| Von Economo neuron | |
|---|---|
 Cartoon of a normalpyramidal cell (left) compared with a von Economo cell (right) | |
| Details | |
| Location | Anterior cingulate cortex (ACC) andFronto-insular cortex (FI) |
| Shape | Unique spindle-shaped projection neuron |
| Function | Global firing rate regulation and regulation of emotional state |
| Presynaptic connections | Local input to ACC and FI |
| Postsynaptic connections | Frontal andtemporal cortex |
| Anatomical terms of neuroanatomy | |
Von Economo neurons, also calledspindle neurons, are a specific class ofmammaliancorticalneurons characterized by a largespindle-shapedsoma (or body) gradually tapering into a single apicalaxon (theramification thattransmits signals) in one direction, with only a singledendrite (the ramification thatreceives signals) facing opposite. Other cortical neurons tend to have many dendrites, and thebipolar-shaped morphology of von Economo neurons is unique here.
Von Economo neurons are found in two very restricted regions in thebrains ofhominids (humans and othergreat apes): theanterior cingulate cortex (ACC) and thefronto-insular cortex (FI) (which each make up thesalience network). In 2008, they were also found in thedorsolateral prefrontal cortex of humans.[1] Von Economo neurons are also found in the brains of a number ofcetaceans,[2][3][4]African andAsian elephants,[5] and to a lesser extent inmacaque monkeys[6] andraccoons.[7] The appearance of von Economo neurons in distantly relatedclades suggests that they representconvergent evolution – specifically, as an adaptation to accommodate the increasing size of these distantly-related animals' brains.
Von Economo neurons were discovered and first described in 1925 by Austrian psychiatrist and neurologistConstantin von Economo (1876–1931).[8][9]
Von Economo neurons are relatively largecells that may allow rapid communication across the relatively large brains ofgreat apes,elephants, andcetaceans. Although rare in comparison to other neurons, von Economo neurons are abundant, and comparatively large, in humans; they are however three times as abundant incetaceans.[3][10]
The discovery of von Economo neurons in diverse whale species[3][4] has led to the suggestion that they are "a possible obligatory neuronal adaptation in very large brains, permitting fast information processing and transfer along highly specific projections and that evolved in relation to emerging social behaviors."[4]: 254 The apparent presence of these specialized neurons only in highly intelligent mammals may be an example ofconvergent evolution.[11]
Their restriction among the primates to greatapes leads to the hypothesis that they developed no earlier than 15–20 million years ago, prior to the divergence of orangutans from the African great apes. Recently, primitive forms of von Economo neurons have also been discovered in macaque monkey brains[12] and raccoons.[7]
In 1999, American neuroscientistJohn Allman and colleagues at theCalifornia Institute of Technology first published a report on von Economo neurons found in theanterior cingulate cortex (ACC) of hominids but not any other species. Neuronal volumes of ACC von Economo neurons were larger in humans andbonobos than the von Economo neurons of thechimpanzee,gorilla andorangutan.[13]
Allman and his colleagues have delved beyond the level of brain infrastructure to investigate how von Economo neurons function at the superstructural level, focusing on their role as "air traffic controllers for emotions ... at the heart of the human social emotion circuitry, including a moral sense".[14][15] Allman's team proposes that von Economo neurons help channel neural signals from deep within the cortex to relatively distant parts of the brain.[14] Specifically, Allman's team found signals from the ACC are received inBrodmann's area 10, in the frontal polar cortex, where regulation ofcognitive dissonance (disambiguation between alternatives) is thought to occur. According to Allman, this neural relay appears to convey motivation to act, and concerns the recognition of error. Self-control – and avoidance of error – is thus facilitated by the executive gatekeeping function of the ACC, as it regulates the interference patterns of neural signals between these two brain regions.[16]
In humans, intenseemotion activates the anterior cingulate cortex, as it relays neural signals transmitted from theamygdala (a primary processing center for emotions) to thefrontal cortex, perhaps by functioning as a sort of lens to focus the complex texture of neural signal interference patterns.[citation needed] The ACC is also active during demanding tasks requiring judgement and discrimination and when errors are detected by an individual. During difficult tasks, or when experiencing intense love, anger, or lust, activation of the ACC increases. In brain imaging studies, the ACC has specifically been found to be active when mothers hear infants cry, underscoring its role in affording a heightened degree of social sensitivity.
