The putamen (; from

Latin Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...

, meaning "nutshell") is a subcortical

nucleus Nucleus (: nuclei) is a Latin word for the seed inside a fruit. It most often refers to: *Atomic nucleus, the very dense central region of an atom *Cell nucleus, a central organelle of a eukaryotic cell, containing most of the cell's DNA Nucleu ...

with a rounded structure, in the

basal ganglia The basal ganglia (BG) or basal nuclei are a group of subcortical Nucleus (neuroanatomy), nuclei found in the brains of vertebrates. In humans and other primates, differences exist, primarily in the division of the globus pallidus into externa ...

nuclear group. It is located at the base of the

forebrain In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the rostral (forward-most) portion of the brain. The forebrain controls body temperature, reproductive functions, eating, sleeping, and the display of emotions. Ve ...

and above the

midbrain The midbrain or mesencephalon is the uppermost portion of the brainstem connecting the diencephalon and cerebrum with the pons. It consists of the cerebral peduncles, tegmentum, and tectum. It is functionally associated with vision, hearing, mo ...

. The putamen and

caudate nucleus The caudate nucleus is one of the structures that make up the corpus striatum, which is part of the basal ganglia in the human brain. Although the caudate nucleus has long been associated with motor processes because of its relation to Parkinso ...

together form the

dorsal striatum The striatum (: striata) or corpus striatum is a cluster of interconnected nuclei that make up the largest structure of the subcortical basal ganglia. The striatum is a critical component of the motor and reward systems; receives glutamater ...

. Through various pathways, the putamen is connected to the

substantia nigra The substantia nigra (SN) is a basal ganglia structure located in the midbrain that plays an important role in reward and movement. ''Substantia nigra'' is Latin for "black substance", reflecting the fact that parts of the substantia nigra a ...

, the

globus pallidus The globus pallidus (GP), also known as paleostriatum or dorsal pallidum, is a major component of the Cerebral cortex, subcortical basal ganglia in the brain. It consists of two adjacent segments, one external (or lateral), known in rodents simpl ...

, the

claustrum The claustrum (Latin, meaning "to close" or "to shut") is a thin sheet of neurons and supporting glial cells in the brain, that connects to the cerebral cortex and subcortical regions including the amygdala, hippocampus and thalamus. It is locate ...

, and the

thalamus The thalamus (: thalami; from Greek language, Greek Wikt:θάλαμος, θάλαμος, "chamber") is a large mass of gray matter on the lateral wall of the third ventricle forming the wikt:dorsal, dorsal part of the diencephalon (a division of ...

, in addition to many regions of the

cerebral cortex The cerebral cortex, also known as the cerebral mantle, is the outer layer of neural tissue of the cerebrum of the brain in humans and other mammals. It is the largest site of Neuron, neural integration in the central nervous system, and plays ...

. A primary function of the putamen is to regulate movements at various stages such as in preparation and execution; and to influence various types of

learning Learning is the process of acquiring new understanding, knowledge, behaviors, skills, value (personal and cultural), values, Attitude (psychology), attitudes, and preferences. The ability to learn is possessed by humans, non-human animals, and ...

. It employs

GABA GABA (gamma-aminobutyric acid, γ-aminobutyric acid) is the chief inhibitory neurotransmitter in the developmentally mature mammalian central nervous system. Its principal role is reducing neuronal excitability throughout the nervous system. GA ...

acetylcholine Acetylcholine (ACh) is an organic compound that functions in the brain and body of many types of animals (including humans) as a neurotransmitter. Its name is derived from its chemical structure: it is an ester of acetic acid and choline. Par ...

, and

enkephalin An enkephalin is a pentapeptide involved in regulating nociception (pain sensation) in the body. The enkephalins are termed endogenous ligands, as they are internally derived (and therefore endogenous) and bind as ligands to the body's opioid ...

to perform its functions. The putamen also plays a role in

neurodegenerative disease A neurodegenerative disease is caused by the progressive loss of neurons, in the process known as neurodegeneration. Neuronal damage may also ultimately result in their death. Neurodegenerative diseases include amyotrophic lateral sclerosis, mul ...

s, such as

Parkinson's disease Parkinson's disease (PD), or simply Parkinson's, is a neurodegenerative disease primarily of the central nervous system, affecting both motor system, motor and non-motor systems. Symptoms typically develop gradually and non-motor issues become ...

