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The honeycomb maze provides a novel test to study hippocampal-dependent spatial navigation

Nature volume 554, pages 102105 (01 February 2018) | Download Citation


Here we describe the honeycomb maze, a behavioural paradigm for the study of spatial navigation in rats. The maze consists of 37 platforms that can be raised or lowered independently. Place navigation requires an animal to go to a goal platform from any of several start platforms via a series of sequential choices. For each, the animal is confined to a raised platform and allowed to choose between two of the six adjacent platforms, the correct one being the platform with the smallest angle to the goal-heading direction. Rats learn rapidly and their choices are influenced by three factors: the angle between the two choice platforms, the distance from the goal, and the angle between the correct platform and the direction of the goal. Rats with hippocampal damage are impaired in learning and their performance is affected by all three factors. The honeycomb maze represents a marked improvement over current spatial navigation tests, such as the Morris water maze1,2,3, because it controls the choices of the animal at each point in the maze, provides the ability to assess knowledge of the goal direction from any location, enables the identification of factors influencing task performance and provides the possibility for concomitant single-cell recording.

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  1. 1.

    Spatial localization does not require the presence of local cues. Learn. Motiv. 12, 239–260 (1981)

  2. 2.

    , , & Place navigation impaired in rats with hippocampal lesions. Nature 297, 681–683 (1982)

  3. 3.

    Developments of a water-maze procedure for studying spatial learning in the rat. J. Neurosci. Methods 11, 47–60 (1984)

  4. 4.

    & The Hippocampus as a Cognitive Map (Oxford Univ. Press, 1978)

  5. 5.

    & The hippocampus as a spatial map. Preliminary evidence from unit activity in the freely-moving rat. Brain Res. 34, 171–175 (1971)

  6. 6.

    Head-direction cells in the deep cell layers of the dorsal presubiculum in freely moving rats. Abstr. Soc. Neurosci. 10, 599 (1984)

  7. 7.

    , & Head-direction cells recorded from the postsubiculum in freely moving rats. I. Description and quantitative analysis. J. Neurosci. 10, 420–435 (1990)

  8. 8.

    , , , & Microstructure of a spatial map in the entorhinal cortex. Nature 436, 801–806 (2005)

  9. 9.

    , , , & Boundary vector cells in the subiculum of the hippocampal formation. J. Neurosci. 29, 9771–9777 (2009)

  10. 10.

    , , , & Representation of geometric borders in the entorhinal cortex. Science 322, 1865–1868 (2008)

  11. 11.

    in The Hippocampus Book (eds et al.) Ch. 8, 471–544 (Oxford Univ. Press, 2007)

  12. 12.

    , & Place cells, grid cells, and memory. Cold Spring Harb. Perspect. Biol. 7, a021808 (2015)

  13. 13.

    How do animals actually solve the T maze? Behav. Neurosci. 115, 850–860 (2001)

  14. 14.

    , & Hippocampal connections and spatial discrimination. Brain Res. 139, 295–308 (1978)

  15. 15.

    & Remembrance of places passed: spatial memory in rats. J. Exp. Psychol. Anim. Behav. Process. 2, 97–116 (1976)

  16. 16.

    Memory deficits associated with senescence: a neurophysiological and behavioral study in the rat. J. Comp. Physiol. Psychol. 93, 74–104 (1979)

  17. 17.

    in Brain and Space (ed. ) 273–295 (Oxford Univ. Press, 1991)

  18. 18.

    in Language and Space (eds et al.) 277–316 (MIT Press, 1996)

  19. 19.

    , & Hippocampal representation in place learning. J. Neurosci. 10, 3531–3542 (1990)

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We thank M. Bertelli, A. Hastings, D. Howett, N. Khan, P. Mumford, A. O’Leary, B. Potter, S. Richards and R. Wu for their contribution to this work, and D. Farquharson and D. Halpin for their input to the design, building and testing of maze prototypes. This work was supported by grants from the Wellcome Trust and the Gatsby Charitable Foundation to J.O. R.A.W. is an MRC Clinical Research Training Fellow, J.K. is a Wellcome Trust/Royal Society Sir Henry Dale Fellow and is supported by the Kavli Foundation Dream Team project and the Isaac Newton Trust. D.C. is funded by the Cambridge NIHR Biomedical Research Centre and by the Wellcome Trust.

Author information

Author notes

    • Ruth A. Wood
    • , Marius Bauza
    •  & Julija Krupic

    These authors contributed equally to this work.


  1. Sainsbury Wellcome Centre, UCL, London W1T 4JG, UK

    • Ruth A. Wood
    • , Marius Bauza
    • , Stephen Burton
    •  & John O’Keefe
  2. Department of Physiology, Development and Neuroscience, University of Cambridge, Cambridge CB2 3DY, UK

    • Julija Krupic
  3. Cell & Developmental Biology, UCL, London WC1E 6BT, UK

    • Julija Krupic
    •  & John O’Keefe
  4. DZNE German Centre for Neurodegenerative Diseases, Bonn 53127, Germany

    • Andrea Delekate
  5. Department of Clinical Neurosciences, University of Cambridge, Cambridge CB2 2PY, UK

    • Dennis Chan


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J.O. conceived the maze and the study. J.O., M.B., J.K. and S.B. were instrumental in designing, building and testing prototypes of the maze. J.K. and M.B. designed the custom-made software used to operate the maze. R.A.W. designed testing schedule 1 for the control experiment, and A.D. and J.O. designed testing schedule 2 for the lesion experiment. A.D. and S.B. performed the hippocampal lesion and sham lesion surgeries. R.A.W. acquired the behavioural data. R.A.W. and A.D. performed the histology for the lesion experiment, and R.A.W. measured hippocampal lesion volumes. R.A.W. conducted the data management and performed the statistical analyses. J.K., S.B. and J.O. collected the single-unit data and J.K. and J.O. analysed these data. J.O. and R.A.W. wrote the manuscript, with contributions to later drafts from all other authors.

Competing interests

The authors declare no competing financial interests.

Corresponding author

Correspondence to John O’Keefe.

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    Supplementary Information

    This file contains a Supplementary Discussion and additional references.

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  1. 1.

    Control rat navigating the Honeycomb Maze

    A control rat making a series of choices as it navigates to the goal on the Honeycomb Maze.

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