Do Zombies Dream of Undead Sheep? A Neuroscientific View of the Zombie Brain Timothy Verstynen & Bradley Voytek. Princeton University Press (2014). * Winner of the 2015 PROSE award for Biomedicine & Neuroscience.


"Level set trees for applied statistics" Brian Kent (Ph.D., CMU Statistics, 2012) (pdf)

"A biologically motivated synthesis of accumulator and reinforcement-learning models for describing adaptive decision-making" Kyle Dunovan (Ph.D., Pitt Psychology, 2016) (pdf)

"Cortical and behavioral signatures of response binding during sequential skill learning" Patrick Beukema (Ph.D., Pitt Neuroscience, 2018) (pdf)

"A corticostriatal mechanism for integrating subjective value into processes of risky decision-making" Kevin Jarbo (Ph.D., CMU Psychology, 2018) (pdf)


Peterson, E. J., & Verstynen, T. D. (2019). A way around the exploration-exploitation dilemma. bioRxiv, 671362. (pdf)

Agarwal, A., Kumar, V., Dunovan, K., Peterson, E., Verstynen, T., & Sycara, K. (2018). Better Safe than Sorry: Evidence Accumulation Allows for Safe Reinforcement Learning. arXiv preprint arXiv:1809.09147. (pdf)

Millette, A., Lynch, B., & Verstynen, T. (2017). Abdominal adiposity negatively associates with the rate of long term sequential skill learning. BioRxiv, 186742. (pdf)

Verstynen, T., & Sabes, P. N. (2011). An analysis of the emergence of adaptive Bayesian priors from Hebbian learning in a simple attractor network model. arXiv preprint arXiv:1106.2977. (pdf)

Journal Publications & Chapters


Banihashemi, L., Peng, C. W., Verstynen, T., Wallace, M. L., Lamont, D. N., Alkhars, H. M., ... & Germain, A. (2021). Opposing relationships of childhood threat and deprivation with stria terminalis white matter. Human brain mapping, 42(8), 2445-2460. (pdf)

Rubin, J. E., Vich, C., Clapp, M., Noneman, K., & Verstynen, T. (2021). The credit assignment problem in cortico‐basal ganglia‐thalamic networks: A review, a problem and a possible solution. European Journal of Neuroscience, 53(7), 2234-2253. (pdf)

Rasero, J., Sentis, A. I., Yeh, F. C., & Verstynen, T. (2021). Integrating across neuroimaging modalities boosts prediction accuracy of cognitive ability. PLoS computational biology, 17(3), e1008347. (pdf)

Skrzynski, C. J., Creswell, K. G., Verstynen, T., Bachrach, R. L., & Chung, T. (2021). The influence of negative mood on solitary drinking preference: An experiment with young adult solitary drinkers. PloS one, 16(2), e0247202. (pdf)


Verstynen, T., Dunovan, K., Walsh, C., Kuan, C. H., Manuck, S. B., & Gianaros, P. J. (2020). Adiposity covaries with signatures of asymmetric feedback learning during adaptive decisions. Social Cognitive and Affective Neuroscience, 15(10), 1145-1156. (pdf)

Jarbo, K., Colaço, D., & Verstynen, T. (2020). Contextual framing of loss impacts harm avoidance during risky spatial decisions. Journal of Behavioral Decision Making, 33(5), 657-670. (pdf)

Peterson, E., Müyesser, N. A., Verstynen, T., & Dunovan, K. (2020). Combining Imagination and Heuristics to Learn Strategies that Generalize. Neurons, Behavior, Data analysis, and Theory, 3(4), 1-19. (pdf)

Gianaros, P. J., Kraynak, T. E., Kuan, D. C., Gross, J. J., McRae, K., Hariri, A. R., ... & Verstynen, T. D. (2020). Affective brain patterns as multivariate neural correlates of cardiovascular disease risk. Social cognitive and affective neuroscience, 15(10), 1034-1045. (pdf)

Vich, C., Dunovan, K., Verstynen, T., & Rubin, J. (2020). Corticostriatal synaptic weight evolution in a two-alternative forced choice task. Communications in Nonlinear Science and Numerical Simulation, 105048. (pdf)


Bañuelos, C., & Verstynen, T. (2019). How structural brain network topologies associate with cognitive abilities in a value-based decision-making task. Impulse: The Premier Journal for Undergraduate Publications in the Neurosciences. (pdf)

Erickson, K. I., Grove, G. A., Burns, J. M., Hillman, C. H., Kramer, A. F., McAuley, E., ... & Huang, H. (2019). Investigating gains in neurocognition in an Intervention Trial of Exercise (IGNITE): Protocol. Contemporary clinical trials, 85, 105832. (pdf)

