Thalamocortical radiations

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Thalamocortical radiations
Gray685.png
Deep dissection of brain-stem. Lateral view. (Thalamocortical fibers labeled at center top.)
Corticothalamic Pathway.jpg
Tractography of thalamocortical radiations
Details
From Thalamus
To Cortex
Identifiers
Latin radiatio thalamocorticalis,
tractus thalamocorticalis
NeuroNames 1337
Anatomical terms of neuroanatomy

In neuroanatomy, thalamocortical radiations, also known as thalamocortical fibres, are the efferent fibres that project from the thalamus to distinct areas of the cerebral cortex. [1] They form fibre bundles that emerge from the lateral surface of the thalamus. [2]

Contents

Structure

Thalamocortical fibers (TC fibres) have been referred to as one of the two constituents of the isothalamus, the other being microneurons. Thalamocortical fibers have a bush or tree-like appearance as they extend into the internal capsule and project to the layers of the cortex. The main thalamocortical fibers extend from different nuclei of the thalamus and project to the visual cortex, somatosensory (and associated sensori-motor) cortex, and the auditory cortex in the brain. Thalamocortical radiations also innervate gustatory and olfactory pathways, as well as pre-frontal motor areas. Visual input from the optic tract is processed by the lateral geniculate nucleus of the thalamus, auditory input in the medial geniculate nucleus, and somatosensory input in the ventral posterior nucleus of the thalamus. Thalamic nuclei project to cortical areas of distinct architectural organization and relay the processed information back to the area of original activity in the thalamus via corticothalamic fibers (CT fibres). [3] The thalamic reticular nucleus (TRN) receives incoming signals via corticothalamic pathways and regulates activity within the thalamus accordingly. [4] Cortico-thalamic feedback neurons are mostly found in layer VI of the cortex. [5] Reciprocal CT projections to the thalamus are of a higher order than, and synapse with, the TRN in much greater number than do thalamocortical projections to cortex. [6] This suggests that the cortex has a much bigger role in top down processing and regulation of thalamic activity than do the processes originating in thalamic interneurons. Large-scale frequency oscillations and electrical rhythms have also been shown to regulate TC activity for long periods of time, as is evident during the sleep cycle. [7] Other evidence suggests CT modulation of TC rhythms can occur over different time scales, adding even more complexity to their function. [5]

Relay cells

Thalamic interneurons process sensory information and signal different regions of the thalamic nuclei. These nuclei extend to relay cells, which in turn innervate distinct areas of the cortex via thalamocortical fibers. Either specifically or nonspecifically, TC relay cells project specifically to organized areas of the cortex directly and nonspecifically project to large areas of cortex through the innervation of many interconnected collateral axons. [8] According to Jones (2001) there are two primary types of relay neurons in the thalamus of primates–core cells and matrix cells–each creating distinct pathways to various parts and layers throughout the cerebral cortex. [6] Matrix cells of the thalamus, or calbindin-immuno-reactive neurons (CIR neurons), are widely distributed and diffusely dispersed in each of the nuclei of the dorsal thalamus. In comparison, parvalbumin immuno-reactive neurons (PIR neurons) can be found only in principal sensory and motor relay nuclei, and in the pulvinar nuclei as well as the intralaminar nuclei. The PIR neurons cluster together creating "densely terminating afferent fibers…forming a core imposed on a diffuse background matrix of PIR cells" (Jones 2001). PIR cells tend to project upon the cerebral cortex and terminate in an organized topographic manner in specifically localized zones (in deep layer III and in the middle layer IV). In contrast, CIR cells have dispersed projections wherein various adjacent cells connect to non-specific different cortical areas. CIR axons seem to terminate primarily in the superficial layers of the cortex: layers I, II, and upper III. [6]

Function

Thalamocortical signaling is primarily excitatory, causing the activation of corresponding areas of the cortex, but is mainly regulated by inhibitory mechanisms. The specific excitatory signaling is based upon glutamatergic signaling, and is dependent on the nature of the sensory information being processed. Recurrent oscillations in thalamocortical circuits also provide large-scale regulatory feedback inputs to the thalamus via GABAergic neurons that synapse in the TRN. [9] In a study done by Gibbs, Zhang, Shumate, and Coulter (1998) it was found that endogenously released zinc blocked GABA responses within the TC system specifically by interrupting communication between the thalamus and the connected TRN. [9] Computational neuroscientists are particularly interested in thalamocortical circuits because they represent a structure that is disproportionally larger and more complex in humans than other mammals (when body size is taken into account), which may contribute to humans' special cognitive abilities. [10] Evidence from one study (Arcelli et al. 1996) offers partial support to this claim by suggesting that thalamic GABAergic local circuit neurons in mammalian brains relate more to processing ability compared to sensorimotor ability, as they reflect an increasing complexity of local information processing in the thalamus. [8] It is proposed that core relay cells and matrix cells projecting from the dorsal thalamus allow for synchronization of cortical and thalamic cells during "high-frequency oscillations that underlie discrete conscious events", [6] though this is a heavily debated area of research.