History

The word "putamen" is from

, referring to that which "falls off in pruning", from "putare", meaning "to prune, to think, or to consider". Most

MRI Magnetic resonance imaging (MRI) is a medical imaging technique used in radiology to generate pictures of the anatomy and the physiological processes inside the body. MRI scanners use strong magnetic fields, magnetic field gradients, and rad ...

research was focused broadly on the basal ganglia as a whole, for reasons such as poor resolution or hemorrhage within the putamen. Later studies have focused on the basal ganglia and relevant brain-behavior relationships. In the 1970s, the first single-unit recordings were carried out with

monkey Monkey is a common name that may refer to most mammals of the infraorder Simiiformes, also known as simians. Traditionally, all animals in the group now known as simians are counted as monkeys except the apes. Thus monkeys, in that sense, co ...

s, monitoring pallidal

neuronal A neuron (American English), neurone (British English), or nerve cell, is an excitable cell that fires electric signals called action potentials across a neural network in the nervous system. They are located in the nervous system and help to ...

activity related to movement. Since then, more extensive neuronal tracing, stimulation, and imaging research methods including functional MRI, and diffusion weighted MRI that allow for investigation of the putamen have been developed.

Anatomy

Globus pallidus and putamen - very low mag

The putamen is a structure in the

. Along with the

it forms the

. 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 The internal capsule is a paired white matter structure, as a two-way nerve tract, tract, carrying afferent nerve fiber, ascending and efferent nerve fiber, descending axon, fibers, to and from the cerebral cortex. The internal capsule is situate ...

, passing through the middle. The putamen, together with the

, makes up the

lentiform nucleus The lentiform nucleus (or lentiform complex, lenticular nucleus, or lenticular complex) are the putamen (laterally) and the globus pallidus (medially), collectively. Due to their proximity, these two structures were formerly considered one, howev ...

. The putamen is the outermost portion of the

. These are a group of nuclei in the brain that are interconnected with the

, and

brainstem The brainstem (or brain stem) is the posterior stalk-like part of the brain that connects the cerebrum with the spinal cord. In the human brain the brainstem is composed of the midbrain, the pons, and the medulla oblongata. The midbrain is conti ...

. The basal ganglia include the

striatum The striatum (: striata) or corpus striatum is a cluster of interconnected nuclei that make up the largest structure of the subcortical basal ganglia. The striatum is a critical component of the motor and reward systems; receives glutamat ...

nucleus accumbens The nucleus accumbens (NAc or NAcc; also known as the accumbens nucleus, or formerly as the ''nucleus accumbens septi'', Latin for ' nucleus adjacent to the septum') is a region in the basal forebrain rostral to the preoptic area of the hypo ...

, and the subthalamic nucleus. In

mammals A mammal () is a vertebrate animal of the class Mammalia (). Mammals are characterised by the presence of milk-producing mammary glands for feeding their young, a broad neocortex region of the brain, fur or hair, and three middle e ...

, the basal ganglia are associated with

motor control Motor control is the regulation of movements in organisms that possess a nervous system. Motor control includes conscious voluntary movements, subconscious muscle memory and involuntary reflexes, as well as instinctual taxes. To control ...

cognition Cognition is the "mental action or process of acquiring knowledge and understanding through thought, experience, and the senses". It encompasses all aspects of intellectual functions and processes such as: perception, attention, thought, ...

emotion Emotions are physical and mental states brought on by neurophysiology, neurophysiological changes, variously associated with thoughts, feelings, behavior, behavioral responses, and a degree of pleasure or suffering, displeasure. There is ...

, and domain-general functions important for executive functioning as well as support for domain-specific languages. The basal ganglia are located bilaterally, and have rostral, and caudal divisions. The putamen is located in the rostral division as part of the

. The basal ganglia receive input from the cerebral cortex, via the striatum. Striatum Structural MRI

Caudate nucleus

The caudate works with the putamen to receive the input from

. Collectively, they can be considered the "entrance" to the basal ganglia. Projections from the putamen reach the caudate directly via the caudolenticular grey bridges. The putamen and caudate are jointly connected with the

, however the caudate outputs more densely to the

pars reticulata while the putamen sends more afferents to the internal

Substantia nigra

The substantia nigra 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 to control head and

eye movement Eye movement includes the voluntary or involuntary movement of the eyes. Eye movements are used by a number of organisms (e.g. primates, rodents, flies, birds, fish, cats, crabs, octopus) to fixate, inspect and track visual objects of inte ...

s. The SNpc produces dopamine, which is crucial for movements. The SNpc degenerates during

Globus pallidus

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

Physiology

Types of pathways

The putamen (and striatum in general) has numerous, parallel circuits that allow for cortico-subcortico-cortico communication loops. These have been described, broadly, as the direct, indirect, and hyper direct pathways. GABAergic projections of the putamen have an inhibitory effect on the thalamus. Thalamic projections from the centromedian and parafascicular nuclei have an excitatory effect on the putamen. Unlike the thalamus, which has broad reciprocal connectivity, cortical projections with the putamen are afferent, thus sending information as opposed to receiving it. Cortical communication is accomplished via multi-fiber pathways as outlined previously (i.e. via other subcortical structures).

