Putamen
Encyclopedia
"Putamen" is also a botanical
Botany
Botany, plant science, or plant biology is a branch of biology that involves the scientific study of plant life. Traditionally, botany also included the study of fungi, algae and viruses...

 term for the stone in a fruit
Fruit
In broad terms, a fruit is a structure of a plant that contains its seeds.The term has different meanings dependent on context. In non-technical usage, such as food preparation, fruit normally means the fleshy seed-associated structures of certain plants that are sweet and edible in the raw state,...

, such as a peach
Peach
The peach tree is a deciduous tree growing to tall and 6 in. in diameter, belonging to the subfamily Prunoideae of the family Rosaceae. It bears an edible juicy fruit called a peach...

.


The putamen is a round structure located at the base of the forebrain (telencephalon
Telencephalon
The cerebrum or telencephalon, together with the diencephalon, constitutes the forebrain. The cerebrum is the most anterior region of the vertebrate central nervous system. Telencephalon refers to the embryonic structure, from which the mature cerebrum develops...

). The putamen and caudate nucleus
Caudate nucleus
The caudate nucleus is a nucleus located within the basal ganglia of the brains of many animal species. The caudate nucleus is an important part of the brain's learning and memory system.-Anatomy:...

 together form the dorsal striatum
Dorsal striatum
The dorsal striatum, corpus striatum or striated body is a compound structure consisting of the caudate nucleus, and the lentiform nucleus...

. It is also one of the structures that comprises the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

. Through various pathways, the putamen is connected to the substantia nigra
Substantia nigra
The substantia nigra is a brain structure located in the mesencephalon that plays an important role in reward, addiction, and movement. Substantia nigra is Latin for "black substance", as parts of the substantia nigra appear darker than neighboring areas due to high levels of melanin in...

 and globus pallidus
Globus pallidus
The globus pallidus also known as paleostriatum, is a sub-cortical structure of the brain. Topographically, it is part of the telencephalon, but retains close functional ties with the subthalamus - both of which are part of the extrapyramidal motor system...

. The main function of the putamen is to regulate movements and influence various types of learning
Learning
Learning is acquiring new or modifying existing knowledge, behaviors, skills, values, or preferences and may involve synthesizing different types of information. The ability to learn is possessed by humans, animals and some machines. Progress over time tends to follow learning curves.Human learning...

. It employs dopamine
Dopamine
Dopamine is a catecholamine neurotransmitter present in a wide variety of animals, including both vertebrates and invertebrates. In the brain, this substituted phenethylamine functions as a neurotransmitter, activating the five known types of dopamine receptors—D1, D2, D3, D4, and D5—and their...

 to perform its functions. The putamen also plays a role in degenerative neurological disorders
Degenerative disease
A degenerative disease, also called neurodegenerative disease, is a disease in which the function or structure of the affected tissues or organs will progressively deteriorate over time, whether due to normal bodily wear or lifestyle choices such as exercise or eating habits...

, such as Parkinson's disease
Parkinson's disease
Parkinson's disease is a degenerative disorder of the central nervous system...

.

History

The word "putamen" is from Latin
Latin
Latin is an Italic language originally spoken in Latium and Ancient Rome. It, along with most European languages, is a descendant of the ancient Proto-Indo-European language. Although it is considered a dead language, a number of scholars and members of the Christian clergy speak it fluently, and...

, referring to that which "falls off in pruning", from "puto", meaning "to prune". It is pronounced pyu-ta'men.

Until recently, very few studies were conducted that were focused specifically on the putamen. However, many studies have been done on the basal ganglia and the interactions among the brain structures it comprises. In the 1970s, the first single unit recordings
Single-unit recording
In neurophysiology and neurology, single-unit recording is the use of an electrode to record the electrophysiological activity from a single neuron.-History:...

 were done with monkey
Monkey
A monkey is a primate, either an Old World monkey or a New World monkey. There are about 260 known living species of monkey. Many are arboreal, although there are species that live primarily on the ground, such as baboons. Monkeys are generally considered to be intelligent. Unlike apes, monkeys...

s monitoring pallidal neuron
Neuron
A neuron is an electrically excitable cell that processes and transmits information by electrical and chemical signaling. Chemical signaling occurs via synapses, specialized connections with other cells. Neurons connect to each other to form networks. Neurons are the core components of the nervous...

 activity related to movement.

