Prosencephalic afferents to the mediodorsal thalamic nucleus

The afferent projections from the prosencephalon to the mediodorsal thalamic nucleus (MD) were studied in the cat by use of the method of retrograde transport of horseradish peroxidase (HRP). Cortical and subcortical prosencephalic structures project bilaterally to the MD. The cortical afferents originate mainly in the ipsilateral prefrontal cortex. The premotor, prelimbic, anterior limbic, and insular agranular cortical areas are also origins of consistent projections to the MD. The motor cortex, insular granular area, and some other cortical association areas may be the source of cortical connections to the MD. The subcortical projections originate principally in the ipsilateral rostral part of the reticular thalamic nucleus and the rostral lateral hypothalamic area. Other parts of the hypothalamus, the most caudal parts of the thalamic reticular nucleus, the basal prosencephalic structures, the zona incerta, the claustrum, and the entopeduncular and subthalamic nuclei are also sources of projections to the MD. Distinct, but somewhat overlapping areas of the prosencephalon project to the three vertical subdivisions of MD (medial, intermediate, and lateral). The medial band of the MD receives a small number of prosencephalic projections; these arise mainly in the caudal and ventral parts of the prefrontal cortex. Cortical projections also arise in the infralimbic area, while subcortical projections originate in the medial part of the rostral reticular thalamic nucleus and lateral hypothalamic area. The intermediate band of the MD receives the largest number of fibers from the prosencephalon. These arise principally in the intermediate and dorsal part of the lateral and medial surface of the prefrontal cortex, the premotor cortex, and the prelimbic and agranular insular areas. Projections also originate in basal prosencephalic formations (preoptic area, Broca's diagonal band, substantia innominata, and olfactory tubercle), rostral reticular thalamic nucleus, and lateral hypothalamic area. A large number of prosencephalic structures also project to the lateral band of the MD. These are mainly the most dorsal and caudal parts of the lateral and medial surface of the prefrontal cortex, the premotor and motor cortices, and the prelimbic, anterior limbic, and insular areas. Projections arise also in the lateral rostral and caudal parts of the reticular thalamic nucleus, the zona incerta, the lateral and dorsal hypothalamic areas, the claustrum, and the entopeduncular nucleus. These and previous results demonstrate a gradation in the afferent connections to the three subdivisions of the MD. Brain structures related to the olfactory sensory modality and with allocortical formations of the limbic system project principally to the medial band of the MD. The intermediate band of the MD receives subcortical and cortical projections from structures mainly related to the limbic system and cortical regions related to sensory association cortices. The lateral band of the MD receives projections mainly originating in structures related to complex sensory associative processes and to the motor system (especially from brainstem and cortical structures implicated in the regulation of eye movements).     

Topographic organization of projections from the thalamic reticular nucleus to the anterior thalamic nuclei in the rat

We have investigated connections between the thalamic reticular nucleus (TRN) and the anterior thalamic nuclei (ATN) in the rat, following injections of horseradish peroxidase (HRP) into subnuclei of the ATN and different regions of the rostral TRN. Three nonoverlapping groups of neurons in the dorsal part of the ipsilateral rostral TRN project to, and receive reciprocal projections from, specific subnuclei of the ATN. A vertical sheet of neurons in the most dorsal part of the rostral TRN projects to the dorsal half of the posterior subdivision of the anteroventral thalamic nucleus (AVp), the dorsal region of the medial subdivision of the anteroventral thalamic nucleus (AVm), and the dorsolateral part of the rostral anterodorsal thalamic nucleus (AD). Immediately ventral to this part of TRN, but still within its dorsal portion, are a lateral cluster of neurons and a medially located vertical sheet of neurons. The lateral cluster projects to the ventral part of AVp and to the dorsomedial part of rostral AD. The medial sheet projects to the ventral part of AVm, the ventral part of rostral AD, and to the caudal portions of both AV and AD. There appears to be no input to the anteromedial thalamic nucleus (AM) from the TRN. These findings shed new light on the anatomy of the rostral TRN, the ATN, and the connections between the two, and are relevant to emerging hypotheses about the functional organization of the TRN and reticulo-thalamic projections.     

