The motor nuclei and sensory neurons of the IIIrd, IVth, and VIth cranial nerves in the monitor lizard, Varanus exanthematicus.

The motor nuclei of the oculomotor, trochlear, and abducens nerves of the reptile Varanus exanthematicus and the neurons that subserve the sensory innervation of the extraocular muscles were identified and localized by retrograde and anterograde transport of horseradish peroxidase (HRP). The highly differentiated oculomotor nuclear complex, located dorsomedially in the tegmentum of the midbrain, consists of the accessory oculomotor nucleus and the dorsomedial, dorsolateral, intermediate, and ventral subnuclei. The accessory oculomotor nucleus projects ipsilaterally to the ciliary ganglion. The dorsomedial, dorsolateral, and intermediate subnuclei distribute their axons to the ipsilateral orbit, whereas the ventral subnucleus, which innervates the superior rectus muscle, has a bilateral, though predominantly contralateral projection. The trochlear nucleus, which rostrally overlaps the oculomotor nuclear complex, is for the greater part a comma-shaped cell group situated lateral, dorsal, and medial to the medial longitudinal fasciculus. Following HRP application to the trochlear nerve, almost all retrogradely labeled cells were found in the contralateral nucleus. The nuclear complex of the abducens nerve consists of the principal and accessory abducens nuclei, both of which project ipsilaterally. The principal abducens nucleus is located just beneath the fourth ventricle laterally adjacent to the medial longitudinal fasciculus and innervates the posterior rectus muscle. The accessory abducens nucleus has a ventrolateral position in the brainstem in close approximation to the ophthalmic fibers of the descending trigeminal tract. It innervates the retractor bulbi and bursalis muscles. The fibers arising in the accessory abducens muscles form a loop in or just beneath the principal abducens nucleus before they join the abducens nerve root. The afferent fibers conveying sensory information from the extraocular muscles course in the oculomotor nerve and have their perikarya in the ipsilateral trigeminal ganglion, almost exclusively in its ophthalmic portion.

The motor complex and primary projections of the trigeminal nerve in the monitor lizard, Varanus exanthematicus.

The sensory projections and the motor complex of the trigeminal nerve of the reptile Varanus exanthematicus were studied with the methods of anterograde degeneration and anterograde and retrograde axonal transport. The primary afferent fibers diverge in the brainstem into a short ascending and a long descending tract. The former distributes its fibers to the principal sensory trigeminal nucleus, where nerves V1, V2, and V3 are represented along a lateromedial axis. The fibers of the descending tract enter the nucleus of this tract and the reticular formation. Both in the tract and its nucleus, nerves V1, V2 and V3 occupy successively more dorsal positions. A small contingent of nerve V1 fibers course to the accessory abducens nucleus. The descending tract extends caudally into the first and second cervical segments of the spinal cord. The trigeminal motor complex consists of dorsal, ventral, and dorsomedial nuclei. The m. adductor mandibulae externus (the main jaw closer) is represented in the dorsal nucleus, predominantly in its rostral part. The muscles innervated by nerve V3 are represented in the ventral nucleus, mainly in its caudal part. All three divisions of the trigeminal nerve contain peripheral branches of the mesencephalic trigeminal system. Collaterals of the central branches of this system were traced to the ventral motor and the principal sensory trigeminal nuclei.

Primary projections and efferent cells of the VIIIth cranial nerve in the monitor lizard, Varanus exanthematicus.