Anatomy

The putamen is a structure in the forebrain. Along with the caudate nucleus
Caudate nucleus
The caudate nucleus is a nucleus located within the basal ganglia of the brains of many animal species. The caudate nucleus is an important part of the brain's learning and memory system.-Anatomy:...

 it forms the dorsal striatum
Dorsal striatum
The dorsal striatum, corpus striatum or striated body is a compound structure consisting of the caudate nucleus, and the lentiform nucleus...

. The caudate and putamen contain the same types of neurons and circuits - many neuroanatomists consider the dorsal striatum to be a single structure, divided into two parts by a large fiber tract, the internal capsule
Internal capsule
The internal capsule is an area of white matter in the brain that separates the caudate nucleus and the thalamus from the lenticular nucleus. The internal capsule contains both ascending and descending axons....

, passing through the middle. The putamen, together with the globus pallidus
Globus pallidus
The globus pallidus also known as paleostriatum, is a sub-cortical structure of the brain. Topographically, it is part of the telencephalon, but retains close functional ties with the subthalamus - both of which are part of the extrapyramidal motor system...

, makes up the lenticular nucleus. The putamen is the outer most portion of the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

. These are a group of nuclei in the brain that are interconnected with the cerebral cortex
Cerebral cortex
The cerebral cortex is a sheet of neural tissue that is outermost to the cerebrum of the mammalian brain. It plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It is constituted of up to six horizontal layers, each of which has a different...

, thalamus
Thalamus
The thalamus is a midline paired symmetrical structure within the brains of vertebrates, including humans. It is situated between the cerebral cortex and midbrain, both in terms of location and neurological connections...

, and brain stem
Brain stem
In vertebrate anatomy the brainstem is the posterior part of the brain, adjoining and structurally continuous with the spinal cord. The brain stem provides the main motor and sensory innervation to the face and neck via the cranial nerves...

. The other parts of the basal ganglia include the dorsal striatum
Dorsal striatum
The dorsal striatum, corpus striatum or striated body is a compound structure consisting of the caudate nucleus, and the lentiform nucleus...

, substantia nigra
Substantia nigra
The substantia nigra is a brain structure located in the mesencephalon that plays an important role in reward, addiction, and movement. Substantia nigra is Latin for "black substance", as parts of the substantia nigra appear darker than neighboring areas due to high levels of melanin in...

, nucleus accumbens
Nucleus accumbens
The nucleus accumbens , also known as the accumbens nucleus or as the nucleus accumbens septi , is a collection of neurons and forms the main part of the ventral striatum...

, and the subthalamic nucleus
Subthalamic nucleus
The subthalamic nucleus is a small lens-shaped nucleus in the brain where it is, from a functional point of view, part of the basal ganglia system. Anatomically, it is the major part of subthalamus. As suggested by its name, the subthalamic nucleus is located ventral to the thalamus. It is also...

.

In mammals, the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 are associated with motor control
Motor control
Motor control are information processing related activities carried out by the central nervous system that organize the musculoskeletal system to create coordinated movements and skilled actions...

, cognition
Cognition
In science, cognition refers to mental processes. These processes include attention, remembering, producing and understanding language, solving problems, and making decisions. Cognition is studied in various disciplines such as psychology, philosophy, linguistics, and computer science...

, emotion
Emotion
Emotion is a complex psychophysiological experience of an individual's state of mind as interacting with biochemical and environmental influences. In humans, emotion fundamentally involves "physiological arousal, expressive behaviors, and conscious experience." Emotion is associated with mood,...

s, and learning
Learning
Learning is acquiring new or modifying existing knowledge, behaviors, skills, values, or preferences and may involve synthesizing different types of information. The ability to learn is possessed by humans, animals and some machines. Progress over time tends to follow learning curves.Human learning...

