Edvard Moser

Introduction

Edvard Moser (born April 27, 1962) is a Norwegian psychologist and neuroscientist who, with May-Britt Moser, discovered grid cells — neurons in the medial entorhinal cortex that fire at the vertices of a repeating hexagonal lattice as an animal moves through space. The discovery supplied the missing piece of the brain's positioning system: where John O'Keefe's place cells mark where an animal is, grid cells supply the metric that makes distance and direction computable. The three of them shared the 2014 Nobel Prize in Physiology or Medicine for "their discoveries of cells that constitute a positioning system in the brain."

The finding came from a small change in method. Everyone who had looked for spatial firing in the entorhinal cortex had recorded near its ventral end, where the fields are large and fuzzy. The Mosers recorded further toward the dorsal end, and put the rats in a much bigger box. Both decisions were against the grain of the field, and both were necessary.

Early Life and Education

Moser was born in Ålesund, Norway, to German parents who had emigrated in the 1950s. His father, Eduard Paul Moser, trained as a pipe organ builder and came to Norway in 1953 with a friend after being offered work at a pipe organ workshop on the island of Haramsøya; the family later moved to Hareid and then to Ålesund, and built church organs across the country. Moser grew up in a conservative Christian household. His sister, Ingunn Moser, became a sociologist and the founding rector of VID University.

He married May-Britt Moser in 1985, when both were students at the University of Oslo. He was awarded the cand.psychol. degree in psychology in 1990 and then, as a research fellow in the Faculty of Medicine, took the dr.philos. degree in neurophysiology in 1995. He also studied mathematics and statistics, which turned out to matter: the signature of a grid cell is a geometric regularity, and recognising it requires knowing what a lattice looks like. Early in his career he worked under the supervision of Per Andersen, the Norwegian neurophysiologist who had described the hippocampal pathway that carries the name.

Then came the postdoctoral years that set up everything. From 1995 to 1997 he trained with Richard Morris at the Centre for Neuroscience at the University of Edinburgh — Morris being the author of the water maze that had become the standard test of spatial memory — and spent two months as a visiting postdoctoral fellow in O'Keefe's laboratory at University College London. Two months was enough to establish the question the Mosers would spend the next decade answering.

Academic and Professional Career

The Mosers returned to Norway in 1996, joining the Norwegian University of Science and Technology (NTNU) in Trondheim as associate professors of biological psychology. He was promoted to full professor of neuroscience in 1998. What began as two people in a basement became one of the largest neuroscience environments in Europe, built on a sequence of Research Council-funded centres of excellence: the Centre for the Biology of Memory (2002–2012), the Centre for Neural Computation (2012–2022), and the Centre for Algorithms in the Cortex (2023–2033). In 2007 the centre became the Kavli Institute for Systems Neuroscience, with Moser as director. He is also head of the NTNU Institute for Systems Neuroscience.

His awards, before the Nobel, included the Royal Norwegian Society of Sciences and Letters prize for young scientists (1999), the W. Alden Spencer Award (2005), the Betty and David Koetser Award (2006), the Liliane Bettencourt Prize (2006), the Eric K. Fernström Great Nordic Prize (2008), the Louis-Jeantet Prize for Medicine (2011), the Anders Jahre Award (2011, with May-Britt), the Perl-UNC Neuroscience Prize (2012, with May-Britt), and the Louisa Gross Horwitz Prize (2013, with May-Britt and O'Keefe). After 2014 came election as a foreign associate of the National Academy of Sciences, the Körber European Science Prize, and the Grand Cross of the Order of St. Olav. He became an external scientific member of the Max Planck Institute of Neurobiology in 2015.

He and May-Britt announced their divorce in 2016, and both stated that the scientific collaboration would continue. It has: the Moser research environment remains the largest group in the field.

Major Contributions to Psychology

Grid cells

The question the Mosers brought back from London was whether place-cell firing depends on input from outside the hippocampus. The entorhinal cortex is by far the largest input to the hippocampus, and the early studies of it — Quirk and colleagues in 1992 — had found only broad, poorly defined spatial fields. But those studies had all recorded from the ventral end of the structure, and anatomical work had shown that different sectors project to different levels of the hippocampus, with the dorsal hippocampus holding the sharpest place fields. The Mosers, with Marianne Fyhn, Sturla Molden and Menno Witter, recorded from the dorsal end instead.

The result was published in Nature in 2005, with Torkel Hafting as first author. In an enclosure large enough for the pattern to appear, individual entorhinal cells fired not at one location but at many, and those locations formed the vertices of a grid of equilateral triangles. Subsequent work established that the grid has three degrees of freedom — phase, orientation and scale — and that scale is organised topographically along the dorsal-ventral axis, clustering into discrete modules rather than varying smoothly. A cell's grid is stable across environments and, as later work in the lab showed, the correlation structure among grid cells persists into sleep.

The computational implication is what made the discovery famous. A single grid cell signals that the animal is at one of many locations; a set of grids at different scales and orientations uniquely determines position, and the movement of the pattern across the population tracks direction, speed and distance travelled. This is path integration, and it is the mechanism that gives the hippocampal map its metric — the answer to how the brain knows that a landmark is two metres away rather than ten.

The wider entorhinal system

The lab did not stop with the grid. Sargolini and Boccara and others in the group found conjunctive cells that combine grid firing with head direction, border cells that fire along environmental boundaries, and cells responsive to objects — including to the recent absence of an object that had been present. With Li Lu, the Mosers showed that rate remapping in place cells depends on input from the lateral entorhinal cortex, separating the system that encodes position from the system that encodes what is there.