Organization of corticothalamic projections from parietal cortex in cat

Corticothalamic projections from areas 5a, 5b, and 7 of cat parietal cortex were studied with autoradiographic techniques. Each cortical area was identified by its cytoarchitectural characteristics and the patterns of termination were related to the thalamic nuclear groups. Injections of 3H-leucine in cortical area 5a were associated with terminal labeling primarily in the spinal recipient zone of the ventral lateral nucleus (VLsp) and the medial division of the posterior group (POm). The corticothalamic projections of area 5a are loosely topographically organized; medial parts of 5a project heavily to rostral and lateral parts of VLsp and sparsely to POm, while lateral parts of 5a project to more medial and caudal parts of VLsp and heavily to POm. Cortical area 5b projects primarily to the rostral portions of the lateral posterior nucleus (LP). These projections also appear to be topographically organized. The part of area 5b on the marginal gyrus projects to more ventral parts of rostral LP, while area 5b on the middle suprasylvian gyrus projects to more dorsal and lateral parts of rostral LP. Cortical area 7 projects to LP and the pulvinar (Pul). Rostral parts of area 7 project heavily to dorsal and lateral parts of LP and lightly to Pul; more caudal portions of area 7 projects relatively more heavily to Pul. The reticular, central lateral, and paracentral nuclei also receive projections, especially from the suprasylvian gyrus. The results are discussed with regard to putative sensory response characteristics of these cortical areas and to general thalamocortical organization.     

Diverse thalamic projections to the prefrontal cortex in the rhesus monkey.

We studied the sources of thalamic projections to prefrontal areas of nine rhesus monkeys with the aid of retrograde tracers (horseradish peroxidase or fluorescent dyes). Our goal was to determine the proportion of labeled neurons contributing to this projection system by the mediodorsal (MD) nucleus compared to those distributed in other thalamic nuclei, and to investigate the relationship of thalamic projections to specific architectonic areas of the prefrontal cortex. We selected areas for study within both the basoventral (areas 11, 12, and ventral 46) and the mediodorsal (areas 32, 14, 46, and 8) prefrontal sectors. This choice was based on our previous studies, which indicate differences in cortical projections to these two distinct architectonic sectors (Barbas, '88; Barbas and Pandya, '89). In addition, for each sector we included areas with different architectonic profiles, which is also relevant to the connectional patterns of the prefrontal cortices. The results showed that MD included a clear majority (over 80%) of all thalamic neurons directed to some prefrontal cortices (areas 11, 46, and 8); it contributed just over half to some others (areas 12 and 32), and less than a third to area 14. Clusters of neurons directed to basoventral and mediodorsal prefrontal areas were largely segregated within MD: the former were found ventrally, the latter dorsally. However, the most striking findings establish a relationship between thalamic origin and laminar definition of the prefrontal target areas. Most thalamic neurons directed to lateral prefrontal cortices, which are characterized by a high degree of laminar definition (areas 46 and 8), originated in the parvicellular and multiform subdivisions of MD, and only a few were found in other nuclei. In contrast, orbital and medial cortices, which have a low degree of laminar differentiation, were targeted by the magnocellular subdivision of MD and by numerous other limbic thalamic nuclei, including the midline and the anterior. Thus topographic specificity in the origin of thalamic projections increased as the laminar definition of the target area increased. Moreover, the rostrocaudal distribution of labeled neurons in MD and the medial pulvinar also differed depending on the degree of the laminar definition of the prefrontal target areas. The rostral parts of MD and the medial pulvinar projected to the eulaminate lateral prefrontal cortices, whereas their caudal parts projected to orbital and medial limbic cortices. Selective destruction of caudal MD is known to disrupt mnemonic processes in both humans and monkeys, suggesting that this thalamic-limbic prefrontal loop may constitute an important pathway for memory.     

Visual thalamocortical projections in normal and enucleated rats: HRP and fluorescent dye studies

Visual thalamocortical projections of neonatally enucleated and control rats were studied after tracer injections into the striate and peristriate areas of adult pigmented rats. The distribution of retrogradely labeled neurons in the visual thalamic nuclei was mapped after (a) small localized injections of horseradish peroxidase into either area 17, 18, or 18a and (b) simultaneous injections of three different retrograde tracers (fast blue, HRP, and diamidino yellow) into the anterior, medial, and posterior regions of area 17. It was shown in both normal and neonatally enucleated rats, that the dorsal lateral geniculate nucleus projects to the striate cortex (area 17), whereas the laterodorsal thalamic nucleus of the lateral thalamus projects to the medial peristriate area 18, and the lateral posterior thalamic nucleus has a projection to the lateral peristriate area 18a. Additionally, both extrageniculate visual thalamic nuclei project to area 17. Neurons in the dorsoanterior region of the dorsal lateral geniculate nucleus project to the posterior part of area 17, while neurons in the ventroposterior region of the nucleus send their axons to the anterior part of area 17. A similarly inverted projection of anterior and posterior divisions of the lateral posterior thalamic nucleus to visual area 18a was detected. In enucleated rats, the general topography of the projections from the thalamic neurons to the striate and peristriate cortices was indistinguishable from that in the controls. Nonetheless, there was noticeable shrinkage of the dorsal lateral geniculate nucleus and lateral thalamus and a significant decrease in the size of the somata of projecting neurons. Mean somal area of the HRP-labeled neurons in the dorsal lateral geniculate nucleus of enucleated rats was reduced by 19.0% and the mean maximum cell diameter by 14.3% compared with controls.     

