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W O Guldin

Publications and source records attributed to W O Guldin.

15 recordsLinked to original sources

Is there a vestibular cortex?

Very different areas of the primate cortex have been labelled as 'vestibular'. However, no clear concept has emerged as to where and how the vestibular information is processed in the cerebral cortex. On the basis of data from single-unit recordings and tracer studies, the present article gives statistical evidence of the existence of a well-defined vestibular cortical system. Because the data presented here have been verified in three different primate species, it can be predicted that a similar vestibular cortical system also exists in humans.

Animals

Corticofugal connections between the cerebral cortex and brainstem vestibular nuclei in the macaque monkey.

The distribution of cortical efferent connections to brainstem vestibular nuclei was quantitatively analysed by means of retrograde tracer substances injected into different electrophysiologically identified parts of the brainstem vestibular nuclear complex of five Java monkeys (Macaca fascicularis). Three polysensory vestibular areas were found to have a substantial projection to the vestibular nuclei: area 2v located at the tip of the intraparietal sulcus, the parietoinsular vestibular cortex (PIVC) covering the most occipital part of the granular insula (Ig) and the retroinsular area (Ri or reipt), and the dorsolateral part of the somatosensory area 3a ("area 3aV" neck/trunk region). From physiological recording experiments, these three cortical fields were known to contain many neurons responding to stimulation of semicircular canals as well as to optokinetic (area 2v, PIVC) and somatosensory stimuli (PIVC, area 3a). These three regions form the inner cortical vestibular circuit. Besides these polysensory vestibular cortical fields, three other circumscribed cortical regions of the macaque brain were also found to project directly to the brainstem vestibular nuclei: a circumscribed part of the postarcuate premotor cortex (area 6pa), part of the agranular and the adjacent dysgranular cortex located around the cingulate sulcus (area 6c/23c), and a predominantly visual (optokinetic) association field located at the fundus of the lateral sulcus (area T3). These areas are known to have connections with the structures of the inner cortical vestibular circuit. Only a few efferent connections to the brainstem vestibular nuclei were found for the different parts of cytoarchitectonic area 7. Significant differences were found between the efferent innervation patterns of the axons originating in the six cortical areas mentioned and ending in the various compartments of the vestibular nuclear complex. Vestibular nuclei with a dominant output to the gaze motor system of the brainstem receive efferent connections preferably from the parietoinsular vestibular cortex. Vestibular structures with their primary output to skeletomotor centers, however, receive stronger efferent connections from areas 6pa and 3a. The ventrolateral nucleus, which sends efferent axons to both the oculomotor and skeletomotor systems of the brainstem and the spinal cord, also receives its main cortical efferents from the somatomotor area 6 and from area 3aV. Through these connections the cortical somatomotor system may directly influence vestibuloocular and vestibulocollic reflexes. It is speculated that the corticofugal connections to the vestibular brainstem nuclei are predominantly inhibitory, suppressing vestibular reflexes during cortically controlled goal-directed movements.

Animals

Connections from the neocortex to the vestibular brain stem nuclei in the common marmoset.

Monosynaptic projections from the cerebral cortex to the vestibular nuclei were studied in the common marmoset monkey (Callithrix jacchus) by injecting fluorescent dextrans into the brain stem vestibular nuclei. The injection sites were determined by single unit vestibular responses identified later histologically. During the recordings sinewave rotation in pitch, roll or yaw or steady tilt was applied. The most notable loci of labelling were found inside the primary sensory cortex, the cortex deep down along the posterior lateral sulcus, the premotor region and the anterior cingulate cortex. From studies in other primates these cortical areas are known to process vestibular information. Their connections with the vestibular nuclei may serve as an internal feedback modulating the vestibular brain stem activity.

Animals

Corticofugal projections to the vestibular nuclei in squirrel monkeys: further evidence of multiple cortical vestibular fields.

