The striatum, or corpus striatum (also called the striate nucleus), is a

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 Nucle ...

(a cluster of

neuron A neuron, neurone, or nerve cell is an electrically excitable cell that communicates with other cells via specialized connections called synapses. The neuron is the main component of nervous tissue in all animals except sponges and placozoa. N ...

s) in the

subcortical 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. The cerebral cortex mostly consists of the six-layered neocortex, with just 10% consisting of ...

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 extern ...

of the

forebrain In the anatomy of the brain of vertebrates, the forebrain or prosencephalon is the Anatomical terms of location#Directional terms, rostral (forward-most) portion of the brain. The forebrain (prosencephalon), the midbrain (mesencephalon), and hin ...

. The striatum is a critical component of the

motor An engine or motor is a machine designed to convert one or more forms of energy into mechanical energy. Available energy sources include potential energy (e.g. energy of the Earth's gravitational field as exploited in hydroelectric power gen ...

and

reward Reward may refer to: Places * Reward (Shelltown, Maryland), a historic home in Shelltown Maryland * Reward, California (disambiguation) * Reward-Tilden's Farm, a historic home in Chestertown Maryland Arts, entertainment, and media * "Rewa ...

systems; receives

glutamatergic Glutamatergic means "related to glutamate". A glutamatergic agent (or drug) is a chemical that directly modulates the excitatory amino acid (glutamate/ aspartate) system in the body or brain. Examples include excitatory amino acid receptor agonist ...

and

dopaminergic Dopaminergic means "related to dopamine" (literally, "working on dopamine"), dopamine being a common neurotransmitter. Dopaminergic substances or actions increase dopamine-related activity in the brain. Dopaminergic brain pathways facilitate d ...

inputs from different sources; and serves as the primary input to the rest of the basal ganglia. Functionally, the striatum coordinates multiple aspects of

cognition Cognition refers to "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, ...

, including both motor and action

planning Planning is the process of thinking regarding the activities required to achieve a desired goal. Planning is based on foresight, the fundamental capacity for mental time travel. The evolution of forethought, the capacity to think ahead, is consi ...

decision-making In psychology, decision-making (also spelled decision making and decisionmaking) is regarded as the Cognition, cognitive process resulting in the selection of a belief or a course of action among several possible alternative options. It could be ...

motivation Motivation is the reason for which humans and other animals initiate, continue, or terminate a behavior at a given time. Motivational states are commonly understood as forces acting within the agent that create a disposition to engage in goal-dire ...

reinforcement In behavioral psychology, reinforcement is a consequence applied that will strengthen an organism's future behavior whenever that behavior is preceded by a specific antecedent stimulus. This strengthening effect may be measured as a higher freq ...

, and

perception. The striatum is made up of the

caudate nucleus The caudate nucleus is one of the structures that make up the corpus striatum, which is a component of the basal ganglia in the human brain. While the caudate nucleus has long been associated with motor processes due to its role in Parkinson's di ...

and the

lentiform nucleus The lentiform nucleus, or lenticular nucleus, comprises the putamen and the globus pallidus within the basal ganglia. With the caudate nucleus, it forms the dorsal striatum. It is a large, lens-shaped mass of gray matter just lateral to the inter ...

. The lentiform nucleus is made up of the larger

putamen The putamen (; from Latin, meaning "nutshell") is a round structure located at the base of the forebrain (telencephalon). The putamen and caudate nucleus together form the dorsal striatum. It is also one of the structures that compose the basal n ...

, and the smaller

globus pallidus The globus pallidus (GP), also known as paleostriatum or dorsal pallidum, is a subcortical structure of the brain. It consists of two adjacent segments, one external, known in rodents simply as the globus pallidus, and one internal, known in rod ...

. Strictly speaking the globus pallidus is part of the striatum. It is common practice, however, to implicitly exclude the globus pallidus when referring to striatal structures. In

primate Primates are a diverse order of mammals. They are divided into the strepsirrhines, which include the lemurs, galagos, and lorisids, and the haplorhines, which include the tarsiers and the simians (monkeys and apes, the latter including huma ...

s, the striatum is divided into a ventral striatum, and a dorsal striatum, subdivisions that are based upon function and connections. The

ventral Standard anatomical terms of location are used to unambiguously describe the anatomy of animals, including humans. The terms, typically derived from Latin or Greek language, Greek roots, describe something in its standard anatomical position. Th ...

striatum consists of the

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 hypotha ...

and the

olfactory tubercle The olfactory tubercle (OT), also known as the tuberculum olfactorium, is a multi-sensory processing center that is contained within the olfactory cortex and ventral striatum and plays a role in reward cognition. The OT has also been shown to ...

