cerebral cortex
| Dictionary: cerebral cortex |
n.
The extensive outer layer of gray matter of the cerebral hemispheres, largely responsible for higher brain functions, including sensation, voluntary muscle movement, thought, reasoning, and memory.
| World of the Body: cerebral cortex |
Seen from the outside, the most obvious component of the human brain is the intricately folded cerebral cortex that covers the pair of cerebral hemispheres, which conceal most of the rest of the brain. The convolutions, or gyri, of the cortex, and the fissures or sulci that separate them, vary enormously from brain to brain, and from one hemisphere to the other in each individual. True cerebral cortex is found in the brains of fishes, reptiles, and birds, but is a major feature of the brain in all mammals. In humans it is relatively enormous in size, even compared with our closest relatives, chimpanzees and gorillas. The dominance of the cortex in the human brain led Thomas Willis, the eminent Oxford physician, to propose, in 1664, that it is the seat of ‘higher’ mental processes, such as perception, memory, and will.
The cerebral cortex is the thin outer cloak of grey matter that covers the external surface of the cerebral hemispheres, like the ‘bark’ of a tree, which is indeed what its Latin root refers to. The total area of cortex in man is estimated at nearly a square meter; it is about 4 mm thick, and it contains 10 000 million or more nerve cells (neurons). The number of synapses (connections between nerve fibres and other neurons) is even more staggering — there are, on average, around 10 000 synapses on every cortical neuron.
When thin sections are viewed under a microscope, with appropriate staining of cell bodies, the cortex is seen to consist of several distinct layers. The almost universally adopted layering scheme is that proposed by the late nineteenth-century German anatomist Korbinian Brodmann. Most of the cortex, the so-called neocortex, has a 6-layered structure, but some areas of the hemisphere are covered with simpler cortex with fewer layers, which is believed to be representative of a relatively primitive stage of brain development.
Cortical neurons and their connections
Cortical neurons are basically classified as pyramidal and non-pyramidal cells. A pyramidal cell can be recognized by a single, fairly thick process, the apical dendrite, which sprouts out of the top of the cell body and extends up toward the cortical surface. The other dendrites (branches of the cell body that receive incoming information from the fibres of other neurons), called basal dendrites, form a skirt around the lower part of the cell body. All dendrites bear large numbers of spines, small excrescences on which incoming nerve fibres terminate to form synapses. Pyramidal cells receive incoming nerve fibres from the thalamus and from other areas of cortex, as well as from nearby neurons. The axon of a pyramidal cell (the process that conveys impulses away from the neuron to other nerve cells) can extend a long way, for example more than a meter down to the spinal cord, as well as sending branches to other cortical neurons.
The axon of a typical pyramidal cell can make thousands of synapses on other neurons. When the cell fires an impulse it sweeps along all the branches of the axon to reach the synapses at the terminals, where it triggers the release of the excitatory neurotransmitter glutamate. This affects receptor molecules in the membrane of the target neuron in such a way that it becomes more permeable to sodium ions, which rush into the cell, making the interior more positively charged. This depolarization increases the probability that the target cell will itself fire off an impulse.
If the target cell of the axon of a pyramidal neuron lies below the cerebral hemispheres, in the brain stem or spinal cord, it is termed a projection axon. If it goes to other cortical areas in the same hemisphere, it is an association axon, while if it innervates cells in the cortex of the opposite hemisphere it is called commissural.
Other neurons in the cortex are all non-pyramidal cells — a very heterogeneous group. They are called interneurons because they have relatively short axons that make local connections and do not leave the cortex itself. Non-pyramidal cells can be either excitatory or inhibitory. If the latter, they commonly use the important inhibitory substance GABA (gamma-aminobutyric acid) as the transmitter at their synapses. Non-pyramidal cells, like pyramidal cells, receive axons from the thalamus, from other cortical areas, and from other local interneurons.
The outermost layer of the neocortex, layer I, consists of a mesh of axons and dendrites with very few cell bodies. The other layers consist of pyramidal and non-pyramidal neurons in varying proportions. Layer IV contains a high proportion of non-pyramidal cells, and receives most of the incoming fibres from the thalamus. Layers V and VI have particularly large pyramidal cells that project to subcortical centres, such as the spinal cord and thalamus.