Connections of the subthalamic nucleus with ventral striatopallidal parts of the basal ganglia in the rat

The present study was undertaken to establish the precise anatomical relationship of the subthalamic nucleus (STh) with limbic lobe-afferented parts of the basal ganglia in the rat. The anterograde tracer Phaseolus vulgaris-leucoagglutinin (PHA-L), injected in the STh, the globus pallidus, the ventral pallidum, the ventral striatum, and the parafascicular thalamic nucleus, and the retrograde tracers Fluoro-Gold (FG) and cholera toxin B (CTb), injected in the globus pallidus, the ventral pallidum, the ventral striatum, and the ventral mesencephalon, were used for this purpose. The results of these tracing experiments confirm the general notion of reciprocal connections between the STh and pallidal areas. Thus the dorsomedial part of the STh is connected with the subcommisural ventral pallidum, whereas a more ventral and lateral part of the medial STh is related to the medial globus pallidus. The lateral hypothalamic area, directly adjacent to the STh, containing neurons with a morphology quite similar to those in the STh, projects to parts of the ventral pallidum related to the olfactory tubercle. The reciprocal projection from this pallidal area to subthalamic regions appears to be very sparse. The medial STh sends strong projections to the medial part of the entopeduncular nucleus and the adjacent lateral hypothalamic area. Sparser projections from the medial STh reach the rostral and medial part of the caudate-putamen and the nucleus accumbens. The nucleus accumbens sends a very sparse projection back to the medial STh. The projections of the medial STh to the ventral mesencephalon appear also to be topographically organized. The lateral hypothalamus and a few cells in the most medial part of the STh project to the ventral tegmental area, whereas progressively more lateral parts of the ventral mesencephalon, in particular the substantia nigra, receive input from successively more lateral and caudal parts of the STh. In addition, a number of STh fibers reach the midbrain extrapyramidal area. The lateral part of the parafascicular thalamic nucleus projects to the lateral part of the STh, whereas parafascicular neurons medial to the fasciculus retroflexus project to the dorsomedial portion of the STh. The medial part of the STh and the adjacent lateral hypothalamus are intimately connected with limbic parts of the basal ganglia in a way similar and parallel to the connections of the lateral STh with motor-related parts of the basal ganglia. These findings suggest a role for the STh in nonmotor functions of the basal ganglia.     

Anatomical investigation of projections from thalamus to posterior parietal cortex in the rhesus monkey: a WGA-HRP and fluorescent tracer study.

The parietothalamic projections have been shown to be heterogeneous and appear to be a reflection of the detailed architectonic parcellation of the parietal lobe. In the present study WGA-HRP injections were placed in the different subdivisions of the posterior parietal cortex of the rhesus monkey to determine whether a similarly complex pattern also exists in the thalamocortical pathway. Additionally, in an attempt to determine whether there is an intranuclear specificity of projections from individual thalamic nuclei to different subdivisions of the parietal lobe, multiple retrograde fluorescent tracers were injected into the rostral to caudal sectors of the parietal lobe of the same animal. Different subdivisions of the parietal lobe appear to receive different sets of thalamic input. Thus the superior parietal lobule (SPL) projections are derived from more lateral regions in the thalamus, arising predominantly from the lateral posterior (LP) and pulvinar oralis (PO) nuclei, with additional contributions from the pulvinar lateralis (PL) and pulvinar medialis (PM) nuclei. The inferior parietal lobule (IPL), by contrast, receives its projections from more medial thalamic regions, its main thalamic input originating from PM, and aided by LP, PL, and PO. Both the SPL and IPL also receive projections from the mediodorsal (MD), ventroposterior, ventrolateral, intralaminar, and limbic nuclei, albeit from different components within these nuclei. A topographical arrangement also exists in the thalamic projections to the rostral versus the caudal subdivisions of both the SPL and the IPL. Thus, in the SPL, the ventral posterolateral nucleus, pars oralis (VPLo), ventral lateral nucleus, pars oralis (VLo), and ventral lateral nucleus, pars medialis (VLm) project to rostral regions, whereas the PM and limbic nuclei, anteroventral (AV), and anteromedial (AM), project to area PGm on the medial convexity of the SPL. With respect to projections to the IPL, the ventral posteromedial (VPM) and PO nuclei project to rostral regions, whereas the limbic nuclei lateral dorsal (LD), AM and AV project only to the caudal most area, Opt. A rostrocaudal difference is reflected also within certain nuclei (LP, PO, and PM) that project to the SPL or IPL. Thus rostral parietal subdivisions receive projections from ventral regions within these thalamic nuclei, whereas caudal parietal afferents arise from the dorsal parts of these nuclei. Intervening cortical levels receive projections from intermediate positions within the nuclei. It therefore seems that the increasing architectonic and functional complexity as one moves from rostral to caudal in the SPL and IPL appear to be reflected in the thalamic afferents.(ABSTRACT TRUNCATED AT 400 WORDS)     

