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Investigating the cognitive neuroscience of learning, memory, and decision-making

Home: Overview




CIDEGENT Principal Investigator at Universitat Jaume I 

Behavioral, computational, & neuroimaging investigation of memory and decision-making systems.  Focusing on how map-like learning mechanisms could be a widespread phenomenon in the brain beyond spatial cognition.


Postdoc at Universitat Jaume I

The role of episodic memory in navigating the prosocial world


PhD Student at Universitat Jaume I

Relating social and spatial anchoring biases



PhD Student at Universitat Jaume I

Boundary and reference point use in memory-guided decision making



PhD Student at Kavli Institute
(co-supervised by Christian Doeller)

Transfer of learning between spatial and non-spatial domains

Research Assistant

Cognitive computational models of the hippocampal formation

Visiting PhD Student

Computational models of memory-guided fictive planning

Home: Lab Members


  Please send a message to for details about postdoctoral, PhD, research assistant, or other research opportunities in the group.


We investigate how people use past experience to make decisions, plan for the future, and imagine potential scenarios


We build on the Nobel Prize winning discovery of spatially modulated neurons in the hippocampal formation to investigate how such neural computations can inform memory and decision-making in other domains


We study how ‘map-like’ coding of decision variables may inform learning by providing coordinates and boundaries that can be translated between different frames of reference

Home: Research


Please see Raphael Kaplan's Google Scholar page for copies of some of our publications

A Ghouse & R Kaplan(Under Review) Social content is prioritized in episodic memory. PsyArXiv

I Polti#, M Nau#, R Kaplan, V Van Wassenhove, CF Doeller (Under Review) Encoding of task regularities links grid-like signals to human timing behavior. bioRxiv

D Stendardi, L Giacometti Giordani, S Gambino, R Kaplan, E Ciaramelli (2023) Who am I really? The ephemerality of self-knowledge following vmPFC damage. Neuropscyhologia, Available online

I Polti#, M Nau#, R Kaplan, V Van Wassenhove, CF Doeller (2022) Rapid encoding of task regularities in the human hippocampus guides sensorimotor timing. eLife, Nov 1;11:e79027. #=equal contribution

S Arzy & R Kaplan (2022) Transforming social perspectives with cognitive maps. Social Cognitive and Affective Neuroscience

D Kuhrt#, N St. John#, JS Bellmund*, R Kaplan*, CF Doeller* (2021)  An immersive first-person navigation task for abstract knowledge acquisition. Scientific Reports, 11(1):5612.                                                                     *=jointly supervised work.

R Kaplan, A Tauste Campo, D Bush, J King, A Principe, R Koster, M Ley-Nacher, R Rocamora, KJ Friston (2020) Human hippocampal theta oscillations reflect sequential dependencies during spatial planning. Cognitive Neuroscience, 11(3):122-31.

RA Adams#, D Bush#, F Zhen#, SS Meyer, R Kaplan, S Orfanos, TR Marques, O Howes, N Burgess (2020) Impaired theta phase coupling underlies frontotemporal dysconnectivity in schizophrenia. Brain, 143(4):1261-77.

R Kaplan & KJ Friston (2019) Entorhinal transformations in abstract frames of reference. PLoS Biology, 17(5): e3000230.

R Kaplan & KJ Friston (2018) Planning and navigation as active inference. Biological Cybernetics, 112:323-342

R Kaplan, J King, R Koster, WD Penny, N Burgess, & KJ Friston (2017) The neural representation of prospective choice during spatial planning and decisions. PLoS Biology, 15:e1002588.

R Kaplan#, NW Schuck#, CF Doeller (2017) The role of mental maps in decision-making. Trends Neurosci, 40:256-259

R Kaplan#, D Bush#, J Bisby, AJ Horner, SS Meyer, & N Burgess (2017) Medial prefrontal-medial temporal theta phase coupling in dynamic spatial imagery. Journal of Cognitive Neuroscience, 29(3):507-519.

R Kaplan, MH Adhikari, R Hindriks, D Mantini, Y Murayama, NK Logothetis, & G Deco (2016) Hippocampal sharp-wave ripples influence selective activation of the default mode network. Current Biology, 26(5):686-91.
Preview article by M Walker & E Robertson (2016) Current Biology 26(6):239-41

R Kaplan (2014) Endogenous fMRI default mode network fluctuations both positively and negatively correlate with individual transfer of learning. Front Syst Neurosci, 8:229.

R Kaplan, AJ Horner, PA Bandettini, CF Doeller, & N Burgess (2014) Human hippocampal processing of environmental novelty during spatial navigation. Hippocampus, 24(7):740-50.

R Kaplan#, DR Bush#, M Bonnefond, PA Bandettini, GR Barnes, CF Doeller, & N Burgess (2014) Medial prefrontal theta phase coordination during spatial memory retrieval. Hippocampus, 24(6):656-65.

R Kaplan, CF Doeller, GR Barnes, V Litvak, E Duzel, PA Bandettini, & N Burgess (2012) Movement-related theta rhythm in humans: coordinating self-directed hippocampal learning. PLoS Biology, 10(2):e1001267

BR Cornwell, SC Muller, R Kaplan, C Grillon, & M Ernst (2012) Anxiety, a benefit and a detriment to cognition: Behavioral and magnetoencephalographic evidence from a mixed-saccade task. Brain & Cognition, 78(3):257-67

CF Doeller & R Kaplan (2011) Parahippocampal Cortex: translating vision into space. Current Biology, 21:589-91

BR Cornwell, RP Alvarez, S Lissek, R Kaplan, M Ernst, & C Grillon (2011) Anxiety overrides the blocking effects of high perceptual load on amygdala reactivity to threat-related distractors. Neuropsychologia, 5:1363-8.

RP Alvarez, G Chen, J Bodurka, R Kaplan, & C Grillon (2011) Phasic and sustained fear in humans elicits distinct patterns of brain activity. NeuroImage, 55: 389-400.

Home: Publications

Group Alumni

Ignacio Polti(PhD student)-Now a postdoc in the Moser lab at the Kavli Institute for Systems Neuroscience.

Natalie St John(RA)-Now a PhD student in the Navarro Schroeder lab at the Kavli Institute for Systems Neuroscience.

Stelios Orfanos(Visiting Clinical Researcher)-Psychiatrist at South West London and St George's Mental Health NHS Trust

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