You can also find a full list of publications on my Google Scholar Profile.


Nitsch, A., Garvert, M. M., Bellmund, J. L. S., Schuck, N. W., & Doeller, C. F. (2023). Grid-like entorhinal representation of an abstract value space during prospective decision making. bioRxiv.


Everyday decisions require us to predict how valuable different choice options will be in the future. Prior studies have identified a cognitive map in the hippocampal-entorhinal system that encodes relationships between states and enables prediction of future states, but does not inherently convey value during prospective decision making. Here, we investigated whether the entorhinal cortex integrates relational information about changing values by representing an abstract value space. To this end, we combined fMRI with a prospective decision making task that required participants to track and predict changing values of two choice options in a sequence. Such a sequence formed a trajectory through an underlying two-dimensional value space. Our results show that participants successfully integrated and extrapolated changes along the two value dimensions. Participants’ choice behavior was explained by a prospective reinforcement learning model and the degree to which they updated values over time correlated with self-reported navigational abilities and preferences. Crucially, while participants traversed the abstract value space, the entorhinal cortex exhibited a grid-like representation, with the phase of the hexadirectional fMRI signal (i.e., the orientation of the estimated grid) being aligned to the most informative axis through the value space. A network of brain regions, including the ventromedial prefrontal cortex (vmPFC), tracked the prospective value difference between options and the occipital-temporal cortex represented the more valuable option. These findings suggest that the entorhinal grid system might support the prediction of future values by representing a cognitive map, which might be used to generate lower-dimensional signals of the value difference between options and their identities for choices. Thus, these findings provide novel insight for our understanding of cognitive maps as a mechanism to guide prospective decision making in humans.


Bellmund, J. L. S., Deuker, L., Montijn, N. D., & Doeller, C. F. (2022). Mnemonic construction and representation of temporal structure in the hippocampal formation. Nature Communications, 13(1).


The hippocampal-entorhinal region supports memory for episodic details, such as temporal relations of sequential events, and mnemonic constructions combining experiences for inferential reasoning. However, it is unclear whether hippocampal event memories reflect temporal relations derived from mnemonic constructions, event order, or elapsing time, and whether these sequence representations generalize temporal relations across similar sequences. Here, participants mnemonically constructed times of events from multiple sequences using infrequent cues and their experience of passing time. After learning, event representations in the anterior hippocampus reflected temporal relations based on constructed times. Temporal relations were generalized across sequences, revealing distinct representational formats for events from the same or different sequences. Structural knowledge about time patterns, abstracted from different sequences, biased the construction of specific event times. These findings demonstrate that mnemonic construction and the generalization of relational knowledge combine in the hippocampus, consistent with the simulation of scenarios from episodic details and structural knowledge.


Kuhrt, D., St. John, N. R., Bellmund, J. L. S., Kaplan, R., & Doeller, C. F. (2021). An immersive first-person navigation task for abstract knowledge acquisition. Scientific Reports, 11(1).


Advances in virtual reality (VR) technology have greatly benefited spatial navigation research. By presenting space in a controlled manner, changing aspects of the environment one at a time or manipulating the gain from different sensory inputs, the mechanisms underlying spatial behaviour can be investigated. In parallel, a growing body of evidence suggests that the processes involved in spatial navigation extend to non-spatial domains. Here, we leverage VR technology advances to test whether participants can navigate abstract knowledge. We designed a two-dimensional quantity space—presented using a head-mounted display—to test if participants can navigate abstract knowledge using a first-person perspective navigation paradigm. To investigate the effect of physical movement, we divided participants into two groups: one walking and rotating on a motion platform, the other group using a gamepad to move through the abstract space. We found that both groups learned to navigate using a first-person perspective and formed accurate representations of the abstract space. Interestingly, navigation in the quantity space resembled behavioural patterns observed in navigation studies using environments with natural visuospatial cues. Notably, both groups demonstrated similar patterns of learning. Taken together, these results imply that both self-movement and remote exploration can be used to learn the relational mapping between abstract stimuli.


Bellmund, J. L. S. (2020). Piecing Together Cognitive Maps One Dimension at a Time. Neuron, 107(6), 996–999.