Projections

The majority of thalamocortical fibers project to layer IV of the cortex, wherein sensory information is directed to other layers where they either terminate or connect with axons collaterally depending on type of projection and type of initial activation. [11] Activation of the thalamocortical neurons relies heavily on the direct and indirect effects of glutamate, which causes excitatory postsynaptic potentials (EPSPs) at terminal branches in the primary sensory cortices. [12]

Somatosensory areas

Primarily, thalamocortical somatosensory radiation from the VPL, VPM and LP nuclei extends to the primary and secondary somatosensory areas, terminating in cortical layers of the lateral postcentral gyrus. S1 receives parallel thalamocortical radiations from the posterior medial nucleus and the VPN. [13] Projections from the VPN to the postcentral gyrus account for the transfer of sensory information concerning touch and pain. Several studies indicate that parallel innervations to S1 and also S2 via thalamocortical pathways result in the processing of nociceptive and non-nociceptive information. Non-specific projections to sensori-motor areas of the cortex may in part have to do with the relationship between non-noci-receptive processing and motor functions. [14] Past research shows a link between S1 and M1, creating a thalamocortical sensori-motor circuit. When this circuit becomes disrupted symptoms are produced similar to those that accompany Multiple sclerosis, suggesting thalamocortical rhythms are involved in regulating sensori-motor pathways in a highly specialized manner. [15] TC-CT rhythms evident during sleep act to inhibit these thalamocortical fibers so as to maintain the tonic cycling of low frequency waves and the subsequent suppression of motor activity. [7]

Visual areas

The lateral geniculate nucleus and the pulvinar nuclei project to and terminate in V1, and carry motor information from the brain stem as well as other sensory input from the optic tract. The visual cortex connects with other sensory areas which allows for the integration of cognitive tasks such as selective and directed attention, and pre-motor planning, in relation to the processing of incoming visual stimuli. [16] Models of the pulvinar projections to the visual cortex have been proposed by several imaging studies, though their mapping has been difficult due to the fact that pulvinar subdivisions are not conventionally organized and have been difficult to visualize using structural MRI. [16] Evidence from several studies supports the idea that the pulvinar nuclei and superior colliculus receive descending projections from CT fibers while TC fibers extending from the LGN carry visual information to the various areas of the visual cortex near the calcarine fissure.

Auditory areas

Thalamocortical axons project primarily from the medial geniculate nucleus via the sublenticular region of the internal capsule, and terminate in an organized topographic manner in the transverse temporal gyri. [17] MMGN radiations terminate in specific locations while thalamocortical fibers from the VMGN terminate in nonspecific clusters of cells and form collateral connections to neighboring cells. [18] Research done by staining the brains of macaque monkeys reveals projections from the ventral nucleus mainly terminating in layers IV and IIIB, with some nonspecific clusters of PIR cells terminating in layers I, II, IIIA, and VI. Fibers from the dorsal nuclei were found to project more directly to the primary auditory area, with most axons terminating in layer IIIB. The magnocellular nucleus projected a small amount of PIR cells with axons mainly terminating in layer 1, though large regions of the middle cortical layers were innervated through collaterally connected CIR neurons. [19] Past research suggests that the thalamocortical-auditory pathway may be the only neural correlate that can explain a direct translation of frequency information to the cortex via specific pathways. [20]

Motor areas

The primary motor cortex receives terminating thalamocortical fibers from the VL nucleus of the thalamus. This is the primary pathway involved in the transference of cerebellar input to the primary motor cortex. The VA projects widely across the inferior parietal and premotor cortex. Other Non-specific thalamocortical projections, those that originate in the dorsal-medial nuclei of the thalamus, terminate in the prefrontal cortex and have subsequent projections to associative premotor areas via collateral connections. [21] The cortico-basal ganglia-thalamo-cortical loop has been traditionally associated with reward-learning and though has also been noted by some researchers to have a modulatory effect on thalamocortical network functioning–this is due to inherent activation of the premotor areas connecting the VA nucleus with the cortex. [21]

