Wikipedia

Thalamus

This article is about the portion of thebrain.For the British video game developer, seeThalamus Ltd.For the botanical structure, seeReceptacle (botany).

Thethalamus(pl.:thalami;fromGreekθάλαμος,"chamber" ) is a large mass ofgray matteron the lateral walls of thethird ventricleforming thedorsalpart of thediencephalon(a division of theforebrain).Nerve fibersproject out of the thalamus to thecerebral cortexin all directions, known as thethalamocortical radiations,allowinghub-likeexchanges of information. It has several functions, such as the relaying ofsensoryandmotorsignals to the cerebral cortex[1][2]and the regulation ofconsciousness,sleep,andalertness.[3][4]

Thalamus
Thalamus marked (MRIcross-section)
Visual depiction of basic thalamus
Details
Part ofDiencephalon
PartsSeeList of thalamic nuclei
ArteryPosterior cerebral arteryand branches
Identifiers
Latinthalamus dorsalis
MeSHD013788
NeuroNames300
NeuroLexIDbirnlex_954
TA98A14.1.08.101
A14.1.08.601
TA25678
TEE5.14.3.4.2.1.8
FMA62007
Anatomical terms of neuroanatomy

Anatomically, it is a paramedian symmetrical structure of two halves (left and right), within thevertebratebrain, situated between the cerebral cortex and themidbrain.It forms duringembryonic developmentas the main product of the diencephalon, as first recognized by the Swissembryologistand anatomistWilhelm His Sr.in 1893.[5]

Anatomy

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The thalamus is a paired structure of gray matter about four centimetres long, located in theforebrainwhich is superior to themidbrain,near the center of the brain with nerve fibers projecting out to the cerebral cortex in all directions. In fact, almost all thalamic neurons (with the notable exception of thethalamic reticular nucleus[6]) project to the cerebral cortex, and every region of the cortex so far studied has been found to innervate the thalamus.[7]

Each of the thalami may be subdivided into at least 30nuclei,giving a total of at least 60 for the whole thalamus.[4][8]

Estimates of the volume of the whole thalamus vary. A post-mortem study of 10 people with average age 71 years found average volume 13.68 cm.[9]In a study of 12 healthy males with average age 17 years, MRI scans showed mean whole thalamus volume 8.68cm.[10]

The medial surface of the thalamus constitutes the upper part of the lateral wall of thethird ventricle,and is connected to the corresponding surface of the opposite thalamus by a flattened gray band, theinterthalamic adhesion.

The lateral part of the thalamus is theneothalamus,thephylogeneticallynewest part of the thalamus, which includes thelateral nuclei,thepulvinar nucleiand themedialandlateral geniculate nuclei.[11][12]

The surface of the thalamus is covered by two layers ofwhite matter,thestratum zonalecovers the dorsal surface, and theexternal medullary laminacovers the lateral surface. (This stratum zonale should not be confused with thestratum zonaleof thesuperior colliculus.) Within the thalamus theinternal medullary laminadivides the nuclei into anterior, medial, and lateral groups.[13][14]

Derivatives of thediencephaloninclude the dorsally-locatedepithalamus(essentially thehabenulaand annexes) and the perithalamus (prethalamus) containing thezona incertaand the thalamic reticular nucleus. Due to their differentontogeneticorigins, the epithalamus and the perithalamus are formally distinguished from the thalamus proper. Themetathalamusis made up of the lateral geniculate and medial geniculate nuclei.[15][16]

The thalamus comprises a system oflamellae(made up ofmyelinatedfibers) that separate different thalamic subparts. Other areas are defined by distinct clusters ofneurons,such as theperiventricular nucleus,the intralaminar elements, the "nucleus limitans", and others.[17]These latter structures, different in structure from the major part of the thalamus, have been grouped together into theallothalamusas opposed to theisothalamus.[18]This distinction simplifies the global description of the thalamus.

