in press

Liang M, Zheng J, Isham E, Ekstrom AD (2021). Common and distinct roles of frontal midline theta and occipital alpha oscillations in coding temporal intervals and spatial distances.
Journal of Cognitive Neuroscience (accepted). Biorxiv:


Preciado, C. E., Starrett, M. J., & Ekstrom, A. D. (2021). Assessment of a short, focused training to reduce symptoms of cybersickness. Presence, 27(4), 361-377.

Ekstrom A.D. (2021). Regional variation in neurovascular coupling and why we still lack a Rosetta Stone. Philosophical Transactions of the Royal Society, 376, 20190634. [pdf]


Starrett, M. J., McAvan, A. S., Huffman, D. J., Stokes, J. D., Kyle, C. T., Smuda, D. N., Kolarik, B. S., Laczko, J., & Ekstrom A. D. (in press). Landmarks: A solution for spatial navigation and memory experiments in virtual reality. Behavior Research Methods. [Open Access Article]

Ekstrom A.D. & Yonelinas A.P. (2020). Precision, binding, and the hippocampus: Precisely what are we are talking about? Neuropsychologia, 138, 107341.

Harootonian SK, Wilson RC, Hejtmánek L, Ziskin EM, Ekstrom AD (2020) Path integration in large-scale space and with novel geometries: Comparing vector addition and encoding-error models. PLOS Computational Biology 16(5): e1007489. [pdf]

Ekstrom A.D. (2020). Cognitive neuroscience: Why do we get lost when we are stressed? Current Biololgy, 30, R430-R450. [pdf]

Hejtmanek L., Starrett M., Ferrer E. & Ekstrom A.D. (2020). How much of what we learn in virtual reality transfers to real-world navigation? Multisensory Research, 1-25. doi:10.1163/22134808-20201445


Huffman D.J. & Ekstrom A.D. (2019). A modality-independent network underlies retrieval of large-scale spatial environments in the human brain. Neuron, 104(3):611-622.e7. [pdf]

Ekstrom A.D., Harootonian S.K. & Huffman D.L. (2019). Commentary: Grid coding, spatial representation, and navigation: Should we assume an isomorphism? Hippocampus, 1-11. [pdf]

Izadi A., Pevzner A., Lee D.J., Ekstrom A.D., Shahlaie K., Gurkoff G.G. (2019). Medial Septal Stimulation Increases Seizure Threshold and Improves Cognition in Epileptic Rats. Brain Stimulation, 12(3), 735-742. doi:10.1016/j.brs.2019.01.005

Yonelinas A.P., Ranganath C.R., Ekstrom A.D. & Wiltgen B.L. (2019). A Contextual Binding Theory of Episodic Memory: Systems Consolidation Reconsidered. Nature Reviews Neuroscience, 20(6), 364-375. doi:10.1038/s41583-019-0150-4

Kyle C.T., Stokes J.D., Bennett J., Meltzer J. Permenter M.R., Voyt J.A., Ekstrom A.D. and Barnes C.A. (2019). Cytoarchitectonically-driven MRI atlas of nonhuman primate hippocampus: preservation of subfield volumes in aging. Hippocampus, 29(5), 409-421. doi:10.1002/hipo.22809

Peacock, C. E. & Ekstrom, A. D. (2019). Verbal cues flexibly transform spatial representations in human memory. Memory, 27(4), 465-479. doi: 10.1080/09658211.2018.1520890 [accepted manuscript]

Starrett, M. J., Stokes, J. D., Huffman, D. J., Ferrer, E., & Ekstrom, A. D. (2019). Learning-dependent evolution of spatial representations in large-scale virtual environments. Journal of Experimental Psychology: Learning, Memory, and Cognition, 45(3), 497-514. doi:10.1037/xlm0000597 [accepted manuscript]

Schedlbauer, A.M. & Ekstrom, A. D. (2019). Flexible network community organization during encoding and retrieval of spatiotemporal episodic memories. Network Neuroscience, 3(4), 10710-1093. [pdf]


Huffman D.J. & Ekstrom A.D. (2018). Which way is the bookstore? A closer look at the judgments of relative directions task. Spatial Cognition and Computation. 1-37.