. The

dorsal Dorsal (from Latin ''dorsum'' ‘back’) may refer to: * Dorsal (anatomy), an anatomical term of location referring to the back or upper side of an organism or parts of an organism * Dorsal, positioned on top of an aircraft's fuselage * Dorsal co ...

striatum consists of the

and the

. A

white matter White matter refers to areas of the central nervous system (CNS) that are mainly made up of myelinated axons, also called tracts. Long thought to be passive tissue, white matter affects learning and brain functions, modulating the distribution ...

nerve tract A nerve tract is a bundle of nerve fibers (axons) connecting nuclei of the central nervous system. In the peripheral nervous system this is known as a nerve, and has associated connective tissue. The main nerve tracts in the central nervous syste ...

(the

internal capsule The internal capsule is a white matter structure situated in the inferomedial part of each cerebral hemisphere of the brain. It carries information past the basal ganglia, separating the caudate nucleus and the thalamus from the putamen and the g ...

) in the dorsal striatum separates the

and the

. Anatomically, the term ''striatum'' describes its striped (striated) appearance of grey-and-white matter.

Structure

The striatum is the largest structure of the

. The striatum is divided into a ventral and a dorsal subdivision, based upon function and connections. The ventral striatum is composed of the

and the

. The nucleus accumbens is made up of the

nucleus accumbens core 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 hypotha ...

and the

nucleus accumbens shell 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 hypotha ...

, which differ by neural populations. The olfactory tubercle receives input from the

olfactory bulb The olfactory bulb (Latin: ''bulbus olfactorius'') is a grey matter, neural structure of the vertebrate forebrain involved in olfaction, the sense of odor, smell. It sends olfactory information to be further processed in the amygdala, the orbitof ...

but has not been shown to play a role in processing smell. In non-primate species, the

islands of Calleja The islands of Calleja (; IC, ISC, or IClj) are a group of neural granule cells located within the ventral striatum in the brains of most animals. This region of the brain is part of the limbic system, where it aids in the reinforcing effects of ...

are included. The ventral striatum is associated with the limbic system and has been implicated as a vital part of the

circuitry An electronic circuit is composed of individual electronic components, such as resistors, transistors, capacitors, inductors and diodes, connected by conductive wires or traces through which electric current can flow. It is a type of electrical ...

for decision making and reward-related behavior. The dorsal striatum is composed of the

and the

Staining Staining is a technique used to enhance contrast in samples, generally at the microscopic level. Stains and dyes are frequently used in histology (microscopic study of biological tissues), in cytology (microscopic study of cells), and in the ...

can differentiate the striatum into two distinct compartments of

striosome The striosomes (also referred to as ''patches'') are one of two complementary chemical compartments within the striatum (the other compartment is known as the matrix) that can be visualized by staining for immunocytochemical markers such as acetyl ...

s or ''patches'', and a surrounding

matrix Matrix most commonly refers to: * ''The Matrix'' (franchise), an American media franchise ** ''The Matrix'', a 1999 science-fiction action film ** "The Matrix", a fictional setting, a virtual reality environment, within ''The Matrix'' (franchis ...

; this is particularly evident on the components of

acetylcholinesterase Acetylcholinesterase (HGNC symbol ACHE; EC 3.1.1.7; systematic name acetylcholine acetylhydrolase), also known as AChE, AChase or acetylhydrolase, is the primary cholinesterase in the body. It is an enzyme Enzymes () are proteins that a ...

and

calbindin Calbindins are three different calcium-binding proteins: calbindin, calretinin and S100G. They were originally described as vitamin D-dependent calcium-binding proteins in the intestine and kidney in the chick and mammals. They are now classified ...

. More studies have been carried out on the dorsal striatum but the compartments have also been identified in the ventral striatum. In the dorsal striatum striosomes make up 10-15 per cent of the striatal volume.

Cell types

Types of cells in the striatum include: *

Medium spiny neurons Medium spiny neurons (MSNs), also known as spiny projection neurons (SPNs), are a special type of GABAergic inhibitory cell representing 95% of neurons within the human striatum, a basal ganglia structure. Medium spiny neurons have two primary ...

(MSNs), which are the principal neurons of the striatum. They are

GABAergic In molecular biology and physiology, something is GABAergic or GABAnergic if it pertains to or affects the neurotransmitter GABA. For example, a synapse is GABAergic if it uses GABA as its neurotransmitter, and a GABAergic neuron produces GABA. A ...

and, thus, are classified as inhibitory neurons. Medium spiny projection neurons comprise 95% of the total neuronal population of the human striatum. Medium spiny neurons have two characteristic types:

D1-type The D1-like receptors are a subfamily of dopamine receptors that bind the endogenous neurotransmitter dopamine. The D1-like subfamily consists of two G protein–coupled receptors that are coupled to Gs and mediate excitatory neurotransmission ...

