Human Motor Thalamus Reconstructed in 3D from Continuous Sagittal Sections with Identified Subcortical Afferent Territories.

Classification and delineation of the motor-related nuclei in the human thalamus have been the focus of numerous discussions for a long time. Difficulties in finding consensus have for the most part been caused by paucity of direct experimental data on connections of individual nuclear entities. Kultas-Ilinsky et al. (2011) showed that distribution of glutamic acid decarboxylase isoform 65 (GAD65), the enzyme that synthesizes inhibitory neurotransmitter γ-aminobutyric acid, is a reliable marker that allows to delineate connectionally distinct nuclei in the human motor thalamus, namely the territories innervated by nigral, pallidal, and cerebellar afferents. We compared those immunocytochemical staining patterns with underlying cytoarchitecture and used the latter to outline the three afferent territories in a continuous series of sagittal Nissl-stained sections of the human thalamus. The 3D volume reconstructed from the outlines was placed in the Talairach stereotactic coordinate system relative to the intercommissural line and sectioned in three stereotactic planes to produce color-coded nuclear maps. This 3D coordinate-based atlas was coregistered to the Montreal Neurological Institute (MNI-152) space. The current report proposes a simplified nomenclature of the motor-related thalamic nuclei, presents images of selected histological sections and stereotactic maps illustrating topographic relationships of these nuclei as well as their relationship with adjacent somatosensory afferent region. The data are useful in different applications such as functional MRI and diffusion tractography. The 3D dataset is publicly available under an open license and can also be applicable in clinical interventions in the thalamus.

A multicontrast approach for comprehensive imaging of substantia nigra.

We characterize the contrast behavior of substantia nigra (SN) in both magnetization transfer (MT) imaging, which is believed to be sensitive to neuromelanin (NM), and susceptibility weighted imaging (SWI). Images were acquired with a MT prepared dual echo gradient echo sequence. The first echo was taken as the MT contrast image and the second was used to generate the SWI image. SN volumes were segmented from these two types of images using a thresholding method. The spatial and signal characteristics of the extracted SWI and MT volumes were compared. Both images showed the presence of SN but the volumes of the SN identified in the two are spatially incongruent. The MT volume was more caudal than the SWI volume and with only a 12% overlap between the two volumes. Considering the SN volumes in each hemisphere separately, the average distances between the centers of mass of the volumes from the two types images are 5.1±1.1mm and 4.1±1.2mm, respectively. The frequency offsets (homodyne filtered phase/echo time) for the volumes derived from MT (NM) images and SWI images are 0.09±0.32radians/s and -1.12±0.57radians/s (p<0.0001), respectively. The MT contrasts for the two volumes are 0.16±0.02 and 0.10±0.03 (p<0.001), respectively. Our results indicate that the two contrasts are sensitive to different portions of the SN, with MT seeing the more caudal portion of the SN than SWI, likely due to variations of NM and iron content in the SN. Despite the small overlap, these regions are complementary. Our results provide a new understanding of the contrast behavior of the SN in the two imaging approaches commonly used to image it and indicate that using both may yield a more comprehensive visualization of the SN.

Can the chronic administration of the combination of buprenorphine and naloxone block dopaminergic activity causing anti-reward and relapse potential?

Opiate addiction is associated with many adverse health and social harms, fatal overdose, infectious disease transmission, elevated health care costs, public disorder, and crime. Although community-based addiction treatment programs continue to reduce the harms of opiate addiction with narcotic substitution therapy such as methadone maintenance, there remains a need to find a substance that not only blocks opiate-type receptors (mu, delta, etc.) but also provides agonistic activity; hence, the impetus arose for the development of a combination of narcotic antagonism and mu receptor agonist therapy. After three decades of extensive research, the federal Drug Abuse Treatment Act 2000 (DATA) opened a window of opportunity for patients with addiction disorders by providing increased access to options for treatment. DATA allows physicians who complete a brief specialty-training course to become certified to prescribe buprenorphine and buprenorphine/naloxone (Subutex, Suboxone) for treatment of patients with opioid dependence. Clinical studies indicate that buprenorphine maintenance is as effective as methadone maintenance in retaining patients in substance abuse treatment and in reducing illicit opioid use. With that stated, we must consider the long-term benefits or potential toxicity attributed to Subutex or Suboxone. We describe a mechanism whereby chronic blockade of opiate receptors, in spite of only partial opiate agonist action, may ultimately block dopaminergic activity causing anti-reward and relapse potential. While the direct comparison is not as yet available, toxicity to buprenorphine can be found in the scientific literature. In considering our cautionary note in this commentary, we are cognizant that, to date, this is what we have available, and until such a time when the real magic bullet is discovered, we will have to endure. However, more than anything else this commentary should at least encourage the development of thoughtful new strategies to target the specific brain regions responsible for relapse prevention.