. The basal ganglia are located on the left and right sides of the brain, and have rostral and caudal divisions. The putamen is located in the rostral division as part of the striatum
Striatum
The striatum, also known as the neostriatum or striate nucleus, is a subcortical part of the forebrain. It is the major input station of the basal ganglia system. The striatum, in turn, gets input from the cerebral cortex...

. The basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 receive input from the cerebral cortex, via the striatum
Striatum
The striatum, also known as the neostriatum or striate nucleus, is a subcortical part of the forebrain. It is the major input station of the basal ganglia system. The striatum, in turn, gets input from the cerebral cortex...

.

The putamen is interconnected with the following structures:

Caudate Nucleus

The caudate works with the putamen to receive the input from cerebral cortex
Cerebral cortex
The cerebral cortex is a sheet of neural tissue that is outermost to the cerebrum of the mammalian brain. It plays a key role in memory, attention, perceptual awareness, thought, language, and consciousness. It is constituted of up to six horizontal layers, each of which has a different...

. They can be considered the "entrance" to the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

. The nucleus accumbens
Nucleus accumbens
The nucleus accumbens , also known as the accumbens nucleus or as the nucleus accumbens septi , is a collection of neurons and forms the main part of the ventral striatum...

 and medial caudate receive input from frontal cortex and limbic regions. The putamen and caudate are jointly connected with the substantia nigra
Substantia nigra
The substantia nigra is a brain structure located in the mesencephalon that plays an important role in reward, addiction, and movement. Substantia nigra is Latin for "black substance", as parts of the substantia nigra appear darker than neighboring areas due to high levels of melanin in...

, but most of their output goes to the globus pallidus
Globus pallidus
The globus pallidus also known as paleostriatum, is a sub-cortical structure of the brain. Topographically, it is part of the telencephalon, but retains close functional ties with the subthalamus - both of which are part of the extrapyramidal motor system...

.

Substantia Nigra

The substantia nigra
Substantia nigra
The substantia nigra is a brain structure located in the mesencephalon that plays an important role in reward, addiction, and movement. Substantia nigra is Latin for "black substance", as parts of the substantia nigra appear darker than neighboring areas due to high levels of melanin in...

 contains two parts: the substantia nigra pars compacta (SNpc) and the substantia nigra pars reticulata (SNpr). The SNpc obtains input from the putamen and caudate, and sends information back. The SNpr also obtains input from the putamen and caudate. However, it sends the input outside the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 to control head and eye movements. The SNpc produces dopamine, which is crucial for movements. The SNpc is the part which degenerates during Parkinson's disease
Parkinson's disease
Parkinson's disease is a degenerative disorder of the central nervous system...

.

Globus Pallidus

The globus pallidus
Globus pallidus
The globus pallidus also known as paleostriatum, is a sub-cortical structure of the brain. Topographically, it is part of the telencephalon, but retains close functional ties with the subthalamus - both of which are part of the extrapyramidal motor system...

 contains two parts: the globus pallidus externa (GPe) and the globus pallidus interna (GPi). Both regions acquire input from the putamen and caudate and communicate with the subthalamic nucleus. However, mostly the GPi sends the inhibitory output from the basal ganglia to the thalamus. The GPi also sends a few projections to parts of the midbrain, which have been assumed to affect posture control.

