016) Inhibition of vasopressin V1a receptors in the medioventral bed nucleus of the stria terminalis has sex- and context-specific anxiogenic effects. Neuropharmacology. 59-68. (Epub ahead of print). pdf

Smith AL, Freeman SM, Barnhart TE, Abbott DH, Ahlers EO, Kukis DL, Bales KL, Goodman MM, Young LJ. (2016) Initial investigation of three selective and potent small molecule oxytocin receptor PET ligands in New World monkeys. Bioorg Med Chem Lett. 3370-5. (Epub 2016 May 7). pdf

Freeman SM, Smith AL, Goodman MM, Bales KL. (2016) Selective localization of oxytocin receptors and vasopressin 1a receptors in the human brainstem. Social Neuroscience (Epub ahead of print). pdf

Freeman SM, Samineni S, Allen PC, Stockinger D, Bales KL, Hwa GG, Roberts JA. (2016) Plasma and CSF oxytocin levels after intranasal and intravenous oxytocin in awake macaques. Psychoneuroendocrinology 66: 185-194. pdf

Blevins JE, Thompson BW, Anekonda VT, Ho JM, Graham JL, Roberts ZS, Hwang BH, Ogimoto K, Wolden-Hanson TH, Nelson J, Kaiyala KJ, Havel PJ, Bales KL, Morton GJ, Schwartz MW, Baskin DG. (2016) Chronic CNS Oxytocin Signaling Preferentially Induces Fat Loss in High Fat Diet-Fed Rats by Enhancing Satiety Responses and Increasing Lipid Utilization. American Journal of Physiology (in press). pdf


Steinman MQ, Duque-Wilckens N, Greenberg GD, Hao R, Campi KL, Laredo SA, Laman-Maharg A, Manning CE, Doig IE, Lopez EM, Walch K, Bales KL, Trainor BC. (2015) Sex-Specific Effects of Stress on Oxytocin Neurons Correspond With Responses to Intranasal Oxytocin. Biological Psychiatry (in press). pdf

Fisher-Phelps ML, Mendoza SP, Serna S, Griffin LL, Schaefer TJ, Jarcho MR, Ragen BJ, Goetze LR, Bales KL. (2015) Laboratory simulations of mate-guarding as a component of the pair-bond in male titi monkeys, Callicebus cupreus. American Journal of Primatology (online before print). pdf

Díaz-Muñoz SL, Bales KL. (2015) “Monogamy” in primates: Variability, trends, and synthesis: Introduction to special issue on primate monogamy. American Journal of Primatology (online before print). pdf

Carp, S.B., Rothwell, E.S., Bourdon, A., Freeman, S.M., Ferrer, E., Bales, K.L. (2015) Development of a partner preference test that differentiates between established pair bonds and other relationships in socially monogamous titi monkeys (Callicebus cupreus).  American Journal of Primatology (online before print). pdf

Bales, K.L., Saltzman, W. (2015) Fathering in rodents: Neurobiological substrates and consequences for offspring.  Hormones and Behavior (online before print). pdf

Seelke, A.M., Perkeybile, A.M., Grunewald, R., Bales, K.L., Krubitzer, L.A. (2015) Individual differences in cortical connections of somatosensory cortex are associated with parental rearing style in prairie voles (Microtus ochrogaster). Journal of Comparative Neurology (online before print). pdf

Kinsley, C.H., Bales, K., Bardi, M., Stolzenberg, D. (2015) Reproductive experiential regulation of cognitive and emotional resilience. Neuroscience and Biobehavioral Reviews (online before print). pdf

Muth, C., Bales, K.L., Hinde, K., Maninger, N., Mendoza, S.P., Ferrer, E. (2015) Alternative models for small samples in psychological research: applying linear mixed effects models and generalized estimating equations to repeated measures data.  Educational and Psychological Measurement 1-24 (online before print). pdf

Perkeybile, A.M., Delaney-Busch, N., Hartman, S., Grimm, K.J., Bales, K.L. (2015) Intergenerational transmission of alloparental behavior and oxytocin and vasopressin receptor distribution in the prairie vole.  Frontiers in Behavioral Neuroscience 9:191. pdf

Perkeybile, A.M., Bales, K.L. (2015) Early rearing experience is associated with vasopressin immunoreactivity to an acute non-social stressor in the prairie vole. Physiology & Behavior 147: 149-156. pdf

