Wen X, Hassani AS, Neumann A, Tiesinga P & Womelsdorf T (2025) Two Distinct Attentional Priorities Guide Exploratory and Exploitative Gaze in Parallel. p. 1-27, bioRxiv. doi: 10.1101/2025.09.20.677503 pdf

Tiesinga P, Womelsdorf T (2025) Parallel Belief States account for Learning and Updating of Attentional Priorities in Multidimensional Environments. 1-24, bioRxiv. doi: 10.1101/2025.09.03.674042 pdf

Gerrity CG, Treuting RL, Peters RA & Womelsdorf T (2025) Neuronal Decoding of Decisions in Multidimensional Feature Space Using a Gated Recurrent Variational Autoencoder. 1-12. bioRxiv. doi: 10.1101/2025.08.20.671126 pdf

Treuting RL, Banaie Boroujeni K, Gerrity CG, Neumann A, Tiesinga P & Womelsdorf T (2025) Adaptive Reinforcement Learning is causally supported by Anterior Cingulate Cortex and Striatum. Neuron; 113, 1–16. https://doi.org/10.1016/j.neuron.2025.05.018 pdf

Cheung SM; Neumann A, Womelsdorf T (2025) Assessing Attentiveness and Cognitive Engagement across Tasks using Video-based Action Understanding in Non-Human Primates. 1–28, bioRxiv doi: https://doi.org/10.1101/2025.05.31.657183

Wen X, Malchin LA, Womelsdorf T (2025) A Toolbox for Generating Multidimensional 3-D Objects with Fine-Controlled Feature Space: Quaddle 2.0. Behavioral Research Methods, 1–22, doi: 10.1101/2024.12.19.629479

Hassani SA, Tiesinga P, & Womelsdorf T (2024) Noradrenergic alpha-2a Receptor Stimulation Enhances Prediction Error Signaling and Updating of Attention Sets in Anterior Cingulate Cortex and Striatum. Nature Communication. 15:9905. pdf

Dockum A, Phipps MA, Newman M, Kusunose J, Treuting RL, Neumann A, Womelsdorf T, Caskey C (2024) Image Guidance of a Phased Array Transducer for Neuromodulation with a Small Focal Volume. IEEE Transactions on Ultrasonics, Ferroelectrics, and Frequency Control (UFFC-JS) (pp. 1:5).

Banaie Boroujeni K, Womelsdorf T (2023) Routing States Transition During Oscillatory Bursts and Attention States. Neuron 111(18), 2929–2944. doi:https://doi.org/10.1016/j.neuron.2023.06.012

Watson M, Traczewski N, Dunghana S, Banaie Boroujeni K, Neumann A, Wen X, Womelsdorf T (2023) A multi-task platform for profiling cognitive and motivational constructs in humans and nonhuman primates. 1–48; bioRxiv; https://www.biorxiv.org/content/10.1101/2023.11.09.566422v1

Hassani SA, Neumann A, Russell J, Jones CK, Womelsdorf T (2023) M₁ selective muscarinic allosteric modulation enhances cognitive flexibility and effective salience in nonhuman primates. PNAS, www.pnas.org/doi/10.1073/pnas.2216792120.

Sigona MK, Manuel TJ, Phipps MA, Banaie Boroujeni K, Treuting RL, Womelsdorf T, Caskey CF (2023) Generating Patient-Specific Acoustic Simulations for Transcranial Focused Ultrasound Procedures based on Optical Tracking Information. IEEE Open Journal of Ultrasonics, Ferroelectrics, and Frequency Control. In press.

Hassani SA, Lendor S, Neumann A, Sinha Roy K, Banaie Boroujeni K, Hoffman K, Pawliszyn J, Womelsdorf T (2023) Dose-dependent dissociation of pro-cognitive effects of donepezil on attention and cognitive flexibility in rhesus monkeys. Biological Psychiatry GOS. 3(1), 68–77.

Banaie Boroujeni K, Sigona MK, Treuting RL, Manuel TJ, Caskey CF, Womelsdorf T (2022) The anterior cingulate cortex causally supports flexible learning under motivationally challenging and cognitively demanding conditions. PLoS Biology. doi: https://doi.org/10.1101/2021.08.04.455080.