Starrett, M. J., & Ekstrom, A. D. (2018). Perspective: Assessing the flexible acquisition, integration, and deployment of human spatial representations and information. Frontiers in Human Neuroscience, 12(281). doi:10.3389/fnhum.2018.00281 [pdf]

Liang, M., Starrett, M. J., & Ekstrom, A. D. (2018). Dissociation of frontal-midline delta-theta and posterior alpha oscillations: A mobile EEG study. Psychophysiology, 55:e13090. doi:10.1111/psyp.13090 [pdf]

Kim, K, Schedlbauer, A, Rollo, M, Karunakaran, S, Ekstrom AD, & Tandon, N. (2018). Network-based brain stimulation selectively impairs spatial retrieval. Brain Stimulation, 11(1): 213-221. doi:10.1016/j.brs.2017.09.016 [pdf]

Kolarik, BS, Baer, T, Shahalaie, K, Yonelinas, AP, & Ekstrom, AD (2018). Close but no cigar: Spatial precision deficits following medial temporal lobe lesion s provide novel insight into theoretical models of navigation and memory. Hippocampus 28(1): 31-41. doi:10.1002/hipo.22801 [pdf]

Dimsdale-Zucker, H. R., Ritchey, M., Ekstrom, A. D., Yonelinas, A. P., & Ranganath, C. (2018). CA1 and CA3 differentially support spontaneous retrieval of episodic contexts within human hippocampal subfields. Nature Communications, 9(1). doi:10.1038/s41467-017-02752-1 [pdf]

Bouffard, N, Stokes, JD, Dramer, HJ, & Ekstrom, AD. (2018). Temporal encoding strategies result in boosts to final free recall comparable to spatial ones. Memory and Cognition 46(1): 17-31. doi:10.3758/s13421-017-0742-z [pdf]


Ekstrom, AD, Huffman, DJ, & Starrett, M. (2017). Interacting networks of brain regions underlie human spatial navigation: A review and novel synthesis of the literature. Journal of Neurophysiology, 188(6): 3328-3344. doi:10.1152/jn.00531.2017 pdf

Ekstrom, AD, & Ranganath, C. (2017). Space, time, and episodic memory: The hippocampus is all over the cognitive map. Hippocampus. doi:10.1002/hipo.22750 [epub ahead of print] pdf

Ekstrom, AD, & Isham, EA. (2017). Human spatial navigation: representations across dimensions and scales. Current Opinion in Behavioral Science, 17, 84-89. doi:10.1016/j.cobeha.2017.06.005 pdf

Bohbot VD, Copara MS, Gotman J, & Ekstrom AD. (2017). Low-frequency theta oscillations in the human hippocampus during real-world and virtual navigation. Nature Communications, 8, 14415. DOI: 10.1038/ncomms14415. pdf

Lieberman JS, Kyle CT, Schedlbauer A, Stokes J, & Ekstrom AD. (2017). A Tale of Two Temporal Coding Strategies: Common and Dissociable Brain Regions Involved in Recency versus Associative Temporal Order Retrieval Strategies. Journal of Cognitive Neuroscience, 29(4), 739-754. DOI: 10.1162/jocn_a_01081. pdf


Arnold AE, Iaria G, & Ekstrom AD. (2016). Mental simulation of routes during navigation involves adaptive temporal compression. Cognition, 157, 14-23. doi: 10.1016/j.cognition.2016.08.009 pdf

Kim K, Ekstrom AD, & Tandon N. (2016). A network approach for modulating memory processes via direct and indirect brain stimulation: Toward a causal approach for the neural basis of memory. Neurobiology of Learning and Memory, 134 Pt. A, 162-177. DOI: pdf

Kolarik BS, Shahlaie K, Hassan A, Borders AA, Kaufman KC, Gurkoff GE, Yonelinas AP, & Ekstrom AD. (2016). Impairments in precision, rather than spatial strategy, characterize performance on the virtual Morris water maze: a case study. Neuropsychologia, 80, 90-101. DOI: 10.1016/j.neuropsychologia.2015.11.013 pdf

Vass LK, Copara MS, Seyal M, Shahlaie K, Tomaszewski Farias S, Shen PY, & Ekstrom AD. (2016). Oscillations Go the Distance: Low-Frequency Human Hippocampal Oscillations Code Spatial Distance in the Absence of Sensory Cues during Teleportation. Neuron, 89(6), 1180-1186. DOI: 10.1016/j.neuron.2016.01.045 Article: pdf Supplement: pdf


Ekstrom AD. Why vision is important to how we navigate. (2015). Hippocampus, 25(6), 731-735. DOI: 10.1002/hipo.22449 pdf

Kyle CT, Smuda DN, Hassan AS, & Ekstrom AD. (2015). Roles of human hippocampal subfields in retrieval of spatial and temporal context. Behavioral Brain Research, 278, 549-558. DOI: 10.1016/j.bbr.2014.10.034 pdf