Types of Pathways

In order to control movement, the putamen must interact with the other structures that make up the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

. These include the caudate nucleus and globus pallidus. These two structures, along with the putamen, interact through a series of direct and indirect inhibitory pathways. The direct pathway consists of two inhibitory pathways from the putamen to the substantia nigra and the internal region of the globus pallidus. This pathway uses the neurotransmitter
Neurotransmitter
Neurotransmitters are endogenous chemicals that transmit signals from a neuron to a target cell across a synapse. Neurotransmitters are packaged into synaptic vesicles clustered beneath the membrane on the presynaptic side of a synapse, and are released into the synaptic cleft, where they bind to...

s dopamine
Dopamine
Dopamine is a catecholamine neurotransmitter present in a wide variety of animals, including both vertebrates and invertebrates. In the brain, this substituted phenethylamine functions as a neurotransmitter, activating the five known types of dopamine receptors—D1, D2, D3, D4, and D5—and their...

, GABA
Gabâ
Gabâ or gabaa, for the people in many parts of the Philippines), is the concept of a non-human and non-divine, imminent retribution. A sort of negative karma, it is generally seen as an evil effect on a person because of their wrongdoings or transgressions...

 and substance P
Substance P
In the field of neuroscience, substance P is a neuropeptide: an undecapeptide that functions as a neurotransmitter and as a neuromodulator. It belongs to the tachykinin neuropeptide family. Substance P and its closely related neuropeptide neurokinin A are produced from a polyprotein precursor...

. The indirect pathway consists of three inhibitory pathways that go from the putamen and caudate nucleus to the external region of the globus pallidus. This pathway uses dopamine
Dopamine
Dopamine is a catecholamine neurotransmitter present in a wide variety of animals, including both vertebrates and invertebrates. In the brain, this substituted phenethylamine functions as a neurotransmitter, activating the five known types of dopamine receptors—D1, D2, D3, D4, and D5—and their...

, GABA
Gabâ
Gabâ or gabaa, for the people in many parts of the Philippines), is the concept of a non-human and non-divine, imminent retribution. A sort of negative karma, it is generally seen as an evil effect on a person because of their wrongdoings or transgressions...

 and enkephalin
Enkephalin
An enkephalin is a pentapeptide involved in regulating nociception in the body. The enkephalins are termed endogenous ligands, or specifically endorphins, as they are internally derived and bind to the body's opioid receptors. Discovered in 1975, two forms of enkephalin were revealed, one...

. When there is interplay between the direct and indirect pathways, involuntary movement
Movement disorder
Movement disorders include:* Akathisia * Akinesia * Associated Movements * Athetosis...

s occur.

Dopamine

One of the main neurotransmitters that is regulated by the putamen is dopamine
Dopamine
Dopamine is a catecholamine neurotransmitter present in a wide variety of animals, including both vertebrates and invertebrates. In the brain, this substituted phenethylamine functions as a neurotransmitter, activating the five known types of dopamine receptors—D1, D2, D3, D4, and D5—and their...

. When a cell body of a neuron
Neuron
A neuron is an electrically excitable cell that processes and transmits information by electrical and chemical signaling. Chemical signaling occurs via synapses, specialized connections with other cells. Neurons connect to each other to form networks. Neurons are the core components of the nervous...

 (in the putamen or caudate nuclei) fires an action potential
Action potential
In physiology, an action potential is a short-lasting event in which the electrical membrane potential of a cell rapidly rises and falls, following a consistent trajectory. Action potentials occur in several types of animal cells, called excitable cells, which include neurons, muscle cells, and...

, dopamine is released from the presynaptic terminal. Since projections from the putamen and caudate nuclei modulate
Neuromodulation
In Neuromodulation several classes of neurotransmitters regulate diverse populations of central nervous system neurons...

 the dendrites of the substantia nigra
Substantia nigra
The substantia nigra is a brain structure located in the mesencephalon that plays an important role in reward, addiction, and movement. Substantia nigra is Latin for "black substance", as parts of the substantia nigra appear darker than neighboring areas due to high levels of melanin in...

, the dopamine influences the substantia nigra, which affects motor planning. This same mechanism is involved in drug addiction. In order to control the amount of dopamine in the synaptic cleft, and the amount of dopamine binding to post synaptic terminals, presynaptic dopaminergic neurons function to reuptake
Reuptake
Reuptake, or re-uptake, is the reabsorption of a neurotransmitter by a neurotransmitter transporter of a pre-synaptic neuron after it has performed its function of transmitting a neural impulse....

 the excess dopamine.

