Orientation of lizards in a Morris water-maze: roles of the sun compass and the parietal eye.

The present study examined for the first time whether a Morris water-maze can be used to explore compass and other orientation mechanisms in the ruin lizard Podarcis sicula. In the open field, during sunny days, lizards were individually trained to swim from the center of the water maze onto a hidden platform (the goal), positioned at the periphery of the maze in a single compass direction. The goal was invisible because it was placed just beneath the water surface and the water was rendered opaque. The results showed that lizards learn to swim directly towards the hidden goal under the sun in the absence of visual feature cues. We further examined whether the observed orientation response would be due to lizards learning the spatial position of the goal relative to the sun's azimuth, i.e. to the use of a time-compensated sun compass. Lizards reaching learning criteria were subjected to 6 h clock-shift (fast or slow), and tested for goal orientation in the Morris water-maze. Results demonstrated that the learned orientation response is mediated by a time-compensated sun compass. Further investigations provided direct evidence that in ruin lizards an intact parietal eye is required to perform goal orientation under the sun inside a Morris water-maze, and that other brain photoreceptors, like the pineal or deep brain photoreceptors, are not involved in orientation.