MSNs and

D2-type The D2-like receptors are a subfamily of dopamine receptors that bind the endogenous neurotransmitter dopamine. The D2-like subfamily consists of three G-protein coupled receptors that are coupled to Gi/Go and mediate inhibitory neurotransmission ...

MSNs. A subpopulation of MSNs contain both D1-type and D2-type receptors, with approximately 40% of striatal MSNs expressing both

DRD1 Dopamine receptor D1, also known as DRD1. It is one of the two types of D1-like receptor family - receptors D1 and D5. It is a protein that in humans is encoded by the DRD1 gene. Tissue distribution D1 receptors are the most abundant kind of do ...

and

DRD2 Dopamine receptor D2, also known as D2R, is a protein that, in humans, is encoded by the ''DRD2'' gene. After work from Paul Greengard's lab had suggested that dopamine receptors were the site of action of antipsychotic drugs, several groups, in ...

mRNA In molecular biology, messenger ribonucleic acid (mRNA) is a single-stranded molecule of RNA that corresponds to the genetic sequence of a gene, and is read by a ribosome in the process of Protein biosynthesis, synthesizing a protein. mRNA is ...

. *

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. F ...

interneurons Interneurons (also called internuncial neurons, relay neurons, association neurons, connector neurons, intermediate neurons or local circuit neurons) are neurons that connect two brain regions, i.e. not direct motor neurons or sensory neurons. In ...

release acetylcholine, which has a variety of important effects in the striatum. In humans, other primates, and rodents, these interneurons respond to salient environmental stimuli with stereotyped responses that are temporally aligned with the responses of dopaminergic neurons of 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 app ...

. The large aspiny cholinergic interneurons themselves are affected by dopamine through D5 dopamine receptors. Dopamine also directly controls communication between cholinergic interneurons. * There are many types of GABAergic interneurons. The best known are

parvalbumin Parvalbumin (PV) is a calcium-binding protein with low molecular weight (typically 9-11 kDa). In humans, it is encoded by the ''PVALB'' gene. It is not a member of the albumin family; it is named for its size (''parv-'', from Latin ''parvus'' smal ...

expressing interneurons, also known as fast-spiking interneurons, which participate in powerful feedforward inhibition of principal neurons. Also, there are GABAergic interneurons that express

tyrosine hydroxylase Tyrosine hydroxylase or tyrosine 3-monooxygenase is the enzyme responsible for catalyzing the conversion of the amino acid L-tyrosine to L-3,4-dihydroxyphenylalanine (L-DOPA). It does so using molecular oxygen (O2), as well as iron (Fe2+) and t ...

somatostatin Somatostatin, also known as growth hormone-inhibiting hormone (GHIH) or by several other names, is a peptide hormone that regulates the endocrine system and affects neurotransmission and cell proliferation via interaction with G protein-couple ...

nitric oxide synthase Nitric oxide synthases () (NOSs) are a family of enzymes catalyzing the production of nitric oxide (NO) from L-arginine. NO is an important cellular signaling molecule. It helps modulate vascular tone, insulin secretion, airway tone, and perista ...

and neuropeptide-y. Recently, two types of neuropeptide-y expressing GABAergic interneurons have been described in detail, one of which translates synchronous activity of cholinergic interneurons into inhibition of principal neurons. These

s of the striatum are not distributed evenly. There are two regions of

neurogenesis Neurogenesis is the process by which nervous system cells, the neurons, are produced by neural stem cells (NSCs). It occurs in all species of animals except the porifera (sponges) and placozoans. Types of NSCs include neuroepithelial cells (NECs) ...

in the brain – the

subventricular zone The subventricular zone (SVZ) is a region situated on the outside wall of each lateral ventricle of the vertebrate brain. It is present in both the embryonic and adult brain. In embryonic life, the SVZ refers to a secondary proliferative zone ...

in the

lateral ventricle The lateral ventricles are the two largest ventricular system, ventricles of the brain and contain cerebrospinal fluid (CSF). Each cerebral hemisphere contains a lateral ventricle, known as the left or right ventricle, respectively. Each lateral ...

s, and the

dentate gyrus The dentate gyrus (DG) is part of the hippocampal formation in the temporal lobe of the brain, which also includes the hippocampus and the subiculum. The dentate gyrus is part of the hippocampal trisynaptic circuit and is thought to contribute ...

in the

hippocampal formation The hippocampal formation is a compound structure in the Temporal lobe#Medial temporal lobe, medial temporal lobe of the brain. It forms a c-shaped bulge on the floor of the temporal horn of the Lateral ventricles, lateral ventricle. There is no ...