Other Neurotransmitters

The putamen also plays a role in modulation
Neuromodulation
In Neuromodulation several classes of neurotransmitters regulate diverse populations of central nervous system neurons...

 of other neurotransmitters. It releases GABA
Gabâ
Gabâ or gabaa, for the people in many parts of the Philippines), is the concept of a non-human and non-divine, imminent retribution. A sort of negative karma, it is generally seen as an evil effect on a person because of their wrongdoings or transgressions...

, enkephalin
Enkephalin
An enkephalin is a pentapeptide involved in regulating nociception in the body. The enkephalins are termed endogenous ligands, or specifically endorphins, as they are internally derived and bind to the body's opioid receptors. Discovered in 1975, two forms of enkephalin were revealed, one...

, substance P
Substance P
In the field of neuroscience, substance P is a neuropeptide: an undecapeptide that functions as a neurotransmitter and as a neuromodulator. It belongs to the tachykinin neuropeptide family. Substance P and its closely related neuropeptide neurokinin A are produced from a polyprotein precursor...

, and acetylcholine
Acetylcholine
The chemical compound acetylcholine is a neurotransmitter in both the peripheral nervous system and central nervous system in many organisms including humans...

. It receives serotonin
Serotonin
Serotonin or 5-hydroxytryptamine is a monoamine neurotransmitter. Biochemically derived from tryptophan, serotonin is primarily found in the gastrointestinal tract, platelets, and in the central nervous system of animals including humans...

 and glutamate. The majority of these neurotransmitters play a role in motor control.

Function: Motor Skills

While the putamen has many functions, it has been concluded that it has no specific specialization. However, since the putamen is interconnected with so many other structures, it works in conjunction with them to control many types of motor skills. These include controlling motor learning, motor performance and tasks, motor preparation, specifying amplitudes of movement, and movement sequences.

Some neurologists hypothesize that the putamen also plays a role in the selection of movement (e.g. Tourette Syndrome
Tourette syndrome
Tourette syndrome is an inherited neuropsychiatric disorder with onset in childhood, characterized by multiple physical tics and at least one vocal tic; these tics characteristically wax and wane...

) and the "automatic" performance of previously learned movements (e.g. Parkinson's disease
Parkinson's disease
Parkinson's disease is a degenerative disorder of the central nervous system...

).

In one study it was found that the putamen controls limb movement. The goal of this study was to determine whether particular cell activity in the putamen of primates was related to the direction of limb movement or to the underlying pattern of muscular activity. Two monkeys were trained to perform tasks that involved the movement of loads. The tasks were created so that movement could be distinguished from muscle activity. Neurons in the putamen were selected for monitoring only if they were related both to the task and to arm movements outside the task. It was shown that 50% of the neurons that were monitored were related to the direction of movement, independent of the load.

Another study was done to investigate movement extent and speed using PET
Positron emission tomography
Positron emission tomography is nuclear medicine imaging technique that produces a three-dimensional image or picture of functional processes in the body. The system detects pairs of gamma rays emitted indirectly by a positron-emitting radionuclide , which is introduced into the body on a...

 mapping of regional cerebral blood flow in 13 humans. Movement tasks were performed with a joystick
Joystick
A joystick is an input device consisting of a stick that pivots on a base and reports its angle or direction to the device it is controlling. Joysticks, also known as 'control columns', are the principal control in the cockpit of many civilian and military aircraft, either as a center stick or...

-controlled cursor
Cursor (computers)
In computing, a cursor is an indicator used to show the position on a computer monitor or other display device that will respond to input from a text input or pointing device. The flashing text cursor may be referred to as a caret in some cases...