In order to describe the central relations of both the afferent and efferent components of the VIIIth cranial nerve in one reptile, the methods of anterograde and retrograde axonal transport and anterograde degeneration were used to study the vestibular and cochlear projections and the efferent system of this nerve in Varanus exanthematicus. On the basis of cresyl violet and Klüver-Barrera staining, five vestibular nuclei, four cochlear nuclei, and two clusters of small cells which could not be designated as strictly auditory or vestibular are distinguished. The vestibular nuclei include the nucleus dorsolateralis, nucleus ventrolateralis, nucleus tangentialis, nucleus ventromedialis, and nucleus descendens. The well-developed cochlear nuclear complex includes the nucleus angularis, nuclei magnocellulares medialis and lateralis, and nucleus laminaris. The two cell clusters are located dorsolaterally in the brainstem just ventrolateral to the acoustic tubercle. The primary afferent vestibular fibers coursing in the anterior VIIIth nerve root distribute to the ventral portions of all vestibular nuclei except nucleus ventromedialis, whereas the fibers coursing in the posterior root project to the dorsal portions of these nuclei. In nucleus ventromedialis fibers of both roots do not segregate into ventral and dorsal portions. Other targets of the vestibular fibers are the two cell clusters, the granular layer of the ipsilateral cerebellum, the reticular formation, and the descending trigeminal tract and its nucleus. The primary cochlear fibers coursing in the posterior root terminate in nucleus angularis, nuclei magnocellulares medialis and lateralis, and the inner cell strand of nucleus laminaris. The efferent system is, ipsi- and contralaterally in the brainstem, composed of ventral and dorsal cell groups that extend from the level of the principal abducens nucleus caudally where they overlap with the facial motor nucleus. The fibers, which originate from the contralaterally located efferent cells, course beneath the IVth ventricle to exit the brainstem on the ipsilateral side.

Tectal connections in Python reticulatus.

The origins of the axons terminating in the mesencephalic tectum in Python reticulatus were examined by unilateral tectal injections of horseradish peroxidase. Retrogradely labeled cells were observed bilaterally throughout the spinal cord in all subdivisions of the trigeminal system, with the exception of nucleus principalis, which showed labeled cells only on the ipsilateral side. Labeling of the reticular formation occurred bilaterally in nucleus reticularis inferior magnocellularis, nucleus reticularis lateralis, nucleus reticularis, and the mesencephalic reticular formation. The tectum also receives bilateral projections from the dorsal tegmental field, the nucleus of the lateral lemniscus, and nucleus isthmi, and ipsilateral projections from nucleus profundus mesencephali. A few labeled cells were found ipsilaterally in the locus coeruleus and in nuclei vestibulares ventrolateralis and ventromedialis. In the diencephalon labeled cells were observed ipsilaterally in nucleus ventrolateralis thalami, nucleus ventromedialis thalami, nucleus suprapeduncularis, and in the dorsal and ventral lateral geniculate nuclei. Bilateral labeling was observed in nucleus periventricularis hypothalami. Furthermore, labeling was ipsilaterally present in the ventral telencephalic areas. The tectum in Python reticulatus receives a wide variety of afferent connections which confirm the role of the tectum as an integration center of visual and exteroceptive information.

Ascending connections to the forebrain in the Tegu lizard.

The ascending connections to the striatum and the cortex of the Tegu lizard, Tupinambis nigropunctatus, were studied by means of anterograde fiber degeneration and retrograde axonal transport. The striatum receives projections by way of the dorsal peduncle of the lateral forebrain bundle from four dorsal thalamic nuclei: nucleus rotundus, nucleus reuniens, the posterior part of the dorsal lateral geniculate nucleus and nucleus dorsomedialis. The former three nuclei project to circumscribed areas of the dorsal striatum, whereas nucleus dorsomedialis has a distribution to the whole dorsal striatum. Other sources of origin to the striatum are the mesencephalic reticular formation, substantia nigra and nucleus cerebelli lateralis. With the exception of the latter afferentation all these projections are ipsilateral. The ascending connections to the pallium originate for the major part from nucleus dorsolateralis anterior of the dorsal thalamus. The fibers course in both the medial forebrain bundle and the dorsal peduncle of the lateral forebrain bundle and terminate ipsilaterally in the middle of the molecular layer of the small-celled part of the mediodorsal cortex and bilaterally above the intermediate region of the dorsal cortex. The latter area is reached also by fibers from the septal area. The large-celled part of the mediodorsal cortex receives projections from nucleus raphes superior and the corpus mammillare.