Neuroblast In vertebrates, a neuroblast or primitive nerve cell is a postmitotic cell that does not divide further, and which will develop into a neuron after a migration phase. In invertebrates such as ''Drosophila,'' neuroblasts are neural progenitor cells ...

s that form in the lateral ventricle adjacent to the striatum, integrate in the striatum. This has been noted in the human striatum following an

ischemic stroke A stroke is a medical condition in which poor blood flow to the brain causes cell death. There are two main types of stroke: ischemic, due to lack of blood flow, and hemorrhagic, due to bleeding. Both cause parts of the brain to stop functionin ...

. Injury caused to the striatum stimulates the migration of neuroblasts from the subventricular zone, to the striatum, where they differentiate into adult neurons. The normal passage of SVZ neuroblasts is to the

but this traffic is diverted to the striatum after an ischemic stroke. However, few of the new developed neurons survive.

Inputs

The largest connection is from the

cortex Cortex or cortical may refer to: Biology * Cortex (anatomy), the outermost layer of an organ ** Cerebral cortex, the outer layer of the vertebrate cerebrum, part of which is the ''forebrain'' *** Motor cortex, the regions of the cerebral cortex i ...

, in terms of cell axons. Many parts of the

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 ...

innervate A nerve is an enclosed, cable-like bundle of nerve fibers (called axons) in the peripheral nervous system. A nerve transmits electrical impulses. It is the basic unit of the peripheral nervous system. A nerve provides a common pathway for the e ...

the dorsal striatum. The cortical

pyramidal neurons Pyramidal cells, or pyramidal neurons, are a type of multipolar neuron found in areas of the brain including the cerebral cortex, the hippocampus, and the amygdala. Pyramidal neurons are the primary excitation units of the mammalian prefrontal cor ...

projecting to the striatum are located in layers II-VI, with the most dense projections come from layer V. They end mainly on the

dendritic spine A dendritic spine (or spine) is a small membranous protrusion from a neuron's dendrite that typically receives input from a single axon at the synapse. Dendritic spines serve as a storage site for synaptic strength and help transmit electrical si ...

s of the spiny neurons. They are

, exciting striatal neurons. The striatum is seen as having its own internal microcircuitry. The ventral striatum receives direct input from multiple regions in 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. The cerebral cortex mostly consists of the six-layered neocortex, with just 10% consisting of ...

and limbic structures such as the

amygdala The amygdala (; plural: amygdalae or amygdalas; also '; Latin from Greek, , ', 'almond', 'tonsil') is one of two almond-shaped clusters of nuclei located deep and medially within the temporal lobes of the brain's cerebrum in complex verteb ...

thalamus The thalamus (from Greek θάλαμος, "chamber") is a large mass of gray matter located in the dorsal part of the diencephalon (a division of the forebrain). Nerve fibers project out of the thalamus to the cerebral cortex in all directions, ...

, and

hippocampus The hippocampus (via Latin from Greek , 'seahorse') is a major component of the brain of humans and other vertebrates. Humans and other mammals have two hippocampi, one in each side of the brain. The hippocampus is part of the limbic system, a ...

, as well as the

entorhinal cortex The entorhinal cortex (EC) is an area of the brain's allocortex, located in the medial temporal lobe, whose functions include being a widespread network hub for memory, navigation, and the perception of time.Integrating time from experience in the ...

and the

inferior temporal gyrus The inferior temporal gyrus is one of three gyri of the temporal lobe and is located below the middle temporal gyrus, connected behind with the inferior occipital gyrus; it also extends around the infero-lateral border on to the inferior surface o ...

. Its primary input is to the

system. Additionally, the

mesolimbic pathway The mesolimbic pathway, sometimes referred to as the reward pathway, is a dopaminergic pathway in the brain. The pathway connects the ventral tegmental area in the midbrain to the ventral striatum of the basal ganglia in the forebrain. The ventral ...

projects from the

ventral tegmental area The ventral tegmental area (VTA) (tegmentum is Latin for ''covering''), also known as the ventral tegmental area of Tsai, or simply ventral tegmentum, is a group of neurons located close to the midline on the floor of the midbrain. The VTA is the ...

to the

of the ventral striatum. Another well-known afferent is the

nigrostriatal The nigrostriatal pathway is a bilateral dopaminergic pathway in the brain that connects the substantia nigra pars compacta (SNc) in the midbrain with the dorsal striatum (i.e., the caudate nucleus and putamen) in the forebrain. It is one of the fo ...

connection arising from the neurons of the

pars compacta. While cortical axons synapse mainly on spine heads of spiny neurons, nigral axons synapse mainly on spine shafts. In primates, the thalamostriatal afferent comes from the central median-parafascicular complex of the

(see

primate basal ganglia system The basal ganglia form a major brain system in all species of vertebrates, but in primates (including humans) there are special features that justify a separate consideration. As in other vertebrates, the primate basal ganglia can be divided into ...