Basal ganglia and thalamic input from neurons located within the ventral tier cell cluster region of the substantia nigra pars compacta in the rat.

The most caudally located dopaminergic (DA) ventral tier neurons of the substantia nigra pars compacta (SNc) form typical cell clusters that are deeply embedded in the substantia nigra pars reticulata (SNr). Here we examine the efferent projections of 35 neurons located in the SNr region where these SNc cell clusters reside. The neuronal cell body was injected with biotinylated dextran amine so as to trace each complete axon in the sagittal or the coronal plane. Electrophysiological guidance guaranteed that the tracer was ejected among neurons displaying a typical SNc discharge pattern. Furthermore, double immunofluorescence and immunohistochemical labeling ensured that the tracer deposits were placed within the DA cell clusters. Three types of projection neurons occurred in the SNc ventral tier cell cluster region: type I neurons, projecting to basal ganglia; type II neurons, targeting both the basal ganglia and thalamus; and type III neurons, projecting only to the thalamus. The striatum was targeted by most of the type I and II neurons and the innervation reached both the striosome/subcallosal streak and matrix compartments. Many nigrostriatal fibers provided collaterals to the globus pallidus and, less frequently, to the subthalamic nucleus. At a thalamic level, type II and III neurons preferentially targeted the reticular, ventral posterolateral, and ventral medial nuclei. Our results reveal that the SNr region where DA ventral tier cell clusters reside harbors neurons projecting to the basal ganglia and/or the thalamus, thus suggesting that neurodegeneration of nigral neurons in Parkinson's disease might affect various extrastriatal basal ganglia structures and multiple thalamic nuclei.

Overlap of nigrothalamic terminals and thalamostriatal neurons in the feline lateralis medialis-suprageniculate nucleus.

The lateralis medialis-suprageniculate nucleus (LM-Sg) of the feline posterior thalamus is a relay nucleus with a clear visuomotor function. In this study, we examined the distribution of axon terminals of the nigral afferent to the LM-Sg following injection of an anterograde tracer, biocytin, into the substantia nigra pars reticulata, and the distribution of the thalamostriatal projection neurons in the LM-Sg following the injection of wheat germ agglutinin conjugated with horseradish peroxidase (WGA-HRP) as a retrograde tracer into the caudate nucleus. The biocytin-labeled terminal-like puncta were located in the ventromedial portion of this nucleus in such a way that most of the labeled elements took the form of swellings having boutons in places, while a minority appeared in clusters of 3-5 large terminal-like puncta. The retrograde WGA-HRP-labeled neurons were also found in the ventromedial part of the LM-Sg, and the distributions of labeled nigrothalamic axon terminals and labeled thalamostriatal projection neurons therefore overlapped in this region. The present results indicate that the nigral afferent may make synaptic contacts directly with the thalamostriatal projection neurons within the LM-Sg.

Improved segmentation of deep brain grey matter structures using magnetization transfer (MT) parameter maps.

Basal ganglia and brain stem nuclei are involved in the pathophysiology of various neurological and neuropsychiatric disorders. Currently available structural T1-weighted (T1w) magnetic resonance images do not provide sufficient contrast for reliable automated segmentation of various subcortical grey matter structures. We use a novel, semi-quantitative magnetization transfer (MT) imaging protocol that overcomes limitations in T1w images, which are mainly due to their sensitivity to the high iron content in subcortical grey matter. We demonstrate improved automated segmentation of putamen, pallidum, pulvinar and substantia nigra using MT images. A comparison with segmentation of high-quality T1w images was performed in 49 healthy subjects. Our results show that MT maps are highly suitable for automated segmentation, and so for multi-subject morphometric studies with a focus on subcortical structures.

The cortico-basal ganglia integrative network: the role of the thalamus.

