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




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.


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

Transfer of learning between spatial and non-spatial domains


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

Role of temporal coding in human episodic and spatial memory

 My group at the Universitat Jaume I in Castellón de la Plana, Spain has openings for technicians/research assistants and PhD positions for Spanish citizens/EU passport holders. Please contact me if interested. My email is




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 learning and decision-making in other domains


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



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

I Polti#, M Nau#, R Kaplan, V Van Wassenhove, CF Doeller (Under Review) Hippocampus and striatum encode distinct task regularities that guide human timing behavior. bioRxiv, #=equal contribution

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.