). This afferent is glutamatergic. The participation of truly intralaminar neurons is much more limited. The striatum also receives afferents from other elements of the basal ganglia such as the

subthalamic nucleus The subthalamic nucleus (STN) is a small lens-shaped nucleus in the brain where it is, from a functional point of view, part of the basal ganglia system. In terms of anatomy, it is the major part of the subthalamus. As suggested by its name, the ...

(glutamatergic) or the

external globus pallidus The external globus pallidus (GPe or lateral globus pallidus) combines with the internal globus pallidus (GPi) to form the globus pallidus, an anatomical subset of the basal ganglia. Globus pallidus means "pale globe" in Latin, indicating its appea ...

(

Targets

The primary outputs of the ventral striatum project to the

ventral pallidum The ventral pallidum (VP) is a structure within the basal ganglia of the brain. It is an output nucleus whose fibres project to thalamic nuclei, such as the ventral anterior nucleus, the ventral lateral nucleus, and the medial dorsal nucleus. The ...

, then the

medial dorsal nucleus The medial dorsal nucleus (or dorsomedial nucleus of thalamus) is a large nucleus in the thalamus. It is believed to play a role in memory. Structure It relays inputs from the amygdala and olfactory cortex and projects to the prefrontal cortex a ...

of the

, which is part of the

frontostriatal circuit Frontostriatal circuits are neural pathways that connect frontal lobe regions with the basal ganglia (striatum) that mediate motor, cognitive, and behavioural functions within the brain. They receive inputs from dopaminergic, serotonergic, noradrene ...

. Additionally, the ventral striatum projects to the

, and substantia nigra pars reticulata. Some of its other outputs include projections to the

extended amygdala The extended amygdala is a macrostructure in the brain that is involved in reward cognition and defined by connectivity and neurochemical staining. It includes the central medial amygdala, sublenticular substantia innominata, and the bed nucleus ...

lateral hypothalamus The lateral hypothalamus (LH), also called the lateral hypothalamic area (LHA), contains the primary orexinergic nucleus within the hypothalamus that widely projects throughout the nervous system; this system of neurons mediates an array of cognit ...

, and

pedunculopontine nucleus The pedunculopontine nucleus (PPN) or pedunculopontine tegmental nucleus (PPT or PPTg) is a collection of neurons located in the upper pons in the brainstem. It lies caudal to the substantia nigra and adjacent to the superior cerebellar peduncle. ...

. Striatal outputs from both the dorsal and ventral components are primarily composed of

medium spiny neuron Medium spiny neurons (MSNs), also known as spiny projection neurons (SPNs), are a special type of GABAergic inhibitory cell representing 95% of neurons within the human striatum, a basal ganglia structure. Medium spiny neurons have two primary ...

s (MSNs), a type of

projection neuron The projection fibers consist of efferent and afferent fibers uniting the cortex with the lower parts of the brain and with the spinal cord. In human neuroanatomy, bundles of axons (nerve fibers) called tracts, within the brain, can be catego ...

, which have two primary

phenotype In genetics, the phenotype () is the set of observable characteristics or traits of an organism. The term covers the organism's morphology or physical form and structure, its developmental processes, its biochemical and physiological proper ...

s: "indirect" MSNs that express

D2-like receptor The D2-like receptors are a subfamily of dopamine receptors that bind the endogenous neurotransmitter dopamine. The D2-like subfamily consists of three G-protein coupled receptors that are coupled to Gi/Go and mediate inhibitory neurotransmission ...

s and "direct" MSNs that express

D1-like receptor The D1-like receptors are a subfamily of dopamine receptors that bind the endogenous neurotransmitter dopamine. The D1-like subfamily consists of two G protein–coupled receptors that are coupled to Gs and mediate excitatory neurotransmission ...

s. The main nucleus of the basal ganglia is the striatum which projects directly to the globus pallidus via a pathway of

striatopallidal fibers The striatopallidal fibres, also Wilson's pencils, pencil fibres of Wilson, and pencils of Wilson, are prominent myelinated fibres that connect the striatum to the globus pallidus. Their distinctive appearance allows the putamen to be identified ...

. The striato-pallidal pathway has a whitish appearance due to the myelinated fibers. This projection comprises successively the external globus pallidus (GPe), the internal globus pallidus (GPi), the

pars compacta The pars compacta (SNpc) is a portion of the ''substantia nigra'', located in the midbrain. It is formed by dopaminergic neurons and located medial to the pars reticulata. Parkinson's disease is characterized by the death of dopaminergic neurons ...

of the

(SNc), and the

pars reticulata The pars reticulata (SNpr) is a portion of the substantia nigra and is located lateral to the pars compacta. Most of the neurons that project out of the pars reticulata are inhibitory GABAergic neurons (i.e., these neurons release GABA, which is ...

of substantia nigra (SNr). The neurons of this projection are inhibited by GABAergic synapses from the dorsal striatum. Among these targets, the GPe does not send axons outside the system. Others send axons to the

superior colliculus In neuroanatomy, the superior colliculus () is a structure lying on the roof of the mammalian midbrain. In non-mammalian vertebrates, the homologous structure is known as the optic tectum, or optic lobe. The adjective form ''tectal'' is commonly ...