The thalamus is a critical component of the frontal cortical-basal ganglia-thalamic circuits that mediate motivation and emotional drive, planning and cognition for the development and expression of goal-directed behaviors. Each functional region of the frontal cortex is connected with specific areas of each basal ganglia (BG) structure and of the thalamus. In addition, the thalamus sends a massive, topographically organized projection directly to the striatum. Tract-tracing and physiological experiments have indicated a general topographic organization of the cortical-BG-thalamic loops and supported a model of BG function based on parallel and segregated pathways. However, the learning and execution of appropriate behavioral responses require integration of inputs related to emotional, cognitive, and motor cortical functions. Our recent data indicate that integration may occur via non-reciprocal connections between the striatum and substantia nigra and within "hot spots" of convergence between corticostriatal projections from different functional regions. Similarly, integration may exist in the thalamus. There are non-reciprocal connections between the thalamus and cortex via thalamocortical projections that terminate in the superficial and deep cortical layers. These terminals can influence different functional cortical areas that, in turn, project to the striatum and back to the thalamus. In addition, a non-reciprocal corticothalamic projection terminates in thalamic regions that are parts of other circuits. Finally, 'hot spots' of convergence between terminals from different cortical regions may also occur in the thalamus as is seen in the striatum. Thus, via several different pathways, the thalamus may serve as an important center of integration of networks that underlie the ability to modulate behaviors.

Somatotopy in the basal ganglia: experimental and clinical evidence for segregated sensorimotor channels.

Growing experimental and clinical evidence supports the notion that the cortico-basal ganglia-thalamo-cortical loops proceed along parallel circuits linking cortical and subcortical regions subserving the processing of sensorimotor, associative and affective tasks. In particular, there is evidence that a strict topographic segregation is maintained during the processing of sensorimotor information flowing from cortical motor areas to the sensorimotor areas of the basal ganglia. The output from the basal ganglia to the motor thalamus, which projects back to neocortical motor areas, is also organized into topographically segregated channels. This high degree of topographic segregation is demonstrated by the presence of a well-defined somatotopic organization in the sensorimotor areas of the basal ganglia. The presence of body maps in the basal ganglia has become clinically relevant with the increasing use of surgical procedures, such as lesioning or deep brain stimulation, which are selectively aimed at restricted subcortical targets in the sensorimotor loop such as the subthalamic nucleus (STN) or the globus pallidus pars interna (GPi). The ability to ameliorate the motor control dysfunction without producing side effects related to interference with non-motor circuits subserving associative or affective processing requires the ability to target subcortical areas particularly involved in sensorimotor processing (currently achieved only by careful intraoperative microelectrode mapping). The goal of this article is to review current knowledge about the somatotopic segregation of basal ganglia sensorimotor areas and outline in detail what is known about their body maps.

Peroperative transcranial sonography for electrode placement into the targeted subthalamic nucleus of patients with Parkinson disease: technical note.

BACKGROUND: In the search for a better preoperative knowledge of the position of probes and electrodes, we assessed the feasibility and the usefulness of transcranial sonography during surgery for the implantation of stimulation electrodes into the subthalamic nucleus (STN) of patients with Parkinson's disease. METHODS: Transcranial sonography was carried out during stereotactic surgery in 8 patients with Parkinson's disease who had a suitable temporal bone window on the side receiving the electrode. Test stimulation parameters were 130 Hz, 0.1 ms, up to 0 to 4.5 V. RESULTS: The test probe with a diameter of 0.8 mm was visualized through the temporal preauricular window. The correct anatomic position of the electrode tip could be indirectly assessed thanks to the topographic relationship of the STN with the hyperechogenic substantia nigra and the nucleus ruber. The tip position of the final electrode was easily documented. A laterality of 10.5 to 11.5 mm, verified by teleradiographic ventriculography and plain films, was correlated with the best response of symptoms of Parkinson's disease to electrical impulses delivered to the STN. CONCLUSIONS: Transcranial sonography is easily feasible during stereotactic surgery. In combination with the clinical effects of electrostimulation on the symptoms of Parkinson's disease and with stereotactic x-ray images, it enables the assessment and the documentation of the correct position of implanted STN electrodes in real time.

The basal ganglia: anatomy, physiology, and pharmacology.