Beukema, P., Diedrichsen, J., & Verstynen, T. (2019). Binding during sequence learning does not alter cortical representations of individual actions. Journal of Neuroscience, 2669-18. (pdf)

Ashourvan, A., Telesford, Q. K., Verstynen, T., Vettel, J. M., & Bassett, D. S. (2019). Multi-scale detection of hierarchical community architecture in structural and functional brain networks. PloS one, 14(5), e0215520.(pdf)

Dunovan, K., Vich, C., Clapp, M., Verstynen, T., & Rubin, J. (2019). Reward-driven changes in striatal pathway competition shape evidence evaluation in decision-making. PLoS computational biology, 15(5), e1006998. (pdf)

Bansal, K., Garcia, J. O., Tompson, S. H., Verstynen, T., Vettel, J. M., & Muldoon, S. F. (2019). Cognitive chimera states in human brain networks. Science Advances, 5(4), eaau8535. (pdf)

Dunovan, K., & Verstynen, T. (2019). Errors in action timing and inhibition facilitate learning by tuning distinct mechanisms in the underlying decision process. Journal of Neuroscience, 39(12), 2251-2264. (pdf)


Peterson, E., Myesser, N. A., Verstynen, T., & Dunovan, K. (2018). Keep it stupid simple. CoRR. (pdf)

Balakrishnan, S., Choe, Y. J., Singh, A., Vettel, J., & Verstynen, T. (2018). Local White Matter Architecture Defines Functional Brain Dynamics. In 2018 IEEE International Conference on Systems, Man, and Cybernetics (SMC) (pp. 595-602). IEEE. (pdf) * Co-winner of Franklin V. Taylor Memorial Award

Yeh, F. C., Panesar, S., Fernandes, D., Meola, A., Yoshino, M., Fernandez-Miranda, J. C., ... & Verstynen, T. (2018). Population-averaged atlas of the macroscale human structural connectome and its network topology. NeuroImage, 178, 57-68. (pdf)

Porter, A., Leckie, R., & Verstynen, T. (2018). White matter pathways as both a target and mediator of health behaviors. Annals of the New York Academy of Sciences, 1428(1), 71-88. (pdf)

Jarbo, K., Flemming, R., & Verstynen, T. D. (2018). Sensory uncertainty impacts avoidance during spatial decisions. Experimental brain research, 236(2), 529-537. (pdf)

Beukema, P., & Verstynen, T. (2018). Predicting and binding: interacting algorithms supporting the consolidation of sequential motor skills. Current opinion in behavioral sciences, 20, 98-103.(pdf)

Powell, M. A., Garcia, J. O., Yeh, F. C., Vettel, J. M., & Verstynen, T. (2018). Local connectome phenotypes predict social, health, and cognitive factors. Network neuroscience, 2(1), 86-105. (pdf)


Larsen, B., Verstynen, T. D., Yeh, F. C., & Luna, B. (2017). Developmental changes in the integration of affective and cognitive corticostriatal pathways are associated with reward-driven behavior. Cerebral Cortex, 28(8), 2834-2845. (pdf)

Vettel, J. M., Cooper, N., Garcia, J. O., Yeh, F. C., & Verstynen, T. D. (2017). White Matter Tractography and Diffusion‐Weighted Imaging. eLS, 1-9. (pdf)

Gianaros, P. J., Sheu, L. K., Uyar, F., Koushik, J., Jennings, J. R., Wager, T. D., ... & Verstynen, T. D. (2017). A brain phenotype for stressor‐evoked blood pressure reactivity. Journal of the American Heart Association, 6(9), e006053. (pdf)

Lynch, B., Beukema, P., & Verstynen, T. (2017). Differentiating visual from response sequencing during long-term skill learning. Journal of cognitive neuroscience, 29(1), 125-136. (pdf)


Muraskin, J., Dodhia, S., Lieberman, G., Garcia, J. O., Verstynen, T., Vettel, J. M., ... & Sajda, P. (2016). Brain dynamics of post‐task resting state are influenced by expertise: Insights from baseball players. Human brain mapping, 37(12), 4454-4471. (pdf)

Yeh, F. C., Vettel, J. M., Singh, A., Poczos, B., Grafton, S. T., Erickson, K. I., ... & Verstynen, T. D. (2016). Quantifying differences and similarities in whole-brain white matter architecture using local connectome fingerprints. PLoS computational biology, 12(11), e1005203.(pdf)