. Two others comprise the output to the thalamus, forming two separate channels: one through the internal segment of the globus pallidus to the ventral oralis nuclei of the thalamus and from there to the cortical

supplementary motor area The supplementary motor area (SMA) is a part of the motor cortex of primates that contributes to the control of movement. It is located on the midline surface of the hemisphere just in front of (anterior to) the primary motor cortex leg representa ...

and another through the substantia nigra to the ventral anterior nuclei of the thalamus and from there to the

frontal cortex The frontal lobe is the largest of the four major lobes of the brain in mammals, and is located at the front of each cerebral hemisphere (in front of the parietal lobe and the temporal lobe). It is parted from the parietal lobe by a groove betwe ...

and the occulomotor cortex.

Blood supply

Deep penetrating striate arteries supply blood to the striatum. These arteries include the

recurrent artery of Heubner The recurrent artery of Heubner, Heubner's artery or distal medial striate artery is an artery in the head. It is named after the German paediatrician Otto Heubner. It is a branch of the anterior cerebral artery. Its vascular territory is the ant ...

arising from the

anterior cerebral artery The anterior cerebral artery (ACA) is one of a pair of cerebral arteries that supplies oxygenated blood to most midline portions of the frontal lobes and superior medial parietal lobes of the brain. The two anterior cerebral arteries arise from t ...

, and the

lenticulostriate arteries The lenticulostriate arteries, anterolateral central arteries, or antero-lateral ganglionic branches are a group of small arteries arising from the initial part M1 of the middle cerebral artery that supply the basal ganglia. Structure The lenticul ...

arising from the

middle cerebral artery The middle cerebral artery (MCA) is one of the three major paired cerebral artery, cerebral arteries that supply blood to the cerebrum. The MCA arises from the internal carotid artery and continues into the lateral sulcus where it then branches an ...

Function

The ventral striatum, and the

in particular, primarily mediates

, cognition,

, and

motivational salience Motivational salience is a cognitive process and a form of attention that ''motivates'' or propels an individual's behavior towards or away from a particular object, perceived event or outcome. Motivational salience regulates the intensity of be ...

, whereas the dorsal striatum primarily mediates cognition involving

motor function Motor control is the regulation of movement in organisms that possess a nervous system. Motor control includes reflexes as well as directed movement. To control movement, the nervous system must integrate multimodal sensory information (both f ...

, certain

executive functions In cognitive science and neuropsychology, executive functions (collectively referred to as executive function and cognitive control) are a set of cognitive processes that are necessary for the cognitive control of behavior: selecting and succe ...

(e.g.,

inhibitory control Inhibitory control, also known as response inhibition, is a cognitive process – and, more specifically, an executive function – that permits an individual to inhibit their impulses and natural, habitual, or dominant behavioral res ...

and

impulsivity In psychology, impulsivity (or impulsiveness) is a tendency to act on a whim, displaying behavior characterized by little or no forethought, reflection, or consideration of the consequences. Impulsive actions are typically "poorly conceived, prema ...

), and stimulus-response learning; there is a small degree of overlap, as the dorsal striatum is also a component of the

reward system The reward system (the mesocorticolimbic circuit) is a group of neural structures responsible for incentive salience (i.e., "wanting"; desire or craving for a reward and motivation), associative learning (primarily positive reinforcement and class ...

that, along with the

, mediates the encoding of new motor programs associated with future reward acquisition (e.g., the conditioned motor response to a reward cue).

Metabotropic A metabotropic receptor, also referred to by the broader term G-protein-coupled receptor, is a type of membrane receptor that initiates a number of metabolic steps to modulate cell activity. The nervous system utilizes two types of receptors: met ...

dopamine receptors Dopamine receptors are a class of G protein-coupled receptors that are prominent in the vertebrate central nervous system (CNS). Dopamine receptors activate different effectors through not only G-protein coupling, but also signaling through diffe ...

are present both on spiny neurons and on cortical axon terminals.

Second messenger Second messengers are intracellular signaling molecules released by the cell in response to exposure to extracellular signaling molecules—the first messengers. (Intercellular signals, a non-local form or cell signaling, encompassing both first me ...

cascades triggered by activation of these dopamine receptors can modulate pre- and postsynaptic function, both in the short term and in the long term. In humans, the striatum is activated by stimuli associated with reward, but also by

aversive In psychology, aversives are unpleasant stimuli that induce changes in behavior via negative reinforcement or positive punishment. By applying an aversive immediately before or after a behavior the likelihood of the target behavior occurring in ...

novel A novel is a relatively long work of narrative fiction, typically written in prose and published as a book. The present English word for a long work of prose fiction derives from the for "new", "news", or "short story of something new", itsel ...

