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Reading list: virtual mazes and human navigation

Taking the water maze to people.

By the watermaze.org editors · Updated

Virtual versions of the water maze let researchers test people with the same logic as the rodent task.

  • Astur et al. (1998). A characterization of performance by men and women in a virtual Morris water task: a large and reliable sex difference. Behavioural Brain Research 93(1-2):185-190. doi:10.1016/s0166-4328(98)00019-9
    Astur, Ortiz and Sutherland tested men and women in a virtual version of the Morris water task. Men located the hidden platform more efficiently than women, and the study established the virtual Morris water task as a human analogue of the rodent test.
  • Astur et al. (2002). Humans with hippocampus damage display severe spatial memory impairments in a virtual Morris water task. Behavioural Brain Research 132(1):77-84. doi:10.1016/s0166-4328(01)00399-0
    Astur, Taylor, Mamelak, Philpott and Sutherland tested patients with unilateral hippocampal resections in a virtual Morris water task. The patients showed severe spatial navigation impairments compared with controls, supporting the task's validity as a hippocampus-dependent test in humans.
  1. Astur et al. (1998). A characterization of performance by men and women in a virtual Morris water task: a large and reliable sex difference. Behavioural Brain Research 93(1-2):185-190. doi:10.1016/s0166-4328(98)00019-9 Classic
    Astur, Ortiz and Sutherland tested men and women in a virtual version of the Morris water task. Men located the hidden platform more efficiently than women, and the study established the virtual Morris water task as a human analogue of the rodent test.
  2. Burgess et al. (2002). The Human Hippocampus and Spatial and Episodic Memory. Neuron 35(4):625-641. doi:10.1016/s0896-6273(02)00830-9 Classic
    Burgess, Maguire and O'Keefe review neuropsychological, behavioural and neuroimaging evidence for the role of the human hippocampus in spatial and episodic memory. Several of the studies discussed use virtual reality to test navigation and memory in humans.
  3. Astur et al. (2002). Humans with hippocampus damage display severe spatial memory impairments in a virtual Morris water task. Behavioural Brain Research 132(1):77-84. doi:10.1016/s0166-4328(01)00399-0 Classic
    Astur, Taylor, Mamelak, Philpott and Sutherland tested patients with unilateral hippocampal resections in a virtual Morris water task. The patients showed severe spatial navigation impairments compared with controls, supporting the task's validity as a hippocampus-dependent test in humans.
  4. Ekstrom et al. (2003). Cellular networks underlying human spatial navigation. Nature 425(6954):184-188. doi:10.1038/nature01964 Classic
    Ekstrom and colleagues recorded single neurons in patients navigating a virtual town. Cells in the hippocampus responded at specific locations, providing direct evidence for place-like coding in humans during virtual navigation.
  5. Hölscher et al. (2005). Rats are able to navigate in virtual environments. Journal of Experimental Biology 208(3):561-569. doi:10.1242/jeb.01371 Classic
    Hölscher and colleagues built a virtual reality set-up designed for the wide-angle vision of rats. They showed that rats can be trained to navigate to goals in a virtual environment, establishing the feasibility of rodent VR navigation.
  6. Vorhees & Williams (2006). Morris water maze: procedures for assessing spatial and related forms of learning and memory. Nature Protocols 1(2):848-858. doi:10.1038/nprot.2006.116 Classic
    Vorhees and Williams give a detailed protocol for the Morris water maze, covering acquisition, probe (reference memory), reversal and cued trials. They explain how search patterns and cued trials help separate spatial learning from performance factors such as swimming ability or vision.
  7. Harvey et al. (2009). Intracellular dynamics of hippocampal place cells during virtual navigation. Nature 461(7266):941-946. doi:10.1038/nature08499 Classic
    Harvey and colleagues combined whole-cell recordings in head-restrained mice with a virtual linear track. They identified the subthreshold membrane dynamics of hippocampal place cells during virtual navigation.
  8. Doeller et al. (2010). Evidence for grid cells in a human memory network. Nature 463(7281):657-661. doi:10.1038/nature08704 Classic
    Doeller, Barry and Burgess found a grid-cell-like signal in human fMRI during virtual navigation, with sixfold directional modulation in entorhinal cortex. This gave evidence for grid cells in a human memory network.
  9. Vorhees & Williams (2014). Assessing Spatial Learning and Memory in Rodents. ILAR Journal 55(2):310-332. doi:10.1093/ilar/ilu013 Classic
    Vorhees and Williams review methods for assessing spatial learning and memory in rodents, contrasting allocentric and egocentric navigation. The Morris water maze is treated in detail, alongside other mazes and their strengths and limitations.
  10. Epstein et al. (2017). The cognitive map in humans: spatial navigation and beyond. Nature Neuroscience 20(11):1504-1513. doi:10.1038/nn.4656 Classic
    Epstein, Patai, Julian and Spiers review evidence that the human brain uses map-like codes for spatial navigation, similar to place, grid, border and head-direction cells in rodents. They discuss how these cognitive maps are used during navigation and beyond.
  11. Coughlan et al. (2018). Spatial navigation deficits — overlooked cognitive marker for preclinical Alzheimer disease?. Nature Reviews Neurology 14(8):496-506. doi:10.1038/s41582-018-0031-x Classic
    Coughlan, Laczó, Hort, Minihane and Hornberger review spatial navigation deficits as an early cognitive marker of Alzheimer's disease. They argue that navigation tests, including virtual reality tasks, may detect preclinical changes earlier than episodic memory tests.
  12. Thornberry et al. (2021). Virtual Morris water maze: opportunities and challenges. Reviews in the Neurosciences 32(8):887-903. doi:10.1515/revneuro-2020-0149 Classic
    Thornberry, Cimadevilla and Commins review the virtual Morris water maze for studying human spatial navigation. From over 200 VR navigation papers they found the virtual water maze to be the most widely used task, but with inconsistent set-ups and procedures, and they call for standardised protocols.

HVS Image describes how animal paradigms are simulated in humans using virtual reality, and a real-space dry water maze for humans.

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