The primate mediodorsal (MD) nucleus and its projection to the frontal lobe

The frontal lobe projections of the mediodorsal (MD) nucleus of the thalamus were examined in rhesus monkey by transport of retrograde markers injected into one of nine cytoarchitectonic regions (Walker's areas 6, 8A, 9, 10, 11, 12, 13, 46, and Brodmann's area 4) located in the rostral third of the cerebrum. Each area of prefrontal, premotor, or motor cortex injected was found to receive a topographically unique thalamic input from clusters of cells in specific subdivisions within MD. All of the prefrontal areas examined also receive topographically organized inputs from other thalamic nuclei including, most prominently, the ventral anterior (VA) and medial pulvinar nuclei. Conversely, and in agreement with previous findings, MD projects to areas of the frontal lobe beyond the traditional borders of prefrontal cortex, such as the anterior cingulate and supplementary motor cortex. The topography of thalamocortical neurons revealed in coronal sections through VA, MD, and pulvinar is circumferential. In the medial part of MD, for example, thalamocortical neurons shift from a dorsal to a ventral position for cortical targets lying medial to lateral along the ventral surface of the lobe; neurons in the lateral MD move from a ventral to a dorsal position, for cortical areas situated lateral to medial on the convexity of the hemisphere. The aggregate evidence for topographic specificity is supported further by experiments in which different fluorescent dyes were placed in multiple areas of the frontal lobe in each of three cases. The results show that very few, if any, thalamic neurons project to more than one area of cortex. The widespread cortical targets of MD neurons together with evidence for multiple thalamic inputs to prefrontal areas support a revision of the classical hodological definition of prefrontal cortex as the exclusive cortical recipient of MD projections. Rather, the prefrontal cortex is defined by multiple specific relationships with the thalamus.     

Thalamic connections of the cortex of the superior temporal sulcus in the rhesus monkey

The thalamocortical connections of the superior temporal sulcus (STS) were studied by means of the WGA-HRP retrograde tracing technique. The results indicate that the distribution of thalamic projections varies along the rostral-caudal dimension of the STS. Thus the rostral portion of the upper bank receives input primarily from the medialmost portion of the medial pulvinar (PM) nucleus. The middle region of the upper bank receives projections from medial and central portions of the PM nucleus, and also from the oral pulvinar, limitans, suprageniculate, medial geniculate, and dorsomedial nuclei. The cortex of the caudal portion of the upper bank has basically similar thalamic input; however, the projections from the PM nucleus originate in central and lateral portions. Additionally, there are projections from the lateral pulvinar (PL), ventroposterolateral, central lateral, parafascicular, and paracentral nuclei. In contrast to the dorsal bank, the cortex of the ventral bank of the STS receives somewhat different and less extensive thalamic input. The rostral portion of the lower bank receives projections only from the ventromedial sector of the PM nucleus, whereas the middle portion of the lower bank receives projections from the PL and the inferior pulvinar nuclei as well as from the PM nucleus. The upper bank of the STS, on the basis of physiological and anatomical studies (Jones and Powell, '70; Seltzer and Pandya, '78; Gross et al., '81; Baylis et al., '87), has been shown to contain multimodal areas. The present data indicate that the multimodal region of the STS has a preferential relationship with the central sector of the PM nucleus.     

