According to the vertebrate literature, I got the impression that head-direction cells are always anchored to an allocentric reference frame. This can easily be tested with freely moving animals. But what is about tethered animals that can freely rotate around their body axis but can`t change their actual position. Are neurons encoding the animals heading automatically defined as allocentric ones? Or how could you test with a tethered approach, if the neurons encode heading in a allocentric reference frame or in an egocentric one?