Banaie Boroujeni K, Watson MR, Womelsdorf T (2022) Gains and Losses Differentially Regulate Learning at Low and High Attentional Load. Journal of Cognitive Neuroscience. 34(10), 1952–1971.

Womelsdorf T, Watson MR, Tiesinga P (2022) Learning at variable attentional load requires cooperation between working memory, meta-learning and attention-augmented reinforcement learning. Journal of Cognitive Neuroscience. 34(1), 79–107.

Banaie Boroujeni K, Tiesinga P, Womelsdorf T (2021) Interneuron Specific Gamma Synchronization Indexes Cue Uncertainty and Prediction Errors in Lateral Prefrontal and Anterior Cingulate Cortex. eLife. doi: 10.7554/eLife.69111.

Womelsdorf T, Thomas C, Neumann A, Watson MR, Banaie Boroujeni K, Hassani SA, Parker J, Hoffman KL (2021) A Kiosk Station for the Assessment of Multiple Cognitive Domains and Cognitive Enrichment of Monkeys. Frontiers in Behavioral Neuroscience. https://doi.org/10.3389/fnbeh.2021.721069.

Alexander WH, Womelsdorf T (2021) Interactions of medial and lateral prefrontal cortex in hierarchical predictive coding. Frontiers in Computational Neuroscience. https://doi.org/10.3389/fncom.2021.605271.

Womelsdorf T (2021) Translating Expectation into Visual Selection through a Beta Synchronous Fronto-Parietal Neural Subnetwork. Neuron. 109(1), 8–10. https://doi.org/10.1016/j.neuron.2020.12.011.

Voloh B, Oemisch M, Womelsdorf T (2020) Phase of Firing Coding of Learning Variables across the Fronto-Striatal Network during Feature Learning. Nature Communications. 11:4669, 1–16. https://doi.org/10.1101/2019.12.12.874859.

Banaie Boroujeni K, Oemisch M, Hassani SA, Womelsdorf T (2020) Fast Spiking Interneuron Activity in Primate Striatum Tracks Learning of Attention Cues. PNAS, Proceedings of the National Academy of Sciences, USA. 117(30), 18049–58. https://doi.org/10.1073/pnas.2001348117.

Javitt DC, Siegel S, Spencer K, Uhlhaas P, Mathalon D, Hong E, Martinez A, Ehlers C, Abbas AI, Teichert T, Lakatos P, Womelsdorf T (2020) A Roadmap for Development of Neuro-Oscillations as Translational Biomarkers in Neuropsychopharmacology. Neuropsychopharmacology. (in press).

Onorato I, Neuenschwander S, Lima B, Rocha KS, Broggini AC, Uran C, Spyropoulos G, Womelsdorf T, Fries P, Niell C, Singer W, Vinck M (2020) A distinct class of bursting neurons with strong gamma synchronization and stimulus selectivity in monkey V1. Neuron. (in press).

Voloh B, Watson MR, König S, Womelsdorf T (2020) MAD saccade: statistically robust saccade threshold estimation via the median absolute deviation. Journal of Eye Movement Research. 12(8):3.

Banaie Boroujeni K, Tiesinga P, Womelsdorf T (2020) Adaptive spike-artifact removal from local field potentials uncovers prominent beta and gamma band neuronal synchronization. Journal of Neuroscience Methods. doi: 10.1016/j.jneumeth.2019.108485.

Azimi M, Oemisch M, Womelsdorf T (2019) Dissociation of Nicotinic alpha-7 and alpha-4/beta2 Sub-receptor Agonists for Enhancing Learning and Attentional Filtering in Nonhuman Primates. Psychopharmacology. 237, 997–1010.

Hassani SA, Lendor S, Boyaci E, Pawliszyn J, Womelsdorf T (2019) Multi-Neuromodulator Measurements across Fronto-Striatal Network Areas of the Behaving Macaque using Solid-Phase Microextraction. Journal of Neurophysiology. (in press). doi: 10.1152/jn.00321.2019.