Clinical significance

Absence seizures

Thalamocortical radiations have been researched extensively in the past due to their relationship with attention, wakefulness, and arousal. Past research has shown how an increase in spike-and-wave activity within the TC network can disrupt normal rhythms involved with the sleep-wakefulness cycle, ultimately causing absence seizures and other forms of epileptic behavior. Burst firing within a part of the TC network stimulates GABA receptors within the thalamus causing moments of increased inhibition, leading to frequency spikes, which offset oscillation patterns. [22] [23] Another study done on rats suggests during spike-and-wave seizures, thalamic rhythms are mediated by local thalamic connections, while the cortex controls the synchronization of these rhythms over extended periods of time. Thalamocortical dysrhythmia is a term associated with spontaneously reoccurring low frequency spike-and-wave activity in the thalamus, which causes symptoms normally associated with impulse control disorders such as obsessive compulsive disorder, Parkinson's disease, attention deficit hyperactivity disorder, and other forms of chronic psychosis. [24] Other evidence has shown how reductions in the distribution of connections of nonspecific thalamocortical systems is heavily associated with loss of consciousness, as can be seen with individuals in a vegetative state, or coma. [25]

Prefrontal lobotomy

The bilateral interruption or severing of the connection between thalamocortical radiations the medial and anterior thalamic nuclei results in a prefrontal lobotomy, which causes a drastic personality change and a subdued behavioral disposition without cortical injury. [26]

Research

Evolutionary theories of consciousness

Theories of consciousness have been linked to thalamocortical rhythm oscillations in TC-CT pathway activity. One such theory, the dynamic core theory of conscious experience, proposes four main pillars in support of conscious awareness as a consequence of dorsal thalamic activity: [27]

  1. the results of cortical computations underlay consciousness
  2. vegetative states and general anesthetics work primarily to disrupt normal thalamic functioning
  3. the anatomy and physiology of the thalamus implies consciousness
  4. neural synchronization accounts for the neural basis of consciousness.

This area of research is still developing, and most current theories are either partial or incomplete.

Related Research Articles

<span class="mw-page-title-main">Cerebral cortex</span> Outer layer of the cerebrum of the mammalian brain

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 the allocortex. It is separated into two cortices, by the longitudinal fissure that divides the cerebrum into the left and right cerebral hemispheres. The two hemispheres are joined beneath the cortex by the corpus callosum. The cerebral cortex is the largest site of neural integration in the central nervous system. It plays a key role in attention, perception, awareness, thought, memory, language, and consciousness. The cerebral cortex is part of the brain responsible for cognition.

<span class="mw-page-title-main">Thalamus</span> Structure within the brain

The thalamus is a large mass of gray matter located in the dorsal part of the diencephalon. Nerve fibers project out of the thalamus to the cerebral cortex in all directions, known as the thalamocortical radiations, allowing hub-like exchanges of information. It has several functions, such as the relaying of sensory signals, including motor signals to the cerebral cortex and the regulation of consciousness, sleep, and alertness.

<span class="mw-page-title-main">Trigeminal nerve</span> Cranial nerve responsible for the faces senses and motor functions

In neuroanatomy, the trigeminal nerve (lit. triplet nerve), also known as the fifth cranial nerve, cranial nerve V, or simply CN V, is a cranial nerve responsible for sensation in the face and motor functions such as biting and chewing; it is the most complex of the cranial nerves. Its name (trigeminal, from Latin tri- 'three', and -geminus 'twin') derives from each of the two nerves (one on each side of the pons) having three major branches: the ophthalmic nerve (V1), the maxillary nerve (V2), and the mandibular nerve (V3). The ophthalmic and maxillary nerves are purely sensory, whereas the mandibular nerve supplies motor as well as sensory (or "cutaneous") functions. Adding to the complexity of this nerve is that autonomic nerve fibers as well as special sensory fibers (taste) are contained within it.

<span class="mw-page-title-main">Lateral geniculate nucleus</span> Component of the visual system in the brains thalamus

In neuroanatomy, the lateral geniculate nucleus is a structure in the thalamus and a key component of the mammalian visual pathway. It is a small, ovoid, ventral projection of the thalamus where the thalamus connects with the optic nerve. There are two LGNs, one on the left and another on the right side of the thalamus. In humans, both LGNs have six layers of neurons alternating with optic fibers.

