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Von Economo neurons (VENs), also called spindle neurons, are a specific class of
mammalian Mammals () are a group of vertebrate animals constituting the class Mammalia (), characterized by the presence of mammary glands which in females produce milk for feeding (nursing) their young, a neocortex (a region of the brain), fu ...
cortical
neurons A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa ...
characterized by a large spindle-shaped
soma Soma may refer to: Businesses and brands * SOMA (architects), a New York–based firm of architects * Soma (company), a company that designs eco-friendly water filtration systems * SOMA Fabrications, a builder of bicycle frames and other bicycle ...
(or body) gradually tapering into a single apical
axon An axon (from Greek ἄξων ''áxōn'', axis), or nerve fiber (or nerve fibre: see spelling differences), is a long, slender projection of a nerve cell, or neuron, in vertebrates, that typically conducts electrical impulses known as action p ...
(the ramification that ''transmits'' signals) in one direction, with only a single
dendrite Dendrites (from Greek δένδρον ''déndron'', "tree"), also dendrons, are branched protoplasmic extensions of a nerve cell that propagate the electrochemical stimulation received from other neural cells to the cell body, or soma, of the ...
(the ramification that ''receives'' signals) facing opposite. Other cortical neurons tend to have many dendrites, and the bipolar-shaped morphology of von Economo neurons is unique here. Von Economo neurons are found in two very restricted regions in the
brain A brain is an organ (biology), organ that serves as the center of the nervous system in all vertebrate and most invertebrate animals. It is located in the head, usually close to the sensory organs for senses such as Visual perception, vision. I ...
s of
hominid The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ...
s (
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s and other
great apes The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ...
): the
anterior cingulate cortex In the human brain, the anterior cingulate cortex (ACC) is the frontal part of the cingulate cortex that resembles a "collar" surrounding the frontal part of the corpus callosum. It consists of Brodmann areas 24, 32, and 33. It is involve ...
(ACC) and the fronto-insular cortex (FI) (which each make up the salience network). In 2008, they were also found in the
dorsolateral prefrontal cortex The dorsolateral prefrontal cortex (DLPFC or DL-PFC) is an area in the prefrontal cortex of the primate brain. It is one of the most recently derived parts of the human brain. It undergoes a prolonged period of maturation which lasts until adult ...
of humans. Von Economo neurons are also found in the brains of a number of
cetaceans Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
, African and
Asian elephant The Asian elephant (''Elephas maximus''), also known as the Asiatic elephant, is the only living species of the genus '' Elephas'' and is distributed throughout the Indian subcontinent and Southeast Asia, from India in the west, Nepal in t ...
s, and to a lesser extent in
macaque monkey The macaques () constitute a genus (''Macaca'') of gregarious Old World monkeys of the subfamily Cercopithecinae. The 23 species of macaques inhabit ranges throughout Asia, North Africa, and (in one instance) Gibraltar. Macaques are principall ...
s and
raccoon The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of ...
s. The appearance of von Economo neurons in distantly related
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
s suggests that they represent
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
specifically, as an adaptation to accommodate the increasing size of these distantly-related animals' brains. Von Economo neurons were discovered and first described in 1929 by Austrian psychiatrist and neurologist
Constantin von Economo Constantin Freiherr von Economo ( gr, Κωνσταντίνος Οικονόμου; 21 August 1876 – 21 October 1931) was an Austrian psychiatrist and neurologist of Greek descent, born in modern-day Romania (then Ottoman Empire). He is mos ...
(1876–1931).


Name

Von Economo neurons are named after the Austrian neurologist
Constantin von Economo Constantin Freiherr von Economo ( gr, Κωνσταντίνος Οικονόμου; 21 August 1876 – 21 October 1931) was an Austrian psychiatrist and neurologist of Greek descent, born in modern-day Romania (then Ottoman Empire). He is mos ...
, who first described these neurons in 1926.


Function

Von Economo neurons are relatively large
cell Cell most often refers to: * Cell (biology), the functional basic unit of life Cell may also refer to: Locations * Monastic cell, a small room, hut, or cave in which a religious recluse lives, alternatively the small precursor of a monastery ...
s that may allow rapid communication across the relatively large brains of
great ape The Hominidae (), whose members are known as the great apes or hominids (), are a taxonomic family of primates that includes eight extant species in four genera: '' Pongo'' (the Bornean, Sumatran and Tapanuli orangutan); ''Gorilla'' (the ...
s,
elephant Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantida ...
s, and
cetaceans Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
. Although rare in comparison to other neurons, von Economo neurons are abundant, and comparatively large, in humans; they are however three times as abundant in
cetacea Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
ns.