Specialized cortical areas
Korbinian Brodmann also recognized that the cortex is divided into a large number of fields or areas, distinguished by slight differences in the appearance of the layers. He suggested that each anatomically distinctive area is specialized for a particular sensory, motor, or associative function.
Each hemisphere is mainly concerned with the control of muscles and with sensory input from the opposite side of the body, and also with visual and auditory information from the opposite side of the outside world. Hence damage of one hemisphere tends to affect sensation and movement on the opposite side. The cerebral cortex can be seen as the terminus of all the sensory pathways of the nervous system, in the sense that the cortex seems to be needed for conscious perception. Only when information originating in the eyes, the ears, the skin, or any other sensory organ finally reaches the cortex can it then be felt as a subjective experience. Equally, the cortex is the origin of our intended actions. From the motor cortex, axons, especially those from the very large ‘Betz cells’ in layer V, project all the way to the spinal cord to contact motor neurons, which relay the signals out to the muscles. But the cortex is also an integrative organ. Large areas of it are associative in function, meaning that they bring together activity from different sensory and motor systems to make higher-level functions possible, such as speech, memory, and thought. We know from studies of patients who have suffered damage to various parts of the cortex that some of this association cortex is intimately related to our character or personality.
The visual cortex
An area of the neocortex that has been particularly well studied is the primary visual cortex (area 17 of Brodmann), found at the back of the occipital lobe, mainly on the banks of a deep sulcus. Both eyes send signals, via the thalamus, to the visual cortex of both hemispheres, in such a way that each hemisphere receives information about the opposite half of the visual field. Thalamic fibres carrying information from the right eye and the left eye are segregated from each other as they enter the cortex and they form alternating patches of right-eye and left-eye input, about 0.3 mm across, to the cells of layer IV. Since connections between cortical cells mainly run up and down, this has the effect of imposing a pattern of functional ‘columns’ on the cortex, the neurons below any particular point on the cortical surface tending to be dominated functionally by either the right or the left eye. Such ‘columnar’ organization is a characteristic feature of the cerebral cortex, reflecting the segregation of different classes of incoming nerve fibres and the arrangement of connections between cortical neurons.
Area 17 is responsible for basic visual feature detection, but there exist dozens of other, interconnected visual areas in the occipital lobe and even in temporal and parietal lobes. Some are concerned with colour discrimination, or complex pattern recognition, certain cells even responding when the eyes view a stimulus as complex as a face. These areas belong to the association cortex, mentioned above, which is a striking feature of the human brain, permitting the integration and further analysis of simple sensory information to form the basis of meaningful, conscious experience, and the accurate control of action.
The primitive cortex
An example of the more ‘primitive’ cortex described earlier is the hippocampus (Latin for ‘sea-horse’, on account of its appearance in cross-section), which is tucked underneath, on the inner aspect of the temporal lobe. It is unusual in that it has white matter on the outside, and its structure is simple compared with the neocortex, with only three layers. The hippocampus, which receives processed information from much of the association cortex, seems to be involved in short-term conscious memory. It is functionally connected with the hypothalamus and the limbic system, parts of the brain that control basic life functions such as hormonal systems and basic body rhythms and appetites.
So the cerebral cortex is important for an amazing range of functions, from basic drives for self-preservation to the highest levels of consciousness.
— Laurence Garey
See nervous system. See also brain; central nervous system; vision.
| Dental Dictionary: cerebral cortex |
n
A thin layer of gray matter on the surface of the cerebral hemisphere, folded into gyri with about two-thirds of its area buried in fissures. The cerebral cortex integrates higher mental functions, general movement, visceral functions, perception, and behavioral reactions.
| Britannica Concise Encyclopedia: cerebral cortex |
For more information on cerebral cortex, visit Britannica.com.