<span class="mw-page-title-main">Internal capsule</span> White matter structure situated in the inferomedial part of each cerebral hemisphere of the brain

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 globus pallidus. The internal capsule contains both ascending and descending axons, going to and coming from the cerebral cortex. It also separates the caudate nucleus and the putamen in the dorsal striatum, a brain region involved in motor and reward pathways.

<span class="mw-page-title-main">Spinothalamic tract</span> Sensory pathway from the skin to the thalamus

The spinothalamic tract is a part of the anterolateral system or the ventrolateral system, a sensory pathway to the thalamus. From the ventral posterolateral nucleus in the thalamus, sensory information is relayed upward to the somatosensory cortex of the postcentral gyrus.

<span class="mw-page-title-main">Dorsal column–medial lemniscus pathway</span> Sensory spinal pathway

The dorsal column–medial lemniscus pathway (DCML) is a sensory pathway of the central nervous system that conveys sensations of fine touch, vibration, two-point discrimination, and proprioception from the skin and joints. It transmits information from the body to the primary somatosensory cortex in the postcentral gyrus of the parietal lobe of the brain. The pathway receives information from sensory receptors throughout the body, and carries this in nerve tracts in the white matter of the dorsal column of the spinal cord to the medulla, where it is continued in the medial lemniscus, on to the thalamus and relayed from there through the internal capsule and transmitted to the somatosensory cortex. The name dorsal-column medial lemniscus comes from the two structures that carry the sensory information: the dorsal columns of the spinal cord, and the medial lemniscus in the brainstem.

<span class="mw-page-title-main">Barrel cortex</span> Region of the somatosensory cortex in some rodents and other species

The barrel cortex is a region of the somatosensory cortex that is identifiable in some species of rodents and species of at least two other orders and contains the barrel field. The 'barrels' of the barrel field are regions within cortical layer IV that are visibly darker when stained to reveal the presence of cytochrome c oxidase and are separated from each other by lighter areas called septa. These dark-staining regions are a major target for somatosensory inputs from the thalamus, and each barrel corresponds to a region of the body. Due to this distinctive cellular structure, organisation, and functional significance, the barrel cortex is a useful tool to understand cortical processing and has played an important role in neuroscience. The majority of what is known about corticothalamic processing comes from studying the barrel cortex, and researchers have intensively studied the barrel cortex as a model of neocortical column.

<span class="mw-page-title-main">Pretectal area</span> Structure in the midbrain which mediates responses to ambient light

In neuroanatomy, the pretectal area, or pretectum, is a midbrain structure composed of seven nuclei and comprises part of the subcortical visual system. Through reciprocal bilateral projections from the retina, it is involved primarily in mediating behavioral responses to acute changes in ambient light such as the pupillary light reflex, the optokinetic reflex, and temporary changes to the circadian rhythm. In addition to the pretectum's role in the visual system, the anterior pretectal nucleus has been found to mediate somatosensory and nociceptive information.

<span class="mw-page-title-main">Reticular formation</span> Spinal trigeminal nucleus

The reticular formation is a set of interconnected nuclei that are located throughout the brainstem. It is not anatomically well defined, because it includes neurons located in different parts of the brain. The neurons of the reticular formation make up a complex set of networks in the core of the brainstem that extend from the upper part of the midbrain to the lower part of the medulla oblongata. The reticular formation includes ascending pathways to the cortex in the ascending reticular activating system (ARAS) and descending pathways to the spinal cord via the reticulospinal tracts.

<span class="mw-page-title-main">Dentate nucleus</span> Nucleus in the centre of each cerebellar hemisphere

The dentate nucleus is a cluster of neurons, or nerve cells, in the central nervous system that has a dentate – tooth-like or serrated – edge. It is located within the deep white matter of each cerebellar hemisphere, and it is the largest single structure linking the cerebellum to the rest of the brain. It is the largest and most lateral, or farthest from the midline, of the four pairs of deep cerebellar nuclei, the others being the globose and emboliform nuclei, which together are referred to as the interposed nucleus, and the fastigial nucleus. The dentate nucleus is responsible for the planning, initiation and control of voluntary movements. The dorsal region of the dentate nucleus contains output channels involved in motor function, which is the movement of skeletal muscle, while the ventral region contains output channels involved in nonmotor function, such as conscious thought and visuospatial function.

<span class="mw-page-title-main">Medial geniculate nucleus</span>

The medial geniculate nucleus (MGN) or medial geniculate body (MGB) is part of the auditory thalamus and represents the thalamic relay between the inferior colliculus (IC) and the auditory cortex (AC). It is made up of a number of sub-nuclei that are distinguished by their neuronal morphology and density, by their afferent and efferent connections, and by the coding properties of their neurons. It is thought that the MGN influences the direction and maintenance of attention.