Lichenstein, S. D., Verstynen, T., & Forbes, E. E. (2016). Adolescent brain development and depression: a case for the importance of connectivity of the anterior cingulate cortex. Neuroscience & Biobehavioral Reviews, 70, 271-287. (pdf)

Yeh, F. C., & Verstynen, T. D. (2016). Converting multi-shell and diffusion spectrum imaging to high angular resolution diffusion imaging. Frontiers in neuroscience, 10, 418. (pdf)

Lichenstein, S. D., Bishop, J. H., Verstynen, T. D., & Yeh, F. C. (2016). Diffusion capillary phantom vs. human data: outcomes for reconstruction methods depend on evaluation medium. Frontiers in neuroscience, 10, 407. (pdf)

Muraskin, J., Sherwin, J., Lieberman, G., Garcia, J. O., Verstynen, T., Vettel, J. M., & Sajda, P. (2016). Fusing multiple neuroimaging modalities to assess group differences in perception–action coupling. Proceedings of the IEEE, 105(1), 83-100. (pdf)

Barredo, J., Verstynen, T. D., & Badre, D. (2016). Organization of cortico-cortical pathways supporting memory retrieval across subregions of the left ventrolateral prefrontal cortex. Journal of neurophysiology, 116(3), 920-937. (pdf)

Oberlin, L. E., Verstynen, T. D., Burzynska, A. Z., Voss, M. W., Prakash, R. S., Chaddock-Heyman, L., ... & Phillips, S. M. (2016). White matter microstructure mediates the relationship between cardiorespiratory fitness and spatial working memory in older adults. Neuroimage, 131, 91-101. (pdf)

Dunovan, K., & Verstynen, T. (2016). Believer-Skeptic meets Actor-Critic: Rethinking the role of basal ganglia pathways during decision-making and reinforcement learning. Frontiers in neuroscience, 10, 106. (pdf)

Yeh, F. C., Badre, D., & Verstynen, T. (2016). Connectometry: a statistical approach harnessing the analytical potential of the local connectome. Neuroimage, 125, 162-171. (pdf)


Rofey, D. L., Arslanian, S. A., El Nokali, N. E., Verstynen, T., Watt, J. C., Black, J. J., ... & Erickson, K. I. (2015). Brain volume and white matter in youth with type 2 diabetes compared to obese and normal weight, non-diabetic peers: A pilot study. International Journal of Developmental Neuroscience, 46, 88-91. (pdf)

Beukema, P., Yeh, F. C., & Verstynen, T. (2015). In vivo characterization of the connectivity and subcomponents of the human globus pallidus. NeuroImage, 120, 382-393. (pdf)

Dunovan, K., Lynch, B., Molesworth, T., & Verstynen, T. (2015). Competing basal ganglia pathways determine the difference between stopping and deciding not to go. Elife, 4, e08723. (pdf)

Verstynen, T. D. (2015). How form constrains function in the human brain. Emerging Trends in the Social and Behavioral Sciences: An Interdisciplinary, Searchable, and Linkable Resource, 1-16. (pdf)

Fernández-Miranda, J. C., Wang, Y., Pathak, S., Stefaneau, L., Verstynen, T., & Yeh, F. C. (2015). Asymmetry, connectivity, and segmentation of the arcuate fascicle in the human brain. Brain Structure and Function, 220(3), 1665-1680. (pdf)

Jarbo, K., & Verstynen, T. D. (2015). Converging structural and functional connectivity of orbitofrontal, dorsolateral prefrontal, and posterior parietal cortex in the human striatum. Journal of Neuroscience, 35(9), 3865-3878. (pdf)

Molesworth, T., Sheu, L. K., Cohen, S., Gianaros, P. J., & Verstynen, T. D. (2015). Social network diversity and white matter microstructural integrity in humans. Social cognitive and affective neuroscience, 10(9), 1169-1176. (pdf)


Erickson, K. I., Creswell, J. D., Verstynen, T. D., & Gianaros, P. J. (2014). Health neuroscience: defining a new field. Current directions in psychological science, 23(6), 446-453. (pdf)

Schlerf, J., Wiestler, T., Verstynen, T., & Diedrichsen, J. (2014). Big challenges from the little brain—imaging the cerebellum. In Advanced Brain Neuroimaging Topics in Health and Disease-Methods and Applications. IntechOpen. (pdf)

Kent, B. P., Rinaldo, A., Yeh, F. C., & Verstynen, T. (2014). Mapping topographic structure in white matter pathways with level set trees. PloS one, 9(4), e93344. (pdf)

