Neocortex

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For other uses, see Neocortex (disambiguation).
Brain: Neocortex
NeuroNames ancil-754
MeSH Neocortex
NeuroLex ID birnlex_2547
TA A14.1.09.304
FMA FMA:62429
A representative column of neocortex. Cell body layers are labeled on the left, and fiber layers are labeled on the right.

The neocortex (Latin for "new bark" or "new rind"), also called the neopallium ("new mantle") and isocortex ("equal rind"), is a part of the mammalian brain. In humans it is the largest part of the cerebral cortex which covers the two cerebral hemispheres, with the allocortex making up the rest. The neocortex is made up of six layers, labelled from the outer in, I to VI. In humans, the neocortex is involved in higher functions such as sensory perception, generation of motor commands, spatial reasoning, conscious thought and language.[1] There are two types of cortex in the neocortex – the true isocortex and the proisocortex.

Anatomy[edit]

The neocortex is the most developed of the cerebral tissues.[2] The neocortex consists of the grey matter, or neuronal cell bodies and unmyelinated fibers, surrounding the deeper white matter (myelinated axons) in the cerebrum. There are two types of cortex in the neocortex, the proisocortex and the true isocortex. The pro-isocortex is a transitional area between the true isocortex, and the periallocortex (part of the allocortex). It is found in the cingulate cortex (part of the limbic system), in Brodmann's areas 24, 25, 30 and 32, the insula and the parahippocampal gyrus.

The neocortex is smooth in rodents and other small mammals, whereas in primates and other larger mammals it has deep grooves (sulci) and ridges (gyri). These folds allow the surface area of the neocortex to increase far beyond what could otherwise be fit in the same size skull. All human brains have the same overall pattern of main gyri and sulci, although they differ in detail from one person to another. The mechanism by which the gyri form during embryogenesis is not entirely clear, and there are several competing hypotheses that explain gyrification. Axonal tension,[3] cortical buckling,[4] or differences in cellular proliferation rates in different areas of the cortex[5] during embryonic development may contribute to the formation of gyri.

The neocortex contains two primary types of neurons, excitatory pyramidal neurons (~80% of neocortical neurons) and inhibitory interneurons (~20%).[6] The structure of the neocortex is relatively uniform (hence the alternative names "iso-" and "homotypic" cortex), consisting of six horizontal layers segregated principally by cell type and neuronal connections. However, there are many exceptions to this uniformity; for example, the motor cortex lacks layer IV. There is some canonical circuitry within the cortex; for example, pyramidal neurons in the upper layers II and III project their axons to other areas of neocortex, while those in the deeper layers V and VI project primarily out of the cortex, e.g. to the thalamus, brainstem, and spinal cord. Neurons in layer IV receive all of the synaptic connections from outside the cortex (mostly from thalamus), and themselves make short-range, local connections to other cortical layers.[6] Thus, layer IV receives all incoming sensory information and distributes it to the other layers for further processing.

The neurons of the neocortex are also arranged in vertical structures called cortical columns. These are patches of the neocortex with a diameter of about 0.5 mm (and a depth of 2 mm). Each column typically responds to a sensory stimulus representing a certain body part or region of sound or vision. These columns are similar, and can be thought of as the basic repeating functional units of the neocortex. In humans, the neocortex consists of about a half-million of these columns, each of which contains approximately 70,000 neurons.[7]

Function[edit]

The neocortex is derived embryonically from the dorsal telencephalon, which is the rostral part of the forebrain. The neocortex is divided, into regions demarcated by the cranial sutures in the skull above, into frontal, parietal, occipital, and temporal lobes, which perform different functions. For example, the occipital lobe contains the primary visual cortex, and the temporal lobe contains the primary auditory cortex. Further subdivisions or areas of neocortex are responsible for more specific cognitive processes. In humans, the frontal lobe contains areas devoted to abilities that are enhanced in or unique to our species, such as complex language processing localized to the ventrolateral prefrontal cortex (Broca's area).[6] In humans and other primates, social and emotional processing is localized to the orbitofrontal cortex. (See Cerebral cortex and Cerebrum.)