Dopamine

The putamen is part of the nigrostriatal pathway. The Substantia nigra pars compacta, a structure in the brainstem that contains dopaminergic neurons, releases dopamine to the dorsal striatum, which consists of the putamen and caudate nucleus. This dopamine release is highly important for motor control, involving mechanisms for movement promotion (via the Direct Pathway), as well as mechanisms for movement inhibition (via the Indirect Pathway). Striatal neurons expressing D1 type dopaminergic receptors are involved in the Direct Pathway, whereas D2 type dopaminergic receptors are expressed in striatal neurons of the Indirect Pathway. In order to control the amount of dopamine in the

synaptic cleft Chemical synapses are biological junctions through which neurons' signals can be sent to each other and to non-neuronal cells such as those in neuromuscular junction, muscles or glands. Chemical synapses allow neurons to form biological neural ...

, and the amount of dopamine binding to post synaptic terminals, presynaptic dopaminergic neurons function to

reuptake Reuptake is the reabsorption of a neurotransmitter by a neurotransmitter transporter located along the plasma membrane of an axon terminal (i.e., the pre-synaptic neuron at a synapse) or glial cell after it has performed its function of transmi ...

the excess dopamine. The dopaminergic release to the putamen and caudate might be implicated in addictive behaviour as this release seems to play an important role in habit learning.

Other neurotransmitters

The putamen also plays a role in

modulation Signal modulation is the process of varying one or more properties of a periodic waveform in electronics and telecommunication for the purpose of transmitting information. The process encodes information in form of the modulation or message ...

of other neurotransmitters. It releases GABA, enkephalin,

substance P Substance P (SP) is an undecapeptide (a peptide composed of a chain of 11 amino acid residues) and a type of neuropeptide, belonging to the tachykinin family of neuropeptides. It acts as a neurotransmitter and a neuromodulator. Substance P ...

, and

. It receives

serotonin Serotonin (), also known as 5-hydroxytryptamine (5-HT), is a monoamine neurotransmitter with a wide range of functions in both the central nervous system (CNS) and also peripheral tissues. It is involved in mood, cognition, reward, learning, ...

and

glutamate Glutamic acid (symbol Glu or E; known as glutamate in its anionic form) is an α-amino acid that is used by almost all living beings in the biosynthesis of proteins. It is a Essential amino acid, non-essential nutrient for humans, meaning that ...

Function: motor skills

The putamen is interconnected with many other structures, and works in conjunction with them to influence many types of motor behaviors. These include motor planning, learning, and execution, 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 (TS), or simply Tourette's, is a common neurodevelopmental disorder that begins in childhood or adolescence. It is characterized by multiple movement (motor) tics and at least one vocal (phonic) tic. Common tics are blinkin ...

) and the "automatic" performance of previously learned movements (e.g.

). 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 A pet, or companion animal, is an animal kept primarily for a person's company or entertainment rather than as a working animal, livestock, or a laboratory animal. Popular pets are often considered to have attractive/ cute appearances, inte ...

mapping of regional cerebral blood flow in 13 humans. Movement tasks were performed with a

joystick A joystick, sometimes called a flight stick, 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. Also known as the control column, it is the principal control devic ...

-controlled cursor. 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 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 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

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 and

implicit learning Implicit learning is the learning of complex information in an unintentional manner, without awareness of what has been learned. According to Frensch and Rünger (2003) the general definition of implicit learning is still subject to some controvers ...

Reinforcement learning Reinforcement learning (RL) is an interdisciplinary area of machine learning and optimal control concerned with how an intelligent agent should take actions in a dynamic environment in order to maximize a reward signal. Reinforcement learnin ...

is interacting with the environment and catering actions to maximize the outcome. Implicit learning 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 agents are compounds which mimic the action of acetylcholine and/or butyrylcholine. In general, the word " choline" describes the various quaternary ammonium salts containing the ''N'',''N'',''N''-trimethylethanolammonium cation ...

interneurons Interneurons (also called internuncial neurons, association neurons, connector neurons, or intermediate neurons) are neurons that are not specifically motor neurons or sensory neurons. Interneurons are the central nodes of neural circuits, ena ...

that fire during the entire duration of the stimulus and fire at about 0.5–3 impulses per second. Phasic 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 is a medical condition in which poor cerebral circulation, blood flow to a part of the brain causes cell death. There are two main types of stroke: brain ischemia, ischemic, due to lack of blood flow, and intracranial hemorrhage, hemor ...

in order to study

category learning Concept learning, also known as category learning, concept attainment, and concept formation, is defined by Bruner, Goodnow, & Austin (1956) as "the search for and testing of attributes that can be used to distinguish exemplars from non exemplar ...