Oliva, J., Póczos, B., Verstynen, T., Singh, A., Schneider, J., Yeh, F. C., & Tseng, W. Y. (2014). Fusso: Functional shrinkage and selection operator. In Artificial Intelligence and Statistics (pp. 715-723). (pdf)

Verstynen, T. D. (2014). The organization and dynamics of corticostriatal pathways link the medial orbitofrontal cortex to future behavioral responses. Journal of neurophysiology, 112(10), 2457-2469. (pdf) (data)


Ryan, J. P., Sheu, L. K., Verstynen, T. D., Onyewuenyi, I. C., & Gianaros, P. J. (2013). Cerebral blood flow links insulin resistance and baroreflex sensitivity. PloS one, 8(12), e83288. (pdf)

Yeh, F. C., Verstynen, T. D., Wang, Y., Fernández-Miranda, J. C., & Tseng, W. Y. I. (2013). Deterministic diffusion fiber tracking improved by quantitative anisotropy. PloS one, 8(11), e80713. (pdf)

Verstynen, T. D., Weinstein, A., Erickson, K. I., Sheu, L. K., Marsland, A. L., & Gianaros, P. J. (2013). Competing physiological pathways link individual differences in weight and abdominal adiposity to white matter microstructure. Neuroimage, 79, 129-137. (pdf)

Pyles, J. A., Verstynen, T. D., Schneider, W., & Tarr, M. J. (2013). Explicating the face perception network with white matter connectivity. PloS one, 8(4), e61611. (pdf)


Verstynen, T., Phillips, J., Braun, E., Workman, B., Schunn, C., & Schneider, W. (2012). Dynamic sensorimotor planning during long-term sequence learning: the role of variability, response chunking and planning errors. PLoS One, 7(10), e47336. (pdf)

Verstynen, T. D., Weinstein, A. M., Schneider, W. W., Jakicic, J. M., Rofey, D. L., & Erickson, K. I. (2012). Increased body mass index is associated with a global and distributed decrease in white matter microstructural integrity. Psychosomatic medicine, 74(7), 682. (pdf)

Wang, Y., Fernández-Miranda, J. C., Verstynen, T., Pathak, S., Schneider, W., & Yeh, F. C. (2012). Rethinking the role of the middle longitudinal fascicle in language and auditory pathways. Cerebral cortex, 23(10), 2347-2356. (pdf)

Gianaros, P. J., Marsland, A. L., Sheu, L. K., Erickson, K. I., & Verstynen, T. D. (2012). Inflammatory pathways link socioeconomic inequalities to white matter architecture. Cerebral cortex, 23(9), 2058-2071. (pdf)

Fernandez-Miranda, J. C., Pathak, S., Engh, J., Jarbo, K., Verstynen, T., Yeh, F. C., ... & Friedlander, R. (2012). High-definition fiber tractography of the human brain: neuroanatomical validation and neurosurgical applications. Neurosurgery, 71(2), 430-453. (pdf)

Verstynen, T. D., Badre, D., Jarbo, K., & Schneider, W. (2012). Microstructural organizational patterns in the human corticostriatal system. Journal of neurophysiology, 107(11), 2984-2995. (pdf)

Greenberg, A. S., Verstynen, T., Chiu, Y. C., Yantis, S., Schneider, W., & Behrmann, M. (2012). Visuotopic cortical connectivity underlying attention revealed with white-matter tractography. Journal of Neuroscience, 32(8), 2773-2782. (pdf)

Jarbo, K., Verstynen, T., & Schneider, W. (2012). In vivo quantification of global connectivity in the human corpus callosum. Neuroimage, 59(3), 1988-1996. (pdf)

Verstynen, T. D., Lynch, B., Miller, D. L., Voss, M. W., Prakash, R. S., Chaddock, L., ... & Fanning, J. (2012). Caudate nucleus volume mediates the link between cardiorespiratory fitness and cognitive flexibility in older adults. Journal of aging research, 2012. (pdf)


Verstynen, T., & Ivry, R. B. (2011). Network dynamics mediating ipsilateral motor cortex activity during unimanual actions. Journal of cognitive neuroscience, 23(9), 2468-2480. (pdf)

Verstynen, T., & Sabes, P. N. (2011). How each movement changes the next: an experimental and theoretical study of fast adaptive priors in reaching. Journal of Neuroscience, 31(27), 10050-10059. (pdf) Supplemental analysis of learning dynamics (pdf).