, unexpected, or intense

stimuli A stimulus is something that causes a physiological response. It may refer to: * Stimulation ** Stimulus (physiology), something external that influences an activity ** Stimulus (psychology), a concept in behaviorism and perception * Stimulus (eco ...

, and cues associated with such events.

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 ...

evidence suggests that the common property linking these stimuli, to which the striatum is reacting, is salience under the conditions of presentation. A number of other brain areas and circuits are also related to reward, such as frontal areas. Functional maps of the striatum reveal interactions with widely distributed regions of the cerebral cortex important to a diverse range of functions. The interplay between the striatum and 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, BA46, ...

is relevant for behavior, particularly adolescent development as proposed by the

dual systems model The dual systems model, also known as the maturational imbalance model, is a theory arising from developmental cognitive neuroscience which posits that increased risk-taking during adolescence is a result of a combination of heightened reward sensi ...

Clinical significance

Parkinson's disease and other movement disorders

Parkinson's disease Parkinson's disease (PD), or simply Parkinson's, is a long-term degenerative disorder of the central nervous system that mainly affects the motor system. The symptoms usually emerge slowly, and as the disease worsens, non-motor symptoms becom ...

results in loss of dopaminergic innervation to the dorsal striatum (and other basal ganglia) and a cascade of consequences.

Atrophy Atrophy is the partial or complete wasting away of a part of the body. Causes of atrophy include mutations (which can destroy the gene to build up the organ), poor nourishment, poor circulation, loss of hormonal support, loss of nerve supply t ...

of the striatum is also involved in

Huntington's disease Huntington's disease (HD), also known as Huntington's chorea, is a neurodegenerative disease that is mostly inherited. The earliest symptoms are often subtle problems with mood or mental abilities. A general lack of coordination and an unst ...

, and

movement disorder Movement disorder refers to any clinical syndrome with either an excess of movement or a paucity of voluntary and involuntary movements, unrelated to weakness or spasticity. Movement disorders are synonymous with basal ganglia or extrapyramidal d ...

s such as

chorea Chorea (or choreia, occasionally) is an abnormal involuntary movement disorder, one of a group of neurological disorders called dyskinesias. The term ''chorea'' is derived from the grc, χορεία ("dance"; see choreia), as the quick movement ...

choreoathetosis Choreoathetosis is the occurrence of involuntary movements in a combination of chorea (irregular migrating contractions) and athetosis (twisting and writhing). It is caused by many different diseases and agents. It is a symptom of several diseases ...

, and

dyskinesias Dyskinesia refers to a category of movement disorders that are characterized by involuntary muscle movements, including movements similar to tics or chorea and diminished voluntary movements. Dyskinesia can be anything from a slight tremor of t ...

. These have also been described as ''circuit disorders'' of the basal ganglia.

Addiction

Addiction Addiction is a neuropsychological disorder characterized by a persistent and intense urge to engage in certain behaviors, one of which is the usage of a drug, despite substantial harm and other negative consequences. Repetitive drug use o ...

, a disorder of the brain's

, arises through the overexpression of DeltaFosB (ΔFosB), a

transcription factor In molecular biology, a transcription factor (TF) (or sequence-specific DNA-binding factor) is a protein that controls the rate of transcription of genetic information from DNA to messenger RNA, by binding to a specific DNA sequence. The fu ...

, in the

s of the

ventral striatum The striatum, or corpus striatum (also called the striate nucleus), is a nucleus (a cluster of neurons) in the subcortical basal ganglia of the forebrain. The striatum is a critical component of the motor and reward systems; receives glutamate ...

. ΔFosB is an

inducible gene Gene expression is the process by which information from a gene is used in the synthesis of a functional gene product that enables it to produce end products, protein or non-coding RNA, and ultimately affect a phenotype, as the final effect. The ...

which is increasingly expressed in the

as a result of repeatedly

overdosing A drug overdose (overdose or OD) is the ingestion or application of a drug or other substance in quantities much greater than are recommended.

on an addictive drug or overexposure to other addictive stimuli.
Table 1
/ref>

Bipolar disorder

An association has been observed between striatal expression of variants of the

PDE10A cAMP and cAMP-inhibited cGMP 3',5'-cyclic phosphodiesterase 10A is an enzyme that in humans is encoded by the ''PDE10A'' gene. Various cellular responses are regulated by the second messengers cAMP and cGMP. Phosphodiesterases, such as PDE10A, e ...

gene and some

bipolar I disorder Bipolar I disorder (BD-I; pronounced "type one bipolar disorder") is a type of bipolar spectrum disorder characterized by the occurrence of at least one manic episode, with or without mixed or psychotic features. Most people also, at other time ...

patients. Variants of other genes,

DISC1 Disrupted in schizophrenia 1 is a protein that in humans is encoded by the ''DISC1'' gene. In coordination with a wide array of interacting partners, DISC1 has been shown to participate in the regulation of cell proliferation, differentiation, m ...

and GNAS, have been associated with

bipolar II disorder Bipolar II disorder (BP-II) is a mood disorder on the bipolar spectrum, characterized by at least one episode of hypomania and at least one episode of major depression. Diagnosis for BP-II requires that the individual must never have experience ...