. The advantage to using these types of patients is that dopaminergic projections to the

prefrontal cortex In mammalian brain anatomy, the prefrontal cortex (PFC) covers the front part of the frontal lobe of the cerebral cortex. It is the association cortex in the frontal lobe. The PFC contains the Brodmann areas BA8, BA9, BA10, BA11, BA12, ...

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 dependent on working memory. Information-integration tasks are ones wherein 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

lesion A lesion is any damage or abnormal change in the tissue of an organism, usually caused by injury or diseases. The term ''Lesion'' is derived from the Latin meaning "injury". Lesions may occur in both plants and animals. Types There is no de ...

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 in 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 and caudate affect category learning.

Role in "hate circuit"

Tentative studies have suggested that the putamen may play a role in the so-called "

hate Hatred or hate is an intense negative emotional response towards certain people, things or ideas, usually related to opposition or revulsion toward something. Hatred is often associated with intense feelings of anger, contempt, and disgust. Ha ...

circuit" of the brain. A recent study was done by the department of cell and developmental biology at

University College London University College London (Trade name, branded as UCL) is a Public university, public research university in London, England. It is a Member institutions of the University of London, member institution of the Federal university, federal Uni ...

. An

fMRI Functional magnetic resonance imaging or functional MRI (fMRI) measures brain activity by detecting changes associated with blood flow. This technique relies on the fact that cerebral blood flow and neuronal activation are coupled. When an area o ...

was carried out on subjects 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 subcortical areas of the brain implied that the "hate circuit" involves the

superior frontal gyrus In neuroanatomy, the superior frontal gyrus (SFG, also marginal gyrus) is a gyrus – a ridge on the brain's cerebral cortex – which makes up about one third of the frontal lobe. It is bounded laterally by the superior frontal sulcus. The su ...

, the putamen and the insula. It has been theorized that the "putamen plays a role in the perception of

contempt In colloquial usage, contempt usually refers to either the act of despising, or having a general lack of respect for something. This set of emotions generally produces maladaptive behaviour. Other authors define contempt as a negative emotio ...

and

disgust Disgust (, from Latin , ) is an emotional response of rejection or revulsion to something potentially contagious or something considered offensive, distasteful or unpleasant. In ''The Expression of the Emotions in Man and Animals'', Charles D ...

, and may be part of the

motor system The motor system is the set of central nervous system, central and peripheral nervous system, peripheral structures in the nervous system that support motor functions, i.e. movement. Peripheral structures may include skeletal muscles and Efferen ...

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.

Role in transgender individuals

The putamen of transgender women was found to have significantly larger amounts of grey matter compared to the putamen of cisgender men. This possibly suggests that a difference in the putamen may exist between

trans women A trans woman or transgender woman is a woman who was assigned male at birth. Trans women have a female gender identity and may experience gender dysphoria (distress brought upon by the discrepancy between a person's gender identity and their ...

and cisgender men.

Pathology

Parkinson's disease

After discovering the function of the putamen, it has become apparent to neurologists that the putamen and other parts of the basal ganglia play an important role in

and other diseases that involve the degeneration of

neurons A neuron (American English), neurone (British English), or nerve cell, is an membrane potential#Cell excitability, excitable cell (biology), cell that fires electric signals called action potentials across a neural network (biology), neural net ...

. Parkinson's disease 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 and the globus pallidus. In Parkinson's disease the activity in direct pathways to interior globus pallidus decreases and activity in indirect pathways to external globus pallidus increases. It has also been noted that Parkinson's patients have a difficult time with motor planning.

Other diseases and disorders

The following diseases and disorders are linked with the putamen: * Cognitive decline in

Alzheimer's disease Alzheimer's disease (AD) is a neurodegenerative disease and the cause of 60–70% of cases of dementia. The most common early symptom is difficulty in remembering recent events. As the disease advances, symptoms can include problems wit ...

Huntington's disease Huntington's disease (HD), also known as Huntington's chorea, is an incurable neurodegenerative disease that is mostly Genetic disorder#Autosomal dominant, inherited. It typically presents as a triad of progressive psychiatric, cognitive, and ...