Single- and multiple-unit recordings were made from nerve cells located in the different nuclei of the brainstem vestibular nuclear complex (VNC) of anaesthetized squirrel monkeys (Saimiri sciureus) by conventional stereotaxic techniques. After neurons responding to semicircular canal stimulation in a yaw, roll, or pitch direction or to otholith stimulation were identified, small amounts of retrograde tracer substances were deposited at the recording sites. Up to three different tracers were administered to different parts of the VNC in the same animal (Fast Blue, HRP-WGA, and Rhodamine-dextranes). After adequate survival times, the animals were sacrificed. Following histological processing, the cortical grey matter was screened systematically for cells labelled with the retrograde tracers (fluorescence microscopy or light microscopy for HRP processing). Labelled nerve cells which clearly project to the VNC directly were found predominantly in the cytoarchitectonic layer 5 of seven different cortical areas: 1) The parieto-insular vestibular cortex PIVC, which in squirrel monkeys consists mainly of the medial area Ri and parts of the anterior area Ig; 2) area 7ant, which presumably corresponds to the macaque area 2v; 3) area 3aV, a vestibular field of area 3a; 4) the temporal area T3 bordering on area Ri; 5) the premotor area 6a; and 6, 7) the areas 6c and 23c of the anterior cingulate cortex. The PIVC, area 7ant, and area 3aV form the "inner cortical vestibular circuit" (Guldin et al.: J. Comp. Neurol. 326:375-401, '92), while the other cortical areas mentioned also have direct projections to the structures of the inner cortical vestibular circuit. It is speculated that the direct projections of the cortical vestibular structures to the brainstem vestibular nuclei regulate the vestibulo-ocular, the vestibulo-spinal, and the optokinetic reflexes mediated through the VNC, thus preventing counteractions of these reflexes during voluntary, goal-directed head movements or locomotion.

Animals

Cortico-cortical connections and cytoarchitectonics of the primate vestibular cortex: a study in squirrel monkeys (Saimiri sciureus).

The cortical connections of two vestibular fields [parieto-insular vestibular cortex (PIVC) and area 3aV] were studied in the squirrel monkey (Saimiri sciureus) by means of retrograde tracer techniques. Small iontophoretic or pressure injections of horseradish peroxidase (HRP), wheat-germ-HRP, Nuclear Yellow, and Fast Blue were administered to the cytoarchitectonic areas Ri (PIVC), 3aV, the parieto-temporal association area T3, the granular insula (Ig), and the rostral part of area 7 (7ant). The injection sites were physiologically characterized by means of microelectrode recordings and vestibular, optokinetic, or somatosensory stimulation: Area Ri is the region of the parieto-insular vestibular cortex (PIVC) as defined in macaques. The neck-trunk region of area 3a (area 3aV) also contains many neurons responding to stimulation of semicircular canal receptors. Some neurons of area T3 bordering on the PIVC also receive vestibular signals, but most neurons in area T3 responded preferentially to large-field optokinetic stimulation and not to vestibular stimulation. In none of the areas mentioned were responses to otolith stimulation found. The PIVC receives inputs from frontal and parietal cortical areas, especially areas 8a, 6, 3a, 3aV, 2, and 7ant. Area T3 receives signals from the insular and retroinsular cortex, various parts of area 7, visual areas of the parieto-occipital and parieto-temporal regions (area 19) and from a sector of the upper bank of the temporal sulcus (STS-area). The cortical afferents to area 3aV stem from areas 24, 4, 6, 7ant, from other parts of the primary somatosensory cortex, the secondary somatosensory cortex (SII), the retroinsular cortex (Ri), and the granular insula (Ig). In the border region of the areas 2 and 7ant, labelled neurons appeared after injections into both the PIVC and the area 3aV. This region is presumably the homologue to the vestibular area 2v of the macaque brain. In all regions cells within the contralateral cortex were less frequently labelled than cells in the homologous structures of the ipsilateral hemisphere. The cortical system for processing vestibular information about head-in-space movement consists mainly of the reciprocally interconnected areas PIVC and 3aV, and most likely of border regions of area 2 and 7ant. This "inner cortical vestibular circuit" also receives signals from two other cortical sensory systems, the somatosensory-proprioceptive system mediated by the primary somatosensory cortex and the visual movement system (optokinetic or visual flow signals). These visual movement signals reach PIVC via area 19 and area T3.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Thalamic connections of the vestibular cortical fields in the squirrel monkey (Saimiri sciureus).