Origin and topography of fibers contributing to the fornix in macaque monkeys

The distribution of neurons contributing to the fornix was mapped by placing the retrograde tracer horseradish peroxidase (HRP) in polyacrylamide gels in different medial to lateral locations within the fornix of three rhesus monkeys (Macaca mulatta). The HRP was placed from 3 to 5 mm caudal to the descending columns of the fornix. Additional information came from a series of rhesus and cynomolgus monkeys (Macaca fasciculata) with anterograde tracer injections in the medial temporal lobe. The hippocampal formation, including the subiculum and presubiculum, together with the entorhinal cortex (EC) and perirhinal cortex (area 35) contribute numerous axons to the fornix in a topographical manner. In contrast, the lateral perirhinal cortex (area 36) and parahippocampal cortical areas TF and TH only contained a handful of cells labeled via the fornix. The medial fornix originates from cells in the caudal half of the subiculum, the lamina principalis interna of the caudal half of the presubiculum, and from the perirhinal cortex (area 35). The intermediate portion of the fornix (i.e., that part midway between the midline and most lateral parts of the fornix) originates from cells in the rostral half of the subiculum and prosubiculum, the anterior presubiculum (only from the lamina principalis externa), the caudal presubiculum (primarily from lamina principalis interna), the rostral half of CA3, the EC (primarily 28I and 28M), and the perirhinal cortex (area 35). The lateral parts of the fornix arise from the rostral EC (28L only) and the most rostral portion of CA3. Subcortically, the medial septum, nucleus of the diagonal band, supramammillary nucleus, lateral hypothalamus, dorsal raphe nucleus, and the thalamic nucleus reuniens all send projections through the fornix, which presumably terminate in the hippocampus and adjacent parahippocampal region. These results not only help to define those regions that project via the fornix, but also reveal those subcortical projections to the hippocampal formation most likely to rely entirely on nonfornical pathways.     

Corticothalamic connections of the posterior parietal cortex in the rhesus monkey

Corticothalamic connections of posterior parietal regions were studied in the rhesus monkey by using the autoradiographic technique. Our observations indicate that the rostral superior parietal lobule (SPL) is connected with the ventroposterolateral (VPL) thalamic nucleus. In addition, whereas the rostral SPL is connected with the ventrolateral (VL) and lateral posterior (LP) thalamic nuclei, the rostral IPL has connections with the ventroposteroinferior (VPI), ventroposteromedial parvicellular (VPMpc), and suprageniculate (SG) nuclei as well as the VL nucleus. The caudal SPL and the midportion of IPL show projections mainly to the lateral posterior (LP) and oral pulvinar (PO) nuclei, respectively. These areas also have minor projections to the medial pulvinar (PM) nucleus. Finally, the medial SPL and the caudal IPL project heavily to the PM nucleus, dorsally and ventrally, respectively. In addition, the medial SPL has some connections with the LP nucleus, whereas the caudal IPL has projections to the lateral dorsal (LD) nucleus. Furthermore, the caudal and medial SPL and the caudal IPL regions have additional projections to the reticular and intralaminar nuclei-the caudal SPL predominantly to the reticular, and the caudal IPL mainly to the intralaminar nuclei. These results indicate that the rostral-to-caudal flow of cortical connectivity within the superior and inferior parietal lobules is paralleled by a rostral-to-caudal progression of thalamic connectivity. That is, rostral parietal association cortices project primarily to modality-specific thalamic nuclei, whereas more caudal regions project most strongly to associative thalamic nuclei.     

Organization of projections from the gracile, medial cuneate and lateral nuclei in the North American opossum. Horseradish peroxidase study of the cells projecting to the cerebellum, thalamus and spinal cord

Using the horseradish peroxidase technique on the North American opossum, we were able to locate the neurons within the dorsal column and lateral cuneate nuclei which innervate the cerebellum and thalamus as well as those within the dorsal column nuclei which project spinalward. The medial and lateral cuneate nuclei supply axons to the anterior lobe, the paramedian lobule and the pyramis of the cerebellum and the lateral nucleus provides an additional projection to the uvula. The cerebellar projections from these nuclei arise from neurons located rostral to the obex. The thalamic projections from the gracile and medial cuneate nuclei originate from neurons throughout their rostral to caudal extent, although most of them are located just rostral to the obex. Neurons within the lateral cuneate nucleus which innervate the thalamus are found at intermediate rostrocaudal levels where most of them approximate the medial cuneate nucleus. The medial cuneate also projects to at least lumbar levels of the spinal cord in the opossum and neurons giving rise to such connections are found at the level of the obex and caudal to it. Neurons within the dorsal part of the dorsal column nuclei were labelled only after thalamic injections. Our results in the opossum are compared with those obtained in several placental mammals.     