Thalamic nuclei.Metathalamuslabelled MTh.
(Left thalamus viewed from left.)
Nuclei of right thalamus
(viewed from above right)

Thalamic nuclei

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See also:List of thalamic nuclei
Medial nuclei of the left thalamus.
Key: CeMCentral Medial.CLCentral Lateral.CMCentroMedian.MDMedial Dorsal.MV MedioVentral=Reuniens.PfParafascicular.(Lateral view shows sagittal section through left thalamus)
Lateral nuclei of the left thalamus.
Key: VAVentral Anterior.VLVentral Lateral.VMVentral Medial.VPI Ventral PosteroInferior. VPLVentral PosteroLateral.VPMVentral PosteroMedial.(Medial view shows sagittal section through left thalamus.)

The principal subdivision of the thalamus into nucleus groups is the trisection of each thalamus (left and right) by a Y-shapedinternal medullary lamina.This trisection divides each thalamus intoanterior,medialandlateral groupsof nuclei.[8]The medial group is subdivided into themedial dorsal nucleusandmidline group.The lateral group is subdivided intoventral,pulvinar,lateral dorsal,lateral posteriorand metathalamus. The ventral group is further subdivided intoventral anterior,ventral lateralandventral posterior.

The interior medullary lamina is subdivided intointralaminar nuclei.Additional structures are the reticular nucleus (which envelops the lateral thalamus), the stratum zonale,[19]and theinterthalamic adhesion.[20]

Combining these division principles yields the following hierarchy, which is subject to many further subdivisions.[21]

  • anterior group
  • medial group
    • medial dorsal nucleus
    • midline group
  • lateral group
    • ventral group
      • ventral anterior group
      • ventral lateral group
      • ventral posterior group
    • pulvinar group
    • lateral dorsal nucleus
    • lateral posterior nucleus
    • metathalamus
      • lateral geniculate nucleus
      • medial geniculate nucleus
  • intralaminar group
  • reticular nucleus
  • stratum zonale
  • interthalamic adhesion

The term "lateral nuclear group" is used with two meanings. It can mean either the complete set of nuclei in the lateral "third" of the trisection by the lamina, or the subset which excludes the ventral group and the geniculate nuclei.[22][23]

Blood supply

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The thalamus derives its blood supply from a number of arteries: the polar artery (posterior communicating artery), paramedian thalamic-subthalamic arteries, inferolateral (thalamogeniculate) arteries, andposterior (medial and lateral) choroidal arteries.[24][25]These are all branches of theposterior cerebral artery.[8][26]

Some people have theartery of Percheron,which is a rare anatomic variation in which a single arterial trunk arises from the posterior cerebral artery to supply both parts of the thalamus.

Connections

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The thalamus is connected to the spinal cord via thespinothalamic tract

The thalamus has many connections to the hippocampus via themammillothalamic tract.This tract comprises themammillary bodiesandfornix.[27]

The thalamus is connected to thecerebral cortexvia thethalamocortical radiations.[28]

Thespinothalamic tractis a sensory pathway originating in the spinal cord. It transmits information to the thalamus about pain, temperature, itch andcrude touch.There are two main parts: thelateral spinothalamic tract,which transmits pain and temperature, and theanterior (or ventral) spinothalamic tract,which transmits crude touch and pressure.

Function

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The thalamus has multiple functions, and is generally believed to act as a relay station, orhub,relaying information between different subcortical areas and the cerebral cortex.[29]In particular, every sensory system (with the exception of theolfactory system) includes a thalamic nucleus that receives sensory signals and sends them to the associated primary cortical area.[30][31]

For the visual system, for example, inputs from theretinaare sent to thelateral geniculate nucleusof the thalamus, which in turn projects to thevisual cortexin theoccipital lobe.[32]Similarly themedial geniculate nucleusacts as a keyauditoryrelay between theinferior colliculusof themidbrainand theprimary auditory cortex.[citation needed]Theventral posterior nucleusis a keysomatosensoryrelay, which sends touch andproprioceptiveinformation to theprimary somatosensory cortex.[citation needed]In rodents, proprioceptive information of head and whisker movements is integrated already at the thalamic level.[33]

The thalamus is believed to both process sensory information as well as relay it—each of the primary sensory relay areas receives strong feedback connections from the cerebral cortex.[34]