| Sports Science and Medicine: cerebral cortex |
The outer, surface layer of grey matter in the cerebral hemispheres of the forebrain. The cerebral cortex contains motor areas, which control complex motor skills and some involuntary movements; sensory areas, which receive information from the sense organs; and association areas, which are responsible for thought, learning language, and personality. See also prefrontal cortex, premotor cortex, primary motor cortex.
| Health Dictionary: cerebral cortex |
The surface layer of gray tissue of the cerebrum, frequently called the gray matter. The large size of the cerebral cortex in humans distinguishes them from other animals. Specific parts of the cortex control specific functions, including sensation, voluntary muscle movement, thought, reasoning, and memory.
| World of the Mind: cerebral cortex |
The outer layer or 'bark' (from the Greek) of the brain, associated with sensory perception and the higher mental functions. It appeared late in evolution, and is especially developed in man.
(Published 1987)
(Published 1987)
| Wikipedia: Cerebral cortex |
For other uses, see Cortex.
| Brain: Cerebral cortex | ||
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| Location of the cerebral cortex | ||
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| Golgi-stained neurons in the cortex. | ||
| Latin | cortex cerebri | |
| Part of | Telencephalon | |
| NeuroLex ID | birnlex_1494 | |
The cerebral cortex is a structure within the brain that plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It constitutes the outermost layer of the cerebrum. In preserved brains, it has a grey color, hence the name "grey matter". Grey matter is formed by neurons and their unmyelinated fibers, whereas the white matter below the grey matter of the cortex is formed predominantly by myelinated axons interconnecting different regions of the central nervous system. The human cerebral cortex is 2–4 mm (0.08–0.16 inches) thick.
The surface of the cerebral cortex is folded in large mammals, such that more than two-thirds of the cortical surface is buried in the grooves, called "sulci." The phylogenetically most recent part of the cerebral cortex, the neocortex, also called isocortex, is differentiated into six horizontal layers; the more ancient part of the cerebral cortex, the hippocampus (also called archicortex), has at most three cellular layers, and is divided into subfields. Relative variations in thickness or cell type (among other parameters) allow us to distinguish between different neocortical architectonic fields. The geometry of at least some of these fields seems to be related to the anatomy of the cortical folds, and, for example, layers in the upper part of the cortical ridges (called gyri) seem to be more clearly differentiated than in its deeper parts.[1]
Contents |
Development
The cerebral cortex develops from the most anterior part of the neural plate, a specialized part of the embryonic ectoderm. The neural plate folds and closes to form the neural tube. From the cavity inside the neural tube develops the ventricular system, and, from the epithelial cells of its walls, the neurons and glia of the nervous system. The most anterior (frontal) part of the neural tube, the telencephalon, gives rise to the cerebral hemispheres and cortex.
Cortical neurons are generated within the ventricular zone, next to the ventricles. At first, this zone contains "progenitor" cells, which divide to produce glial and neuronal cells [2]. The glial fibers produced in the first divisions of the progenitor cells are radially oriented, spanning the thickness of the cortex from the ventricular zone to the outer, pial surface, and provide scaffolding for the migration of neurons outwards from the ventricular zone. The first divisions of the progenitor cells are symmetric, which duplicates the total number of progenitor cells at each mitotic cycle. Then, some progenitor cells begin to divide asymmetrically, producing one postmitotic cell that migrates along the radial glial fibers, leaving the ventricular zone, and one progenitor cell, which continues to divide until the end of development, when it differentiates into a glial cell or an ependymal cell. The migrating daughter cells become the pyramidal neurons of the cerebral cortex.[3]
The layered structure of the mature cerebral cortex is formed during development. The first pyramidal neurons generated migrate out of the ventricular zone and together with Cajal-Retzius cells form the preplate. Next, a cohort of neurons migrating into the middle of the preplate divides this transient layer into the superficial marginal zone, which will become layer one of the mature neocortex, and the subplate, forming a middle layer called the cortical plate. These cells will form the deep layers of the mature cortex, layers five and six. Later born neurons migrate radially into the cortical plate past the deep layer neurons, and become the upper layers (two to four). Thus, the layers of the cortex are created in an inside-out order. The only exception to this inside-out sequence of neurogenesis occurs in the layer I of primates, in which, contrary to rodents, neurogenesis continues throughout the entire period of corticogenesis.[4]
Laminar pattern
The different cortical layers each contain a characteristic distribution of neuronal cell types and connections with other cortical and subcortical regions. One of the most clear examples of cortical layering is the Stria of Gennari in the primary visual cortex. This is a band of whiter tissue that can be observed with the naked eye in the fundus of the calcarine sulcus of the occipital lobe. The Stria of Gennari is composed of axons bringing visual information from the thalamus into layer four of visual cortex.