. Statistical tests were done to calculate the extent of movements and what regions of the brain the movements correlate to. It was found that "increasing movement extent was associated with parallel increases of rCBF in bilateral basal ganglia (BG; putamen and globus pallidus) and ipsilateral cerebellum." This not only shows that the putamen affects movement but it also shows that the putamen integrates with other structures in order to perform tasks.

One study was done in order to specifically investigate how the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 influences the learning of sequential movements. Two monkeys were trained to press a series of buttons in sequence. The methods used were designed to be able to monitor the well-learned tasks versus the new tasks. Muscimol
Muscimol
Muscimol is the major psychoactive alkaloid present in many mushrooms of the Amanita genus. Unlike psilocin , which is a serotonergic psychedelic and agonist for the 5-HT2A receptor set, muscimol is a potent, selective agonist for the GABAA receptor set and is a deliriant as a opposed...

 was injected into various parts of the basal ganglia, and it was found that "the learning of new sequences became deficient after injections in the anterior caudate and putamen, but not the middle-posterior putamen." This shows that different areas of the striatum
Striatum
The striatum, also known as the neostriatum or striate nucleus, is a subcortical part of the forebrain. It is the major input station of the basal ganglia system. The striatum, in turn, gets input from the cerebral cortex...

 are utilized when performing various aspects of the learning of sequential movements

Role in learning

In many studies, it has become apparent that the putamen plays a role in many types of learning. Some examples are listed below:

Reinforcement and implicit learning

Along with various types of movement, the putamen also affects reinforcement learning
Reinforcement learning
Inspired by behaviorist psychology, reinforcement learning is an area of machine learning in computer science, concerned with how an agent ought to take actions in an environment so as to maximize some notion of cumulative reward...

 and implicit learning
Implicit learning
Implicit learning is learning of complex information in an incidental manner, without awareness of what has been learned. It may require a certain minimal amount of attention and may depend on attentional and working memory mechanisms...

.

Reinforcement learning
Reinforcement learning
Inspired by behaviorist psychology, reinforcement learning is an area of machine learning in computer science, concerned with how an agent ought to take actions in an environment so as to maximize some notion of cumulative reward...

 is interacting with the environment and catering actions to maximize the outcome. Implicit learning
Implicit learning
Implicit learning is learning of complex information in an incidental manner, without awareness of what has been learned. It may require a certain minimal amount of attention and may depend on attentional and working memory mechanisms...

 is a passive process where people are exposed to information and acquire knowledge through exposure. Although the exact mechanisms are not known, it is clear that dopamine and tonically active neurons play a key role here. Tonically active neurons are cholinergic
Cholinergic
The word choline generally refers to the various quaternary ammonium salts containing the N,N,N-trimethylethanolammonium cation. Found in most animal tissues, choline is a primary component of the neurotransmitter acetylcholine and functions with inositol as a basic constituent of lecithin...

 interneurons that fire during the entire duration of the stimulus and fire at about 0.5-3 impulses per second. Tonic neurons are the opposite and only fire an action potential when movement occurs.

Category Learning

One particular study used patients with focal lesions on the basal ganglia (specifically the putamen) due to stroke
Stroke
A stroke, previously known medically as a cerebrovascular accident , is the rapidly developing loss of brain function due to disturbance in the blood supply to the brain. This can be due to ischemia caused by blockage , or a hemorrhage...

 in order to study category learning. The advantage to using these types of patients is that dopaminergic projections to the prefrontal cortex
Prefrontal cortex
The prefrontal cortex is the anterior part of the frontal lobes of the brain, lying in front of the motor and premotor areas.This brain region has been implicated in planning complex cognitive behaviors, personality expression, decision making and moderating correct social behavior...

 are more likely to be intact. Also, in these patients, it is easier to relate specific brain structures to function because the lesion only occurs in a specific place. The goal of this study was to determine whether or not these lesions affect rule-based and information-integration task learning. Rule-based tasks are learned via hypothesis-testing which is dependant on working memory. Information-integration tasks are ones where the accuracy is maximized when information from two sources are integrated at a pre-decisional stage, which follows a procedural-based system.