Watson M, Voloh B, Thomas C, Hasan A, Womelsdorf T (2019) USE: An integrative suite for temporally precise psychophysical experiments in virtual environments in human, nonhuman and artificially intelligent agents. Journal of Neuroscience Methods. 326:108374. doi: 10.1016/j.jneumeth.2019.108374.

Oemisch M, Westendorff S, Azimi M, Hassani SA, Ardid S, Tiesinga P, Womelsdorf T (2019) Feature Specific Prediction Errors and Surprise across Macaque Fronto-Striatal Circuits. Nature Communications. 10(1), 176. doi: 10.1101/266205.

Lendor S, Hassani SA, Boyaci E, Singh V, Womelsdorf T, Pawliszyn J (2019) Development of solid phase microextraction-based miniaturized probe and protocol for extraction of neurotransmitters from brain in vivo. Analytical Chemistry. 91(7), 4896–4905. doi: 10.1021/acs.analchem.9b00995.

Womelsdorf T (2019) Neocortical Cell Classification: Essential Contributions from Electrophysiology. Current Biology. (in press).

Nácher V, Hassani SA, Womelsdorf T (2019) Asymmetric effective connectivity between primate anterior cingulate and lateral prefrontal cortex revealed by electrical microstimulation. Brain Structure and Function. 224(2), 779–793. doi: 10.1007/s00429-018-1806-y.

Watson M, Voloh B, Naghizadeh M, Womelsdorf T (2019) Quaddles: A multidimensional 3D object set with parametrically controlled and customizable features. Behavior Research Methods. 51, 2522–2532. https://doi.org/10.1101/194381.

Womelsdorf T, Hoffman K (2018) Latent Connectivity: Neuronal Oscillations Can Be Leveraged for Transient Plasticity. Current Biology. 28(16), R879–R882. doi: 10.1016/j.cub.2018.06.073.

Chang M, Dian JA, Dufour S, Wang L, Moradi-Chameh H, Ramani M, Zhang L, Carlen PL, Womelsdorf T, Valiante TA (2018) Brief activation of GABAergic interneurons initiates the transition to ictal events through post-inhibitory rebound excitation. Neurobiology of Disease. https://doi.org/10.1016/j.nbd.2017.10.007.

Voloh B, Womelsdorf T (2017) Cell-type specific burst firing interacts with theta and beta activity in prefrontal cortex during attention states. Cerebral Cortex. https://doi.org/10.1093/cercor/bhx287.

Hassani SA, Oemisch M, Balcarras M, Westendorff S, Ardid S, van der Meer MA, Tiesinga P, Womelsdorf T (2017) A computational psychiatry approach identifies how alpha-2A noradrenergic agonist Guanfacine affects feature-based reinforcement learning in the macaque. Scientific Reports. doi: 10.1038/srep40606.

Babapoor-Farrokhran S, Vinck M, Womelsdorf T, Everling S (2017) Theta and beta synchrony coordinate frontal eye fields and anterior cingulate cortex during sensorimotor mapping. Nature Communications. doi: 10.1038/ncomms13967.

Oemisch M, Watson MR, Womelsdorf T, Schubö A (2017) Changes of attention during value-based reversal learning are tracked by N2pc and feedback-related negativity. Frontiers in Neuroscience. 11, 540.

Westendorff S, Kaping D, Everling S, Womelsdorf T (2016) Prefrontal and anterior cingulate cortex neurons encode attentional targets even when they do not apparently bias behavior. Journal of Neurophysiology. (in press). doi: 10.1152/jn.00027.2016. pdf

Balcarras M, Ardid S, Kaping D, Everling S, Womelsdorf T (2016) Attentional selection can be predicted by reinforcement learning of task-relevant stimulus features weighted by value-independent stickiness. Journal of Cognitive Neuroscience. 28(2), 333–349. pdf

Lewis C, Bosmann C, Womelsdorf T, Fries P (2016) Stimulus-induced visual cortical networks are recapitulated by spontaneous local and inter-areal synchronization. PNAS, Proceedings of the National Academy of Sciences, USA. (in press). pdf

Skoblenick K, Womelsdorf T, Everling S (2016) Ketamine alters outcome-related local field potentials in monkey prefrontal cortex. Cerebral Cortex. 26(6), 2743–2752. pdf

Voloh B, Womelsdorf T (2016) A role of phase-resetting in coordinating large scale neural oscillations during attention and goal-directed behavior. Frontiers in Systems Neuroscience. doi: 10.3389/fnsys.2016.00018. pdf.