Ragen, B.J., Freeman, S.M., Laredo, S.A., Mendoza, S.P., Bales, K.L. (2015) µ and ĸ opioid receptor distribution in the monogamous titi monkey (Callicebus cupreus): implications for social behavior and endocrine functioning.  Neuroscience 290: 421-434. pdf

Perkeybile, A.M., Bales, K.L. (2015) Early rearing experience is related to altered aggression and vasopressin production following chronic social isolation in the prairie vole. Behavioural Brain Research 283: 37-46. pdf

Blevins, J.E., Graham, J.L., Morton, G.J., Bales, K.L., Schwartz, M.W., Baskin, D.G., Havel, P.J. (2015) Chronic oxytocin administration inhibits food intake, increases energy expenditure, and produces weight loss in fructose-fed obese rhesus monkeys.  American Journal of Physiology. Regulatory, Integrative and Comparative Physiology 308: R431-R438. pdf

Ragen, B.J., Maninger, N., Mendoza, S.P., Bales, K.L. (2015) The effects of morphine, naloxone, and ĸ opioid manipulation on endocrine functioning and social behavior in monogamous titi monkeys (Callicebus cupreus).  Neuroscience 287: 32-42. pdf


Bales, K.L., Solomon, M., Jacob, S., Crawley, J.N., Silverman, J.L., Larke, R.H., Sahagun, E., Puhger, K.R., Pride, M.C., Mendoza, S.P. (2014) Long-term exposure to intranasal oxytocin in a mouse autism model.  Translational Psychiatry 4:e480. pdf

Weinstein, T.A., Bales, K.L., Maninger, N., Hostetler, C.M., Capitanio, J.P. (2014) Early involvement in friendships predicts later plasma concentrations of oxytocin and vasopressin in juvenile rhesus macaques (Macaca mulatta).  Frontiers in Behavioral Neuroscience 8:295. pdf

Mendoza, A., Ng, J., Bales, K.L., Mendoza, S.P., George, D.A., Smith, D.G., Kanthaswamy, S. (2014) Population genetics of the California National Primate Research Center’s (CNPRC) captive Callicebus cupreus colony.  Primates 56: 37-44. pdf

Freeman, S.M., Walum, H., Inoue, K., Smith, A.L., Goodman, M.M., Bales, K.L.*, Young, L.J.* (2014) Neuroanatomical distribution of oxytocin and vasopressin 1a receptors in the socially monogamous coppery titi monkey (Callicebus cupreus). Neuroscience 273: 12-23.
* denotes co-last authors pdf

Phillips, K.A., Bales, K.L., Capitanio, J.P., Conley, A., Czoty, P.W., t’Hart, B.A., Hopkins, W.D.,   Hu, S.-L., Miller, L.A., Nader, M.A., Nathanielsz, P.W., Rogers, J., Shively, C.A., Voytko, M.L.  (2014) Why primate models matter.  American Journal of Primatology 76:801-827. pdf

Schradin, C., Larke, R.H., Bales, K.L. (2014) Growing up in the family or growing up alone influences behavior and hormones, but not arginine vasopressin receptor 1a expression in male African striped mice. Physiology & Behavior, 129: 205-213. PMID: 24631307 pdf


Millan, M.J., Bales, K.L. (2013) Towards improved animal models for evaluating social cognition and its disruption in schizophrenia: the CNTRICS initiative.  Neuroscience and Biobehavioral Reviews, 37: 2166-2180. PMID: 24090822 pdf

Ragen, B.J., Maninger, N., Mendoza, S.P., Jarcho, M.R., Bales, K.L. (2013) Presence of a pair-mate regulates the behavioral and physiological effects of opioid manipulation in the monogamous titi monkey (Callicebus cupreus).  Psychoneuroendocrinology 48: 2448-2661. PMCID:3812423 pdf

Miller, M., Bales, K.L., Taylor, S.L., Yoon, J., Hostetler, C.M., Carter, C.S., Solomon, M. (2013) Oxytocin and vasopressin in children and adolescents with autism spectrum disorders: sex differences and associations with symptoms. Autism Research 6:91-102. PMCID:3657571 pdf

Yu, G., Yagi, S., Carrion, R. Jr, Chen, E.C., Liu, M., Brasky, K.M., Lanford, R.E., Kelly, K.R., Bales, K.L., Schnurr, D.P., Canfield, D.R., Patterson, J.L., Chiu, C.Y. (2013) Experimental cross-species infection of common marmosets by titi monkey adenovirus. PLOS One. 8(7):e68558. PMCID:3722195 pdf