Contralateral thalamic projections predominantly reach transitional cortices in the rhesus monkey

Connections between the thalamus and the cortex are generally regarded as ipsilateral, even though contralateral connections exist as well in several adult mammalian species. It is not known however, whether contralateral thalamocortical projections reach particular cortices or whether they emanate from specific nuclei. In the rhesus monkey different types of cortices, ranging from transitional to eulaminate, vary in their cortical connectional pattern and may also differ in thier thalamic connections. Because olfactory and transitional prefrontal cortices receive widespread projections, we investaged whether they are the target of projections from the contralateral thalamus as well. With the aid of retrograde tracers, we studied the thalamic projections of primary olfactory (olfactory tubercle and prepiriform cortex) and transitional orbital (areas PAPP, Pro 13) and medial (areas 25, 24, 32) areas, and of eulaminate (areas 11, 12, 9) cortices for comparison. To determine the prevalence of neurons in the contralateral thalamus, we compared them with the ipsilateral in each case. The pattern of ipsilateral thalamic projections differed somewhat among orbital, medial, and olfactory cortices. The mediodorsal nucleus was the predominant source of projections to orbital areas, midline nuclei included consistently about 25% of the thalamic neurons directed to medial transitional cortices, and primary olfactory areas were distinguished by receiving thalamic projections predominantly from neurons in midline and intralaminar nuclei. Notwithstanding some broad differences in the ipsilateral thalamofrontal projections, which appeared to depend on cortical location, the pattern of contralateral projections was thalamus were noted in midline, the magnocellular sector of the mediodorsal nucleus, the anterior medial and intralaminar nuclei, and ranged from 0 to 14% of the ipsilateral; they were directed primarily to olfactory and transitional orbital and medical cortices but rarely projected to eulaminate areas. Several thalamic nuclei projected from both sides to olfactory and transitional areas, but issued only ipsilateral projections to eulaminate areas. Though ipsilateral thalamocortical projections predominate in adult mammalian species, crossed projections are a common feature in development. The results suggest differences in the persistence of contralateral thalamocortical interactions between transitional and eulaminate cortices. © 1994 Wiley-Liss, Inc.     

The lateral ventromedial thalamic nucleus spreads nociceptive signals from the whole body surface to layer I of the frontal cortex

Neurons within the lateral ventromedial thalamic nucleus (VMl) convey selectively nociceptive information from all parts of the body. The present experiments were performed in rats and were designed to determine the organization of cortical projections from VMl neurons. In a first series of experiments, these cells were characterized electrophysiologically and individually labelled in a Golgi-like manner following juxtacellular electrophoresis of biotin-dextran. In a second experimental series, topical applications of the tracers fluorogold and tetramethylrhodamine-labelled dextran were placed into both the rostral-most and caudal areas of layer I of the dorsolateral frontal cortex, respectively. All VMl nociceptive neurons were fusiform and their full dendritic arborizations were bipolar, extending in the lateromedial axis. VMl cells are thus particularly well located to receive widespread nociceptive inputs via a brainstem link, viz. the medullary subnucleus reticularis dorsalis. VMl neurons driven by 'whole body' nociceptive receptive fields project to the rostral part of the layer I of the dorsolateral frontal cortex. These projections are widespread because double-labelling data showed a great number of VMl neurons labelled from both rostral and caudal dorsolateral cortices. The VMl comprises a homogeneous, organized subset of thalamic neurons that allow any signals of pain to modify cortical activity in a widespread manner, by interacting with the entire layer I of the dorsolateral neocortex.     

Organization of thalamic projections in the nucleus accumbens and the caudate nucleus in cats and its relation with hippocampal and other subcortical afferents

The organization of thalamic projections in the nucleus accumbens (NA) and the caudate nucleus of cats and its relation to other subcortical striatal afferents were studied with a retrograde tracing technique by use of lectin-conjugated horseradish peroxidase. The study showed that the paraventricular and medial parafascicular nuclei (PF) of the thalamus project to the medial NA and the parataenial and medial PF project to the lateral NA. The ventral tegmental area and substantia nigra pars dorsalis (SNpd) project to medial and lateral NA. The midline thalamic nuclei, rostral intralaminar nuclei, ventroanterior nucleus, medial and lateral PF, lateral posterior complex, and nucleus limitans project to medial caudate nucleus. The most medial substantia nigra pars compacta (SNpc) and rostral SNpd project to medial caudate nucleus. The center median, ventrolateral, and the central lateral nuclei of thalamus, SNpc, and SNpd project to lateral caudate nucleus. These results suggest that the thalamic and subcortical nuclei known to connect with the limbic and frontal cortices project to NA and medial caudate nucleus. Those thalamic nuclei connected with the motor system project to lateral caudate nucleus. The hippocampus projects selectively to medial NA. The amygdala, raphe, and other mesencephalic nuclei project only to NA and medial caudate nucleus. The organization of hippocampal, amygdala, and other subcortical afferents suggests that NA and caudate nucleus can be separated into medial "limbic" and lateral nonlimbic "sensory-motor" compartments. A brief review of the distribution pattern of some neurotransmitters, neuropeptides, and their receptors and behavior studies provides additional support to the concept that the striatum can be divided into several subcompartments.     