Seven participants with basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 lesion
Lesion
A lesion is any abnormality in the tissue of an organism , usually caused by disease or trauma. Lesion is derived from the Latin word laesio which means injury.- Types :...

s were used in the experiment, along with nine control participants. It is important to note that the caudate was not affected. The participants were tested for each type of learning during separate sessions, so the information processes would not interfere with each other. During each session, participants sat in front of a computer screen and various lines were displayed. These lines were created by using a randomization technique where random samples were taken from one of four categories. For ruled-based testing, these samples were used to construct lines of various length and orientation that fell into these four separate categories. After the stimulus was displayed, the subjects were asked to press 1 of 4 buttons to indicate which category the line fell into. The same process was repeated for information-integration tasks, and the same stimuli were used, except that the category boundaries were rotated 45°. This rotation causes the subject to integrate the quantitative information about the line before determining what category it is in.

It was found that subjects in the experimental group were impaired while performing rule-based tasks, but not information-integration ones. After statistical testing, it was also hypothesized that the brain began using information-integration techniques to solve the rule-based learning tasks. Since rule-based tasks use the hypothesis-testing system of the brain, it can be concluded that the hypothesis-testing system of the brain was damaged/weakened. It is known that the caudate and working memories are part of this system. Therefore, it was confirmed that the putamen is involved category learning, competition between the systems, feed-back processing in rule-based tasks, and is involved in the processing of pre-frontal regions (which relate to working memory and executive functioning). Now it is known that not only the basal ganglia
Basal ganglia
The basal ganglia are a group of nuclei of varied origin in the brains of vertebrates that act as a cohesive functional unit. They are situated at the base of the forebrain and are strongly connected with the cerebral cortex, thalamus and other brain areas...

 and caudate affect category learning.

Role in "hate circuit"

Recent, tentative studies have suggested that the putamen may play a role in the so-called "hate
Hatred
Hatred is a deep and emotional extreme dislike, directed against a certain object or class of objects. The objects of such hatred can vary widely, from inanimate objects to animals, oneself or other people, entire groups of people, people in general, existence, or the whole world...

 circuit" of the brain. A recent study was done in London
London
London is the capital city of :England and the :United Kingdom, the largest metropolitan area in the United Kingdom, and the largest urban zone in the European Union by most measures. Located on the River Thames, London has been a major settlement for two millennia, its history going back to its...

 by the department of cell and developmental biology at University College London
University College London
University College London is a public research university located in London, United Kingdom and the oldest and largest constituent college of the federal University of London...

. An fMRI was done on patients while they viewed a picture of people they hated and people who were "neutral". During the experiment, a "hate score" was recorded for each picture. The activity in sub-cortical areas of the brain implied that the "hate circuit" involves the putamen and the insula
Insular cortex
In each hemisphere of the mammalian brain the insular cortex is a portion of the cerebral cortex folded deep within the lateral sulcus between the temporal lobe and the frontal lobe. The cortical area overlying it towards the lateral surface of the brain is the operculum...

. It has been theorized that the "putamen plays a role in the perception of contempt
Contempt
Contempt is an intensely negative emotion regarding a person or group of people as inferior, base, or worthless—it is similar to scorn. It is also used when people are being sarcastic. Contempt is also defined as the state of being despised or dishonored; disgrace, and an open disrespect or willful...

 and disgust
Disgust
Disgust is a type of aversion that involves withdrawing from a person or object with strong expressions of revulsion whether real or pretended. It is one of the basic emotions and is typically associated with things that are regarded as unclean, inedible, infectious, gory or otherwise offensive...

, and may be part of the motor system
Motor system
The motor system is the part of the central nervous system that is involved with movement. It consists of the pyramidal and extrapyramidal system....

 that's mobilized to take action." It was also found that the amount of activity in the hate circuit correlates with the amount of hate a person declares, which could have legal implications concerning malicious crimes.