Wilson's disease Wilson's disease (also called hepatolenticular degeneration) is a genetic disorder characterized by the excess build-up of copper in the body. Symptoms are typically related to the brain and liver. Liver-related symptoms include vomiting, wea ...

Dementia Dementia is a syndrome associated with many neurodegenerative diseases, characterized by a general decline in cognitive abilities that affects a person's ability to perform activities of daily living, everyday activities. This typically invo ...

with Lewy bodies * Corticobasal degeneration *

Schizophrenia Schizophrenia () is a mental disorder characterized variously by hallucinations (typically, Auditory hallucination#Schizophrenia, hearing voices), delusions, thought disorder, disorganized thinking and behavior, and Reduced affect display, f ...

* Depression *

Attention deficit hyperactivity disorder Attention deficit hyperactivity disorder (ADHD) is a neurodevelopmental disorder characterised by symptoms of inattention, hyperactivity, impulsivity, and emotional dysregulation that are excessive and pervasive, impairing in multiple con ...

Chorea Chorea, or (rarely) choreia, () is an abnormal involuntary movement disorder, characterized by quick movements of the hands or feet. It is one of a group of neurological disorders called dyskinesias. The term ''chorea'' is derived , as the move ...

* Obsessive-Compulsive Disorder *

Kernicterus Kernicterus is a bilirubin-induced brain dysfunction. The term was coined in 1904 by Christian Georg Schmorl. Bilirubin is a naturally occurring substance in the body of humans and many other animals, but it is neurotoxic when its concentration ...

* Other

anxiety disorder Anxiety disorders are a group of mental disorders characterized by significant and uncontrollable feelings of anxiety and fear such that a person's social, occupational, and personal functions are significantly impaired. Anxiety may cause phys ...

In other animals

The putamen in humans is relatively similar in structure and function to other animals. Therefore, many studies on the putamen have been done on animals (

rat Rats are various medium-sized, long-tailed rodents. Species of rats are found throughout the order Rodentia, but stereotypical rats are found in the genus ''Rattus''. Other rat genera include '' Neotoma'' (pack rats), '' Bandicota'' (bandicoo ...

cat The cat (''Felis catus''), also referred to as the domestic cat or house cat, is a small domesticated carnivorous mammal. It is the only domesticated species of the family Felidae. Advances in archaeology and genetics have shown that the ...

s, etc.), as well as humans. However, inter-species variations are indeed observed in mammals, and have been documented for white matter putaminal connectivity. Variation is primarily related to structural connectivity patterns, while somatotopic organization principles are retained. Primate research since the 1980s through to the present has established that cortical regions related to higher-order cognition primarily send afferent neurons to the rostral-most portion of the putamen, while the remainder of this structure in primates primarily serves sensorimotor functions and is densely interconnected with primary and supplementary motor regions.

Additional images

Image:Gray744.png, Coronal section of brain through

anterior commissure The anterior commissure (also known as the precommissure) is a white matter nerve tract, tract (a bundle of axons) connecting the two temporal lobes of the cerebral hemispheres across the midline, and placed in front of the columns of the Fornix o ...

. Image:Telencephalon-Horiconatal.jpg, Horizontal section of right cerebral hemisphere. Image:Constudoverbrain_-_2.png, Brain Image:Human brain frontal (coronal) section description 2.JPG, Human brain frontal (coronal) section Image:Putamen.jpg, Horizontal slice of MRI-image showing the putamen. The other nuclei of the basal ganglia (caudate nucleus and globus pallidus) can be seen as well. File:Slide8gg.JPG, Putamen File:Slide8kk.JPG, Putamen File:Putamen_-_DK_ATLAS.png, Putamen along with other subcortical structures File:Putamen coronal sections.gif, Putamen highlighted in green on coronal T1 MRI images File:Putamen sagittal sections.gif, Putamen highlighted in green on sagittal T1 MRI images File:Putamen transversal sections.gif, Putamen highlighted in green on transversal T1 MRI images

References

External links

* *
Diagram at uni-tuebingen.de
* – "The Visual Pathway from Below" {{Authority control Basal ganglia

History

Anatomy

Caudate nucleus

Substantia nigra

Globus pallidus

Physiology

Types of pathways

Dopamine

Other neurotransmitters

Function: motor skills

Role in learning

Reinforcement and implicit learning

Category learning

Role in "hate circuit"

Role in transgender individuals

Pathology

Parkinson's disease

Other diseases and disorders

In other animals

Additional images

See also

References

External links