The afferent thalamic connections to cortical fields important for control of head movement in space were analysed by intracortical retrograde tracer injections. The proprioceptive/vestibular area 3aV, the neck-trunk region of area 3a, receives two thirds of its thalamic projections from the oral and superior ventroposterior nucleus (VPO/VPS), which is considered as the proprioceptive relay of the ventroposterior complex (Kaas et al., J. Comp. Neurol. 226:211-240, 1984). The parieto-insular vestibular cortex (PIVC, area retroinsularis, Ri) receives its main thalamic input from posterior parts of the ventroposterior complex and from the medial pulvinar. Anatomical evidence is presented that the posterior region of the ventroposterior complex is a special compartment within this principal somatosensory relay complex. The parietotemporal association area T3, mainly involved in visual-optokinetic signal processing, receives a substantial input from the medial, the lateral, and the inferior pulvinar. Dual tracer experiments revealed that about 5% of the thalamic neurons projecting to 3aV were spatially intermingled with neurons projecting to areas PIVC or T3. This spatial intermingling was distributed over small but numerous, circumscribed thalamic regions, called "common patches," which were found mainly in the intralaminar nuclei, the posterior group of thalamic nuclei, and the caudal parts of the ventroposterior complex. The "common patches" may indicate a functional coupling of area 3aV with the PIVC or area T3 on the thalamic level. In control experiments thalamic projections to the granular insula Ig and the anterior part of area 7, two cerebral structures connected with the vestibular cortical areas, were studied. Some overlap in the thalamic relay structures projecting to these areas with those projecting to the vestibular cortices was found. A quantitative evaluation of thalamic regions projecting to different cortical structures was performed by constructing so-called "thalamograms." A scheme was developed that describes the afferent thalamic connections by which vestibular, visual-optokinetic, and proprioceptive signals reach the vestibular cortical areas PIVC and 3aV.

Animals

Cortical projections originating from the cat's insular area and remarks on claustrocortical connections.

The cortical projections originating in the cat's insular cortex and claustrum were investigated with the aid of the horseradish peroxidase retrograde tracing technique. Twenty small injections of horseradish peroxidase were distributed along lateral and medial regions of the hemisphere. Labeling in the insular cortex occurred following all injections except those six situated along the lateral gyrus--that is, within the visual cortex. In the claustrum labeled neurons were found following all injections, except following the injection situated in the posterior temporal area. Claustral labeling was frequently more intense than insular labeling. The injections into the occipital cortex that revealed no insular innervation nevertheless received a considerable number of claustral projections. As the insular cortex itself receives at most a minor projection from the claustrum the differing cortical projection patterns of insula and claustrum have to be considered unrelated. Our findings confirm the view that the claustrum projects to most regions of the cerebral cortex; these projections are at least in part topographically organized. A topographical pattern can also be constructed for the insular cortex, though it is less stringent than for the claustrocortical connections. Both the afferent and efferent connections of the insula show similarities to those of the prefrontal cortex. Nevertheless, the insula differs in that it receives strong input from the sensory associative nuclei of the thalamus. Consequently, and in line with behavioral observations following its ablation, we consider the insula as involved in the temporal structuring of perceived patterns.

Animals

Cortical and thalamic afferent connections of the insular and adjacent cortex of the cat.

