Торлық клеткалар: кеңістіктік бағдарламалау жүйесінің ашылуы

the concept in topology

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. bats, monkeys, and humans. Grid cells were discovered in 2005 by Edvard Moser, May Britt Moser, and their students Torkel Hafting, Marianne Fyhn, and Sturla Molden at the Centre for the Biology of Memory (CBM) in Norway. They were awarded the 2014 Nobel Prize in Physiology or Medicine together with John O'Keefe for their discoveries of cells that constitute a positioning system in the brain. The arrangement of spatial firing fields, all at equal distances from their neighbors, led to a hypothesis that these cells encode a neural representation of Euclidean space. The discovery also suggested a mechanism for dynamic computation of self position based on continuously updated information about position and direction. To detect grid cell activity in a typical rat experiment, an electrode which can record single neuron activity is implanted in the dorsomedial entorhinal cortex and collects recordings as the rat moves around freely in an open arena. The resulting data can be visualized by marking the rat's position on a map of the arena every time that neuron fires an action potential. These marks accumulate over time to form a set of small clusters, which in turn form the vertices of a grid of equilateral triangles. The regular triangle pattern distinguishes grid cells from other types of cells that show spatial firing. By contrast, if a place cell from the rat hippocampus is examined in the same way, then the marks will frequently only form one cluster (one "place field") in a given environment, and even when multiple clusters are seen, there is no perceptible regularity in their arrangement.

In 1971 John O'Keefe and Jonathon Dostrovsky reported the discovery of place cells in the rat hippocampus—cells that fire action potentials when an animal passes through a specific small region of space, which is called the place field of the cell. This discovery, although controversial at first, led to a series of investigations that culminated in the 1978 publication of a book by O'Keefe and his colleague Lynn Nadel called The Hippocampus as a Cognitive Map (a phrase that also appeared in the title of the 1971 paper)—the book argued that the hippocampal neural network instantiates cognitive maps as hypothesized by the psychologist Edward C. Tolman. This theory aroused a great deal of interest, and motivated hundreds of experimental studies aimed at clarifying the role of the hippocampus in spatial memory and spatial navigation. Because the entorhinal cortex provides by far the largest input to the hippocampus, it was clearly important to understand the spatial firing properties of entorhinal neurons. The earliest studies, such as Quirk et al. (1992), described neurons in the entorhinal cortex as having relatively large and fuzzy place fields. But the Mosers thought it was possible that a different result would be obtained if recordings were made from a different part of the entorhinal cortex. The entorhinal cortex is a strip of tissue running along the back edge of the rat brain from the ventral to the dorsal sides. Anatomical studies had shown that different sectors of the entorhinal cortex project to different levels of the hippocampus: the dorsal end of the EC projects to the dorsal hippocampus, the ventral end to the ventral hippocampus. This was relevant because several studies had shown that place cells in the dorsal hippocampus have considerably sharper place fields than cells from more ventral levels. But every study of entorhinal spatial activity before 2004 had made use of electrodes implanted near the ventral end of the EC. Accordingly, together with Marianne Fyhn, Sturla Molden, and Menno Witter, the Mosers set out to examine spatial firing from the different dorsal to ventral levels of the entorhinal cortex. They found that in the dorsal part of medial entorhinal cortex (MEC), cells had sharply defined place fields like in the hippocampus but the cells fired at multiple locations. The arrangement of the firing fields showed hints of regularity, but the size of the environment was too small for spatial periodicity to be visible in this study. The next set of experiments, reported in 2005, made use of a larger environment, which led to the recognition that the cells were actually firing in a hexagonal grid pattern. Grid cells that lie next to one another (i. e., cells recorded from the same electrode) usually show the same grid spacing and orientation, but their grid vertices are displaced from one another by apparently random offsets. But cells recorded from separate electrodes at a distance from one another typically show different grid spacings. Cells located more ventrally (farther from the dorsal border of the MEC) generally have larger firing fields at each grid vertex, and correspondingly greater spacing between the grid vertices. These recordings only extended 3/4 of the way to the ventral tip, so it is possible that even larger grids exist. Such multi scale representations have been shown to be information theoretically desirable. Grid cell activity does not require visual input, since grid patterns remain unchanged when all the lights in an environment are turned off. If the features of an environment are altered less radically, however, the place field pattern may show a lesser degree of change, referred to as "rate remapping", in which many cells alter their firing rates but the majority of cells retain place fields in the same locations as before. This was examined using simultaneous recordings of hippocampal and entorhinal cells, and found that in situations where the hippocampus shows rate remapping, grid cells show unaltered firing patterns, whereas when the hippocampus shows complete remapping, grid cell firing patterns show unpredictable shifts and rotations.

Neural activity in nearly every part of the hippocampal system is modulated by the hippocampal theta rhythm, which has a frequency range of about 6–9 Hz in rats. The entorhinal cortex is no exception: like the hippocampus, it receives cholinergic and GABAergic input from the medial septal area, the central controller of theta. Grid cells, like hippocampal place cells, show strong theta modulation. Most grid cells from layer III do not precess, but their spike activity is largely confined to half of the theta cycle. The grid cell phase precession is not derived from the hippocampus, because it continues to appear in animals whose hippocampus has been inactivated by an agonist of GABA.