The basal ganglia are perceived as important nodes in cortico-subcortical networks involved in the transfer, convergence, and processing of information in motor, cognitive, and limbic domains. How this integration might occur remains a matter of some debate, particularly given the consistent finding in anatomic and physiologic studies of functional segregation in cortico-subcortical loops. More recent theories, however, have raised the notion that modality-specific information might be integrated not spatially, but rather temporally, by coincident processing in discrete neuronal populations. Basal ganglia neurotransmitters, given their diverse roles in motor performance, learning, working memory, and reward-related activity are also likely to play an important role in the integration of cerebral activity. Further work will elucidate this to a greater extent, but for now, it is clear that the basal ganglia form an important nexus in the binding of cognitive, limbic, and motor information into thought and action.

The basal ganglia and motor control.

This paper briefly reviews the functional anatomy of the basal ganglia and their relationships with the thalamocortical system. The basal ganglia, including the striatum, pallidum, subthalamic nucleus, and substantia nigra, are involved in a number of parallel, functionally segregated cortical-subcortical circuits. These circuits support a wide range of sensorimotor, cognitive and emotional-motivational brain functions. A main role of the basal ganglia is the learning and selection of the most appropriate motor or behavioral programs. The internal functional organization of the basal ganglia is very well suited for such selection mechanisms, both in development and in adulthood. The question of whether clumsiness may be, at least in part, attributed to dysfunction of the basal ganglia is discussed in the context of the differential, complementary, or interactive roles of the basal ganglia and the cerebellum in the development of motor control.

[Morphological analysis of information processing in basal ganglia of mammals]. / Morfologicheskii analiz putei provedeniia informatsii v bazal'nykh gangliiakh mlekopitaiushchikh.

The article reviews published and own data on special features of organization of afferent cortical and infracortical correctional systems of the total striopallidaric nuclear complex in various species of mammals. The article focuses on analysis of organization of these flows in a dog being a classical object of behavioural experiments and a rare object of neuromorphological studies. It was discovered that the general trend towards segregated and converged flows of functionally different information at the level of striopallidum is retained in all tested mammals, being an evidence of the fundamental nature of the discussed principles of organization of the projection systems of basal ganglia. The reviewed morphological data confirm reliability of the presented concept of basal ganglia functioning.

Enkephalinergic involvement in substantia nigra in the modulation of hypothalamically-induced predatory attack behavior.

The present study was carried out in five cats which did not attack the rats spontaneously. Predatory attack on an anaesthetized rat was elicited by electrical stimulation of lateral hypothalamus at a mean current strength of 650 microA. The attack was accompanied by minimal affective display and culminated in neck biting. Microinfusions of DAME (delta-alanine methionine enkephaline) in 500 ng dose in substantia nigra facilitated the predatory attack and there was a significant reduction in the threshold current strength for affective display as well as somatomotor components. Microinfusions of naloxone, an opioid antagonist in 1.0 microg dose when DAME effect was at its peak reversed the facilitatory effects and the threshold returned to the control levels within 10 minutes of naloxone infusion at the same locus. Microinfusions of naloxone alone in similar dosage completely blocked the predatory attack response as indicated by an increase in the threshold current strength for somatomotor as well as affective display components. The somatomotor were completely inhibited and could not be elicited even when the current strength was increased to 1000 microA. Control injections of saline in similar volumes (0.5 microl) failed to produce any response Microinfusions of naloxone in lower dose (250 ng) failed to produce any blocking effect. These findings indicate that hypothalamically elicited predatory attack is facilitated by enkephalinergic mechanisms operating at the midbrain level.

Physiologic study of the subthalamic volume.

Deep brain stimulation (DBS) obtains good control of advanced PD symptoms. Chronic stimulation of Stn may alleviate rigidity, dyskinesia and tremor. Anatomical and functional intraoperative mapping are mandatory to obtain careful target localisation. Per-operative macrostimulation was carried out in 22 patients undergoing bilateral DBS in Stn; a volume 6 mm above to 4 mm below Stn was explored. Positive, collateral and adverse effects were recorded every 2 mm. Results obtained during acute stimulation were correlated to anatomical data from stereotactic atlases. Our findings suggest a volume, encompassing the zona incerta, Forel's fields and the lowermost part of anterior thalamus, functionally homogeneous to Stn. In fact, the stimulation of this volume obtains reduction of PD symptoms comparable to Stn.

Contrast optimization of Macaca mulatta basal ganglia in magnetic resonance images at 4.7 Tesla.