The thalamo-cortical and cortico-cortical afferents of the cat's insular cortex were investigated with the retrograde horseradish peroxidase technique. The most prominent loci of thalamic labeling were the suprageniculate nucleus and parts of the posterolateral nucleus. Injections into the anterior part of the insular cortex also resulted in labeled cells in the ventromedial posterior nucleus and in the intralaminar nuclei, while injections into posterior parts revealed projections from the medial and dorsal parts of the medial geniculate nucleus. Only the anterior and most ventral parts of the insular cortex overlying the anterior rhinal sulcus were connected with the mediodorsal nucleus of the thalamus. All injections into the gyrus sylvius anterior showed a specific pattern of cortical afferents: With the exception of the labeling in the prefrontal cortex and the inferotemporal region, the labeled cells were very narrowly restricted to the presylvian, the suprasylvian, and the splenial sulcus. The thalamic neurons projecting to the cortex were generally organized in a bandlike pattern which crossed nuclear borders. The majority of the cortico-cortical connections originated from sulcal areas next to the prefrontal, parietal, and cingulate cortex, that is, next to so-called association cortices. In the light of the present results the role of the insular cortex as a multifunctional association area is discussed, as well as its relation to other cortical centers.

Afferent Pathways

Afferents to the ventral tegmental nucleus of Gudden in the mouse, rat, and cat.

Afferents to the ventral tegmental nucleus of Gudden (VT) were investigated in mice, rats, and cats. Unilateral and bilateral injections or iontophoretical applications of horseradish peroxidase (HRP) were made into the region of the VT. The entire cerebrum was then screened for labeled neurons. Following injections situated principally within the VT, in all three species many retrogradely labeled neurons were observed in the mamillary bodies and the lateral habenular nuclei. Fewer labeled cells were observed in the prefrontal cortex, the basal forebrain, various hypothalamic nuclei, the interpeduncular nucleus, nucleus of the posterior commissure, nucleus of Darkschewitsch and interstitial nucleus of Cajal, vestibular nucleus, and nucleus praepositus hypoglossi. Scant but consistent labeling occurred in the cingular, retrosplenial, and insular cortices, within the medial forebrain bundle, fields of Forel, zona incerta, ventral tegmental area of Tsai, substantia nigra, pretectal area, periaqueductal gray, dorsal tegmental nucleus, locus ceruleus, and raphe complex. Our results show a high similarity in the distribution of afferent connections converging on the VT of mice, rats, and cats. They indicate furthermore that the VT is reached by a variety of cortical and subcortical afferents, which belong either to the limbic system or to brain stem regions related to motor, sensory, and autonomic functions. It is suggested that the VT subserves as a midbrain core structure of the limbic system, which is responsible for the transfer of motor, sensory, and autonomic informations arising within the brain stem to limbic forebrain structures.

Animals

Cortical and thalamic afferent connections of the insular and adjacent cortex of the rat.

Thalamic and cortical afferents to the insular and perirhinal cortex of the rat were investigated. Unilateral injections of horseradish peroxidase (HRP) were made iontophoretically along the rhinal sulcus. HRP injections covered or invaded areas along the rhinal fissure from about the level of the middle cerebral artery to the posterior end of the fissure. The most anterior injection labeled a few cells in the mediodorsal nucleus. More posterior injections labeled neurons in the basal portion of the nucleus ventralis medialis, thus suggesting that this cortical region constitutes the rat's gustatory (insular) cortex. We consider the cortex situated posterior to the gustatory cortex in and above the rhinal sulcus as the core region of the rat's (associative) insular cortex, as this cortex receives afferents from the regions of and between the nuclei suprageniculatus and geniculatus medialis, pars magnocellularis. It includes parts of the cortex termed perirhinal in other studies. The cortex dorsal and posterior to the insular cortex we consider auditory cortex, as it receives afferents from the principal part of the medial geniculate nucleus, and the cortex ventral to the insular cortex (below the fundus of the rhinal sulcus) we consider to constitute the prepiriform cortex, which is athalamic. The posterior part of the perirhinal cortex (area 35) receives afferents from nonspecific thalamic nuclei (midline nuclei). Cortical afferents to the injection loci arise from a number of regions, above all from regions of the medial and sulcal prefrontal cortex. Those injections confined to the projection cortex of the suprageniculate-magnocellular medial geniculate nuclear complex also led to labeling in contralateral prefrontal regions, particularly in area 25 (infralimbic region). A comparison of our results with those on the insular cortex of cats and monkeys suggests that on the basis of thalamocortical connections, topographical relations, and involvements of neurons in information processing and overt behavior, the insular cortex has to be regarded as a heterogeneous region which may be separated into prefrontal insular, gustatory (somatosensory) insular, and associative insular portions.

