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The triune brain is a model of the
evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...
of the
vertebrate Vertebrates () comprise all animal taxa within the subphylum Vertebrata () ( chordates with backbones), including all mammals, birds, reptiles, amphibians, and fish. Vertebrates represent the overwhelming majority of the phylum Chordata, with ...
forebrain In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hindbrain (rhombencephalon) are the three primary ...
and behavior, proposed by the American physician and
neuroscientist A neuroscientist (or neurobiologist) is a scientist who has specialised knowledge in neuroscience, a branch of biology that deals with the physiology, biochemistry, psychology, anatomy and molecular biology of neurons, neural circuits, and glial ...
Paul D. MacLean Paul Donald MacLean (May 1, 1913 – December 26, 2007) was an American physician and neuroscientist who made significant contributions in the fields of physiology, psychiatry, and brain research through his work at Yale Medical School and the ...
in the 1960s. The triune brain consists of the reptilian complex (
basal ganglia The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an ext ...
), the paleomammalian complex (
limbic system The limbic system, also known as the paleomammalian cortex, is a set of brain structures located on both sides of the thalamus, immediately beneath the medial temporal lobe of the cerebrum primarily in the forebrain.Schacter, Daniel L. 2012. ''P ...
), and the neomammalian complex (
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
), viewed each as independently conscious, and as structures sequentially added to the forebrain in the course of evolution. The basal ganglia is in charge of our primal instincts, the limbic system is in charge of our emotions and the neocortex is responsible for objective or rational thoughts. Since the 1970s, in some circles of evolutionary and developmental neuroscience, the concept of the triune brain has been subject to criticism and is regarded as a myth. Due to its longevity, the triune brain idea has also been called "one of the most successful and widespread errors in all of science." as the hypothesis is no longer espoused by the majority of comparative neuroscientists in the post-2000 era. The theory has been embraced by some psychiatrists and at least one leading affective neuroscience researcher. MacLean originally formulated his model in the 1960s and propounded it at length in his 1990 book ''The Triune Brain in Evolution''. The triune brain hypothesis became familiar to a broad popular audience through
Carl Sagan Carl Edward Sagan (; ; November 9, 1934December 20, 1996) was an American astronomer, planetary scientist, cosmologist, astrophysicist, astrobiologist, author, and science communicator. His best known scientific contribution is research on ex ...
's
Pulitzer prize The Pulitzer Prize () is an award for achievements in newspaper, magazine, online journalism, literature, and musical composition within the United States. It was established in 1917 by provisions in the will of Joseph Pulitzer, who had made ...
winning 1977 book ''
The Dragons of Eden ''The Dragons of Eden: Speculations on the Evolution of Human Intelligence'' is a 1977 book by Carl Sagan, in which the author combines the fields of anthropology, evolutionary biology, psychology, and computer science to give a perspective on ho ...
''.


Aspects

;Reptilian complex: The reptilian complex (also known as the "R-complex", "reptilian brain" or "lizard brain") was the name MacLean gave to the
basal ganglia The basal ganglia (BG), or basal nuclei, are a group of subcortical nuclei, of varied origin, in the brains of vertebrates. In humans, and some primates, there are some differences, mainly in the division of the globus pallidus into an ext ...
, structures derived from the floor of the forebrain during development. The term derives from the idea that comparative neuroanatomists once believed that the forebrains of reptiles and birds were dominated by these structures. MacLean proposed that the reptilian complex was responsible for species-typical instinctual behaviours involved in aggression, dominance, territoriality, and ritual displays. ;Paleomammalian complex: This consists of the
septum In biology, a septum (Latin for ''something that encloses''; plural septa) is a wall, dividing a cavity or structure into smaller ones. A cavity or structure divided in this way may be referred to as septate. Examples Human anatomy * Interat ...
,
amygdalae 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 verteb ...
,
hypothalamus The hypothalamus () is a part of the brain that contains a number of small nuclei with a variety of functions. One of the most important functions is to link the nervous system to the endocrine system via the pituitary gland. The hypothalamus ...
, hippocampal complex, and
cingulate cortex The cingulate cortex is a part of the brain situated in the medial aspect of the cerebral cortex. The cingulate cortex includes the entire cingulate gyrus, which lies immediately above the corpus callosum, and the continuation of this in the c ...
. MacLean first introduced the term "
limbic system The limbic system, also known as the paleomammalian cortex, is a set of brain structures located on both sides of the thalamus, immediately beneath the medial temporal lobe of the cerebrum primarily in the forebrain.Schacter, Daniel L. 2012. ''P ...
" to refer to this set of interconnected brain structures in a paper in 1952. MacLean's recognition of the limbic system as a major functional system in the brain was widely accepted among neuroscientists, and is generally regarded as his most important contribution to the field. MacLean maintained that the structures of the limbic system arose early in mammalian evolution (hence "paleomammalian", with '' paleo-'' meaning ''old'') and were responsible for the motivation and emotion involved in feeding, reproductive behaviour, and parental behaviour. ;Neomammalian complex: This consists of the cerebral
neocortex The neocortex, also called the neopallium, isocortex, or the six-layered cortex, is a set of layers of the mammalian cerebral cortex involved in higher-order brain functions such as sensory perception, cognition, generation of motor commands, sp ...
, a structure found uniquely in higher mammals, and especially humans. MacLean regarded its addition as the most recent step in the evolution of the mammalian brain, conferring the ability for language, abstraction, planning, and perception.