The motor nuclei and primary projections of the IXth, Xth, XIth and XIIth cranial nerves in the monitor lizard, Varanus exanthematicus.

The motor nuclei and sensory connections of the IXth, Xth, XIth, and XIIth cranial nerves of the reptile Varanus exanthematicus were studied with the methods of anterograde degeneration and anterograde and retrograde axonal transport. The motor nuclei of nerve IX are located ventrally in the rhombencephalon and are constituted medially by the large-celled glossopharyngeal part of the nucleus ambiguus and laterally by the small-celled nucleus salivatorius inferior. The motor nuclei of nerve X consist of the dorsomedially located dorsal motor nucleus of the vagus and the laterally located vagal part of the nucleus ambiguus. The rostral portion of the latter cell group contains smaller cells than its caudal portion and is rostrally continuous with the nucleus salivatorius inferior of nerve IX. The efferent axons of nerves IX and X arising from the ventrolateral medulla first course dorsomedially, form genua beneath the IVth ventricle, and then exit the brainstem. All primary afferent fibers of nerve IX and the majority of those of nerve X enter the solitary tract. Terminations of vagal fibers were observed in the postvagal portion of the nucleus of the solitary tract, the dorsal motor nucleus of the vagus, and the nucleus of the commissura infima. A small contingent of vagal fibers courses caudally just dorsolateral to the descending trigeminal tract. A separate spinal component of nerve XI could not be found. The bulbar component of this nerve forms part of nerve X and takes its main origin from a detached caudal element of the nucleus ambiguus. The motor nuclear complex of nerve XII consists of a large dorsal nucleus and a small ventral nucleus that extend from the medulla oblongata into the first segment of the cervical spinal cord.

Connections of the parahippocampal cortex in the cat. V. Intrinsic connections; comments on input/output connections with the hippocampus.

The present report is the last in a series of papers on the connectivity of the parahippocampal cortex in the cat, which in this species is considered to be composed of the entorhinal and perirhinal cortices. Injections of anterogradely transported tritiated amino acids and the retrograde tracers HRP, WGA-HRP, fast blue, or nuclear yellow were placed within the limits of the parahippocampal cortex. An analysis was made of the resulting pattern of anterograde labeling and of the distribution of retrogradely labeled neurons within the parahippocampal cortex. It appears that within the parahippocampal cortex of the cat a framework exists, which is composed of longitudinal and transverse connections, organized according to three principles: Medially directed projections originate mostly in superficial layers, whereas laterally directed fibers come from deep layers. The longitudinal connections span the entire rostrocaudal extent of the parahippocampal cortex, whereas the mediolateral extent of the transverse connections is in general more restricted. Based on the organization of these longitudinal and transverse connections four longitudinal zones are recognized. The lateral entorhinal cortex (LEA) projects both within the entorhinal cortex and to the perirhinal cortex, whereas the intrinsic projections of the medial entorhinal cortex (MEA) are confined to the entorhinal cortex. These results are discussed in conjunction with the main organizational features of the afferent and efferent connections of the parahippocampal cortex of the cat. The premise is made that the cytoarchitectonically defined subdivisions of the cortex can be grouped into four areas, each with its own set of fiber connections and subserving different functional roles. A lateral area, constituted by the perirhinal areas 35 and 36, and the caudally adjacent postsplenial cortex, serves as a peripheral area through which the rest of the parahippocampal cortex--i.e., LEA and MEA, and ultimately the hippocampal formation--reciprocally communicates with extensive neocortical, subcortical, and thalamic regions associated with higher-order behavior. The medial part of LEA, constituted by the ventrolateral (VLEA) and ventromedial (VMEA) divisions, has reciprocal connections with the hippocampal formation and with the cortex, partly via the perirhinal cortex, and is connected with a number of subcortical structures such as the amygdala and the striatum.(ABSTRACT TRUNCATED AT 400 WORDS)

A forebrain atlas of the lizard Gekko gecko.

