[1] Studies beginning in the middle of the 20th century confirmed that animals could return directly to a starting point, such as a nest, in the absence of vision and having taken a circuitous outwards journey.
In addition, sensitivity to the Earth's magnetic field for underground animals (e.g., mole rat) can give path integration.
[2] Studies in arthropods, most notably in the Sahara desert ant (Cataglyphis bicolor), reveal the existence of highly effective path integration mechanisms that depend on determination of directional heading (by polarized light or sun position) and distance computations (by monitoring leg movement or optical flow).
This enables a much more fine-grained study of path integration since it is possible to manipulate movement information and see how place and head direction cells respond (a much simpler procedure than training an animal, which is very slow).
The third finding was that neurons in the dorso-medial entorhinal cortex, which feeds information to the place cells in the hippocampus, fire in a metrically regular way across the whole surface of a given environment.