Perkeybile, A.M., Griffin, L.L., Bales, K.L. (2013) Natural variation in early parental care correlates with social behaviors in adolescent prairie voles (Microtus ochrogaster).  Frontiers in Behavioral Neuroscience 7:21. PMCID: 3600544 pdf

Bales, K.L., Perkeybile, A.M., Conley, O.G., Lee, M.H., Guoynes, C.D., Downing, G.M., Yun, C.R., Solomon, M., Jacob, S., Bales, K.L. (2013) Chronic intranasal oxytocin causes long-term impairments in partner preference formation in male prairie voles.  Biological Psychiatry 74: 180-188.  PMCID: 3556198 pdf


Ragen, B.R., Mendoza, S.P., Mason, W.A., Bales, K.L. (2012) Differences in titi monkey (Callicebus cupreus) social bonds affect arousal, affiliation, and response to reward.  American Journal of Primatology 74: 758-769. pdf

Greenberg, G., Van Westerhuyzen, J.A.,  Bales, K.L., Trainor, B.C. (2012) Is it all in the family? The effects of early social structure on neural-behavioral systems of prairie voles (Microtus ochrogaster). Neuroscience 216:46-56. PMCID: 3397474 pdf

Manoli, D.S., Subramanyam, D., Carey, C., Sudin, E., Van Westerhuyzen, J.A., Bales, K.L., Blelloch, R., Shah, N.M. (2012) Generation of induced pluripotent stem cells from the prairie vole.  PLoS One 7:e38119.  PMCID: 3365000 pdf

Hostetler, C.M., Bales, K.L. (2012) DeltaFosB is increased in the nucleus accumbens by amphetamine but not social housing or isolation in the prairie vole. Neuroscience 210:266-274. pdf

Kenkel, W.M., Paredes, J., Yee, J.R., Pournajafi-Nazarloo, H., Bales, K.L., Carter, C.S. (2012) Neuroendocrine and behavioural responses to exposure to an infant in male prairie voles. Journal of Neuroendocrinology. 24:874-886. pdf

Jarcho, M.R., Mendoza, S.P., Bales, K.L. (2012) Hormonal and experiential predictors of infant survivorship and maternal behavior in a monogamous primate (Callicebus cupreus). American Journal of Primatology 74:462-470. pdf

Bales, K.L., Perkeybile, A.M. (2012) Developmental experiences and the oxytocin receptor system. Hormones and Behavior 61:313-319. pdf


Hostetler, C.M., Harkey, S.L., Krzywosinski, T.P., Aragona, B.J., Bales, K.L. (2011) Neonatal exposure to the D1 agonist SKF38393 inhibits pair-bonding in the adult prairie vole. Behavioural Pharmacology 22: 703-710. PMCID: 3174420 pdf

Hostetler, C.M., Kowalczyk, A.S., Griffin, L.L., Bales, K.L. (2011) CART peptide following social novelty in the prairie vole (Microtus ochrogaster). Brain Research 1414:32-40. PMCID: 3176997 pdf

Chen, E.C., Yagi, S., Kelly, K.R., Mendoza, S.P., Maninger, N., Rosenthal, A., Spinner, A., Bales, K.L., Schnurr, D.P., Lerche, N.W, Chiu, C.Y. (2011) Cross-species transmission of a novel adenovirus associated with a fulminant pneumonia outbreak in a titi monkey colony. PLoS Pathogens 7: e1002155. PMCID: 3136464 pdf

Bales, K.L., Boone, E., Epperson, P., Hoffman, G., Carter, C.S. (2011) Are behavioral effects of early experience mediated by oxytocin? Frontiers in Child and Neurodevelopmental Psychiatry  2: 1-12. PMCID: 3098714.  This article was also included as part of the e-book Translational research models and methods for mother-infant interactions and developmental studies, edited by J. Johns, R. Eiden, E.M. Byrnes, S. Moy, J.I. Morrell, L. Mayes, and K. Grewen, 2012, p. 17-28. pdf

Jarcho, M.R., Mendoza, S.P., Mason, W.A., Yang, X., and Bales, K.L. (2011) Intranasal vasopressin affects pair-bonding and peripheral gene expression in male Callicebus cupreus. Genes, Brain, and Behavior 10: 375-383. PMCID: 3086990 pdf