Corticothalamic connections of paralimbic regions in the rhesus monkey

This study addressed the issue of whether paralimbic regions of the cerebral cortex share common thalamic projections. The corticothalamic connections of the paralimbic regions of the orbital frontal, medial prefrontal, cingulate, parahippocampal, and temporal polar cortices were studied with the autoradiographic method in the rhesus monkey. The results revealed that the orbital frontal, medial prefrontal, and temporal polar proisocortices have substantial projections to both the dorsomedial and medial pulvinar nuclei, whereas the anterior cingulate proisocortex (area 24) projects exclusively to the dorsomedial nucleus. These proisocortical areas also have thalamic connections with the intralaminar and midline nuclei. The cortical areas between the proisocortical regions on the one hand and the isocortical areas on the other, that is, the posterior cingulate region (area 23) and the posterior parahippocampal gyrus (areas TF and TH), project predominantly to the dorsal portion of the medial pulvinar nucleus, the anterior nuclear group (AV, AM), and the lateral dorsal (LD) nucleus. Additionally, the posterior cingulate and medial parahippocampal gyri (area TH) have projections to the lateral posterior (LP) nucleus. Thus, it appears that the proisocortical areas, which are characterized by a predominance of infragranular layers and an absence of layer IV, have common thalamic relationships. Likewise, the intermediate paralimbic areas between the proisocortex and isocortical regions, which also have a predominance of infragranular layers but in addition have evidence of a fourth layer, project to the medial pulvinar and to the so-called limbic nuclei, AV, AM, LD, as well as a modality-specific nucleus, LP.     

Topographical organization of the thalamic afferent connections to the motor cortex in the cat

The topographical distribution of the cortical afferent connections to the different subdivisions of the motor cortex (MC) was studied in adult cats. The retrograde axonal transport of horseradish peroxidase technique was used. Small single injections of the enzyme were made in the entire MC, including the hidden regions in the depth of the sulcus cruciatus. The areal location and density of the subsequent thalamic neuronal labeling were evaluated in each case. Comparison of the results obtained in the various cases shows that the following: (1) The ventral anterior-ventral lateral complex is the principal thalamic source of afferents to the MC. (2) The ventral medial, dorsal medial, the different components of the posterior thalamic group (lateral, medial, and ventral posteroinferior and suprageniculate nuclei), and the intralaminar, lateral anterior, lateral intermediate, lateral medial, and anteromedial thalamic nuclei are also thalamic sites in which neural projections to the MC arise. (3) The thalamocortical projections to the MC are sequentially organized. The connections arising from the lateral part of the thalamus end in the region of area 4 that is situated medially in the superior lip of the sulcus cruciatus and in the posterior sigmoid gyrus. The projections originating in the most medial thalamic regions terminate in that region of area 6a beta which is located in the medial part of the inferior lip of the cruciate sulcus, and in the anterior sigmoid gyrus. Moreover, the ventral thalamic areas send connections to the most anteriorly located zones of the MC, while the most dorsal thalamic ones project to the most posteriorly located parts of the MC. (4) This shift in the thalamocortical connections is not restrained by cytoarchitectonic boundaries, either in the thalamus or in the cortex. (5) The populations of thalamocortical cells which project to neighboring MC subdivisions exhibit consistent overlapping among themselves. (6) These findings suggest, moreover, that the basal ganglia and the cerebellar projections to the MC through the thalamus are arranged in a number of parallel pathways, which may occasionally overlap.     

Nigrothalamic projections in the monkey demonstrated by autoradiographic technics.