Parkinson's disease

After discovering the function of the putamen, it has become apparent to neurologists that the putamen and basal ganglia play an important role in Parkinson's disease
Parkinson's disease
Parkinson's disease is a degenerative disorder of the central nervous system...

 and other diseases that involve the degeneration of neurons.

Parkinson's diseases is the slow and steady loss of dopaminergic neurons in substantia nigra pars compacta. In Parkinson's disease the putamen plays a key role because its inputs and outputs are interconnected to the substantia nigra
Substantia nigra
The substantia nigra is a brain structure located in the mesencephalon that plays an important role in reward, addiction, and movement. Substantia nigra is Latin for "black substance", as parts of the substantia nigra appear darker than neighboring areas due to high levels of melanin in...

 and the globus pallidus
Globus pallidus
The globus pallidus also known as paleostriatum, is a sub-cortical structure of the brain. Topographically, it is part of the telencephalon, but retains close functional ties with the subthalamus - both of which are part of the extrapyramidal motor system...

. In Parkinson's diseases the activity in direct pathways to interior globus pallidus decreases and activity in indirect pathways to external globus pallidus increases. Together these actions cause excessive inhibition of the thalamus
Thalamus
The thalamus is a midline paired symmetrical structure within the brains of vertebrates, including humans. It is situated between the cerebral cortex and midbrain, both in terms of location and neurological connections...

. This is why Parkinson’s patients have tremors and have trouble performing voluntary movements. It has also been noted that Parkinson’s patients have a difficult time with motor planning. They must think about everything they do and cannot perform instinctive tasks without focusing on what they are doing.

Other diseases and disorders

The following diseases and disorders are linked with the putamen:
  • Cognitive decline in Alzheimer’s disease
  • Huntington's disease
    Huntington's disease
    Huntington's disease, chorea, or disorder , is a neurodegenerative genetic disorder that affects muscle coordination and leads to cognitive decline and dementia. It typically becomes noticeable in middle age. HD is the most common genetic cause of abnormal involuntary writhing movements called chorea...

  • Wilson's disease
    Wilson's disease
    Wilson's disease or hepatolenticular degeneration is an autosomal recessive genetic disorder in which copper accumulates in tissues; this manifests as neurological or psychiatric symptoms and liver disease...

  • Dementia
    Dementia
    Dementia is a serious loss of cognitive ability in a previously unimpaired person, beyond what might be expected from normal aging...

     with Lewy bodies
  • Corticobasal degeneration
  • Tourette syndrome
    Tourette syndrome
    Tourette syndrome is an inherited neuropsychiatric disorder with onset in childhood, characterized by multiple physical tics and at least one vocal tic; these tics characteristically wax and wane...

  • Schizophrenia
    Schizophrenia
    Schizophrenia is a mental disorder characterized by a disintegration of thought processes and of emotional responsiveness. It most commonly manifests itself as auditory hallucinations, paranoid or bizarre delusions, or disorganized speech and thinking, and it is accompanied by significant social...

  • Depression
  • Attention Deficit Hyperactivity Disorder
  • Chorea
  • Obsessive-Compulsive Disorder
    Obsessive-compulsive disorder
    Obsessive–compulsive disorder is an anxiety disorder characterized by intrusive thoughts that produce uneasiness, apprehension, fear, or worry, by repetitive behaviors aimed at reducing the associated anxiety, or by a combination of such obsessions and compulsions...

  • Other anxiety disorder
    Anxiety disorder
    Anxiety disorder is a blanket term covering several different forms of abnormal and pathological fear and anxiety. Conditions now considered anxiety disorders only came under the aegis of psychiatry at the end of the 19th century. Gelder, Mayou & Geddes explains that anxiety disorders are...

    s

In other animals

The putamen in humans is similar in structure and function to other animals. Therefore, many studies on the putamen have been done on animals (monkeys, rats
RATS
RATS may refer to:* RATS , Regression Analysis of Time Series, a statistical package* Rough Auditing Tool for Security, a computer program...

, etc.), as well as humans.

External links

The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
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