Kyle CT, Stokes JD, Lieberman JS, Hassan AS, & Ekstrom AD. (2015). Successful retrieval of competing spatial environments in humans involves hippocampal pattern separation mechanisms. eLife, 4, e10499. pdf

Stokes J, Kyle C, & Ekstrom AD. (2015). Complementary roles of human hippocampal subfields in differentiation and integration of spatial context. Journal of Cognitive Neuroscience, 27(3), 546–559. doi: 10.1162/jocn_a_00736 pdf

Suthana NA, Parikshak NN, Ekstrom AD, Ison MJ, Knowlton BJ, Bookheimer SY, & Fried I. (2015). Specific responses of human hippocampal neurons are associated with bedtter memory. Proceedings of the National Academy of Sciences, 112(33), 10503-10508. DOI: 10.1073/pnas.1423036112 pdf

Watrous AJ, Fell J, Ekstrom AD, & Axmacher, N. (2014). More than spikes: common oscillatory mechanisms for content specific neural representations during perception and memory. Current Opinion in Neurobiology, 31C, 33–39. DOI: 10.1016/j.conb.2014.07.024 pdf

Yushkevich PA, Amaral RSC, Augustinack JC, Bender AR, Bernstein JD, … Ekstrom A, … Zeineh MM. (2015). Quantitative comparison of 21 protocols for labeling hippocampal subfields and parahippocampal subregions in in vivo MRI: towards a harmonized segmentation protocol. Neuroimage, 111, 526-541. DOI: 10.1016/j.neuroimage.2015.01.004 pdf


Copara MS, Hassan AS, Kyle CT, Libby LA, Ranganath C, & Ekstrom AD. (2014). Complementary roles of human hippocampal subregions during retrieval of spatiotemporal context. Journal of Neuroscience, 34(20), 6834–6842. DOI: 10.1523/JNEUROSCI.5341-13.2014 pdf

Ekstrom AD. (2014). Cognitive neuroscience: navigating human verbal memory. Current Biology, 24(4), R167–R168. DOI: 10.1016/j.cub.2013.12.043 pdf

Ekstrom AD, Arnold AEGF, & Iaria G. (2014). A critical review of the allocentric spatial representation and its neural underpinnings: toward a network-based perspective. Frontiers in Human Neuroscience, 8:803. DOI: 10.3389/fnhum.2014.00803 pdf

Ekstrom AD & Watrous AJ. (2014). Multifaceted roles for low-frequency oscillations in bottom-up and top-down processing during navigation and memory. Neuroimage 85(Pt 2), 667-677. DOI: 10.1016/j.neuroimage.2013.06.049 pdf

Schedlbauer AM, Copara MS, Watrous AJ, & Ekstrom AD. (2014). Multiple interacting brain areas underlie successful spatiotemporal memory retrieval in humans. Scientific Reports, 4, 6431. DOI: 10.1038/srep06431 pdf

Watrous AJ & Ekstrom AD. (2014). The spectro-contextual encoding and retrieval theory of episodic memory. Frontiers in Human Neuroscience, 8:75. DOI: 10.3389/fnhum.2014.00075 pdf

Zhang H, Zherdeva K, & Ekstrom AD. (2014). Different “routes” to a cognitive map: dissociable forms of spatial knowledge derived from route and cartographic map learning. Memory & Cognition, 42(7), 1106–1117. DOI: 10.3758/s13421-014-0418-x pdf


Gruber MJ, Watrous AJ, Ekstrom AD, Ranganath C, & Otten LJ. (2013). Expected reward modulated encoding-related theta activity before an event. Neuroimage, 64, 68-74. DOI: 10.1016/j.neuroimage.2012.07.064 pdf

Lee DJ, Gurkoff GG, Izadi A, Berman RF, Ekstrom AD, Muizelaar P, Lyeth BG, & Shahlaie K. (2013). Medial septal nucleus theta frequency deep brain stimulation improves spatial working memory following traumatic brain injury. Journal of Neurotrauma, 30(2), 131-139. DOI: 10.1089/neu.2012.2646 pdf

Watrous AJ, Lee DJ, Izadi A, Gurkoff GG, Shahlaie K, & Ekstrom AD. (2013). A comparative study of human and rat hippocampal low frequency oscillations during spatial navigation. Hippocampus, 23(8), 656-661. DOI: 10.1002/hipo.22124 pdf

Watrous AJ, Tandon N, Connor CR, Pieters T, & Ekstrom AD.(2013). Frequency-specific increases in network connectivity underlie successful spatiotemporal memory retrieval. Nature Neuroscience, 16(3), 349-356. DOI: 10.1038/nn.3315 pdf