Lewis CM, Bosman CA, Brunet N, Lima B, Roberts MJ, Womelsdorf T, de Weerd P, Neuenschwander S, Singer W, Fries P (2016) Two frequency bands contain the most stimulus-related information in visual cortex. bioRxiv. pdf.

Balcarras M, Womelsdorf T (2016) A flexible mechanism of rule selection enables rapid feature-based reinforcement learning. Frontiers in Neuroscience. doi: 10.3389/fnins.2016.00125. pdf.

Womelsdorf T, Everling S (2015) Long-Range Attention Networks: Circuit Motifs Underlying Endogenously Controlled Stimulus Selection. Trends in Neurosciences. 38(11): 682–700. pdf (or via RG).

Oemisch M, Westendorff S, Everling S, Womelsdorf T (2015) Inter-areal spiketrain correlations of anterior cingulate and dorsal prefrontal cortex during attention shifts. Journal of Neuroscience. 35(38): 13076–13089. pdf.

Leonard TK, Mikkila J, Gerrard JL, Kaping D, Patel S, Eskandar E, Womelsdorf T, Hoffman KL (2015) Sharp wave ripples during visual exploration in the primate hippocampus. Journal of Neuroscience. 35(44): 14771–1482. link to H-lab.

Voloh B, Valiante TA, Everling S, Womelsdorf T (2015) Theta-gamma coordination between anterior cingulate and prefrontal cortex indexes correct attention shifts. PNAS, Proceedings of the National Academy of Sciences, USA. 112: 8457–8462. pdf.

Micheli C, Kaping D, Westendorff S, Valiante TA, Womelsdorf T (2015) Inferior-frontal cortex phase synchronizes with the temporal-parietal junction prior to successful change detection. NeuroImage. 119: 417–431. pdf.

Ardid S, Vinck M, Kaping D, Marquez S, Everling S, Womelsdorf T (2015) Mapping of functionally characterized cell classes onto canonical circuit operations in primate prefrontal cortex. Journal of Neuroscience. 35(7): 2975–2991. pdf.

Brunet N, Bosman C, Roberts M, Oostenveld R, Womelsdorf T, de Weerd P, Fries P (2015) Visual cortical gamma band activity during free viewing of natural images. Cerebral Cortex. 25(4): 918–926. pdf.

Shen C, Ardid S, Kaping D, Westendorff S, Everling S, Womelsdorf T (2015) Anterior cingulate cortex cells identify process-specific errors of attentional control prior to transient prefrontal-cingulate inhibition. Cerebral Cortex. 25(8): 2213–2228. pdf.

Chan J, Koval M, Womelsdorf T, Lomber S, Everling S (2015) Dorsolateral prefrontal cortex deactivation in monkeys reduces preparatory beta and gamma power in the superior colliculus. Cerebral Cortex. 25(12): 4704–4714.

Womelsdorf T, Ardid S, Everling S, Valiante TA (2014) Burst firing synchronizes prefrontal and anterior cingulate cortex during attentional control. Current Biology. 24(22): 2613–2621. pdf.

Womelsdorf T, Valiante TA, Sahin NT, Miller KJ, Tiesinga P (2014) Dynamic circuit motifs underlying rhythmic gain control, gating and integration. Nature Neuroscience. 17: 1031–1039. pdf.

Lipsman N, Nakao T, Kanayama N, Krauss JK, Anderson A, Giacobbe P, Hamani C, Hutchison WD, Dostrovsky JO, Womelsdorf T, Lozano AM, Northoff G (2014) Neural overlap between resting state and self-relevant activity in human subcallosal cingulate cortex – Single unit recording in an intracranial study. Cortex. 60: 139–144.