Afferent Pathways

Heterotopic interhemispheric cortical connections in the rat.

In the rat, contrary to other species, interhemispheric cortical connections have been considered to travel largely between homotopical regions only. Based on iontophoretic injections of horseradish peroxidase, the present study reports extensive heterotopic interhemispheric connections between posterior insular (perirhinal) regions of the lateral part of the hemisphere and anterior prefrontal regions of the medial hemisphere and vice versa. Generally, the areas connected interhemispherically are also connected intrahemispherically. The ratio of contralateral projections appears to be less than one third of the ipsilateral ones.

Animals

Epidural kainate, but not ibotenate, produces lesions in local and distant regions of the brain. A comparison of the intracerebral actions of kainic acid and ibotenic acid.

Epidural placement of ibotenic acid (Ibo) is described as a new technique for easy and successful lesioning of cortical neurons. The method works reliably without apparent damage to passing or underlying fibers and without causing distant lesion effects. In rats, the favorable effects of epidural placement of Ibo on behavior and on neuronal tissue were compared with and found to contrast with those of kainic acid (KA). Survival periods of 2, 4, 6, and 8 days were allowed for animals of each group. The effects of Ibo from Amanita muscaria were compared to those from Amanita pantherina. Local cortical lesions after epidural placement of KA and Ibo were compared with those after subdural placement. The effects resulting from covering a small surface of the dura with Ibo were compared with those resulting from covering a large surface. Epidural placement of KA and Ibo resulted in neuronal damage of the underlying cortex, 0.4 mg of KA leading to approximately the same lesion extent as 0.5 mg of Ibo. Epidural and subdural lesion effects were found to be similar both for KA and Ibo. The most striking difference between epidural application of KA and Ibo was that KA lesions resulted in a considerable number of damaged areas remote from the local cortical lesion, whereas even 10-times higher doses of Ibo (measured in moles) did not produce any observable neuronal damage outside the local lesion. From these results it is suggested that Ibo has to be favored over KA when local neuronal lesions are desired. Furthermore, epidural placement of Ibo is proposed as an effective technique for producing circumscribed cortical lesions.

Animals

No detectable remote lesions following massive intrastriatal injections of ibotenic acid.

Behavioral and anatomical consequences of particularly large intrastriatal injections of ibotenic acid are described. Only in the rat with the largest injection, which encompassed almost the entire frontal lobe, were enduring aphagia and adipsia observed; epileptic attacks were, however, not detectable in this or in any other of the rats. In spite of the massiveness of the lesion, neither remote lesions nor damage to passing fibers was observed. It is therefore suggested to substitute kainic acid by ibotenic acid for the production of local, discrete brain lesions.

Animals

Prefrontal cortex of the mouse defined as cortical projection area of the thalamic mediodorsal nucleus.

In the mouse small amounts of horseradish peroxidase were injected iontophoretically into different portions of the frontal lobe. Out of a large number of mice, 18 with typical injections which together covered most of the anterior half of the cortex were selected, and their retrogradely labeled thalamic cells were described and illustrated in detail. Special emphasis was laid on the cortical projections of the mediodorsal nucleus which were considered to define locus and extent of the prefrontal cortex. The results reveal that topographically the extent of the mouse, prefrontal cortex closely resembles that of the rat, another species of the rodent order. It includes a small area within and dorsal to the rhinal sulcus and a comparatively larger region within the medial half of the anterior cortex. Furthermore, the dorsal tip of the frontal pole is reached by a considerable number of afferents from the mediodorsal nucleus. Whether this last projection also exists in the rat is still disputed. As in the rat, only the sulcal prefrontal cortex seems to be reached by mediodorsal cells alone, whereas the rest of the prefrontal cortical fields have overlapping projections from the anteromedial nucleus. Though a considerable area within the frontal lobe of the mouse is reached by mediodorsal afferents, the significance and usefulness of this definition of the prefrontal cortex is questioned.

Animals