An atlas of the forebrain of the lizard Gekko gecko has been provided, which will serve as the basis for subsequent experimental tracing and immunohistochemical studies. Apart from a strongly developed medial cortex and septal area, the Tokay gecko shows all the main features of the forebrain of the lacertid-type lizards. When its convenience as an experimental animal is also taken into account, this species seems to be very suitable for studying the limbic system in reptiles. The atlas comprises topographical reconstructions of the telencephalon and diencephalon and a series of transverse sections of which the levels have been indicated in the reconstructions. The results obtained in the Gekko are briefly compared with those found in other lizards studied.

Efferent connections of the prelimbic (area 32) and the infralimbic (area 25) cortices: an anterograde tracing study in the cat.

The projections from the caudal part of the medial frontal cortex, encompassing the prelimbic area (PL) and the infralimbic area (IL) (Brodmann's areas 32 and 25, respectively), were studied in the cat with the anterograde autoradiographic tracing technique. The results indicate that the projection fields of IL, in contrast to those of PL, are restricted almost exclusively to limbic structures. Whereas the major thalamic projections from PL reach the mediodorsal, anteromedial, and ventromedial nuclei, the medial part of the lateral posterior nucleus, and the parataenial and reticular nuclei, and weak projections from this area are directed to the nucleus reuniens and other midline nuclei, the nucleus reuniens is the major thalamic termination field of fibers arising from IL. Cortical areas that are reached by fibers originating in PL and, to a lesser degree, also in IL, include more rostral prefrontal areas (areas 8, 6, and 12), the agranular insular, and the rostral perirhinal cortices. In contrast, cortical areas that are more strongly related to IL include the cingulate, retrosplenial, caudal entorhinal, and perirhinal cortices and the subiculum of the hippocampal formation. Another prominent output of PL concerns projections to an extensive medial part of the caudate nucleus and the ventral striatum, whereas fibers from IL only distribute most ventrally in the striatum. In the amygdaloid complex, fibers from PL were found to reach the basolateral, basomedial, and central nuclei, and fibers from IL to distribute to the medial and central nuclei. PL furthermore projects to the claustrum and the endopiriform nucleus. Other structures in the basal forebrain, including the medial septum, the nuclei of the diagonal band, the preoptic area, and the lateral and dorsal hypothalamus are densely innervated by IL and only sparsely by PL. With respect to more caudal parts of the brainstem, projections from PL and IL appeared to be essentially similar. They reach the ventral tegmental area, the periaqueductal gray, the parabrachial nucleus, and in cases of PL injections were followed as far caudally as the pons.

Reciprocal connections of the insular and piriform claustrum with limbic cortex: an anatomical study in the cat.

The connections of the claustrum with non-isocortical limbic and paralimbic cortex in the cat are described, using the anterograde transport of tritiated amino acids and the retrograde transport of various fluorescent tracers and of horseradish peroxidase conjugated to the lectin wheatgerm agglutinin. It could be demonstrated that the claustrum, in addition to its connections with sensory-related areas, is reciprocally and bilaterally connected with widespread limbic and paralimbic cortical regions. These connections are organized such that the area of origin of claustral efferents to a certain cortical region coincides with the area of termination in the claustrum of afferents from that same cortical region. A rostrocaudal topographical organization of the limbic-related connections of the claustrum is not very apparent. However, the results clearly demonstrate a dorsoventral topographical organization in the connections between the claustrum and the cortex. The ventral part of the claustrum has reciprocal connections predominantly with the entorhinal cortex, and possibly with the anterior olfactory nucleus and the prepiriform cortex. A more dorsally located part of the claustrum is preferentially connected with the orbitofrontal, the insular, the perirhinal, the anterior limbic, and the cingular cortices, and with parts of the subicular complex. The most dorsal portion of the claustrum is more heavily connected with parasensory and sensory cortices. It is concluded that the traditional subdivision of the claustrum into two discrete nuclei, i.e. the insular claustrum connected with the isocortex, and the piriform claustrum or endopiriform nucleus connected with the allocortex, does not reflect the actual organization of the cortical connections of the claustrum. The present data provide a more differentiated view, such that the ventral portion of the claustrum is reciprocally connected mainly with the olfactory-related cortices and the entorhinal cortex, whereas the cortical connections of progressively more dorsal parts of the claustrum gradually shift from limbic and paralimbic towards parasensory and sensory cortical connections. The significance of these findings is discussed in the light of a possible function of the claustrum in relation to corticocortical integration and memory processing.