Lipsman N, Kaping D, Westendorff S, Lozano AM, Womelsdorf T (2014). Beta coherence within human ventromedial prefrontal cortex precedes affective value-based choices. NeuroImage, 85(2):769–778. PDF

Womelsdorf T, Landau AN, Fries P (2014). Attentional Selection through Rhythmic Synchronization at Multiple Frequencies. In The Cognitive Neurosciences V (Ed. M. Gazzaniga). MIT Press, Cambridge, MA.

Phillips J, Vinck M, Everling S, Womelsdorf T (2014). A long-range fronto-parietal 5–10 Hz network predicts “top-down” controlled guidance in a task-switch paradigm. Cerebral Cortex, 24(8):1996–2008. PDF

Vinck M, Womelsdorf T, Buffalo B, Desimone R, Fries P (2013). Attentional modulation of cell-class specific gamma-band synchronization in awake monkey area V4. Neuron, 80(4):1077–1089. PDF

Womelsdorf T, Westendorff S, Ardid S (2013). Subnetwork selection in deep cortical layers is mediated by beta-oscillation dependent firing. Frontiers in Systems Neuroscience. doi:10.3389/fnsys.2013.00025. PDF

Hutchison RM, Womelsdorf T, Allen EA, Bandettini PA, Calhoun V, Corbetta M, Della Penna S, Duyn J, Glover G, Gonzalez-Castillo J, Handwerker DA, Keilholz S, Kiviniemi V, Leopold DG, de Pasquale F, Sporns O, Walter M, Chang C (2013). Dynamic functional connectivity: Promises, issues, and interpretations. NeuroImage. doi:10.1016/j.neuroimage.2013.05.079. PDF

Womelsdorf T, Bosman CA, Fries P (2013). Selective Neuronal Synchronization and Attentional Stimulus Selection in Visual Cortex. In The New Visual Neurosciences (Eds. JS Werner & LM Chalupa), Chapter 71, 1013–1030. MIT Press.

Laxton AW, Neimat JS, Davis KD, Womelsdorf T, Hutchison WD, Dostrovsky JO, Hamani C, Mayberg HS, Lozano AM (2013). Neuronal coding of implicit emotion categories in human subgenual prefrontal cortex. Biological Psychiatry. doi:10.1016/j.biopsych.2013.03.029.

Vinck M, Womelsdorf T, Fries P (2013). Gamma oscillations and information transmission. In Principles of Neural Coding (Eds. RQ Quiroga & S Panzeri), Chapter 23.

Maris E, Womelsdorf T, Desimone R, Fries P (2013). Rhythmic neuronal synchronization in visual cortex entails spatial phase relation diversity that is modulated by stimulation and attention. NeuroImage. doi:S1053-8119(13)00127-4.

Womelsdorf T, Lima B, Vinck M, Neuenschwander S, Oostenveld R, Singer W, Fries P (2012). Orientation selectivity and noise correlation in awake monkey V1 are modulated by the gamma cycle. PNAS, 109(11):4302–4307.

Bosman CA, Schoffelen JM, Bastos A, Oostenveld R, Womelsdorf T, Brunet N, Rubehn B, Stieglitz T, de Weerd P, Fries P (2012). Stimulus selection through selective synchronization between monkey visual areas. Neuron, 75:875–888.

Hutchison RM, Womelsdorf T, Gati SS, Menon RS, Everling S (2012). Resting-state networks show dynamic functional connectivity in awake humans and anesthetized macaques. Human Brain Mapping. doi:10.1002/hbm.22058.

Kaping D, Vinck M, Hutchison RM, Everling S, Womelsdorf T (2011). Specific contributions of ventromedial, anterior cingulate, and lateral prefrontal cortex for attentional selection and stimulus valuation. PLoS Biology, 9(12):e1001224. doi:10.1371/journal.pbio.1001224. PDF

Vinck M, Battaglia F, Womelsdorf T, Pennartz CMA (2011). Improved measures of phase-coupling between spikes and the local field potential. Journal of Computational Neuroscience. doi:10.1007/s10827-011-0374-4.