To determine whether high field MRI could distinguish among the different regions of the basal ganglia, the brains of two Macaca mulatta monkeys were explored in vivo using a 4.7 T MR imager. Gradient-echo (GE) and spin-echo images were acquired with proton-density, T1 and T2* weightings. Five GE images with increased susceptibility effects were generated using a GESFID sequence, from which T2* maps were also reconstructed. The first echo of the GESFID sequence (TE = 12.6 ms) produced the best contrast-to-noise ratio (C/N) between the pallidum and the putamen, the pallidum and the thalamus, the substantia nigra and the surrounding white matter, and the substantia nigra and the subthalamic nucleus. An increased T2*-weighting (TE = 37.2 ms) was necessary to maximize C/N between the putamen and the surrounding white matter, and between the subthalamic nucleus and the surrounding white matter. A dual GE sequence with a short TE ( approximately 10 ms) and a longer one ( approximately 30 ms) thus effectively localizes basal ganglia subregions at 4.7 T.

Compartmental organization and chemical profile of dopaminergic and GABAergic neurons in the substantia nigra of the rat.

The substantia nigra (SN) is a midbrain center composed of dopaminergic (DA-) and gamma aminobutyric acid (GABA)ergic (GABA-) neurons. In this study, we investigated the topographical relationship between both cell populations and their chemical profile by using single and double immunostaining for tyrosine hydroxylase (TH), glutamic acid decarboxylase (GAD), cholecystokinin (CCK), calretinin (CR), calbindin (CB), parvalbumin (PV), and nitric oxide synthase (NOS). Our results showed that DA-cells are arranged in two bands, one rostrodorsal that corresponds to the SN pars compacta (SNC), and another caudoventral that corresponds to the SN pars reticulata (SNR) and emits cell bridges that make contact with the rostrodorsal one. In the SNR, GABA-cells are arranged in dorsoventrally elongated clusters that occupy DA-cell free regions. According to cytoarchitectural, topographical, and chemical criteria, we identified ten different cell groups: five dopaminergic ones, and five GABAergic ones. Within DA-cells, we found a cell group in the dorsomedial portion of the SNC which contains CCK, CR, and CB (dmSNC); DA-cells in the SN pars lateralis (SNL) which also contain CCK, CR and CB; DA-cells in the rostral half of the SNC containing CCK and CR (rSNC); DA-cells in the SNR and the caudal half of the SNC which only express CR (cSNC-SNR), and a DA-cell group in the lateral part of the SNC that contains none of the markers studied (lSNC). Within GABA-cells, we distinguished: large GABA-cells in the SNL that contain PV; large GABA-cells in the rostrolateral part of the SNR containing PV and NOS (rlSNR), small GABA-cells in the caudomedial part of the SNR containing PV (cmSNR), and two groups of small GABA-cells in the rostromedial portion of the SNR, one of them containing CR (rmcSNR), and the other containing NOS (rmnSNR). These data suggest that over a compartmental and complementary organization, DA- and GABA-nigral cells form a mosaic of neurochemically different subnuclei which probably differ in their physiological and pharmacological properties and vulnerability to aggression.

Integration and segregation of limbic cortico-striatal loops at the thalamic level: an experimental tracing study in rats.

The frontal lobe and the basal ganglia are involved in a number of parallel, functionally segregated circuits. Information is thought to pass from distinct parts of the (pre)frontal cortex, via the striatum, the pallidum/substantia nigra and the thalamus, back to the premotor/prefrontal cortices. Currently, different views exist as to whether these circuits are to be considered as open or closed loops, as well as to the degree of interconnection between different circuits. The main goal of the present study is to answer some of these questions for the limbic corticostriatal circuits. The latter circuits involve the nucleus accumbens, the ventral pallidum/dorsomedial substantia nigra pars reticulata, the medial parts of the mediodorsal and ventromedial thalamic nuclei and the prefrontal cortex. Within the nucleus accumbens, a core and a shell region are recognized on the basis of anatomical and functional criteria. The shell of the nucleus accumbens projects predominantly to the mediodorsal, the midline and the reticular thalamic nuclei via the ventral pallidum, whereas the core reaches primarily the medial part of the ventromedial thalamic nucleus, the intralaminar and mediodorsal thalamic nuclei via a relay in the dorsomedial substantia nigra pars reticulata. By means of double labeling experiments with injections of anterograde tracers in both the ventral pallidum and the substantia nigra of rats, we were able to demonstrate that circuits involving the shell and the core of the nucleus accumbens remain largely segregated at the level of the thalamus. Only restricted areas of overlap of ventral pallidal and reticular nigral projections occur in the mediodorsal and ventromedial thalamic nuclei, which allows for a limited degree of integration, at the thalamic level, of information passing through the two circuits.

