
Path integration is the method thought to be used by animals for
dead reckoning
In navigation, dead reckoning is the process of calculating the current position of a moving object by using a previously determined position, or fix, and incorporating estimates of speed, heading (or direction or course), and elapsed time. T ...
.
History
Charles Darwin
Charles Robert Darwin ( ; 12 February 1809 – 19 April 1882) was an English Natural history#Before 1900, naturalist, geologist, and biologist, widely known for his contributions to evolutionary biology. His proposition that all speci ...
first postulated an inertially-based
navigation
Navigation is a field of study that focuses on the process of monitoring and controlling the motion, movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navig ...
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 movement (or ''
idiothetic'') cues is 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 coo ...
s. Other cues probably include
proprioception
Proprioception ( ) is the sense of self-movement, force, and body position.
Proprioception is mediated by proprioceptors, a type of sensory receptor, located within muscles, tendons, and joints. Most animals possess multiple subtypes of propri ...
(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. In addition, sensitivity to the
Earth's magnetic field
Earth's magnetic field, also known as the geomagnetic field, is the magnetic field that extends from structure of Earth, Earth's interior out into space, where it interacts with the solar wind, a stream of charged particles emanating from ...
for underground animals (e.g.,
mole rat Mole-rat or mole rat can refer to several groups of burrowing Old World rodents:
* Bathyergidae, a family of about 20 hystricognath species in six genera from Africa also called blesmols.
*''Heterocephalus glaber'', the naked mole-rat.
* Spalacidae ...
) can give path integration.
Mechanism
Studies in
arthropods
Arthropods ( ) are invertebrates in the phylum Arthropoda. They possess an arthropod exoskeleton, exoskeleton with a cuticle made of chitin, often Mineralization (biology), mineralised with calcium carbonate, a body with differentiated (Metam ...
, 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.
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 inferior horn of the lateral ventricle. Typically, the hippocampal formation is said to included the dent ...
, called
place cell
A place cell is a kind of pyramidal neuron in the hippocampus that becomes active when an animal enters a particular place in its environment, which is known as the place field. Place cells are thought to act collectively as a cognitive represe ...
s, respond to the position of the animal.
The second, in the early 1990s, is that
neuron
A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...
s in neighboring regions (including anterior
thalamus
The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...
and post-
subiculum
The subiculum (Latin for "support") also known as the subicular complex, or subicular cortex, is the most inferior component of the hippocampal formation. It lies between the entorhinal cortex and the CA1 hippocampal subfield.
The subicular com ...
), 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 t ...
, 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
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 is d ...
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. It is available either as loose leaf paper or bound in notebooks or graph books.
It is commonly found in mathe ...
, 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 difficul ...
References
Bibliography
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PDF*
{{DEFAULTSORT:Path Integration
Orientation (geometry)
Cognitive neuroscience
Animal migration
Spatial cognition