The ACC is a relatively ancient cortical region and is involved with manyautonomic functions, including motor and digestive functions, while also playing a role in the regulation of blood pressure and heart rate. Significantolfactory andgustatory capabilities of the ACC and fronto-insular cortex appear to have been usurped, during recent evolution, to serve enhanced roles related to higher cognition – ranging from planning and self-awareness to role-playing and deception. The diminished olfactory function of humans, compared with other primates, may be related to the fact that von Economo neurons located at crucial neural network hubs have only two dendrites rather than many, resulting in reduced neurological integration.[citation needed]
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At aSociety for Neuroscience meeting in 2003, Allman reported on von Economo neurons his team found in another brain region, the fronto-insular cortex, a region which appears to have undergone significant evolutionary adaptations in mankind – perhaps as recently as 100,000 years ago.
This fronto-insular cortex is closely connected to theinsula, a region that is roughly the size of a thumb in each hemisphere of the human brain. The insula and fronto-insular cortex are part of theinsular cortex, wherein the elaborate circuitry associated withspatial awareness are found, and whereself-awareness and the complexities of emotion are thought to be generated and experienced. Moreover, this region of the right hemisphere is crucial tonavigation and perception of three-dimensional rotations.
The largest number of ACC von Economo neurons are found in humans, fewer in the gracile great apes, and fewest in the robust great apes. In both humans and bonobos they are often found in clusters of 3 to 6 neurons. They are found in humans,chimpanzees,gorillas,orangutans, somecetaceans, andelephants.[17]: 245 While total quantities of ACC von Economo neurons were not reported by Allman in his seminal research report (as they were in a later report describing their presence in the frontoinsular cortex, below), his team's initial analysis of the ACC layer V inhominids revealed an average of ~9 von Economo neurons per section fororangutans (rare, 0.6% of section cells), ~22 forgorillas (frequent, 2.3%), ~37 forchimpanzees (abundant, 3.8%), ~68 for bonobos (abundant/clusters, 4.8%), ~89 for humans (abundant/clusters, 5.6%).[18]
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All of the primates examined had more von Economo neurons in the fronto-insula of the right hemisphere than in the left. In contrast to the higher number of von Economo neurons found in the ACC of the gracile bonobos andchimpanzees, the number of fronto-insular von Economo neurons was far higher in the cortex of robustgorillas (no data fororangutans was given). An adult human had 82,855 such cells, a gorilla had 16,710, a bonobo had 2,159, and a chimpanzee had a mere 1,808 – despite the fact that chimpanzees and bonobos are great apes most closely related to humans.
Von Economo neurons have been located in thedorsolateral prefrontal cortex of humans[1] and elephants.[5] In humans they have been observed in higher concentration inBrodmann area 9 (BA9) – mostly isolated, or in clusters of 2, while inBrodmann area 24 (BA24) they have been found mostly in clusters of 2–4.[1]
Abnormal von Economo neuron development may be linked to several psychotic disorders, typically those characterized by distortions of reality, disturbances of thought, disturbances of language, and withdrawal from social contact.[citation needed] Altered von Economo neuron states have been implicated in bothschizophrenia andautism, but research into these correlations remains at a very early stage.[citation needed]Frontotemporal dementia involves loss of mostly von Economo neurons.[19] An initial study suggested thatAlzheimer's disease specifically targeted von Economo neurons; this study was performed with end-stage Alzheimer brains in which cell destruction was widespread, but later it was found that Alzheimer's disease does not affect the von Economo neurons.[citation needed]