Evolutionary significance

The discovery of von Economo neurons in diverse whale species 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." The apparent presence of these specialized neurons only in highly intelligent mammals may be an example of
convergent evolution Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last com ...
. Their restriction among the primates to great apes 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 and raccoons.


In the anterior cingulate cortex

In 1999, American neuroscientist
John Allman John Morgan Allman is an American neuroscientist at the California Institute of Technology in Pasadena, California, active in the fields of primates, cognition and evolutionary neuroscience. Life He graduated from University of Chicago with a P ...
and colleagues at the
California Institute of Technology The California Institute of Technology (branded as Caltech or CIT)The university itself only spells its short form as "Caltech"; the institution considers other spellings such a"Cal Tech" and "CalTech" incorrect. The institute is also occasional ...
first published a report on von Economo neurons found in the
anterior cingulate cortex In the human brain, the anterior cingulate cortex (ACC) is the frontal part of the cingulate cortex that resembles a "collar" surrounding the frontal part of the corpus callosum. It consists of Brodmann areas 24, 32, and 33. It is involve ...
(ACC) of hominids but not any other species. Neuronal volumes of ACC von Economo neurons were larger in
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s and
bonobo The bonobo (; ''Pan paniscus''), also historically called the pygmy chimpanzee and less often the dwarf chimpanzee or gracile chimpanzee, is an endangered great ape and one of the two species making up the genus '' Pan,'' the other being the comm ...
s than the von Economo neurons of the
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...
,
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four ...
and
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genu ...
. 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». Allman's team proposes that von Economo neurons help channel neural signals from deep within the cortex to relatively distant parts of the brain. Specifically, Allman's team found signals from the ACC are received in Brodmann's area 10, in the frontal polar cortex, where regulation of
cognitive dissonance In the field of psychology, cognitive dissonance is the perception of contradictory information, and the mental toll of it. Relevant items of information include a person's actions, feelings, ideas, beliefs, values, and things in the environmen ...
(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-controland avoidance of erroris thus facilitated by the executive gatekeeping function of the ACC, as it regulates the interference patterns of neural signals between these two brain regions. In humans, intense
emotion Emotions are mental states brought on by neurophysiological changes, variously associated with thoughts, feelings, behavioral responses, and a degree of pleasure or displeasure. There is currently no scientific consensus on a definitio ...
activates the anterior cingulate cortex, as it relays neural signals transmitted from the
amygdala The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex ver ...
(a primary processing center for emotions) to the
frontal cortex The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere (in front of the parietal lobe and the temporal lobe). It is parted from the parietal lobe by a groove be ...
, perhaps by functioning as a sort of lens to focus the complex texture of neural signal interference patterns. 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 many autonomic functions, including motor and digestive functions, while also playing a role in the regulation of blood pressure and heart rate. Significant
olfactory The sense of smell, or olfaction, is the special sense through which smells (or odors) are perceived. The sense of smell has many functions, including detecting desirable foods, hazards, and pheromones, and plays a role in taste. In humans, ...
and gustatory capabilities of the ACC and fronto-insular cortex appear to have been usurped, during recent evolution, to serve enhanced roles related to higher cognitionranging from planning and self-awareness to role-playing and deception. The diminished olfactory function of humans, compared to 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.


In the fronto-insular cortex

At a
Society for Neuroscience The Society for Neuroscience (SfN) is a professional society, headquartered in Washington, DC, for basic scientists and physicians around the world whose research is focused on the study of the brain and nervous system. It is especially well kn ...
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 mankindperhaps as recently as 100,000 years ago. This fronto-insular cortex is closely connected to the insula, 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 the
insular cortex The insular cortex (also insula and insular lobe) is a portion of the cerebral cortex folded deep within the lateral sulcus (the fissure separating the temporal lobe from the parietal and frontal lobes) within each hemisphere of the mammalian b ...
, wherein the elaborate circuitry associated with
spatial awareness Spatial contextual awareness consociates contextual information such as an individual's or sensor's location, activity, the time of day, and proximity to other people or objects and devices.Chen, Guanling, and David Kotz. 2000. A Survey of Context- ...
are found, and where
self-awareness In philosophy of self, self-awareness is the experience of one's own personality or individuality. It is not to be confused with consciousness in the sense of qualia. While consciousness is being aware of one's environment and body and lifest ...
and the complexities of emotion are thought to be generated and experienced. Moreover, this region of the right hemisphere is crucial to
navigation Navigation is a field of study that focuses on the process of monitoring and controlling the movement of a craft or vehicle from one place to another.Bowditch, 2003:799. The field of navigation includes four general categories: land navigation ...
and perception of three-dimensional rotations.