Staining cross-sections of the cortex to reveal the position of neuronal cell bodies and the intracortical axon tracts allowed neuroanatomists in the early 20th century to produce a detailed description of the laminar structure of the cortex in different species. After the work of Korbinian Brodmann (1909), the neurons of the cerebral cortex are grouped into six main layers, from outside (pial surface) to inside (white matter):
- The molecular layer I, which contains few scattered neurons and consists mainly of extensions of apical dendritic tufts of pyramidal neurons and horizontally-oriented axons, as well as glial cells[5]. Some Cajal-Retzius and spiny stellate neurons can be found here. Inputs to the apical tufts are thought to be crucial for the ‘‘feedback’’ interactions in the cerebral cortex involved in associative learning and attention.[6] While it was once thought that the input to layer I came from the cortex itself,[7] it is now realized that layer I across the cerebral cortex mantal receives substantial input from ‘‘matrix’’ or M-type thalamus cells[8] (in contrast to ‘‘core’’ or C-type that go to layer IV).[9]
- The external granular layer II, which contains small pyramidal neurons and numerous stellate neurons
- The external pyramidal layer III, which contains predominantly small and medium-size pyramidal neurons, as well as non-pyramidal neurons with vertically-oriented intracortical axons; layers I through III are the main target of interhemispheric corticocortical afferents, and layer III is the principal source of corticocortical efferents
- The internal granular layer IV, which contains different types of stellate and pyramidal neurons, and is the main target of thalamocortical afferents from thalamus type C neurons.[9] as well as intra-hemispheric corticocortical afferents
- The internal pyramidal layer V, which contains large pyramidal neurons (such as the Betz cells in the primary motor cortex); it is the principal source of subcortical efferents
- The multiform layer VI, which contains few large pyramidal neurons and many small spindle-like pyramidal and multiform neurons; layer VI sends efferent fibers to the thalamus, establishing a very precise reciprocal interconnection between the cortex and the thalamus.[10]
It is important to note that the cortical layers are not simply stacked one over the other; there exist characteristic connections between different layers and neuronal types, which span all the thickness of the cortex. These cortical microcircuits are grouped into cortical columns and minicolumns, the latter of which have been proposed to be the basic functional units of cortex.[11] In 1957, Vernon Mountcastle showed that the functional properties of the cortex change abruptly between laterally adjacent points; however, they are continuous in the direction perpendicular to the surface. Later works have provided evidence of the presence of functionally distinct cortical columns in the visual cortex (Hubel and Wiesel, 1959),[12] auditory cortex and associative cortex.
Cortical areas that lack a layer IV are called agranular. Cortical areas that have only a rudimentary layer IV are called dysgranular[13]. Information processing within each layer is determined by different temporal dynamics with that in the layers II/III having a slow 2Hz oscillation while that in layer V having a fast 10-15 Hz one.[14]
Connections of the cerebral cortex
The cerebral cortex is connected to various subcortical structures such as the thalamus and the basal ganglia, sending information to them along efferent connections and receiving information from them via afferent connections. Most sensory information is routed to the cerebral cortex via the thalamus. Olfactory information, however, passes through the olfactory bulb to the olfactory cortex (piriform cortex). The vast majority of connections are from one area of the cortex to another rather than to subcortical areas; Braitenberg and Schüz (1991) put the figure as high as 99%.[15]
The cortex is commonly described as comprising three parts: sensory, motor, and association areas.
