Path integration
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Path integration is the method thought to be used by animals for
dead reckoning In navigation, dead reckoning is the process of calculating current position of some moving object by using a previously determined position, or fix, and then incorporating estimates of speed, heading direction, and course over elapsed time. ...
.


History

Charles Darwin Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all species of life have descended ...
first postulated an inertially-based
navigation Navigation is a field of study that focuses on the process of monitoring and controlling the movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navigation ...
system in animals in 1873.The Complete Work of Charles Darwin Online. ''Origin of Certain Instincts'': full text and facsimile
Retrieved 28 February 2012
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 Vision, Visions, or The Vision may refer to: Perception Optical perception * Visual perception, the sense of sight * Visual system, the physical mechanism of eyesight * Computer vision, a field dealing with how computers can be made to gain und ...
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 movements (or '' idiothetic'') cues are information from the vestibular organs, which detect movement in the three
dimension In physics and mathematics, the dimension of a mathematical space (or object) is informally defined as the minimum number of coordinates needed to specify any point within it. Thus, a line has a dimension of one (1D) because only one coord ...
s. Other cues probably include
proprioception Proprioception ( ), also referred to as kinaesthesia (or kinesthesia), is the sense of self-movement, force, and body position. It is sometimes described as the "sixth sense". Proprioception is mediated by proprioceptors, mechanosensory neurons ...
(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 Optical flow or optic flow is the pattern of apparent motion of objects, surfaces, and edges in a visual scene caused by the relative motion between an observer and a scene. Optical flow can also be defined as the distribution of apparent veloci ...
(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. In addition, sensitivity to the earth's magnetic field for underground animals (e.g., mole rat) can give path integration.


Mechanism

Studies in
arthropods Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin, ...
, 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). In mammals, three important discoveries shed light on this issue. The first, in the early 1970s, is that neurons in the
hippocampal formation The hippocampal formation is a compound structure in the medial temporal lobe of the brain. It forms a c-shaped bulge on the floor of the temporal horn of the lateral ventricle. There is no consensus concerning which brain regions are encompassed ...
, called place cells, respond to the position of the animal. The second, in the early 1990s, is that
neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
s in neighboring regions (including anterior
thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all direct ...
and post-
subiculum The subiculum (Latin for "support") is the most inferior component of the hippocampal formation. It lies between the entorhinal cortex and the CA1 subfield of the hippocampus proper. The subicular complex comprises a set of related structures ...
), 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 The entorhinal cortex (EC) is an area of the brain's allocortex, located in the medial temporal lobe, whose functions include being a widespread network hub for memory, navigation, and the perception of time.Integrating time from experience in th ...
, 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 A grid cell is a type of neuron within the entorhinal cortex that fires at regular intervals as an animal navigates an open area, allowing it to understand its position in space by storing and integrating information about location, distance, and ...
looks very much like a
hexagonal In geometry, a hexagon (from Greek , , meaning "six", and , , meaning "corner, angle") is a six-sided polygon. The total of the internal angles of any simple (non-self-intersecting) hexagon is 720°. Regular hexagon A '' regular hexagon'' has ...
ly organized sheet of
graph paper Graph paper, coordinate paper, grid paper, or squared paper is writing paper that is printed with fine lines making up a regular grid. The lines are often used as guides for plotting graphs of functions or experimental data and drawing curves ...
, 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. David Redish states that "The carefully controlled experiments of Mittelstaedt and Mittelstaedt (1980) and Etienne (1987) have demonstrated conclusively that this ability ath integration in mammalsis a consequence of integrating internal cues from vestibular signals and motor efferent copy".Redish, 1999. p 69.


See also

* Cognitive map *
Motion perception Motion perception is the process of inferring the speed and direction of elements in a scene based on visual, vestibular and proprioceptive inputs. Although this process appears straightforward to most observers, it has proven to be a difficult ...


References


Bibliography

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* {{DEFAULTSORT:Path Integration Orientation (geometry) Cognitive neuroscience Animal migration Spatial cognition