Cortical afferents of the nucleus accumbens in the cat, studied with anterograde and retrograde transport techniques.

The cortical afferentation of the nucleus accumbens in the cat was studied with the aid of retrograde tracing techniques. Retrograde experiments were carried out with horseradish peroxidase or one of the fluorescent tracers Bisbenzimid, Nuclear Yellow and Fast Blue. In the anterograde experiments [3H]leucine and [35S]methionine were used as tracers. Following injections in the nucleus accumbens, retrogradely-labelled cells were found in the medial frontal cortex, the anterior olfactory nucleus, the posterior part of the insular cortex, the endopiriform nucleus, the amygdalo-hippocampal area, the entorhinal and perirhinal cortices and the subiculum of the hippocampal formation. In the medial frontal cortex most of the labelled cells were found in layers III and V of the prelimbic area (area 32 of Brodmann), but retrogradely-filled neurons were also present in the infralimbic area and in the caudoventral part of the lateral bank of the proreal gyrus. Retrogradely-labelled cells in the entorhinal and perirhinal cortices were located in the deep cellular layers. Following large injections in the nucleus accumbens, retrograde labelling in the subiculum extended from the most dorsal, septal pole to the most ventral, temporal pole. Injections of anterograde tracers were placed in the frontal cortex, the entorhinal and perirhinal cortices and the hippocampal formation. The prelimbic area was found to project via the internal capsule to mainly the rostral half of the nucleus accumbens, whereas in the caudal half of the nucleus only a lateral region receives frontal cortical fibres. Following injections in the infralimbic area only fibres passing through the nucleus accumbens were labelled. Afferents from the entorhinal and perirhinal cortices reach the nucleus accumbens by way of the external capsule and terminate mainly in a ventral zone of the nucleus accumbens. Afferents from the entorhinal area are distributed to the entire accumbens, whereas the termination field of the perirhinal afferents is largely restricted to the lateral part of the nucleus accumbens. Both the frontal cortex and the entorhinal and perirhinal cortices appear to project also to the nucleus caudatus and the tuberculum olfactorium. These cortical areas also project to the contralateral striatum. Both anterograde and retrograde tracing experiments demonstrated a topographical relationship between the subiculum and the nucleus accumbens. The ventral pole of the subiculum projects via the fornix to the medial part of the caudal half of the nucleus accumbens and to a small dorsomedial area in its rostral half. Successively more dorsal portions in the subiculum project to successively more ventrolateral parts in the rostral nucleus accumbens. The projection from the hippocampus was found to extend also to the tuberculum olfactorium. The results of the present study do not provide unambiguous criteria for the delimitation of the nucleus accumbens in the cat.

The anatomical relationship of the prefrontal cortex with the striatopallidal system, the thalamus and the amygdala: evidence for a parallel organization.