Concentrations


Anterior cingulate cortex

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
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s,
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...
s,
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four ...
s,
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genu ...
s, some
cetacean Cetacea (; , ) is an infraorder of aquatic mammals that includes whales, dolphins, and porpoises. Key characteristics are their fully aquatic lifestyle, streamlined body shape, often large size and exclusively carnivorous diet. They propel th ...
s, and
elephants Elephants are the largest existing land animals. Three living species are currently recognised: the African bush elephant, the African forest elephant, and the Asian elephant. They are the only surviving members of the family Elephantidae ...
. 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 in hominids revealed an average of ~9 von Economo neurons per section for
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genu ...
s (rare, 0.6% of section cells), ~22 for
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four ...
s (frequent, 2.3%), ~37 for
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...
s (abundant, 3.8%), ~68 for bonobos (abundant/clusters, 4.8%), ~89 for
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
s (abundant/clusters, 5.6%).


Fronto-insular cortex

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 and
chimpanzee The chimpanzee (''Pan troglodytes''), also known as simply the chimp, is a species of great ape native to the forest and savannah of tropical Africa. It has four confirmed subspecies and a fifth proposed subspecies. When its close relative t ...
s, the number of fronto-insular von Economo neurons was far higher in the cortex of robust
gorilla Gorillas are herbivorous, predominantly ground-dwelling great apes that inhabit the tropical forests of equatorial Africa. The genus ''Gorilla'' is divided into two species: the eastern gorilla and the western gorilla, and either four ...
s (no data for
orangutan Orangutans are great apes native to the rainforests of Indonesia and Malaysia. They are now found only in parts of Borneo and Sumatra, but during the Pleistocene they ranged throughout Southeast Asia and South China. Classified in the genu ...
s was given). An adult
human Humans (''Homo sapiens'') are the most abundant and widespread species of primate, characterized by bipedalism and exceptional cognitive skills due to a large and complex brain. This has enabled the development of advanced tools, cultu ...
had 82,855 such cells, a gorilla had 16,710, a bonobo had 2,159, and a chimpanzee had a mere 1,808despite the fact that chimpanzees and bonobos are great apes most closely related to humans.


Dorsolateral prefrontal cortex

Von Economo neurons have been located in the
Dorsolateral prefrontal cortex The dorsolateral prefrontal cortex (DLPFC or DL-PFC) is an area in the prefrontal cortex of the primate brain. It is one of the most recently derived parts of the human brain. It undergoes a prolonged period of maturation which lasts until adult ...
of humans and elephants. In humans they have been observed in higher concentration in
Brodmann area 9 Brodmann area 9, or BA9, refers to a cytoarchitecturally defined portion of the frontal cortex in the brain of humans and other primates. It contributes to the dorsolateral and medial prefrontal cortex. Functions The area is involved in sho ...
(BA9)mostly isolated or in clusters of 2, while in
Brodmann area 24 Brodmann area 24 is part of the anterior cingulate in the human brain. Human In the human this area is known as ventral anterior cingulate area 24, and it refers to a subdivision of the cytoarchitecturally defined cingulate cortex region of cereb ...
(BA24) they have been found mostly in clusters of 2–4.


Clinical significance

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. Altered von Economo neuron states have been implicated in both
schizophrenia Schizophrenia is a mental disorder characterized by continuous or relapsing episodes of psychosis. Major symptoms include hallucinations (typically hearing voices), delusions, and disorganized thinking. Other symptoms include social w ...
and
autism The autism spectrum, often referred to as just autism or in the context of a professional diagnosis autism spectrum disorder (ASD) or autism spectrum condition (ASC), is a neurodevelopmental condition (or conditions) characterized by difficulti ...
, but research into these correlations remains at a very early stage.
Frontotemporal dementia Frontotemporal dementia (FTD), or frontotemporal degeneration disease, or frontotemporal neurocognitive disorder, encompasses several types of dementia involving the progressive degeneration of frontal and temporal lobes. FTDs broadly present as ...
involves loss of mostly von Economo neurons. An initial study suggested that Alzheimer'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 doesn't affect the von Economo neurons.


References

;General References *


External links


TaipeiTimes.com
{spaced ndashKnow Thyself and Others
"Well-wired whales"
Michael Balter (2006) ScienceNOW Daily News. 27 November

''Smithsonian'', June 2009 Central nervous system neurons