Interactions between structures

The triune brain model argues that these structures are relatively independent from one another, but that they are still connected to each other in some form or another. The model views different cognitive behaviors as caused by three different entities instead of one. The reptilian complex is said to control all of the instinctual and impulsive actions, while the neomammalian complex is responsible for keeping the primitive instincts constrained. An example is controlling the impulse of eating. It seems that if one is hungry, then that means the reptilian complex is commanding the body to eat. However, an individual has the rational choice not to eat when hungry, and this rational thought is said to be controlled by the neomammalian complex. The model thus suggest that these two (and three depending on the situation) structures are in a perpetual battle to control the body. These interactions between the neocortex and the reptilian brain often seem competitive as the conscious thought generated by the neocortex can supress the primitive thoughts generated by the reptilian complex. Thus, the model suggests that the interactions between structures are not constructive, but that they are conflicting due to the anatomical separation of the brain. This separation of structures proposed an underlying difference between consciousness and unconscious behaviour and argued that the reason why humans are such intelligent and conscious species is due to the not-so-common neocortex that they possess, unlike most other animals. This detachment contributes to the idea that the three complexes interact with each other separately rather than a single construct interacting with itself.


Status of the model

MacLean originally formulated the triune brain hypothesis in the 1960s, drawing on comparative neuroanatomical work done by
Ludwig Edinger } Ludwig Edinger (13 April 1855 – 26 January 1918) was an influential German anatomist and neurologist and co-founder of the University of Frankfurt. In 1914 he was also appointed the first German professor of neurology. Edinger was born ...
, Elizabeth C. Crosby and Charles Judson Herrick early in the twentieth century. Striedter, G. F. (2005) ''Principles of Brain Evolution.'' Sinauer Associates. The 1980s saw a rebirth of interest in comparative neuroanatomy, motivated in part by the availability of a variety of new neuroanatomical techniques for charting the circuitry of animal brains. Subsequent findings according to human brain evolution expert Dr. Terrence Deacon, have refined the traditional neuroanatomical ideas upon which MacLean based his hypothesis. Deacon mentioned that 'the evolutionary addition of different parts of the brain is simply not realistic. However, all the parts of the brain were already existing, they were just further developed upon as the homosapien species evolved and gained life experiences.' For example, the basal ganglia (structures derived from the floor of the forebrain and making up MacLean's reptilian complex) were shown to take up a much smaller portion of the forebrains of reptiles and birds (together called
sauropsids Sauropsida ("lizard faces") is a clade of amniotes, broadly equivalent to the class Reptilia. Sauropsida is the sister taxon to Synapsida, the other clade of amniotes which includes mammals as its only modern representatives. Although early syn ...
) than previously supposed, and to exist in
amphibians Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...
and
fish Fish are Aquatic animal, aquatic, craniate, gill-bearing animals that lack Limb (anatomy), limbs with Digit (anatomy), digits. Included in this definition are the living hagfish, lampreys, and Chondrichthyes, cartilaginous and bony fish as we ...
as well as
mammals 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), fur o ...
and sauropsids. Because the basal ganglia are found in the forebrains of all modern vertebrates, they most likely date to the common evolutionary ancestor of the vertebrates, more than 500 million years ago, rather than to the origin of reptiles. Some recent behavioral studies do not support the traditional view of sauropsid behavior as stereotyped and ritualistic (as in MacLean's reptilian complex). Birds have been shown to possess highly sophisticated cognitive abilities, such as the toolmaking of the
New Caledonian crow The New Caledonian crow (''Corvus moneduloides'') is a medium-sized member of the family Corvidae, native to New Caledonia. The bird is often referred to as the 'qua-qua' due to its distinctive call. It eats a wide range of food, including man ...
and the language-like categorization abilities of the
grey parrot The grey parrot (''Psittacus erithacus''), also known as the Congo grey parrot, Congo African grey parrot or African grey parrot, is an Old World parrot in the family Psittacidae. The Timneh parrot ''(Psittacus timneh)'' once was identified as ...
. Structures of the limbic system, which MacLean proposed arose in early mammals, have now been shown to exist across a range of modern vertebrates. The "paleomammalian" trait of parental care of offspring is widespread in birds and occurs in some fishes as well. Thus, like the basal ganglia, the evolution of these systems presumably dates to a common vertebrate ancestor.Butler, A. B. and Hodos, W. Comparative Vertebrate Neuroanatomy: Evolution and Adaptation, Wiley Finally, recent studies based on paleontological data or comparative anatomical evidence strongly suggest that the neocortex was already present in the earliest emerging mammals. In addition, although non-mammals do not have a neocortex in the true sense (that is, a structure comprising part of the forebrain roof, or pallium, consisting of six characteristic layers of neurons), they possess pallial regions, and some parts of the pallium are considered homologous to the mammalian neocortex. While these areas lack the characteristic six neocortical layers, birds and reptiles generally possess three layers in the dorsal pallium (the homolog of the mammalian neocortex). The
telencephalon The cerebrum, telencephalon or endbrain is the largest part of the brain containing the cerebral cortex (of the two cerebral hemispheres), as well as several subcortical structures, including the hippocampus, basal ganglia, and olfactory bulb. ...
of birds and mammals makes neuroanatomical connections with other telecencephalic structures like those made by neocortex. It mediates similar functions such as perception, learning and memory, decision making, motor control, conceptual thinking.