In 2018 Moser's group reported a network in the lateral entorhinal cortex that encodes the time elapsed within an experience, in a form analogous to the spatial code — a step toward the claim that the entorhinal cortex supplies a general framework for organising experience rather than a specifically spatial one.

What grid cells do from moment to moment

The 2005 result, and most of what followed it, treated the grid as a stable readout: the animal is here, and this cell says so. In 2025 the laboratory qualified that picture. Recording with Neuropixels 2.0 probes at millisecond resolution, Abraham Vollan, Richard Gardner and the two Mosers reported in Nature that the population does not hold a steady position signal but fires a sweep roughly ten times a second, running outward from the animal's location and alternating about 30 degrees to the right and then the left, each sweep lasting about 125 milliseconds — one theta cycle. In a prey-chase task the sweeps unhooked from head direction, narrowed into a beam and pointed at the target before the animal itself turned toward it. A 2026 paper in Science from the same group showed that the sweeps are state-dependent: they track a moving goal, reverse during backward locomotion, and persist through REM sleep, with the alternation traced to direction-coded cells in the parasubiculum.

The consequence for the metric account is direct. Grid fields measured across a session are partly the time-average of this scanning, and removing the sweep component leaves them smaller and more precise. Moser's own description of the older recordings is that the entire dynamic was merged into "the large, fat grid fields we know" — which means the hexagonal lattice was never the finest-grained description available, only the finest one the electrodes of 2005 could resolve. It is a useful reminder that the grid was characterised with a particular instrument, and that a better instrument changed what the cell was seen to be doing.

Major Publications

  • Hafting, T., Fyhn, M., Molden, S., Moser, M.-B., & Moser, E. I. (2005). Microstructure of a spatial map in the entorhinal cortex. Nature, 436(7052), 801–806.
  • Fyhn, M., Molden, S., Witter, M. P., Moser, E. I., & Moser, M.-B. (2004). Spatial representation in the entorhinal cortex. Science, 305(5688), 1258–1264.
  • Fyhn, M., Hafting, T., Treves, A., Moser, M.-B., & Moser, E. I. (2007). Hippocampal remapping and grid realignment in entorhinal cortex. Nature, 446(7132), 190–194.
  • Moser, E. I., Moser, M.-B., & McNaughton, B. L. (2017). Spatial representation in the hippocampal formation: A history. Nature Neuroscience, 20(11), 1448–1464.
  • Tsao, A., Moser, M.-B., & Moser, E. I. (2013). Traces of experience in the lateral entorhinal cortex. Current Biology, 23(5), 399–405.
  • Moser, E. I., & Moser, M.-B. (2013). Grid cells and neural coding in high-end cortices. Neuron, 80(3), 765–774.
  • Vollan, A. Z., Gardner, R. J., Moser, M.-B., & Moser, E. I. (2025). Left–right-alternating theta sweeps in entorhinal–hippocampal maps of space. Nature, 639(8056), 995–1005.

Impact and Legacy

Grid cells changed the standing of the cognitive map. Tolman's claim that the brain holds a map of the environment had been argued on behavioural grounds since the 1940s; O'Keefe gave it a cellular basis in 1971; the Mosers gave it a coordinate system, and with it a plausible mechanism for how a map can be computed from self-motion rather than read off the senses. The discovery also gave Alzheimer's disease research a target: the entorhinal cortex is among the earliest regions to show pathology, and it is now standard to ask whether deficits in spatial navigation can serve as an early behavioural marker.

The interpretation is nonetheless contested on several fronts, and the contests are substantive rather than rhetorical.

The first is whether the grid code does what the metric story requires. Almost all grid cell data come from rats and mice running in small, flat, empty enclosures — exactly the conditions in which a hexagonal lattice is easiest to see. In larger and more complex environments, and in the three-dimensional spaces that bats and, arguably, primates inhabit, the regularity degrades; the field is still arguing about whether the grid is a general-purpose metric or an artefact of a very particular experimental situation. Related work on boundary vector and border cells supports an alternative in which spatial firing is built up from the geometry of the environment rather than from an internally generated lattice.

The second is how much of the story transfers to humans. Human grid cells were reported in 2013, but the evidence is necessarily indirect — you cannot implant electrodes in a healthy person and ask them to walk around, so the data are functional imaging and the rare intracranial recordings from patients undergoing evaluation for epilepsy. The representations look similar, and the inference is reasonable, but it rests on methods with far poorer spatial and temporal resolution than a tetrode in a rat.

The third concerns the cognitive map as a whole. The dominant account holds that place cells encode location and little else, but a substantial body of data now shows place-cell firing is modulated by the shape of the environment, running speed, elapsed time, and the current goal of the task, and in some paradigms by which direction the animal faces. In primates, hippocampal neurons often behave more like spatial view cells, firing when the animal looks at a place rather than occupies it — which implies an egocentric rather than allocentric frame. If the hippocampus is coding something richer and more task-dependent than position, then the relationship between grid cells and place cells is more complicated than the feed-forward metric account suggests.

Finally, the 2014 prize rewarded three people for a discovery that involved many more, and the allocation has been discussed since. O'Keefe's 1971 finding was the foundation and the other half of the prize acknowledged that; equally, the place-cell literature of the 1970s and 1980s, and the long argument about whether the hippocampus was spatial at all, involved investigators who did not share the award. Moser himself has written the history in a way that credits the lineage generously.

Learn More

References

  • Hafting, T., Fyhn, M., Molden, S., Moser, M.-B., & Moser, E. I. (2005). Microstructure of a spatial map in the entorhinal cortex. Nature, 436(7052), 801-806.

Last updated: 2026-10-20

Menu
Loading