Place cells that fire in one spot, grid cells that tile space in triangles: how John O'Keefe and May-Britt and Edvard Moser discovered the brain's positioning system and shared the 2014 Nobel Prize.
O'Keefe, the Mosers and the Brain's GPS

In plain English
John O'Keefe found neurons in the rat hippocampus that fire only when the animal is in a particular place. Decades later May-Britt and Edvard Moser found 'grid cells' nearby that fire in a repeating triangular pattern, like map coordinates. Together they form the brain's built-in navigation system.
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How do you know where you are? In 1971 John O'Keefe, an American-British neuroscientist at University College London, recorded from single neurons in the hippocampus of rats exploring an enclosure. Some cells fired only when the rat was in one particular spot β and different cells preferred different spots. He called them place cells.
A cognitive map
In 1978 O'Keefe and Lynn Nadel argued, in The Hippocampus as a Cognitive Map, that the hippocampus builds an internal map of the environment β reviving an idea the psychologist Edward Tolman had proposed in the 1940s from rats' maze behaviour. Place cells re-map when the animal enters a new environment, building a fresh map for each.
Grid cells
The Norwegian couple May-Britt Moser and Edvard Moser, who had trained with O'Keefe, looked for where the hippocampus gets its spatial information. In 2005, in the entorhinal cortex, they found neurons that fire at many locations β and those locations form a strikingly regular triangular grid tiling the whole floor. These grid cells behave like a coordinate system, with grids of different sizes stacked at different depths.
Along with head-direction cells (discovered by James Ranck in the 1980s and studied by Jeffrey Taube), which fire when the head points a certain way, and border cells that fire near walls, they form a positioning system that works even in the dark.
Why it matters
O'Keefe and the Mosers shared the 2014 Nobel Prize. The entorhinal cortex is one of the first regions damaged in Alzheimer's disease, which may explain why getting lost is an early symptom. And similar grid-like codes have been found for abstract "spaces" of concepts, suggesting the brain may reuse its map-making machinery for thinking.
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Graph paper in your head
Grid cells are like sheets of graph paper laid over the world at different scales β some with fine squares, some coarse. By combining which grid points you're currently on, the brain can pin down your position, the way a GPS combines signals from several satellites.
Key takeaways
- Place cells in the hippocampus fire in specific locations (O'Keefe, 1971).
- Grid cells in the entorhinal cortex fire in a triangular lattice (Mosers, 2005).
- Together with head-direction and border cells they form the brain's navigation system.
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