The neocortex has also been shown to play an influential role in sleep, memory and learning processes. Semantic memories appear to be stored in the neocortex, specifically the anterolateral temporal lobe of the neocortex.[8] It is also involved in instrumental conditioning; responsible for transmitting sensory information and information about plans for movement to the basal ganglia.[8] The firing rate of neurons in the neocortex also has an effect on slow-wave sleep. When the neurons are at rest and are hyperpolarizing, a period of inhibition occurs during a slow oscillation, called the down state. When the neurons of the neocortex are in the excitatory depolarizing phase and are firing briefly at a high rate, a period of excitation occurs during a slow oscillation, called the up state.[8]

There is still much to learn about the roles the neocortex has in the neurological processes exemplified in human behavior. To further understand the vital role the neocortex plays in human cognition, IBM’s computational model of the human brain was created that simulated the electrochemistry of the neocortex. The super computer, the Blue Brain Project, was created to improve understanding of the processes of perception, learning and memory and gain further knowledge about mental health disorders.[9]

Clinical Significance[edit]

Lesions that develop in neurodegenerative disorders, such as Alzheimer’s disease, interrupt the transfer of information from the sensory neocortex to the prefrontal neocortex. This disruption of sensory information contributes to the progressive symptoms seen in neurogenetive disorders such as changes in personality, decline in cognitive abilities, and dementia.[10] Damage to the neocortex of the anterolateral temporal lobe results in semantic dementia, which is the loss of memory of factual information (semantic memories). These symptoms can also be replicated by transcranial magnetic stimulation of this area. If damage is sustained to this area, patients do not develop anterograde amnesia and are able to recall episodic information.[11]

Evolution[edit]

The neocortex is the newest part of the cerebral cortex to evolve (prefix neo meaning new); the other part of the cerebral cortex is the allocortex. The cellular organization of the allocortex is different from the six-layered neocortex. In humans, 90% of the cerebral cortex is neocortex.[12]

For a species to develop a larger neocortex, the brain must too evolve in size so that it’s large enough to support the region. Body size, basal metabolic rate and life history are factors affecting brain evolution and the coevolution of neocortex size and group size.[13] The neocortex increased in size in response to pressures for greater cooperation and competition in early ancestors. With the size increase, there was greater voluntary inhibitory control of social behaviors resulting in increased social harmony.[14]

The six-layer cortex appears to be a distinguishing feature of mammals; it has been found in the brains of all mammals, but not in any other animals.[1] There is some debate,[15][16] however, as to the cross-species nomenclature for neocortex. In avians, for instance, there are clear examples of cognitive processes that are thought to be neocortical in nature, despite the lack of the distinctive six-layer neocortical structure.[17] In a similar manner, reptiles, such as turtles, have primary sensory cortices. A consistent, alternative name has yet to be agreed upon.

Neocortex ratio[edit]

The neocortex ratio of a species is the ratio of the size of the neocortex to the rest of the brain. A high neocortex ratio is thought to correlate with a number of social variables such as group size and the complexity of social mating behaviors.[18] (See Dunbar's number) Humans have a large neocortex as a percentage of total brain matter when compared with other mammals. For example, there is only a 30:1 ratio of neocortical gray matter to the size of the medulla in the brainstem of chimpanzees, while the ratio is 60:1 in humans.[19]

See also[edit]

External links[edit]

References[edit]