Zhang H & Ekstrom AD. (2013). Human Neural Systems Underlying Rigid and Flexible Forms of Allocentric Spatial Representation. Human Brain Mapping, 34(5), 1070-1087. DOI: 10.1002/hbm.21494 pdf


Kern KC, Ekstrom AD, Suthana NA, Giesse BS, Montag M, Arshanapalli A, Bookheimer SY, & Sicotte NL. (2012). Fornix damage limits verbal memory functional compensation in multiple sclerosis. Neuroimage, 59(3), 2932-40. DOI: 10.1016/j.neuroimage.2011.09.071 pdf

Libby LA, Ekstrom AD, Ragland JD, & Ranganath C. (2012). Differential connectivity of perirhinal and parahippocampal cortices within human hippocampal subregions revealed by high-resolution functional imaging. Journal of Neuroscience, 32(19), 6550-6560. DOI: 10.1523/JNEUROSCI.3711-11.2012 pdf

Staba RJ, Ekstrom AD, Suthana NA, Burggren A, Fried I, Engel J Jr, & Bookheimer SY. (2012). Gray matter loss correlates with mesial temporal lobe neuronal hyper-excitability inside the human seizure-onset zone. Epilepsia, 53(1), 25-34. DOI: 10.1111/j.1528-1167.2011.03333.x pdf

Zhang H, Copara M, & Ekstrom AD. (2012). Differential Recruitment of Brain Networks Following Route and Cartographic Map Learning of Spatial Environments. PLoS ONE. 7(9), e44886. DOI: 10.1371/journal.pone.0044886 pdf


Addante RJ, Watrous AJ, Yonelinas AP, Ekstrom AD, & Ranganath C. (2011). Prestimulus theta activity predicts correct source memory retrieval. Proceedings of the National Academy of Sciences, 108(26), 10702-10707. DOI: 10.1073/pnas.1014528108 pdf

Ekstrom AD, Copara MS, Isham EA, Wang W, & Yonelinas AP. (2011). Dissociable networks involved in spatial and temporal order source retrieval. Neuroimage, 56, 1803-1813. DOI: 10.1016/j.neuroimage.2011.02.033 pdf

Suthana N, Ekstrom AD, Moshirvaziri S, Knowlton B, & Bookheimer S. (2011). Dissociations within human hippocampal subregions during encoding and retrieval of spatial information. Hippocampus, 21(7), 694-701. DOI: 10.1002/hipo.20833 pdf

Watrous A, Fried I, & Ekstrom AD. (2011). Behavioral correlates of human hippocampal delta and theta oscillations during navigation. Journal of Neurophysiology, 105, 1747-1755. DOI: 10.1152/jn.00921.2010 pdf


Donix M, Burggren AC, Suthana NA, Siddarth P, Ekstrom AD, Krupa AK, Jones M, Martin-Harris L, Ercoli LM, Miller KJ, Small GW, & Bookheimer SY. (2010). Family history of Alzheimer’s disease and hippocampal structure in healthy people. The American Journal of Psychiatry, 167(11), 1399-1406. DOI: 10.1176/appi.ajp.2010.09111575 pdf

Donix M, Burggren AC, Suthana NA, Siddarth P, Ekstrom AD, Krupa AK, Jones M, Rao A, Martin-Harris L, Ercoli LM, Miller KJ, Small GW, & Bookheimer SY. (2010). Longitudinal changes in medial temporal cortical thickness in normal subjects with the APOE-4 polymorphism. Neuroimage, 53(1), 37-43. DOI: 10.1016/j.neuroimage.2010.06.009 pdf

Ekstrom, AD. (2010). How and when the fMRI BOLD signal relates to underlying neural activity: The danger in dissociation. Brain Research Reviews, 62(2), 233-244. DOI: 10.1016/j.brainresrev.2009.12.004 pdf

Ekstrom AD. (2010). Navigation in Virtual Space: Psychological and Neural Aspects. In Encyclopedia of Behavioral Neuroscience (eds. Koob, Thompson and Le Moal). pdf

Jacobs J, Kahana MJ, Ekstrom AD, Mollison MV, & Fried I. (2010). A sense of direction in human entorhinal cortex. Proceedings of the National Academy of Sciences of the United States of America, 107(14), 6487-6492. DOI: 10.1073/pnas.0911213107 pdf

Jacobs J, Korolev IO, Caplan JB, Ekstrom AD, Litt B, Baltuch G, Fried I, Schulze-Bonhage A, Madsen JR, & Kahana MJ. (2010). Right-lateralized brain oscillations in human spatial navigation. Journal of Cognitive Neuroscience, 22(5), 824-836. DOI: 10.1162/jocn.2009.21240 pdf