Laugero, K.D., Smilowitz, J.T., German, J.B., Jarcho, M.R., Mendoza, S.P., and Bales, K.L. (2011) Plasma omega 3 polyunsaturated fatty acid status and monounsaturated fatty acids are altered by chronic social stress and predict endocrine responses to acute stress in titi monkeys.  Prostaglandins, Leukotrienes, and Essential Fatty Acids 84:71-78. PMID: 21211954 pdf


Stone, A.I., and Bales, K.L. (2010) Intergenerational transmission of the behavioral consequences of early experience in prairie voles.  Behavioural Processes 84: 732-738.  PMCID: 20457234 pdf

Hostetler, C.M., Harkey, S.L., and Bales, K.L. (2010) D2 antagonist during development decreases anxiety and infanticidal behavior in adult female prairie voles (Microtus ochrogaster). Behavioral Brain Research 210: 127-130. PMCID: 20152865 pdf

Stone, A.I., Mathieu, D., Griffin, L.L., Bales, K.L. (2010) Alloparenting experience affects future parental behavior and reproductive success in prairie voles. Behavioural Processes 83: 8-15. PMCID: 19732810 pdf

Campi, K.L., Bales,K.L., Grunewald, R.,  Krubitzer, L.A. (2010) Connections of auditory and visual cortex in the prairie vole (Microtus ochrogaster): Evidence for multisensory processing in primary sensory areas. Cerebral Cortex. 20: 89-108. PMCID: 19395525 pdf


Campbell, J.C., Laugero, K.D., van Westerhuyzen, J.A., Hostetler, C.M., Cohen, J.D., Bales, K.L. (2009) Costs of pair-bonding and parenting in male prairie voles (Microtus ochrogaster). Physiology & Behavior 98: 367-373. PMCID: 19576326 pdf

Zhang, Y., Xiao, Y., Bales, K.L. (2009) Primate social systems, scent-marking, and their applications in mobile and static sensor networks.  International Journal of Sensor Networks 5: 210-222.

Carter, C.S., Boone, E.M., Pournajafi-Nazarloo, H., Bales, K.L. (2009) The consequences of early experiences and exposure to oxytocin and vasopressin are sexually dimorphic. Developmental Neuroscience 31: 332-341. PMCID: 19546570 pdf


Carter, C.S.., Boone, E.M.,  Bales, K.L. (2008) Early experience and the developmental programming of oxytocin and vasopressin.  In Neurobiology of the Parental Brain, ed. R.S. Bridges. New York: Academic Press, pp. 417-433. pdf

Chau, M.J., Stone, A.I., Mendoza, S.P., Bales, K.L. (2008) Is play behavior sexually dimorphic in monogamous species? Ethology 114: 989-998. pdf

Samuel, P.A., Hostetler, C.M., Bales, K.L. (2008) Urocortin II increases spontaneous parental behavior in prairie voles (Microtus ochrogaster). Behavioural Brain Research 186: 284-288. PMCID: 2279148 pdf


Bales, K.L., Mason, W.A., Catana, C., Cherry, S.R., and Mendoza, S.P. (2007) Neural correlates of pair-bonding in a monogamous primate. Brain Research 1184: 245-253.  PMCID: 2387250 pdf

Bales, K.L., van Westerhuyzen, J.A., Lewis-Reese, A.D., Grotte, N., Lanter, J.A., Carter, C.S. (2007) Oxytocin has dose-dependent developmental effects on pair-bonding and alloparental care in female prairie voles.  Hormones and Behavior 52: 274-279.  PMCID: 1978481 pdf

Bales, K.L., Lewis-Reese, A.D., Pfeifer, L.A., Kramer, K.M., and Carter, C.S. (2007) Early Experience Affects the Traits of Monogamy in a Sexually Dimorphic Manner.  Developmental Psychobiology 49: 335-342. pdf

Campi, K., Carlen, S.J., Bales, K.L., and Krubitzer, L. (2007) The Organization of Sensory Neocortex in Prairie Voles (Microtus ochrogaster).  Journal of Comparative Neurology 502: 414-426. pdf

Bales, K.L., Plotsky, P.M., Young, L.J., Lim, M.M., Grotte, N., Ferrer, E., and Carter, C.S.(2007) Neonatal Oxytocin Manipulations Have Long-lasting, Sexually Dimorphic Effects on Vasopressin Receptors. Neuroscience 144: 38-45. pdf


