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Reading list

Reading list: measures, strategies and statistics

How to turn a swim path into a result.

By the watermaze.org editors · Updated

These papers are about method: which measures to use, how to classify search strategies and how to analyse probe trials.

  • Gallagher et al. (1993). Severity of spatial learning impairment in aging: Development of a learning index for performance in the Morris water maze. Behavioral Neuroscience 107(4):618-626. doi:10.1037/0735-7044.107.4.618
    Gallagher, Burwell and Burchinal studied the variability of spatial learning impairment in aged rats in the water maze. They developed a learning index based on proximity to the platform location during probe trials, which is the basis of the Gallagher proximity measures.
  • Garthe et al. (2009). Adult-Generated Hippocampal Neurons Allow the Flexible Use of Spatially Precise Learning Strategies. PLoS ONE 4(5):e5464. doi:10.1371/journal.pone.0005464
    Garthe, Behr and Kempermann analysed water maze search strategies in mice with reduced adult hippocampal neurogenesis. They classified swim paths into strategies and found that new neurons were needed for the flexible use of precise spatial strategies, especially after the goal moved.
  • Maei (2009). What is the most sensitive measure of water maze probe test performance?. Frontiers in Integrative Neuroscience 3. doi:10.3389/neuro.07.004.2009
    Maei and colleagues compared water maze probe test measures: quadrant and zone occupancy, platform crossings and average proximity to the former platform location. Using more than 1600 mouse probe tests, they found that mean proximity to the former platform location consistently detected group differences better than the other measures.
  • Blokland et al. (2004). A detailed analysis of rats’ spatial memory in a probe trial of a Morris task. Behavioural Brain Research 154(1):71-75. doi:10.1016/j.bbr.2004.01.022
  • Han et al. (2024). The probe trial in a water maze test should not exceed two times. Research Square Platform LLC . doi:10.21203/rs.3.rs-4134660/v1
  1. Treit & Fundytus (1988). Thigmotaxis as a test for anxiolytic activity in rats. Pharmacology Biochemistry and Behavior 31(4):959-962. doi:10.1016/0091-3057(88)90413-3 Classic
    Treit and Fundytus showed that rats tested in an open field spend most of their time near the walls (thigmotaxis), and that anxiolytic drugs reduce this tendency. They proposed thigmotaxis as a simple test for anxiolytic activity.
  2. Simon et al. (1994). Thigmotaxis as an index of anxiety in mice. Influence of dopaminergic transmissions. Behavioural Brain Research 61(1):59-64. doi:10.1016/0166-4328(94)90008-6 Classic
    Simon, Dupuis and Costentin proposed the degree of thigmotaxis (staying close to the walls) during early open-field exploration as an index of anxiety in mice. They validated it with anxiogenic and anxiolytic drugs and examined the role of dopaminergic transmission.
  3. 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.
  4. Brody & Holtzman (2006). Morris water maze search strategy analysis in PDAPP mice before and after experimental traumatic brain injury. Experimental Neurology 197(2):330-340. doi:10.1016/j.expneurol.2005.10.020 Classic
    Brody and Holtzman classified water maze search strategies in PDAPP mice before and after experimental traumatic brain injury. Injury and genotype shifted mice away from spatial strategies, and differences in which strategies were used accounted for most of the differences in performance between groups.
  5. Garthe et al. (2009). Adult-Generated Hippocampal Neurons Allow the Flexible Use of Spatially Precise Learning Strategies. PLoS ONE 4(5):e5464. doi:10.1371/journal.pone.0005464 Classic
    Garthe, Behr and Kempermann analysed water maze search strategies in mice with reduced adult hippocampal neurogenesis. They classified swim paths into strategies and found that new neurons were needed for the flexible use of precise spatial strategies, especially after the goal moved.
  6. Maei (2009). What is the most sensitive measure of water maze probe test performance?. Frontiers in Integrative Neuroscience 3. doi:10.3389/neuro.07.004.2009 Classic
    Maei and colleagues compared water maze probe test measures: quadrant and zone occupancy, platform crossings and average proximity to the former platform location. Using more than 1600 mouse probe tests, they found that mean proximity to the former platform location consistently detected group differences better than the other measures.
  7. 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.
  8. Gehring et al. (2015). Detailed classification of swimming paths in the Morris Water Maze: multiple strategies within one trial. Scientific Reports 5(1). doi:10.1038/srep14562 Classic
    Gehring, Luksys, Sandi and Vasilaki proposed a semi-automated method that splits water maze swim paths into segments and classifies each segment by strategy. They showed that animals often use several strategies within a single trial.
  9. Hawthorne (2026). Baker-Hawthorne Direct Path Measures in Morris Water Maze Analysis. HVS Image Software Ltd . doi:10.5281/zenodo.21870806 Classic
    This HVS technical note defines the Baker-Hawthorne Direct Path Measures for Morris water maze analysis, which identify different forms of initial goal-directed navigation towards the target or its immediate vicinity. It covers their implementation, interpretation and limitations, and applies to rodent water maze and human virtual reality navigation tasks.
  10. Hawthorne (2026). Inactivity or Slow Movement (Floating) in Morris Water Maze Analysis. HVS Image Software Ltd . doi:10.5281/zenodo.21797205 Classic
    This HVS technical note defines the inactivity (floating or slow movement) measure used in Morris water maze analysis. It explains how to interpret inactivity alongside other measures to separate cognitive impairment from non-cognitive influences, for rodent and human virtual reality tasks.
  11. Anisman & McIntyre (2002). Conceptual, Spatial, and Cue Learning in the Morris Water Maze in Fast or Slow Kindling Rats: Attention Deficit Comorbidity. The Journal of Neuroscience 22(17):7809-7817. doi:10.1523/jneurosci.22-17-07809.2002
  12. Woolley et al. (2010). Sex differences in human virtual water maze performance: Novel measures reveal the relative contribution of directional responding and spatial knowledge. Behavioural Brain Research 208(2):408-414. doi:10.1016/j.bbr.2009.12.019

Two of the less common measures here are explained on hvsimage.com: the Blokland measure and the Kelly test.

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