The thalamus also plays an important role in regulating states ofsleep,andwakefulness.[35]Thalamic nuclei have strong reciprocal connections with the cerebral cortex, formingthalamo-cortico-thalamic circuitsthat are believed to be involved withconsciousness.[36][37]The thalamus plays a major role in regulating arousal, the level of awareness, and activity. Damage to the thalamus can lead to permanentcoma.[38]

The role of the thalamus in the more anteriorpallidalandnigralterritories in thebasal gangliasystem disturbances is recognized but still poorly understood. The contribution of the thalamus to vestibular or totectalfunctions is almost ignored. The thalamus has been thought of as a "relay" that simply forwards signals to the cerebral cortex. Newer research suggests that thalamic function is more selective.[39]Many different functions are linked to various regions of the thalamus. This is the case for many of the sensory systems (except for the olfactory system), such as theauditory,somatic,visceral,gustatoryandvisual systemswhere localized lesions provoke specific sensory deficits. A major role of the thalamus is support of motor and language systems, and much of the circuitry implicated for these systems is shared.

The thalamus isfunctionally connectedto thehippocampus[40]as part of the extended hippocampal system at the thalamic anterior nuclei.[41]With respect to spatial memory and spatial sensory datum they are crucial for human episodic event memory.[42][43]The thalamic region's connection to themedial temporal lobeprovides differentiation of the functioning of recollective and familiarity memory.[27]

The neuronal information processes necessary for motor control were proposed as a network involving the thalamus as a subcortical motor center.[44]Through investigations of the anatomy of the brains of primates[45]the nature of the interconnected tissues of thecerebellumto the multiple motor cortices suggested that the thalamus fulfills a key function in providing the specific channels from the basal ganglia and cerebellum to the cortical motor areas.[46][47]In an investigation of thesaccadeandantisaccade[48]motor response in three monkeys, the thalamic regions were found to be involved in the generation of antisaccade eye-movement (that is, the ability to inhibit the reflexive jerking movement of the eyes in the direction of a presented stimulus).[49]

Recent research suggests that the mediodorsal thalamus (MD) may play a broader role in cognition. Specifically, the mediodorsal thalamus may "amplify the connectivity (signaling strength) of just the circuits in the cortex appropriate for the current context and thereby contribute to the flexibility (of the mammalian brain) to make complex decisions by wiring the many associations on which decisions depend into weakly connected cortical circuits."[50]Researchers found that "enhancing MD activity magnified the ability of mice to" think, "[50]driving down by more than 25 percent their error rate in deciding which conflicting sensory stimuli to follow to find the reward. "[51]

Development

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The thalamic complex is composed of the perithalamus (or prethalamus, previously also known as ventral thalamus), the mid-diencephalic organiser (which forms later thezona limitans intrathalamica(ZLI) ) and the thalamus (dorsal thalamus).[52][53]The development of the thalamus can be subdivided into three steps.[54] The thalamus is the largest structure deriving from the embryonicdiencephalon,the posterior part of the forebrain situated between the midbrain and the cerebrum.

Early brain development

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Afterneurulation,the early developmental stage (primordium) of theprethalamusand the thalamus is induced within theneural tube.Data from different vertebrate model organisms support a model in which the interaction between twotranscription factors,Fez and Otx, is of decisive importance. Fez is expressed in the prethalamus, and functional experiments show that Fez is required for prethalamus formation.[55][56]Posteriorly,OTX1andOTX2abut the expression domain of Fez and are required for proper development of the thalamus.[57][58]

Formation of progenitor domains

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Early in thalamic development two progenitor domains form, a caudal domain, and a rostral domain. The caudal domain gives rise to all of the glutamatergic neurons in the adult thalamus while the rostral domain gives rise to all of the GABAergic neurons in the adult thalamus.[59]

The formation of the mid-diencephalic organiser (MDO)

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At the interface between the expression domains of Fez and Otx, the mid-diencephalic organizer (MDO, also called the ZLI organiser) is induced within the thalamicanlage.The MDO is the central signalling organizer in the thalamus. A lack of the organizer leads to the absence of the thalamus. The MDO matures from ventral to dorsal during development. Members of thesonic hedgehog(SHH) family and of theWntfamily are the main principal signals emitted by the MDO.

Besides its importance as signalling center, the organizer matures into the morphological structure of thezona limitans intrathalamica(ZLI).