Womelsdorf T, Fries P (2011). Rhythmic neuronal synchronization subserves selective attentional processing. In Characterizing Consciousness: From Cognition to the Clinic (Eds. S. Dehaene & Y. Christen). Springer Berlin Heidelberg, pp.109–132.

Womelsdorf T (2011). Deciphering the attentional search engine in the brain. Trends in Cognitive Sciences, 15(10):451–452.

Hutchison RM, Womelsdorf T, Gati J, Leung LS, Menon RS, Everling S (2011). Resting-state connectivity identifies distinct functional networks in macaque cingulate cortex. Cerebral Cortex. doi:10.1093/cercor/bhr181.

Womelsdorf T, Johnston K, Vinck M, Everling S (2010). Theta activity in anterior cingulate cortex predicts task-rules and their adjustments following errors. PNAS, 107(11):5248–5253.

Vinck M, Lima B, Womelsdorf T, Oostenveld R, Singer W, Neuenschwander S, Fries P (2010). Gamma phase shifting in awake monkey visual cortex. Journal of Neuroscience, 30(4):9471–9480.

Womelsdorf T, Vinck M, Leung S, Everling S (2010). Selective theta synchronization of choice-relevant information subserves goal-directed behavior. Frontiers in Human Neuroscience, 107(11):5248–5253.

Vinck M, van Wingerden M, Womelsdorf T, Fries P, Pennartz CMA (2010). The pairwise phase consistency: A bias-free measure of rhythmic neuronal synchronization. NeuroImage, 51(1):112–122.

Bosman CA, Womelsdorf T, Desimone R, Fries P (2009). A microsaccadic rhythm modulates gamma-band synchronization and behavior. Journal of Neuroscience, 29(30):9471–9480.

Womelsdorf T, Fries P (2009). Selective attention through selective neuronal synchronization. In The Cognitive Neurosciences IV (Ed. M. Gazzaniga). MIT Press, Cambridge, MA.

Bosman C, Womelsdorf T (2008). Neuronal signatures of selective attention: Synchronization and gain modulation as mechanisms for selective sensory information processing. In Dynamical Approaches to Brain Function: Neurobiological, Clinical and Methodological Trends (Eds. D. Cosmelli & F. Aboitiz). Springer, Berlin, Germany.

Womelsdorf T, Anton-Erxleben K, Treue S (2008). Receptive field shift and shrinkage in macaque area MT through attentional gain modulation. Journal of Neuroscience, 28(36):8934–8944.

Fries P, Womelsdorf T, Oostenveld R, Desimone R (2008). The effects of visual stimulation and selective visual attention on rhythmic neuronal synchronization in macaque area V4. Journal of Neuroscience, 28(18):4823–4835.

Tzvetanov T, Womelsdorf T (2008). Predicting human perceptual decisions by decoding neuronal information profiles. Biological Cybernetics, 98(5):397–411.

Womelsdorf T, Schoffelen JM, Oostenveld R, Singer W, Desimone R, Engel AK, Fries P (2007). Modulation of neuronal interactions through neuronal synchronization. Science, 316:1609–1612.

Womelsdorf T, Fries P (2007). The role of neuronal synchronization in selective attention. Current Opinion in Neurobiology, 17:154–160.

Womelsdorf T, Anton-Erxleben K, Pieper F, Treue S (2006). Dynamic shifts of visual receptive fields in cortical area MT by spatial attention. Nature Neuroscience, 9:1156–1160.

Womelsdorf T, Fries P, Mitra PP, Desimone R (2006). Gamma-band synchronization in visual cortex predicts speed of change detection. Nature, 439:733–736.

Womelsdorf T, Fries P (2006). Neuronal coherence during selective attentional processing and sensory–motor integration. Journal of Physiology, Paris, 100:182–193.

Tzvetanov T, Womelsdorf T, Niebergall R, Treue S (2006). Feature-based attention influences contextual interactions during motion repulsion. Vision Research, 46:3651–3658.

Womelsdorf T, Eysel UT, Kisvarday ZF (2001). Comparison of orientation maps obtained with different number of stimulus orientations. NeuroImage, 13:1131–1139.