Miller, K., Bales, K.L., Ramos, J., and Dietz, J.M. (2006) Energy Intake, Energy Expenditure, and Reproductive Costs of Female Wild Golden Lion Tamarins (Leontopithecus rosalia). American Journal of Primatology 68: 1037-1053. pdf

Tardif, S., Bales, K.L., Williams, L., Moeller, E.L., Abbott, D., Schultz-Darken, N., Mendoza, S., Mason, W., Bourgeois, S., and Ruiz, J. (2006)  Preparing New World Monkeys for Laboratory Research. ILAR Journal 47: 307-315. pdf

Bales, K.L., Kramer, K.M., Lewis-Reese, A.D., and Carter, C.S. (2006) Effects of Stress on Parental Care are Sexually Dimorphic in Prairie Voles. Physiology & Behavior 87: 424-429. pdf

Bales, K.L., French, J.A., McWilliams, J., Lake, R.A., Dietz, J.M. (2006) Effects of Social Status, Age, and Season on Androgen and Cortisol Levels in Wild Male Golden Lion Tamarins (Leontopithecus rosalia). Hormones and Behavior 49:88-95. pdf


Bales, K.L., French, J.A., Hostetler, C.M., Dietz, J.M. (2005) Social and Reproductive Factors Affecting Cortisol Levels in Wild Female Golden Lion Tamarins (Leontopithecus rosalia). American Journal of Primatology 67: 25-35. pdf


Bales, K.L., Abdelnabi, M., Cushing, B.S., Ottinger, M.A., and Carter, C.S. (2004) Effects of Neonatal Oxytocin Manipulations on Male Reproductive Potential in Prairie Voles. Physiology & Behavior 81: 519-526. pdf

Bales, K.L., Kim, A.J., Lewis-Reese, A.D, and Carter, C.S. (2004) Both Oxytocin and Vasopressin May Influence Alloparental Care in Male Prairie Voles.  Hormones and Behavior 45: 354-361. pdf

Tardif, S.D., and Bales, K.L. (2004) Relations Among Birth Condition, Maternal Condition, and Post-natal Growth in Captive Common Marmoset Monkeys (Callithrix jacchus). American Journal of Primatology 62:83-94. pdf

Bales, K.L., Pfeifer, L. A., and Carter, C.S. (2004) Sex Differences and Developmental Effects of Manipulations of Oxytocin on Alloparenting and Anxiety in Prairie Voles. Developmental Psychobiology 44:123-131. pdf


French, J.A., Bales, K.L., Baker, A.J., and Dietz, J.M. (2003) Endocrine Monitoring of Dominant and Subordinate Females in Free-Ranging Golden Lion Tamarins. International Journal of Primatology 24:1281- 1299. pdf

Bales, K., and Carter, C.S. (2003) Sex Differences and Developmental Effects of Oxytocin on Aggression and Social Behavior in Prairie Voles (Microtus ochrogaster). Hormones and Behavior, special issue on Aggressive and Violent Behavior 44:178-184. pdf

Bales, K., and Carter, C.S. (2003) Developmental Exposure to Oxytocin Facilitates Partner Preferences in Male Prairie Voles (Microtus ochrogaster). Behavioral Neuroscience 117:854-859. pdf

1997 – 2002

Bales, K., French, J.A., and J.M. Dietz (2002). Explaining Variation in Maternal Care in a Cooperatively Breeding Mammal. Animal Behaviour   63:453-461. pdf

Bales, K., O’Herron, M., Baker, A.J., and J.M. Dietz (2001) Sources of Variability in Numbers of Live Births in Wild Golden Lion Tamarins (Leontopithecus rosalia). American Journal of Primatology 54:211-221. pdf

Bales, K., Dietz, J., Baker, A., Miller, K., and S.D. Tardif (2000). The Effects of Allocaregivers on Fitness of Infants and Parents in Callitrichid Primates.  Folia Primatologica 71:27-38.

Tardif, S.D., Jaquish, C., Layne, D., Bales, K., Power, M., Power, R., and Oftedal, O. (1998).  Growth Variation in Common Marmoset Monkeys (Callithrix jacchus) fed a Purified Diet: Relation to Care-giving and Weaning Behaviors. Laboratory Animal Science 48:264-269. pdf

Tardif, S.D., and K. Bales (1997) Is Infant Carrying a Courtship Strategy in Captive Callitrichids? Animal Behaviour 53: 1001 – 1007. pdf


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