MR imaging of human brain at 3.0 T: preliminary report on transverse relaxation rates and relation to estimated iron content.

PURPOSE: To determine the transverse relaxation rates R2 and R2' from several gray matter regions and from frontal cortical white matter in healthy human brains in vivo and to determine the relationship between relaxation rates and iron concentration [Fe]. MATERIALS AND METHODS: Six healthy adults aged 19-42 years underwent thin-section gradient-echo sampling of free induction decay and echo magnetic resonance (MR) imaging at 3.0 T. Imaging covered the mesencephalon and basal ganglia. RESULTS: Relaxation rates (mean +/- SD) were highest in globus pallidus (R2 = 25.8 seconds-1 +/- 1.1, R2' = 12.0 seconds-1 +/- 2.1) and lowest in prefrontal cortex (R2 = 14.4 seconds-1 +/- 1.8, R2' = 3.4 seconds-1 +/- 1.1). Frontal white matter measurements were as follows: R2 = 18.0 seconds-1 +/- 1.2 and R2' = 3.9 seconds-1 +/- 1.2. For gray matter, both R2 and R2' showed a strong correlation (r = 0.92, P < .001 and r = 0.90, P < .001, respectively) with [Fe]. Although the slopes of the regression lines for R2' versus [Fe] and for R2 versus [Fe] were similar, the iron-independent component of R2' (2.2 seconds-1 +/- 0.6), the value when [Fe] = 0, was much less than that of R2 (12.7 seconds-1 +/- 0.7). CONCLUSION: The small iron-independent component R2', as compared with that of R2, is consistent with the hypothesis that R2' has higher iron-related specificity.

The distribution of neurons in the substantia nigra pars reticulata with input from the motor, premotor and prefrontal areas of the cerebral cortex in monkeys.

We investigated the distribution of neurons in the substantia nigra pars reticulata (SNr) which received cortical input. The activities of single SNr neurons were studied extracellulary in awake monkeys. SNr neurons showed excitatory and/or inhibitory responses to cortical stimulation. These responses were considered to be mediated by the subthalamic nucleus and striatum, respectively. The neurons receiving inhibitory input from the motor, premotor and supplementary motor areas (Motor-related cortical areas) were located in the lateral part of the SNr, whereas those with input from the medial, dorsal and orbital areas of the prefrontal cortex (PFmdo) were frequently found in the rostro-medial part of this nucleus. SNr neurons with inhibitory input from the ventral periprincipal area (PSv) were mainly distributed in the intermedio-lateral portion, with some degree of overlap with input from other cortical areas. The distribution of the excitatory input was almost similar to that of inhibitory one, but the excitatory input from the PSv was much stronger than that from the PFmdo. Some SNr neurons receiving cortical input were proved to project to the thalamus. Our results support the existence of several parallel organization of the cortico-basal ganglia loop circuits [G.E. Alexander, M.R. DeLong, P.L. Strick, Parallel organization of functionally segregated circuits linking basal ganglia and cortex, Ann. Rev. Neurosci., 9, 1986, pp. 357-381.], but interaction between the loops can not be ignored.

Overlapping territories between the thalamostriatal and nigrothalamic projections in cats.

Fluorescent tracers were injected into different regions of the caudate nucleus and HRP-WGA in the substantia nigra of the cat in order to analyse the thalamic distribution of retrogradely labelled thalamostriatal neurones and anterogradely labelled nigrothalamic terminals within the thalamus. Overlapping thalamic territories between the thalamostriatal neurones projecting to areas of the caudate nucleus and the nigrothalamic connections were observed in the rostral nuclei of the central thalamic group (ventral anterior nucleus, ventral anterior-ventral lateral complex and ventral medial nucleus) and, more restricted, in the rostral (rhomboid, paracentral, ventral lateral, dorsal mediodorsal nuclei) and caudal intralaminar nuclei (centromedian-parafascicular complex). This study provides evidence of the existence of thalamic areas in which the input and output of the basal ganglia converge.