Autism spectrum disorder

Autism spectrum disorder The autism spectrum, often referred to as just autism or in the context of a professional diagnosis autism spectrum disorder (ASD) or autism spectrum condition (ASC), is a neurodevelopmental disorder, neurodevelopmental condition (or conditions) ...

(ASD) is characterized by cognitive inflexibility and poor understanding of social systems. This inflexible behavior originates in defects in the pre-frontal cortex as well as the striatal circuits. The defects in the striatum seem to specifically contribute to the motor, social and communication impairments seen in ASD patients. In mice which have an ASD-like phenotype induced via the overexpression of the eukaryotic initiation of translation factor 4E, it has been shown that these defects seem to stem from the reduced ability to store and process information in the striatum, which leads to the difficulty seen in forming new motor patterns, as well as disengaging from existing ones.

Dysfunction

Dysfunction in the ventral striatum can lead to a variety of disorders, most notably, depression and obsessive-compulsive disorder. Because of its involvement in reward pathways, the ventral striatum has also been implicated in playing a critical role in addiction. It has been well established that the ventral striatum is strongly involved in mediating the reinforcing effects of drugs, especially stimulants, through dopaminergic stimulation.

History

In the seventeenth and eighteenth centuries, the term "corpus striatum" was used to designate many distinct, deep, infracortical elements of the hemisphere. Etymologically it is derived from (Latin) "striatus" = "grooved, striated" and the English "striated" = having parallel lines or grooves on the surface. In 1876

David Ferrier Sir David Ferrier FRS (13 January 1843 – 19 March 1928) was a pioneering Scottish neurologist and psychologist. Ferrier conducted experiments on the brains of animals such as monkeys and in 1881 became the first scientist to be prosecuted u ...

contributed decades of research to the subject; concluding that the corpus striatum was vital in the "organization and generation of voluntary movement". In 1941, Cécile and

Oskar Vogt Oskar Vogt (6 April 1870, in Husum – 30 July 1959, in Freiburg im Breisgau) was a German physician and neurologist. He and his wife Cécile Vogt-Mugnier are known for their extensive cytoarchetectonic studies on the brain. Personal life He wa ...

simplified the nomenclature by proposing the term ''striatum'' for all elements in the

built with striatal elements: the

, the

, and the fundus striati, which is the ventral part linking the two preceding together ventrally to the inferior part of the

. The term ''neostriatum'' was forged by comparative anatomists comparing the subcortical structures between vertebrates, because it was thought to be a phylogenetically newer section of the corpus striatum. The term is still used by some sources, including

Medical Subject Headings Medical Subject Headings (MeSH) is a comprehensive controlled vocabulary for the purpose of indexing journal articles and books in the life sciences. It serves as a thesaurus that facilitates searching. Created and updated by the United States N ...

Other animals

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...

s the term used was the ''paleostriatum augmentatum'', and in the new avian terminology listing (as of 2002) for ''neostriatum'' this has been changed to the ''

nidopallium The nidopallium, meaning nested pallium, is the region of the avian brain that is used mostly for some types of executive functions but also for other higher cognitive tasks. The region was renamed nidopallium in 2002 during the Avian Brain Nomen ...

''. In non-primate species, the

are included in the ventral striatum.

Additional images

File:Striatum coronal sections.gif, Striatum highlighted in green on coronal T1 MRI images File:Striatum sagittal sections.gif, Striatum highlighted in green on sagittal T1 MRI images File:Striatum transversal sections.gif, Striatum highlighted in green on transversal T1 MRI images

References

External links

* * * * https://web.archive.org/web/20131029195257/http://www.nimh.nih.gov/images/news-items/r1_braindorsal1.jpg * https://web.archive.org/web/20090914200329/http://www.hnl.bcm.tmc.edu/fmri.html {{Authority control Cerebrum Basal ganglia Addiction

Structure

Cell types

Inputs

Targets

Blood supply

Function

Clinical significance

Parkinson's disease and other movement disorders

Addiction

Bipolar disorder

Autism spectrum disorder

Dysfunction

History

Other animals

See also

Additional images

References

External links