Recent findings in primates indicate that the connections of the frontal lobe, the basal ganglia, and the thalamus are organized in a number of parallel, functionally segregated circuits. In the present account, we have focused on the organization of the connections between the prefrontal cortex, the basal ganglia and the mediodorsal thalamic nucleus in the rat. It is concluded that in this species, in analogy with the situation in primates, a number of parallel basal ganglia-thalamocortical circuits exist. Furthermore, data are presented indicating that the projections from particular parts of the amygdala and from individual nuclei of the midline and intralaminar thalamic complex to the prefrontal cortex and the striatum are in register with the arrangements in the parallel circuits. These findings emphasize that the functions of the different subregions of the prefrontal cortex cannot be considered separately but must be viewed as components of the integrative functions of the circuits in which they are involved.

Further studies on the cortical connections of the Tegu lizard.

The efferent fiber connections of the caudal half of the cerebral cortex, the lateral cortex and the pallial thickening were studied using the Nauta-Gygax and Fink-Heimer techniques. The following observations were made, (1) In the caudal half of the hemisphere corticoseptal and corticohypothalamic fibers originate from the small-celled part of the mediodorsal cortex and the thickened caudal part of the dorsal cortex in its whole mediolateral extent. (2) The dorsal cortex in the middle of the hemisphere projects by way of both the pre- and postcommissural fornices. Its rostral pole distributes its fibers solely to the postcommissural fornix, whereas its caudal part projects via the precommissural fornix. (3) The posterior pallial commissure carries fibers that arise caudally in the small-celled part of the mediodorsal cortex and terminate in the contralateral ventral cortex. (4) Projections to the dorsal striatum originate from the lateral cortex, the dorsal cortex and the superficial portion of the pallial thickening. In addition, the latter two zones project to the nucleus accumbens. (5) The deep portion of the pallial thickening projects to the ventral striatum.

Mixed-junction axon terminals on identified principal abducens motoneurons in the monitor lizard.

Motoneurons in the principal abducens nucleus of the monitor lizard Varanus exanthematicus were identified by retrograde labeling following application of horseradish peroxidase to the abducens nerve. The ultrastructure and synaptology of thirty labeled neurons were studied. We observed a type of axon terminal which forms mixed junctions with the cell bodies and the initial axon segments of labeled motoneurons. The juxtaposed membranes of the terminals and the motoneurons display gap junctions and small asymmetric synaptic specializations. The mixed-junction terminals contain spherical synaptic vesicles which are located immediately adjacent to the synaptic junction. They may originate from local circuit neurons or from neurons extrinsic to the principal abducens nucleus.

Discriminant analysis of the localization of aggression-inducing electrode placements in the hypothalamus of male rats.

Over 400 sites in the hypothalami of 270 male CPB/WE-zob rats were electrically stimulated in order to induce fights between males. The localization of electrodes inducing fights seems to differ from the localization of electrodes in which no fights can be induced. The differences in localization were detected and tested by a non-parametric discriminant analysis. The results were plotted by computer in a stereotaxic atlas of the hypothalamus of the CPB/WE strain. The method delimits areas within the hypothalamus where the probability to induce aggression is high, intermediate or low. Moreover, the procedure allows discrimination between areas where the thresholds for attack behaviour are generally lower than elsewhere and where the fiercest forms of attack are induced. None of the areas delimited coincide with a classical subdivision of the hypothalamus. Parts of the perifornical, anterior, lateral and ventromedial hypothalamus seem to be involved. The methods developed here may help to relate stimulation-induced aggression to other characteristics of the 'aggressive' area which cannot be obtained directly from fighting rats such as cytological, endocrinological, biochemical or physiological data. In addition, the procedure may help to settle disputes on the specificity of the localization of neural substrates of other stimulation-induced behaviours. The methods to discriminate between overlapping 3-dimensional reconstructions validated here for aggressive responses, can also be applied to other types of stereotaxic data and other types of effects, such as electrical, hormonal or other physiological responses. They may be especially useful if the localization of the neural population involved is not yet known, and unknown current-spread or diffusion of substances complicates the interpretation of stereotaxic data.