In spite of repeated demonstrations by degeneration technics, nigrothalamic fibers have been regarded with some skepticism. Attempts were made to trace nigral efferent projections in the monkey by autoradiographic technics. Tritiated amino acids (L-leucine, L-lysine and L-proline), injected into portions of the substantia nigra (SN), labeled cells in four regions, designated as, (1) rostrolateral, (2) caudolateral, (3) rostromedial and (4) central. Rostrolateral nigral neurons transported radioactive label preferentially and abundantly to thalamic nuclei; localized isotope was found in parts of three thalamic nuclei, the medial part of the ventral lateral nucleus (VLm), the magnocellular part of the ventral anterior nucleus (VAmc), and the paralaminar part of the dorsomedial nucleus (DMpl)9 Lateral neurons in the caudal half of the SN transmitted radioactive label to the same thalamic nuclei as rostrolateral nigral neuron. Isotope transported to portions of the striatum was modest and localized. Radioactive label taken up by large cells in the caudal third of the SN was transported to portions of the striatum, but not to thalamic nuclei. Labeled nigral neurons in the medial two-thirds of the rostral half of the SN, and in the middle third of the central part of the SN, transported the label mainly to parts of the caudate nucleus and putamen. In these animals modest radioactive label was seen in VLm and VAmc, but not in other thalamic nuclei. There was no evidence that nigral neurons project to the subthalamic nucleus. No radioactive transport from nigral neurons was detected in the superior colliculus, the midbrain tegmentum, or the red nucleus, and none was transported to more caudal brain stem nuclei. Nigrothalamic fibers arise particularly from cells in rostral and lateral parts of the substantia nigra. While some cells in other parts of the nigra project to thalamic nuclei, these appear scattered and less numerous. Large cells in caudal parts of the SN do not project to thalamic nuclei. These observations confirm nigrothalamic projections to VLm and VAmc, and identify a new nigral projection to part of the dorsomedial nucleus of the thalamus (DMpl). No nigral efferent fibers project to any of the intralaminar thalamic nuclei.     

Origin of the neurotensinergic innervation of the rat basal forebrain studied by retrograde transport of cholera toxin

Basal forebrain cholinergic neurons project to the hippocampus and cerebral cortex where they play an important role in cortical activation, attention, and memory. These neurons have been shown to express functional neurotensin receptors and to receive a dense neurotensinergic innervation. In the present study, we investigated the origin of this innervation by combining retrograde transport of cholera toxin with immunohistochemical detection of neurotensin. After injection of cholera toxin in the anterior substantia innominata and diagonal band of Broca, retrogradely labelled cells were widely distributed throughout forebrain limbic structures. Only a small proportion of these cells, located (by decreasing order of frequency) in the lateral septum, medial preoptic area, rostral hypothalamus, nucleus accumbens, and rostral basal forebrain, were dually labelled for neurotensin. After injection of cholera toxin in the posterior substantia innominata and magnocellular preoptic nucleus, retrogradely labelled cells were detected throughout the limbic forebrain and ponto-mesencephalic tegmentum. Here again, only a small proportion of these cells, located (by decreasing order of frequency) in the nucleus accumbens, lateral septum, rostral basal forebrain, hypothalamus, bed nucleus of the stria terminalis, supramammilliary nucleus, ventral tegmental area, and raphe complex co-localized neurotensin. In view of the burst generating properties of neurotensin on basal forebrain cholinergic neurons, our results suggest that neurotensin projections may be part of the septo-hippocampo-septal loop regulating hippocampal theta activity. More caudally, neurotensin axons originating from the lateral hypothalamus and pontomesencephalic tegmentum may contribute to the contingent of ascending reticular formation fibers involved in the regulation of the sleep-wake cycle. J. Comp. Neurol. 391:30–41, 1998. © 1998 Wiley-Liss, Inc.     

Thalamic projections to the posteromedial cortex in the macaque

The medial parietal, posterior cingulate, and retrosplenial cortices collectively constitute a region of cortex referred to as the posteromedial cortices (PMC). In an effort to shed light on the neuroanatomical organization of the PMC, we undertook a study to identify and analyze the thalamocortical connections of these cortices. Retrograde tracer injections were placed in the posterior cingulate (PCC), retrosplenial (RSC), medial parietal cortices (MPC), and posterior cingulate sulcus (PCS), and the labeling patterns within the thalamus were analyzed. Three afferent projection patterns were observed to the PMC from the thalamus: a PCC/RSC pattern that involved the anterior thalamic nuclei, an MPC pattern that involved the lateral posterior and pulvinar nuclei, and a PCS pattern that involved the ventral thalamic nuclei. Additionally, a shared pattern of projections from the anterior intralaminar nuclei (AILN) and posterior thalamic nuclei (PTN) to all cortical regions of the PMC was observed. Our findings suggest that distinct regions within the PMC are supplied by distinctive patterns of thalamic input, but also share common projections from intralaminar and posterior thalamic sources. In addition, we relate our findings to functional abnormalities in aging and dementia, and address a domain-like pattern of thalamocortical labeling of the PMC that is drawn selectively and collectively from multiple thalamic nuclei.