A cortical column, also called hypercolumn, macrocolumn, functional column or sometimes cortical module, is a group of neurons in the cortex of the brain that can be successively penetrated by a probe inserted perpendicularly to the cortical surface, and which have nearly identical receptive fields. Neurons within a minicolumn (microcolumn) encode similar features, whereas a hypercolumn "denotes a unit containing a full set of values for any given set of receptive field parameters". A cortical module is defined as either synonymous with a hypercolumn (Mountcastle) or as a tissue block of multiple overlapping hypercolumns.
Contents
- Mammalian cerebral cortex
- Columnar functional organization
- Hubel and Wiesel studies
- Number of Cortical columns
- References
The columnar hypothesis states that the cortex is composed of discrete, modular columns of neurons, characterized by a consistent connectivity profile.
It is still unclear what precisely is meant by the term, and it does not correspond to any single structure within the cortex. It has been impossible to find a canonical microcircuit that corresponds to the cortical column, and no genetic mechanism has been deciphered that designates how to construct a column. However, the columnar organization hypothesis is currently the most widely adopted to explain the cortical processing of information.
Mammalian cerebral cortex
The mammalian cerebral cortex, the grey matter encapsulating the white matter, is composed of layers. The human cortex is roughly 2.4 mm thick. The number of layers is the same in all mammals, but varies throughout the cortex. In the neocortex 6 layers can be recognized although many regions lack one or more layers, fewer layers are present in the archipallium and the paleopallium.
Columnar functional organization
The columnar functional organization, as originally framed by Vernon Mountcastle, suggests that neurons that are horizontally more than 0.5 mm (500 µm) from each other do not have overlapping sensory receptive fields, and other experiments give similar results: 200–800 µm. Various estimates suggest there are 50 to 100 cortical minicolumns in a hypercolumn, each comprising around 80 neurons.
An important distinction is that the columnar organization is functional by definition, and reflects the local connectivity of the cerebral cortex. Connections "up" and "down" within the thickness of the cortex are much denser than connections that spread from side to side.
Hubel and Wiesel studies
David Hubel and Torsten Wiesel followed up on Mountcastle's discoveries in the somatic sensory cortex with their own studies in vision. A part of the discoveries that resulted in them winning the 1981 Nobel Prize was that there were cortical columns in vision as well, and that the neighboring columns were also related in function in terms of the orientation of lines that evoked the maximal discharge. Hubel and Wiesel followed up on their own studies with work demonstrating the impact of environmental changes on cortical organization, and the sum total of these works resulted in their Nobel Prize.
Number of Cortical columns
There are about 100,000,000 cortical minicolumns in the neo-cortex with up to 110 neurons each. There may be more if the columns can overlap, as suggested by Tsunoda et al..