The zona incerta (ZI) is a horizontally elongated region of gray matter in the subthalamus below the thalamus. Its connections project extensively over the brain from the cerebral cortex down into the spinal cord.

<span class="mw-page-title-main">Cochlear nucleus</span> Two cranial nerve nuclei of the human brainstem

The cochlear nuclear (CN) complex comprises two cranial nerve nuclei in the human brainstem, the ventral cochlear nucleus (VCN) and the dorsal cochlear nucleus (DCN). The ventral cochlear nucleus is unlayered whereas the dorsal cochlear nucleus is layered. Auditory nerve fibers, fibers that travel through the auditory nerve carry information from the inner ear, the cochlea, on the same side of the head, to the nerve root in the ventral cochlear nucleus. At the nerve root the fibers branch to innervate the ventral cochlear nucleus and the deep layer of the dorsal cochlear nucleus. All acoustic information thus enters the brain through the cochlear nuclei, where the processing of acoustic information begins. The outputs from the cochlear nuclei are received in higher regions of the auditory brainstem.

<span class="mw-page-title-main">Primate basal ganglia</span>

The basal ganglia form a major brain system in all species of vertebrates, but in primates there are special features that justify a separate consideration. As in other vertebrates, the primate basal ganglia can be divided into striatal, pallidal, nigral, and subthalamic components. In primates, however, there are two pallidal subdivisions called the external globus pallidus (GPe) and internal globus pallidus (GPi). Also in primates, the dorsal striatum is divided by a large tract called the internal capsule into two masses named the caudate nucleus and the putamen—in most other species no such division exists, and only the striatum as a whole is recognized. Beyond this, there is a complex circuitry of connections between the striatum and cortex that is specific to primates. This complexity reflects the difference in functioning of different cortical areas in the primate brain.

A topographic map is the ordered projection of a sensory surface, like the retina or the skin, or an effector system, like the musculature, to one or more structures of the central nervous system. Topographic maps can be found in all sensory systems and in many motor systems.

The isothalamus is a division used by some researchers in describing the thalamus.

Recurrent thalamo-cortical resonance is an observed phenomenon of oscillatory neural activity between the thalamus and various cortical regions of the brain. It is proposed by Rodolfo Llinas and others as a theory for the integration of sensory information into the whole of perception in the brain. Thalamocortical oscillation is proposed to be a mechanism of synchronization between different cortical regions of the brain, a process known as temporal binding. This is possible through the existence of thalamocortical networks, groupings of thalamic and cortical cells that exhibit oscillatory properties.

The cerebellothalamic tract or the tractus cerebellothalamicus, is part of the superior cerebellar peduncle. It originates in the cerebellar nuclei, crosses completely in the decussation of the superior cerebellar peduncle, bypasses the red nucleus, and terminates in posterior division of ventral lateral nucleus of thalamus. The ventrolateral nucleus has different divisions and distinct connections, mostly with frontal and parietal lobes. The primary motor cortex and premotor cortex get information from the ventrolateral nucleus projections originating in the interposed nucleus and dentate nuclei. Other dentate nucleus projections via thalamic pathway transmit information to prefrontal cortex and posterior parietal cortex. The cerebellum sends thalamocortical projections and in addition may also send connections from the thalamus to association areas serving cognitive and affective functions.

<span class="mw-page-title-main">Medial pulvinar nucleus</span>

Medial pulvinar nucleus is one of four traditionally anatomically distinguished nuclei of the pulvinar of the thalamus. The other three nuclei of the pulvinar are called lateral, inferior and anterior pulvinar nuclei.

References

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  21. 1 2 Arslan O. neuroanatomical basis of clinical neurology.
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  25. ↑ Zhou; et al. (2011). "specific and nonspecific thalamocortical functional connectivity in normal and vegetative states". Consciousness and Cognition. 20 (2): 257–268. doi:10.1016/j.concog.2010.08.003. PMC   3056940 . PMID   21078562.
  26. ↑ Freeman W.; Watts J.K. (1942). "Prefrontal lobotomy; the surgical relief of mental pain". Bulletin of the New York Academy of Medicine. 18 (12): 794–812. PMC   1933933 . PMID   19312295.
  27. ↑ Ward L. M. (2011). "the thalamic dynamic core theory of conscious experience". Consciousness and Cognition. 20 (2): 464–486. doi:10.1016/j.concog.2011.01.007. PMID   21349742. S2CID   6099949.
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