In this issue of Neuron, Park et al., 2020 show that the brain forms unified cognitive maps of relational knowledge. The hippocampal-entorhinal region and medial prefrontal cortices spontaneously combine multiple, distinct rank orders to two-dimensional cognitive maps enabling flexible inference.

Bellmund, J. L. S., Polti, I., & Doeller, C. F. (2020). Sequence Memory in the Hippocampal–Entorhinal Region. Journal of Cognitive Neuroscience, 32(11), 2056–2070.


Episodic memories are constructed from sequences of events. When recalling such a memory, we not only recall individual events, but we also retrieve information about how the sequence of events unfolded. Here, we focus on the role of the hippocampal–entorhinal region in processing and remembering sequences of events, which are thought to be stored in relational networks. We summarize evidence that temporal relations are a central organizational principle for memories in the hippocampus. Importantly, we incorporate novel insights from recent studies about the role of the adjacent entorhinal cortex in sequence memory. In rodents, the lateral entorhinal subregion carries temporal information during ongoing behavior. The human homologue is recruited during memory recall where its representations reflect the temporal relationships between events encountered in a sequence. We further introduce the idea that the hippocampal–entorhinal region might enable temporal scaling of sequence representations. Flexible changes of sequence progression speed could underlie the traversal of episodic memories and mental simulations at different paces. In conclusion, we describe how the entorhinal cortex and hippocampus contribute to remembering event sequences—a core component of episodic memory.

Bellmund, J. L. S., Cothi, W. de, Ruiter, T. A., Nau, M., Barry, C., & Doeller, C. F. (2020). Deforming the metric of cognitive maps distorts memory. Nature Human Behaviour, 4(2), 177–188.


Environmental boundaries anchor cognitive maps that support memory. However, trapezoidal boundary geometry distorts the regular firing patterns of entorhinal grid cells, proposedly providing a metric for cognitive maps. Here we test the impact of trapezoidal boundary geometry on human spatial memory using immersive virtual reality. Consistent with reduced regularity of grid patterns in rodents and a grid-cell model based on the eigenvectors of the successor representation, human positional memory was degraded in a trapezoid environment compared with a square environment—an effect that was particularly pronounced in the narrow part of the trapezoid. Congruent with changes in the spatial frequency of eigenvector grid patterns, distance estimates between remembered positions were persistently biased, revealing distorted memory maps that explained behaviour better than the objective maps. Our findings demonstrate that environmental geometry affects human spatial memory in a similar manner to rodent grid-cell activity and, therefore, strengthen the putative link between grid cells and behaviour along with their cognitive functions beyond navigation.


Bellmund, J. L. S. (2019). Hippocampal-entorhinal codes for space, time and cognition. Dissertation.


How do we remember important events? Focusing on episodic memory, I explored the idea that coding principles discovered in spatial navigation research are shared across cognitive domains. The hippocampal-entorhinal region might support our memory through the mapping of cognitive spaces; allowing us to remember where and when events occurred. Cognitive maps stored in memory can guide behavior through the simulation of possible future trajectories. Combining human imagination with representational similarity analysis of multi-voxel patterns, I show that compass-like representations in the parahippocampal gyrus and grid-like hexadirectional representations in the entorhinal cortex contribute to mental simulation. The regular firing patterns of grid cells in the entorhinal cortex supposedly provide a metric for cognitive maps. Hence, grid pattern distortions might affect cognitive functions. Employing immersive virtual reality technology, I demonstrate that human spatial memory is distorted in a trapezoidal environment — mirroring distortions of grid-cell firing patterns in navigating rodents. These mnemonic distortions persisted in memory outside of the trapezoidal environment. Beyond space, time constitutes a second fundamental dimension for episodic memories. I show that the anterior-lateral entorhinal cortex maps the temporal structure of a learned event sequence and that this temporal map relates to memory recall. Combining findings from rodent electrophysiology, cognitive neuroscience and a theory from cognitive science, I describe a theoretical framework for how the hippocampal-entorhinal region might support memory and other cognitive functions through the mapping of cognitive spaces.

Bellmund, J. L. S., Deuker, L., & Doeller, C. F. (2019). Mapping sequence structure in the human lateral entorhinal cortex. eLife, 8, e45333.


