Path integration
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- See also: path integral
Path integration is the name given to the method thought to be used by animals for dead reckoning.
Charles Darwin and J.J. Murphy first postulated an inertially-based navigation system in animals in 1873. 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. This shows that they can use cues to track distance and direction in order to estimate their position, and hence how to get home. This process was named path integration to capture the concept of continuous integration of movement cues over the journey. Manipulation of inertial cues confirmed that at least one of these movement (or idiothetic) cues is information from the vestibular organs, which detect movement in the three dimensions. Other cues probably include proprioception (information from muscles and joints about limb position), motor efference (information from the motor system telling the rest of the brain what movements were commanded and executed), and optic flow (information from the visual system signaling how fast the visual world is moving past the eyes). Together, these sources of information can tell the animal which direction it is moving, at what speed, and for how long.
The question of how the animal's brain integrates this dynamic information, in order to draw conclusions about position and homing information, remains unanswered. Extensive studies in arthropods, most notably in the desert ant Cataglyphis fortis, reveal the existence of highly effective path integration mechanisms that depend on determination of directional heading (by polarizing cues such as the sun) and distance computations (by, essentially, “counting footsteps”). How these sources of information are integrated is not yet known.
In mammals, three important discoveries promise to shed light on this issue. The first, in the early 1970s, is that neurons in the hippocampal formation, called place cells, respond to the position of the animal. The second, in the early 1990s, is that neurons in neighboring regions (including anterior thalamus and post-subiculum), called head direction cells, respond to the head direction of the animal. 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. The activity patterns of these grid cells looks very much like a hexagonally organized sheet of graph paper, and suggest a possible metric system that place cells can use to compute distances. Whether place and grid cells actually compute a path integration signal remains to be seen, but computational models exist suggesting this is plausible. Certainly, brain damage to these regions seems to impair the ability of animals to path integrate.
[edit] References
- Best PJ, White AM, Minai A (2003) Spatial processing in the brain: the activity of hippocampal place cells. Annu Rev Neurosci 24: 459-486.
- Etienne AS, Jeffery KJ (2004) Path integration in mammals. Hippocampus 14: 180-192.
- McNaughton BL, Battaglia FP, Jensen O, Moser EI, Moser MB (2006) Path integration and the neural basis of the 'cognitive map'. Nat Rev Neurosci 7: 663-678.
- Taube JS (1998) Head direction cells and the neurophysiological basis for a sense of direction. Prog Neurobiol 55: 225-256.
