Shortcut paradigm studies using animals or humans have also shown the significance of landmarks. Gould and Gould (
1982) trained honeybees to fly consistently between their hive and two feeding points. When captured and displaced while en route to a particular feeding point, trained bees were seen to be able to take shortcuts to the right feeder location if sun-related compass information or prominent landmarks were visible from their displaced position (Dyer,
1991; Dyer, Berry, & Richard,
1993; Menzel, Chittila, Eichmuller, Pietsch, & Knoll,
1990; Wehner, Bleuler, Nievergelt, & Shah,
1990). In humans, Foo et al. (
2005) looked at shortcut-taking ability in virtual reality environments. When stable landmarks were available, humans used them to make accurate novel shortcuts. When landmarks were unavailable, subjects were still able to complete novel shortcuts, but performance was comparatively poor, with less accuracy and higher variability in creating novel shortcuts. Results such as these provide confirmation that humans, bees, rodents, and other animals appear to rely heavily on landmarks when they are available for successful navigation (Dyer,
1991; Dyer et al.,
1993; Menzel et al.,
1990; Wehner et al.,
1990). Landmark information appears to be used from the first learning trial to refine path integration. For example, Foo et al. (
2005) showed, in an immersive virtual environment with colored posts as landmarks, that reliance on visual landmarks was immediate and continued throughout the training period. A large body of additional work also shows immediate reliance on landmarks (Collet, Collet, & Wehner,
2001; Collet & Graham,
2004; Etienne, Boulens, Maurer, Rowe, & Siegrist,
2000; Etienne, Maurer, Boulens, Levy, & Rowe,
2004; Foo et al.,
2005; Kearns, Warren, Duchon, & Tarr,
2002; Riecke, van Veen, & Bulthoff,
2002).