“It’s all in their head”: hierarchical exploration of a three-dimensional layered pyramid in rats

  • Zohar Hagbi
  • Alexandra Dorfman
  • Efrat Blumenfeld-Lieberthal
  • David EilamEmail author
Original Paper


Wayfinding in a three-dimensional (3D) environment is intricate, and surface-bounded animals may overcome this complexity by breaking it down into horizontal layers along with the vertical location of each layer. Here, we examined how rats explored a layered pyramid placed in a large open field. We found that exploration presented a hierarchical (or fractal) shape of three types of roundtrips: (1) from the primary home-base to the open-field floor; (2) from the floor up and down the pyramid levels; and (3) from local home-base on each pyramid level. Ascent was slow and interrupted, whereas descent was fast. This difference was a result of level altitude, remaining after data were normalized proportionally to level area. In contrast, the time spent and the distance traveled on each level were dependent on level area, not on level altitude. This structure of spatial behavior accords with multilevel exploration, presenting a relatively independent exploration of each level. The vertical dimension in this experiment thus did not alter the typical spatiotemporal behavior, and the 3D environment was explored by application of the same spatiotemporal approach as that of a horizontal open field. We suggest that this lack of alteration is due to the horizontal posture of the animal’s head and trunk during progression on the pyramid. This behavior also seems to fit the bicoding hypothesis, in which the vertical information is virtually contextual (non-metric), and so, when the rat progresses to a new level, it explores it as a newly accessed horizontal floor area.


Roundtrip Exploration Open field Multilevel Bicoding Quasiplanar model 



This study is dedicated to the memory of Daniel Serruya, who was my first graduate student 30 years ago. Daniel, who published his study on the rock hyrax in 1996, was not only my student, but also became a much loved family friend (DE). We are grateful to Ms. Pazit Zadicario and Roi Gerstel for their help in testing and in data acquisition and analysis, and to Ms. Naomi Paz for language editing. The study was supported by a TAU Vice-President’s internal grant for research encouragement # 0604313192 to DE.

Compliance with ethical standards

Conflict of interest

The authors declare that they have no conflict of interest.

Ethical approval

The experiments and maintenance conditions for the rats were carried out under the regulations and approval of the Institutional Committee for Animal Experimentation at Tel-Aviv University (permit 04-19-009).


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Copyright information

© Springer-Verlag GmbH Germany, part of Springer Nature 2019

Authors and Affiliations

  1. 1.School of ZoologyTel-Aviv UniversityTel AvivIsrael
  2. 2.David Azrieli School of ArchitectureTel-Aviv UniversityTel AvivIsrael

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