  1. ^ a b Lui, J. H.; Hansen, D. V.; Kriegstein, A. R. (2011). "Development and Evolution of the Human Neocortex". Cell 146 (1): 18–36. doi:10.1016/j.cell.2011.06.030. PMC 3610574. PMID 21729779.  edit
  2. ^ Dorland's (2012). Dorland's Illustrated Medical Dictionary (32nd ed.). Elsevier Saunders. p. 1238. ISBN 978-1-4160-6257-8. 
  3. ^ Van Essen, DC (Jan 23, 1997). "A tension-based theory of morphogenesis and compact wiring in the central nervous system.". Nature 385 (6614): 313–8. doi:10.1038/385313a0. PMID 9002514. 
  4. ^ Richman, David (4 July 1975). "Mechanical model of brain convolutional development". Science 189 (4196): 18–21. doi:10.1126/science.1135626. PMID 1135626. 
  5. ^ Ronan, L.; et al. (29 March 2013). "Differential Tangential Expansion as a Mechanism for Cortical Gyrification". Cereb. Cortex. doi:10.1093/cercor/bht082. PMID 23542881. 
  6. ^ a b c Noback; Strominger; Demarest; Ruggiero, Charles; Norman; Robert; David (2005). The Human Nervous System: Structure and Function (Sixth ed.). Totowa, NJ: Humana Press. ISBN 1-59259-730-0. 
  7. ^ bluebrain.epfl.ch
  8. ^ a b c Carlson, Neil (2013). Physiology of Psychology (Eleventh ed.). Pearson. ISBN 978-0-205-239481. 
  9. ^ Haslinger, Kiyrn (September 21, 2005). "Big Blue's Neocortex". Scientific American Mind 16 (3). Retrieved 6 May 2014. 
  10. ^ Braak; Del Tredici; Bohl; Bratzke; Braak, Heiko; Kelly; Jürgen; Hansjürgen; Eva (2000). Annals of the New York academy of sciences, Vol. 911. New York, NY, US: New York Academy of Sciences. ISBN 1-57331-263-0. 
  11. ^ Carlson, Neil (2013). Physiology of Behavior. Pearson. ISBN 978-0-205-23948-1. 
  12. ^ Noback; Strominger; Demarest; Ruggiero, Charles; Norman; Robert; David (2005). The Human Nervous System: Structure and Function. Totowa, NJ: Humana Press. p. 25. ISBN 1-59259-730-0. 
  13. ^ Shultz, Robin Dunbar & Susanne (2007). Understanding primate brain evolution. New York, NY: Oxford University Press. ISBN 978-0-19-921654-3. 
  14. ^ Bjorklund; Kipp, David; Katherine (2002). Social cognition, inhibition, and theory of mind: The evolution of human intelligence. Mahwah, NJ: Lawrence Erlbaum Associate Publishers. ISBN 0-8058-3267-X. 
  15. ^ Jarvis, Erich D.; Güntürkün, Onur; Bruce, Laura; Csillag, András; Karten, Harvey; Kuenzel, Wayne; Medina, Loreta; Paxinos, George et al. (2005). "Opinion: Avian brains and a new understanding of vertebrate brain evolution". Nature Reviews Neuroscience 6 (2): 151–9. doi:10.1038/nrn1606. PMC 2507884. PMID 15685220. 
  16. ^ Reiner, Anton; Perkel, David J.; Bruce, Laura L.; Butler, Ann B.; Csillag, András; Kuenzel, Wayne; Medina, Loreta; Paxinos, George et al. (2004). "Revised nomenclature for avian telencephalon and some related brainstem nuclei". The Journal of Comparative Neurology 473 (3): 377–414. doi:10.1002/cne.20118. PMC 2518311. PMID 15116397. 
  17. ^ Prior, Helmut; Schwarz, Ariane; Güntürkün, Onur (2008). De Waal, Frans, ed. "Mirror-Induced Behavior in the Magpie (Pica pica): Evidence of Self-Recognition". PLoS Biology 6 (8): e202. doi:10.1371/journal.pbio.0060202. PMC 2517622. PMID 18715117. Lay summaryNew Scientist (August 19, 2008). 
  18. ^ Dunbar, R.I.M. (1995). "Neocortex size and group size in primates: A test of the hypothesis". Journal of Human Evolution 28 (3): 287–96. doi:10.1006/jhev.1995.1021. 
  19. ^ Semendeferi, K.; Lu, A.; Schenker, N.; Damasio, H. (2002). "Humans and great apes share a large frontal cortex". Nature Neuroscience 5 (3): 272–6. doi:10.1038/nn814. PMID 11850633.