Maturation and parcellation of the thalamus

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After its induction, the MDO starts to orchestrate the development of the thalamic anlage by release of signalling molecules such as SHH.[60]In mice, the function of signaling at the MDO has not been addressed directly due to a complete absence of thediencephalonin SHH mutants.[61]

Studies in chicks have shown that SHH is both necessary and sufficient for thalamic gene induction.[62]Inzebrafish,it was shown that the expression of two SHH genes, SHH-a and SHH-b (formerly described as twhh) mark the MDO territory, and that SHH signaling is sufficient for the molecular differentiation of both the prethalamus and the thalamus but is not required for their maintenance and SHH signaling from the MDO/alar plate is sufficient for the maturation of prethalamic and thalamic territory while ventral Shh signals are dispensable.[63]

The exposure to SHH leads to differentiation of thalamic neurons. SHH signaling from the MDO induces a posterior-to-anterior wave of expression the proneural geneNeurogenin1in the major (caudal) part of the thalamus, and Ascl1 (formerly Mash1) in the remaining narrow stripe of rostral thalamic cells immediately adjacent to the MDO, and in the prethalamus.[64][65]

This zonation of proneural gene expression leads to the differentiation of glutamatergic relay neurons from the Neurogenin1+ precursors and of GABAergic inhibitory neurons from the Ascl1+ precursors. In fish, selection of these alternative neurotransmitter fates is controlled by the dynamic expression of Her6 the homolog ofHES1.Expression of this hairy-like bHLHtranscription factor,which represses Neurogenin but is required for Ascl1, is progressively lost from the caudal thalamus but maintained in the prethalamus and in the stripe of rostral thalamic cells. In addition, studies on chick and mice have shown that blocking the Shh pathway leads to absence of the rostral thalamus and substantial decrease of the caudal thalamus. The rostral thalamus will give rise to the reticular nucleus mainly whereby the caudal thalamus will form the relay thalamus and will be further subdivided in thethalamic nuclei.[54]

In humans, a common genetic variation in the promoter region of theserotonin transporter(the SERT-long and -short allele:5-HTTLPR) has been shown to affect the development of several regions of the thalamus in adults. People who inherit two shortalleles(SERT-ss) have more neurons and a larger volume in thepulvinarand possibly the limbic regions of the thalamus. Enlargement of the thalamus provides an anatomical basis for why people who inherit two SERT-ss alleles are more vulnerable tomajor depression,post-traumatic stress disorder,and suicide.[66]

Clinical significance

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A thalamus damaged by astrokecan lead tothalamic pain syndrome,[67]which involves a one-sided burning or aching sensation often accompanied bymood swings.Bilateralischemiaof the area supplied by the paramedian artery can cause serious problems includingakinetic mutism,and be accompanied byoculomotorproblems. A related concept isthalamocortical dysrhythmia.The occlusion of theartery of Percheroncan lead to a bilateral thalamus infarction.

Korsakoff syndromestems from damage to themammillary body,themammillothalamic fasciculusor the thalamus.[68][69]

Fatal familial insomniais a hereditarypriondisease in which degeneration of the thalamus occurs, causing the patient to gradually lose their ability to sleep and progressing to a state of totalinsomnia,which invariably leads to death. In contrast, damage to the thalamus can result in coma.

Atrophy of the thalamus is an indicator of the start ofmultiple sclerosis.[70][71]Thalamic volume loss by atrophy, is also significantly shown in sporadicfrontotemporal dementia,noted in the anterior-dorsal thickness.[72]

Microstimulation of the posterior portion of theventral medial thalamic nucleuscan be used to evoke pain, temperature and visceral sensations.[73]

Additional images

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  • Human brain dissection, showing the thalamus.
    (View from above.)
  • Human thalamus along with other subcortical structures, in glass brain.
  • The thalamus in a 360° rotation
  • Diagram showing the anterior and lateralspinothalamic tractswithin thespinal cord
  • Dorsal view
  • Coronal section of lateral and third ventricles
  • Median sagittal section of brain of human embryo of three months
  • Human brain frontal (coronal) section. Thalamus label is 10.
  • Thalamus coronal section
  • Left thalamus viewed from left dorsal posterior

See also

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References

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