Lay interest

The triune model of the mammalian brain is seen as an oversimplified organizing theme by some in the field of comparative neuroscience. It continues to hold public interest because of its simplicity. While inaccurate in many respects as an explanation for brain activity, structure and evolution, it remains a commonly used concept as the "neocortex" represents that cluster of brain structures involved in advanced cognition, including planning, modeling and simulation; the "limbic brain" refers to those brain structures, wherever located, associated with social and nurturing behaviors, mutual reciprocity, and other behaviors and affects that arose during the age of the mammals; and the "reptilian brain" refers to those brain structures related to territoriality, ritual behavior and other "reptile" behaviors. The simple explanatory value makes this approximation engaging and may be a useful level of complexity for high-school students to begin engaging with brain research.
Howard Bloom Howard Bloom (born June 25, 1943) is an American author. He was a music publicist in the 1970s and 1980s for singers and bands such as Prince, Billy Joel, and Styx.The Lucifer Principle'', references the concept of the triune brain in his explanations of certain aspects of human behavior.
Arthur Koestler Arthur Koestler, (, ; ; hu, Kösztler Artúr; 5 September 1905 – 1 March 1983) was a Hungarian-born author and journalist. Koestler was born in Budapest and, apart from his early school years, was educated in Austria. In 1931, Koestler join ...
made MacLean's concept of the triune brain the centerpiece of much of his later work, notably ''
The Ghost in the Machine ''The Ghost in the Machine'' is a 1967 book about philosophical psychology by Arthur Koestler. The title is a phrase (see ghost in the machine) coined by the Oxford philosopher Gilbert Ryle to describe the Cartesian dualist account of the mi ...
''. English novelist
Julian Barnes Julian Patrick Barnes (born 19 January 1946) is an English writer. He won the Man Booker Prize in 2011 with ''The Sense of an Ending'', having been shortlisted three times previously with '' Flaubert's Parrot'', '' England, England'', and ''Art ...
quotes MacLean on the triune brain in the foreword to his 1982 novel ''Before She Met Me''. Peter A. Levine uses the triune brain concept in his book ''Waking the Tiger'' to explain his
somatic experiencing Somatic experiencing (SE) is a form of alternative therapy aimed at treating trauma and stressor-related disorders like PTSD. The primary goal of SE is to modify the trauma-related stress response through bottom-up processing. The Clients’ ...
approach to healing trauma. Glynda-Lee Hoffmann, in her book ''The Secret Dowry of Eve, Women's Role in the Development of Consciousness'', references the triune theory explored by MacLean and goes one step further. Her theory about human behavior, and the problems we create with that behavior, distinguishes the
prefrontal cortex In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, BA13, BA14, BA24, BA25, BA32, BA44, BA45, BA ...
as uniquely different from the rest of the neocortex. The prefrontal cortex, with its agenda of integration, is the part of the brain that can get the other parts to work together for the good of the individual. Hoffmann claims that in many humans the reptilian cortex (agenda: territory and reproduction; in humans that translates to power and sex) is out of control, and the amygdala stokes the fear that leads to more bad behavior. The prefrontal cortex, she believes, is the key to our future if we can harness its power.


References


Further reading

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