Sensory areas
The sensory areas are the areas that receive and process information from the senses. Parts of the cortex that receive sensory inputs from the thalamus are called primary sensory areas. The senses of vision, audition, and touch are served by the primary visual cortex, primary auditory cortex and primary somatosensory cortex. In general, the two hemispheres receive information from the opposite (contralateral) side of the body. For example the right primary somatosensory cortex receives information from the left limbs, and the right visual cortex receives information from the left visual field. The organization of sensory maps in the cortex reflects that of the corresponding sensing organ, in what is known as a topographic map. Neighboring points in the primary visual cortex, for example, correspond to neighboring points in the retina. This topographic map is called a retinotopic map. In the same way, there exists a tonotopic map in the primary auditory cortex and a somatotopic map in the primary sensory cortex. This last topographic map of the body onto the posterior central gyrus has been illustrated as a deformed human representation, the somatosensory homunculus, where the size of different body parts reflects the relative density of their innervation. Areas with lots of sensory innervation, such as the fingertips and the lips, require more cortical area to process finer sensation.
Motor areas
The motor areas are located in both hemispheres of the cortex. They are shaped like a pair of headphones stretching from ear to ear. The motor areas are very closely related to the control of voluntary movements, especially fine fragmented movements performed by the hand. The right half of the motor area controls the left side of the body, and vice versa.
Two areas of the cortex are commonly referred to as motor:
- Primary motor cortex, which executes voluntary movements
- Supplementary motor areas and premotor cortex, which select voluntary movements.
In addition, motor functions have been described for:
- Posterior parietal cortex, which guides voluntary movements in space
- Dorsolateral prefrontal cortex, which decides which voluntary movements to make according to higher-order instructions, rules, and self-generated thoughts.
Association areas
Association areas function to produce a meaningful perceptual experience of the world, enable us to interact effectively, and support abstract thinking and language. The parietal, temporal, and occipital lobes - all located in the posterior part of the cortex - organize sensory information into a coherent perceptual model of our environment centered on our body image. The frontal lobe or prefrontal association complex is involved in planning actions and movement, as well as abstract thought. Our language abilities are localized to the association areas of the parietal-temporal-occipital complex, typically in the left hemisphere. Wernicke's area relates to understanding language while Broca's area relates to its use.
Classification
Based on the differences in lamination the cerebral cortex can be classified into two major groups:
- Isocortex (homotypical cortex), the part of the cortex with six layers.
- Allocortex (heterotypical cortex) the part of the cortex with less than six layers (varying in number). The allocortex includes the olfactory cortex and the hippocampus.
Auxiliary classes are:
- Mesocortex, classification between isocortex and allocortex where layers 2, 3, and 4 are merged
- Proisocortex, Brodmann areas 24, 25, 32
- Periallocortex, comprising cortical areas adjacent to allocortex.
Based on supposed developmental differences the following classification also appears:
- Neocortex or Neopallium, which corresponds to the isocortex.
- Archicortex, which phylogenetically is the oldest cortex
- Paleocortex
In addition, cortex may be classified on the basis of gross topographical conventions into four lobes:
- Occipital Cortex
- Temporal Cortex
- Parietal Cortex
- Frontal Cortex
Cortical thickness
With magnetic resonance brain scanners, it is possible to get a measure for the thickness of the human cerebral cortex and relate it to other measures. One study has found some positive association between the cortical thickness and intelligence.[16] Another study has found that the somatosensory cortex is thicker in migraine sufferers.[17]
See also
- Cortical column
- Frontal lobe
- Limbic system
- List of regions in the human brain
- Microgyrus
- Occipital lobe
- Parietal lobe
- Temporal lobe
- Cerebral hemisphere
- Neocortex
- Subplate
- Brain-computer interface
References
- ^ Welker, W. 1991. Why does the cerebral cortex fissure and fold? Cerebral Cortex, Vol 8b
- ^ Stephen C. Noctor, Alexander C. Flint, Tanily A. Weissman, Ryan S. Dammerman & Arnold R. Kriegstein (2001). "Neurons derived from radial glial cells establish radial units in neocortex". Nature 409 (6821): 714–720. doi:. PMID 11217860. http://www.nature.com/nature/journal/v409/n6821/abs/409714a0.html.
