Path integration is the method thought to be used by animals for dead reckoning.
Path integration is the method thought to be used by animals for dead reckoning.
Charles Darwin first postulated an inertially-based navigation system in animals in 1873. [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. 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. 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). [3]
In mammals, three important discoveries shed light on this.
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.
David Redish states that "The carefully controlled experiments of Mittelstaedt and Mittelstaedt (1980) and Etienne (1987) have demonstrated conclusively that this ability [path integration in mammals] is a consequence of integrating internal cues from vestibular signals and motor efferent copy". [4]
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. The EC is the main interface between the hippocampus and neocortex. The EC-hippocampus system plays an important role in declarative (autobiographical/episodic/semantic) memories and in particular spatial memories including memory formation, memory consolidation, and memory optimization in sleep. The EC is also responsible for the pre-processing (familiarity) of the input signals in the reflex nictitating membrane response of classical trace conditioning; the association of impulses from the eye and the ear occurs in the entorhinal cortex.
The hippocampus is a major component of the brain of humans and other vertebrates. The hippocampus is part of the limbic system, and plays important roles in the consolidation of information from short-term memory to long-term memory, and in spatial memory that enables navigation. The hippocampus is located in the allocortex, with neural projections into the neocortex, in humans as well as other primates. The hippocampus, as the medial pallium, is a structure found in all vertebrates. In humans, it contains two main interlocking parts: the hippocampus proper, and the dentate gyrus.
The limbic system, also known as the paleomammalian cortex, is a set of brain structures located on both sides of the thalamus, immediately beneath the medial temporal lobe of the cerebrum primarily in the forebrain.
The dentate gyrus (DG) is part of the hippocampal formation in the temporal lobe of the brain, which also includes the hippocampus and the subiculum. The dentate gyrus is part of the hippocampal trisynaptic circuit and is thought to contribute to the formation of new episodic memories, the spontaneous exploration of novel environments and other functions.
Adult neurogenesis is the process in which neurons are generated from neural stem cells in the adult. This process differs from prenatal neurogenesis.
In cognitive psychology and neuroscience, spatial memory is a form of memory responsible for the recording and recovery of information needed to plan a course to a location and to recall the location of an object or the occurrence of an event. Spatial memory is necessary for orientation in space. Spatial memory can also be divided into egocentric and allocentric spatial memory. A person's spatial memory is required to navigate in a familiar city. A rat's spatial memory is needed to learn the location of food at the end of a maze. In both humans and animals, spatial memories are summarized as a cognitive map.
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 representation of a specific location in space, known as a cognitive map. Place cells work with other types of neurons in the hippocampus and surrounding regions to perform this kind of spatial processing. They have been found in a variety of animals, including rodents, bats, monkeys and humans.
Idiothetic literally means "self-proposition", and is used in navigation models to describe the use of self-motion cues, rather than allothetic, or external, cues such as landmarks, to determine position and movement. The word is sometimes also spelled "ideothetic". Idiothetic cues include vestibular, optic flow and proprioception. Idiothetic cues are important for the type of navigation known as path integration in which subjects navigate purely using such self-motion cues. This is achieved by an animal through the signals generated by angular and linear accelerations in the course of its exploration. This information generates and updates a vector towards the starting point and an accurate path for return. The spider Cupiennius salei has been known to posses this idiothetic orientation, demonstrating its higher cognitive abilities.
A cognitive map is a type of mental representation used by an individual to order their personal store of information about their everyday or metaphorical spatial environment, and the relationship of its component parts. The concept was introduced by Edward Tolman in 1948. He tried to explain the behavior of rats that appeared to learn the spatial layout of a maze, and subsequently the concept was applied to other animals, including humans. The term was later generalized by some researchers, especially in the field of operations research, to refer to a kind of semantic network representing an individual's personal knowledge or schemas.
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 problem from a computational perspective, and difficult to explain in terms of neural processing.
Head direction (HD) cells are neurons found in a number of brain regions that increase their firing rates above baseline levels only when the animal's head points in a specific direction. They have been reported in rats, monkeys, mice, chinchillas and bats, but are thought to be common to all mammals, perhaps all vertebrates and perhaps even some invertebrates, and to underlie the "sense of direction". When the animal's head is facing in the cell's "preferred firing direction" these neurons fire at a steady rate, but firing decreases back to baseline rates as the animal's head turns away from the preferred direction.
Theta waves generate the theta rhythm, a neural oscillation in the brain that underlies various aspects of cognition and behavior, including learning, memory, and spatial navigation in many animals. It can be recorded using various electrophysiological methods, such as electroencephalogram (EEG), recorded either from inside the brain or from electrodes attached to the scalp.
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 by the term, with some authors defining it as the dentate gyrus, the hippocampus proper and the subiculum; and others including also the presubiculum, parasubiculum, and entorhinal cortex. The hippocampal formation is thought to play a role in memory, spatial navigation and control of attention. The neural layout and pathways within the hippocampal formation are very similar in all mammals.
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 direction. Grid cells have been found in many animals, including rats, mice, bats, monkeys, and humans.
Spatial view cells are neurons in primates' hippocampus; they respond when a certain part of the environment is in the animal's field of view.
Animal navigation is the ability of many animals to find their way accurately without maps or instruments. Birds such as the Arctic tern, insects such as the monarch butterfly and fish such as the salmon regularly migrate thousands of miles to and from their breeding grounds, and many other species navigate effectively over shorter distances.
Hippocampal replay is a phenomenon observed in rats, mice, cats, rabbits, songbirds and monkeys. During sleep or awake rest, replay refers to the re-occurrence of a sequence of cell activations that also occurred during activity, but the replay has a much faster time scale. It may be in the same order, or in reverse. Cases were also found where a sequence of activations occurs before the actual activity, but it is still the same sequence. This is called preplay.
Howard B. Eichenbaum was an American psychologist and neuroscientist who studied the hippocampus. He was a university professor and director of the Center for Memory and Brain at Boston University, having previously worked at Wellesley College. He was the editor-in-chief of the scientific journal Hippocampus.
Phase precession is a neurophysiological process in which the time of firing of action potentials by individual neurons occurs progressively earlier in relation to the phase of the local field potential oscillation with each successive cycle. In place cells, a type of neuron found in the hippocampal region of the brain, phase precession is believed to play a major role in the neural coding of information. John O'Keefe, who later shared the 2014 Nobel Prize in Physiology or Medicine for his discovery that place cells help form a "map" of the body's position in space, co-discovered phase precession with Michael Recce in 1993.
Cyriel Marie Antoine Pennartz is a Dutch neuroscientist serving as professor and head of the Department of Cognitive and Systems Neuroscience at the University of Amsterdam, the Netherlands. He is known for his research on memory, motivation, circadian rhythms, perception and consciousness. Pennartz’ work uses a multidisciplinary combination of techniques to understand the relationships between distributed neural activity and cognition, including in vivo electrophysiology and optical imaging, animal behavior and computational modelling.