Mukamel R, Ekstrom AD, Kaplan J, Iacoboni M, & Fried I. (2010). Single-neuron responses in humans during execution and observation of actions. Current Biology, 20(8), 750-756. DOI: 10.1016/j.cub.2010.02.045 pdf

Suthana NA, Krupa A, Donix M, Burggren A, Ekstrom AD, Jones M, Ercoli LM, Miller KJ, Siddarth P, Small GW, & Bookheimer SY. (2010). Reduced hippocampal CA2, CA3 and dentate gyrus activity in asymptotic people at genetic risk for Alzheimer’s disease. Neuroimage, 53(3), 1077-1084. DOI: 10.1016/j.neuroimage.2009.12.014 pdf


Ekstrom AD, Bazih AJ, Suthana NA, Al-Hakim R, Ogura K, Zeineh M, Burggren AC, & Bookheimer SY. (2009). Advances in high-resolution imaging and computational unfolding of the human hippocampus. Neuroimage, 47(1), 42-49. DOI: 10.1016/j.neuroimage.2009.03.017 pdf

Ekstrom A, Suthana N, Millet D, Fried I, & Bookheimer S. (2009). Correlation between BOLD fMRI and theta-band local field potentials in the human hippocampal area. Journal of Neurophysiology, 101(5), 2668-2678. DOI: 10.1152/jn.91252.2008 pdf

Suthana NA, Ekstrom AD, Moshirvaziri S, Knowlton B, & Bookheimer SY. (2009). Human hippocampal CA1 involvement during allocentric encoding of spatial information. Journal of Neuroscience, 29(34), 10512-10519. DOI: 10.1523/JNEUROSCI.0621-09.2009 pdf


Burggren AC, Zeineh MM, Ekstrom AD, Braskie MN, Thompson PM, Small GW, & Bookheimer SY. (2008). Reduced cortical thickness in hippocampal subregions among cognitively normal Apolipoprotein E e4 carriers. Neuroimage, 41(4), 1177-83. DOI: 10.1016/j.neuroimage.2008.03.039 pdf

Ekstrom A, Suthana N, Behnke E, Salamon N, Bookheimer S, & Fried I. (2008). High-resolution depth electrode localization and imaging in patients with pharmacologically intractable epilepsy. Technical note. Journal of Neurosurgery, 108(4), 812-815. DOI: 10.3171/JNS/2008/108/4/0812 pdf


Ekstrom AD and Bookheimer SY. (2007). Spatial and temporal episodic memory retrieval recruit dissociable functional networks in the human brain. Learning and Memory, 14(10), 645-654. DOI: 10.1101/lm.575107 pdf

Ekstrom A, Viskontas I, Kahana MJ, Jacobs J, Upchurch K, Bookheimer SY, & Fried I. (2007). Contrasting roles of neural firing rate and local field potentials in human memory. Hippocampus, 17(8), 606-617. DOI: 10.1002/hipo.20300 pdf

Jacobs J, Kahana MJ, Ekstrom AD, & Fried I. (2007). Brain oscillations control timing of single-neuron activity in humans. Journal of Neuroscience, 27(14), 3839–3844. DOI: 10.1523/JNEUROSCI.4636-06.2007 pdf

Viskontas I, Ekstrom AD, Wilson CL, & Fried I. (2007). Characterizing interneuron and pyramidal cells in the human medial temporal lobe in vivo using extracellular recordings. Hippocampus, 17(1), 49-57. DOI: 10.1002/hipo.20241 pdf


Ekstrom AD, Caplan JB, Shattuck K, Fried I, & Kahana MJ. (2005). Human hippocampal theta activity during virtual navigation. Hippocampus, 15(7), 881-889. DOI: 10.1002/hipo.20109 pdf


Ekstrom AD, Kahana MJ, Caplan JB, Fields TA, Isham EA, Newman E, & Fried I. (2003). Cellular networks underlying human spatial navigation. Nature, 425(6954), 184-188. DOI: 10.1038/nature01964 pdf


Redish AD, Battaglia FP, Chawla MK, Ekstrom AD, Gerrard JL, Lipa P, Rosenzweig ES, Worley PF, Guzowski JF, McNaughton BL, & Barnes CA. (2001). Independence of firing correlates of anatomically proximate hippocampal pyramidal cells. Journal of Neuroscience, 21(5), 1-6. pdf

Ekstrom AD, Meltzer J, McNaughton BL, & Barnes CA. (2001). NMDA receptor antagonism blocks Experience-Dependent Expansion of Hippocampal Place Fields. Neuron, 31, 631-638. pdf