- ^ P. Rakic (1988). "Specification of cerebral cortical areas". Science 241 (4862): 170–176. doi:. PMID 3291116. http://www.sciencemag.org/cgi/content/abstract/241/4862/170.
- ^ Zecevic N, Rakic P (2001). "Development of layer I neurons in the primate cerebral cortex". J Neurosci. 21: 5607–19. PMID 11466432. http://www.jneurosci.org/cgi/content/full/21/15/5607.
- ^ Shipp, Stewart (2007-06-17). "Structure and function of the cerebral cortex". Current Biology 17 (12): R443–9. doi:. PMID 17580069. http://www.cell.com/current-biology/retrieve/pii/S0960982207011487. Retrieved 2009-02-17.
- ^ Gilbert CD, Sigman M. (2007). Brain states: top-down influences in sensory processing. Neuron. 54(5):677-96. PMID 175534
- ^ Cauller L. (1995). Layer I of primary sensory neocortex: where top-down converges upon bottom-up. Behav Brain Res. 71(1-2):163-70. PMID 8747184
- ^ Rubio-Garrido P, Pérez-de-Manzo F, Porrero C, Galazo MJ, Clascá F. (2009). Thalamic input to distal apical dendrites in neocortical layer 1 is massive and highly convergent. Cereb Cortex. 19(10):2380-95. PMID 19188274 doi:10.1093/cercor/bhn259
- ^ a b Jones EG. (1998). Viewpoint: the core and matrix of thalamic organization. Neuroscience. 85(2):331-45. PMID 9622234
- ^ Creutzfeldt, O. 1995. Cortex Cerebri. Springer-Verlag.
- ^ Mountcastle, V. 1997. The columnar organization of the neocortex. Brain, 120:701-722
- ^ HUBEL DH, WIESEL TN (October 1959). "Receptive fields of single neurones in the cat's striate cortex". J. Physiol. (Lond.) 148: 574–91. PMID 14403679. PMC 1363130. http://www.jphysiol.org/cgi/pmidlookup?view=long&pmid=14403679.
- ^ S.M. Dombrowski , C.C. Hilgetag , and H. Barbas. Quantitative Architecture Distinguishes Prefrontal Cortical Systems in the Rhesus Monkey.Cereb. Cortex 11: 975-988. "...they either lack (agranular) or have only a rudimentary granular layer IV (dysgranular)."
- ^ Sun W, Dan Y. (2009). Layer-specific network oscillation and spatiotemporal receptive field in the visual cortex. Proc Natl Acad Sci U S A. 106:17986–17991 doi:10.1073/pnas.0903962106 PMID 19805197
- ^ Braitenberg, V and Schüz, A 1991. "Anatomy of the Cortex: Statistics and Geometry" NY: Springer-Verlag
- ^ Katherine L. Narr, Roger P. Woods, Paul M. Thompson, Philip Szeszko, Delbert Robinson, Teodora Dimtcheva, Mala Gurbani, Arthur W. Toga and Robert M. Bilder (2007). "Relationships between IQ and Regional Cortical Grey Matter Thickness in Healthy Adults". Cerebral Cortex 17 (9): 2163–2171. doi:. PMID 17118969.
- ^ Alexandre F.M. DaSilva, Cristina Granziera, Josh Snyder and Nouchine Hadjikhani (2007). "Thickening in the somatosensory cortex of patients with migraine". Neurology 69: 1990–1995. doi:. PMID 18025393. News report:
- Catharine Paddock (2007-11-20). "Migraine Sufferers Have Thicker Brain Cortex". Medical News Today. http://www.medicalnewstoday.com/articles/89286.php.
External links
- Webvision - The primary visual cortex Comprehensive article about the structure and function of the primary visual cortex.
- Webvision - Basic cell types Image of the basic cell types of the monkey cerebral cortex.
- Development of the Cerebral Cortex Different topics on cortical development in the form of columns written by leading scientists.
- Cerebral Cortex - Cell Centered Database
- NIF Search - Cerebral Cortex via the Neuroscience Information Framework
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