Remembering event sequences is central to episodic memory and presumably supported by the hippocampal-entorhinal region. We previously demonstrated that the hippocampus maps spatial and temporal distances between events encountered along a route through a virtual city (Deuker et al., 2016), but the content of entorhinal mnemonic representations remains unclear. Here, we demonstrate that multi-voxel representations in the anterior-lateral entorhinal cortex (alEC) — the human homologue of the rodent lateral entorhinal cortex — specifically reflect the temporal event structure after learning. Holistic representations of the sequence structure related to memory recall and the timeline of events could be reconstructed from entorhinal multi-voxel patterns. Our findings demonstrate representations of temporal structure in the alEC; dovetailing with temporal information carried by population signals in the lateral entorhinal cortex of navigating rodents and alEC activations during temporal memory retrieval. Our results provide novel evidence for the role of the alEC in representing time for episodic memory.


Bellmund, J. L. S., Gärdenfors, P., Moser, E. I., & Doeller, C. F. (2018). Navigating cognition: Spatial codes for human thinking. Science, 362(6415), eaat6766.


The hippocampal formation has long been suggested to underlie both memory formation and spatial navigation. We discuss how neural mechanisms identified in spatial navigation research operate across information domains to support a wide spectrum of cognitive functions. In our framework, place and grid cell population codes provide a representational format to map variable dimensions of cognitive spaces. This highly dynamic mapping system enables rapid reorganization of codes through remapping between orthogonal representations across behavioral contexts, yielding a multitude of stable cognitive spaces at different resolutions and hierarchical levels. Action sequences result in trajectories through cognitive space, which can be simulated via sequential coding in the hippocampus. In this way, the spatial representational format of the hippocampal formation has the capacity to support flexible cognition and behavior.

Nau, M., Navarro Schröder, T., Bellmund, J. L. S., & Doeller, C. F. (2018). Hexadirectional coding of visual space in human entorhinal cortex. Nature Neuroscience, 21(2), 188–190.


Entorhinal grid cells map the local environment, but their involvement beyond spatial navigation remains elusive. We examined human functional MRI responses during a highly controlled visual tracking task and show that entorhinal cortex exhibited a sixfold rotationally symmetric signal encoding gaze direction. Our results provide evidence for a grid-like entorhinal code for visual space and suggest a more general role of the entorhinal grid system in coding information along continuous dimensions.


Deuker, L., Bellmund, J. L. S., Navarro Schröder, T., & Doeller, C. F. (2016). An event map of memory space in the hippocampus. ELife, 5, e16534.


The hippocampus has long been implicated in both episodic and spatial memory, however these mnemonic functions have been traditionally investigated in separate research strands. Theoretical accounts and rodent data suggest a common mechanism for spatial and episodic memory in the hippocampus by providing an abstract and flexible representation of the external world. Here, we monitor the de novo formation of such a representation of space and time in humans using fMRI. After learning spatio-temporal trajectories in a large-scale virtual city, subject-specific neural similarity in the hippocampus scaled with the remembered proximity of events in space and time. Crucially, the structure of the entire spatio-temporal network was reflected in neural patterns. Our results provide evidence for a common coding mechanism underlying spatial and temporal aspects of episodic memory in the hippocampus and shed new light on its role in interleaving multiple episodes in a neural event map of memory space.

Bellmund, J. L. S., Deuker, L., Navarro Schröder, T., & Doeller, C. F. (2016). Grid-cell representations in mental simulation. eLife, 5, e17089.


Anticipating the future is a key motif of the brain, possibly supported by mental simulation of upcoming events. Rodent single-cell recordings suggest the ability of spatially tuned cells to represent subsequent locations. Grid-like representations have been observed in the human entorhinal cortex during virtual and imagined navigation. However, hitherto it remains unknown if grid-like representations contribute to mental simulation in the absence of imagined movement. Participants imagined directions between building locations in a large-scale virtual-reality city while undergoing fMRI without re-exposure to the environment. Using multi-voxel pattern analysis, we provide evidence for representations of absolute imagined direction at a resolution of 30° in the parahippocampal gyrus, consistent with the head-direction system. Furthermore, we capitalize on the six-fold rotational symmetry of grid-cell firing to demonstrate a 60° periodic pattern-similarity structure in the entorhinal cortex. Our findings imply a role of the entorhinal grid-system in mental simulation and future thinking beyond spatial navigation.