Verstynen, T. D., & Deshpande, V. (2011). Using pulse oximetry to account for high and low frequency physiological artifacts in the BOLD signal. Neuroimage, 55(4), 1633-1644. (pdf)


Verstynen, T., Jarbo, K., Pathak, S., & Schneider, W. (2010). In vivo mapping of microstructural somatotopies in the human corticospinal pathways. Journal of neurophysiology, 105(1), 336-346. (pdf)

Oliveira, F. T., Diedrichsen, J., Verstynen, T., Duque, J., & Ivry, R. B. (2010). Transcranial magnetic stimulation of posterior parietal cortex affects decisions of hand choice. Proceedings of the National Academy of Sciences, 107(41), 17751-17756. (pdf)

Diedrichsen, J., Verstynen, T., Schlerf, J., & Wiestler, T. (2010). Advances in functional imaging of the human cerebellum. Current opinion in neurology, 23(4), 382-387.(pdf)

Schlerf, J. E., Verstynen, T. D., Ivry, R. B., & Spencer, R. M. (2010). Evidence of a novel somatopic map in the human neocerebellum during complex actions. Journal of neurophysiology, 103(6), 3330-3336. (pdf)

Verstynen, T., Oliver, M., & IVRY, R. B. (2010). Experiencing the future: the influence of self-initiation on temporal perception. Space and Time in Perception and Action. (pdf)


Miller, B. T., Verstynen, T., Johnson, M. K., & D'Esposito, M. (2008). Prefrontal and parietal contributions to refreshing: an rTMS study. Neuroimage, 39(1), 436-440. (pdf)

Esterman, M., Prinzmetal, W., DeGutis, J., Landau, A., Hazeltine, E., Verstynen, T., & Robertson, L. (2008). Voluntary and involuntary attention affect face discrimination differently. Neuropsychologia, 46(4), 1032-1040. (pdf)


Spencer, R. M., Verstynen, T., Brett, M., & Ivry, R. (2007). Cerebellar activation during discrete and not continuous timed movements: an fMRI study. Neuroimage, 36(2), 378-387. (pdf)

Diedrichsen, J.*, Verstynen, T.*, Hon, A., Zhang, Y., & Ivry, R. B. (2007). Illusions of force perception: the role of sensori-motor predictions, visual information, and motor errors. Journal of neurophysiology, 97(5), 3305-3313. *Co-First Authors. (pdf)

Esterman, M., Verstynen, T., & Robertson, L. C. (2007). Attenuating illusory binding with TMS of the right parietal cortex. Neuroimage, 35(3), 1247-1255. (pdf)

Verstynen, T., Spencer, R., Stinear, C. M., Konkle, T., Diedrichsen, J., Byblow, W. D., & Ivry, R. B. (2007). Ipsilateral corticospinal projections do not predict congenital mirror movements: a case report. Neuropsychologia, 45(4), 844-852. (pdf)


Verstynen, T., Konkle, T., & Ivry, R. B. (2006). Two types of TMS-induced movement variability after stimulation of the primary motor cortex. Journal of neurophysiology, 96(3), 1018-1029. (pdf)

Esterman, M., Verstynen, T., Ivry, R. B., & Robertson, L. C. (2006). Coming unbound: disrupting automatic integration of synesthetic color and graphemes by transcranial magnetic stimulation of the right parietal lobe. Journal of cognitive neuroscience, 18(9), 1570-1576. (pdf)


Diedrichsen, J., Verstynen, T., Lehman, S. L., & Ivry, R. B. (2005). Cerebellar involvement in anticipating the consequences of self-produced actions during bimanual movements. Journal of Neurophysiology, 93(2), 801-812. (pdf)

Verstynen, T.*, Diedrichsen, J.*, Albert, N., Aparicio, P., & Ivry, R. B. (2005). Ipsilateral motor cortex activity during unimanual hand movements relates to task complexity. Journal of neurophysiology, 93(3), 1209-1222. *Co-First Authors (pdf)


Diedrichsen, J., Verstynen, T., Hon, A., Lehman, S. L., & Ivry, R. B. (2003). Anticipatory adjustments in the unloading task: Is an efference copy necessary for learning?. Experimental Brain Research, 148(2), 272-276. (pdf)


Tang, A. C., & Verstynen, T. (2002). Early life environment modulates ‘handedness’ in rats. Behavioural brain research, 131(1-2), 1-7.(pdf)


Verstynen, T., Tierney, R., Urbanski, T., & Tang, A. (2001). Neonatal novelty exposure modulates hippocampal volumetric asymmetry in the rat. Neuroreport, 12(14), 3019-3022.(pdf)

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