Overview of the main and accessory olfactory bulb projections in reptiles.

The present account is a review of the main and accessory olfactory bulb projections in reptiles. From previous studies by means of the classical degeneration techniques and recent studies using the autoradiographic method or the Phaseolus vulgarus-leucoagglutinin (PHA-L) tracing technique, it has become evident that, in the brain of reptiles, the vomeronasal and main olfactory systems are segregated. Moreover, the PHA-L studies have demonstrated that the main olfactory bulbs project not only to the anterior olfactory nucleus, the olfactory tubercle, and the lateral cortex, but also to the rostral portions of the external and central amygdaloid nuclei. The latter studies also provided evidence that, apart from a massive projection to the nucleus sphericus, at least in some reptilian species, the accessory olfactory bulbs project to the caudal portion of the central amygdaloid nucleus and the bed nucleus of the stria terminalis. A comparison of the olfactory projections between the various reptilian species studied revealed a considerable variation in the course of the main olfactory bulb efferents that reach the contralateral hemisphere.

Compartmental distribution of ventral striatal neurons projecting to the mesencephalon in the rat.

The ventral striatum is characterized by an intricate neurochemical compartmentation that is reflected in the distribution of most of its afferent fiber systems. In the present study, the compartmental relationships of ventral striatal neurons projecting to the mesencephalon were studied by combining tract tracing with the immunohistochemical localization of leu-enkephalin. Injections of the retrograde tracer cholera toxin subunit B were placed at various sites in the ventral mesencephalon. The anterograde tracer Phaseolus vulgaris leucoagglutinin was injected in single compartments in the rostrolateral part of the nucleus accumbens. The projections from the ventral striatum to the dopaminergic cell groups in the ventral mesencephalon and those to the substantia nigra pars reticulata originate from distinct subpopulations of ventral striatal neurons that respect neurochemically defined compartmental boundaries. In the "shell" of the nucleus accumbens, neurons that project to the dopaminergic cell groups are located outside areas of high cell density and weak enkephalin immunoreactivity (ENK-IR). Rostrolaterally in the "core" of the nucleus accumbens, neurons inside large areas of strong ENK-IR surrounding the anterior commissure project to the dorsomedial part of the substantia nigra pars reticulata, whereas neurons outside these areas innervate the ventral tegmental area and/or the medial part of the substantia nigra pars compacta. By contrast, more caudally in the dorsal part of the nucleus accumbens and in the ventral part of the caudate-putamen, the relationships are reversed: neurons in- or outside small patches of strong ENK-IR project respectively to the pars compacta or the pars reticulata of the substantia nigra. Since the thalamic and cortical afferents of the ventral striatum are compartmentally ordered as well, the present results imply that through the ventral striatal compartments information from disparate combinations of cortical and thalamic sources may be conveyed to distinct mesencephalic targets. The component of the ventral striatomesencephalic system reaching the dopaminergic cell groups A10, A9, and A8 may modulate the dopaminergic input to virtually the entire striatum. The other component can, by way of the pars reticulata of the substantia nigra, participate in nigrothalamic and nigrotectal output pathways of the basal ganglia.

Afferent connections of the striatum and the nucleus accumbens in the lizard Gekko gecko.

The afferent connections of the striatum and the nucleus accumbens of the lizard Gekko gecko were studied with retrograde tracing by means of horseradish peroxidase and Fluoro-Gold and with anterograde tracing by means of Phaseolus vulgaris leukoagglutinin. The striatum receives projections from the cortex, the dorsal ventricular ridge, the lateral amygdaloid nucleus, the globus pallidus, the anterior peduncular nucleus, the ventral tegmental area and substantia nigra, the area ventral to the substantia nigra, and the dorsal thalamus. The nucleus accumbens is projected upon by the cortex, the diagonal band, the ventral pallidum, the lateral preoptic area, the ventral tegmental area, and the dorsal thalamus. The source of the cortical projection to the striatum and the nucleus accumbens is a longitudinal zone in the dorsal cortex that, rostrally in the hemisphere, is located medially and, more caudally, in its middle one third. The medial and rostrolateral areas of the dorsal ventricular ridge each project to the striatum in a vertical zone. The fibers from the caudolateral area of the ridge end in two oblique bands located parallel to the border between the dorsal ventricular ridge and the striatum. The pathways from the mesencephalic tegmentum to the striatum and the nucleus accumbens show a medial to lateral topography. This is similar to the situation in birds, but contrary to that in mammals in which these pathways are extensively interconnected. The specific sensory nuclei of the dorsal thalamus were found to project not only to the dorsal ventricular ridge, but also, and in a topographical fashion, to the striatum. The dorsomedial thalamic nucleus, which innervates the dorsal ventricular ridge, has additional projections to the striatum and the nucleus accumbens. This projection pattern is similar to that of the intralaminar thalamic nuclei of birds and mammals.

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.

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.

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.

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.

The collateral ligaments of the knee joint in the cat and man. Morphological and functional study of the internal arrangement of fibers.

The present investigation of fiber arrangement in the collateral ligaments of the knee was carried out in cats and man in various positions of flexion and extension, without compression load. In all knee joint positions, the fibers of the collateral ligaments are twisted except for the fibers in the meniscal part of the medial collateral ligament which have a parallel arrangement. Furthermore, most of the fibers in the collateral ligaments are taut in all positions of the knee joint in both cat and man. By means of planar models representing different fiber arrangements, the kinematic behavior of the collateral ligaments was analyzed. It appears that a crossed (twisted) arrangement of the fibers is most effective in rotatory movements, whereas a parallel orientation is most effective in translation. Our data further indicate that, in measuring the changes in lengths of ligaments during joint motion, one cannot neglect the internal arrangement of fibers and the geometry of the articular surfaces and menisci.

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.

Mesotocin and vasotocin in the brain of the lizard Gekko gecko. An immunocytochemical study.

The distribution of mesotocin and vasotocin was studied in the brain of the lizard Gekko gecko with antisera specific for either peptide. Both mesotocinergic and vasotocinergic perikarya are found in the paraventricular and supraoptic nuclei of the hypothalamus, whereas vasotocinergic neurons are exclusively present in the bed nucleus of the stria terminalis and in a cell group of the rhombencephalon. The distributional pattern of the mesotocinergic fibers corresponds closely to that of the vasotocinergic fibers. However, throughout the entire brain the mesotocinergic innervation is less dense than the vasotocinergic innervation. No sex differences are present in the mesotocinergic fiber system.

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.

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.

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)

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.

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.

Inputs from the olfactory bulb and olfactory cortex to the entorhinal cortex in the cat. I. Anatomical observations.

The spatial organization and laminar distribution of projections from the olfactory bulb and the anterior (PPCa) and posterior (PPCp) divisions of the prepiriform cortex to the entorhinal cortex were studied with anterograde (3H-leucine) and retrograde (WGA-HRP) tracing techniques. After 3H-leucine injections into the olfactory bulb transported labeling was seen over the lateral entorhinal area, except its most medial part, and over the rostral part of the medial entorhinal area. The labeling covers exclusively layer Ia. The lateral and medial entorhinal areas are also reached by fibers from the prepiriform cortex. The projection to the medial entorhinal area has not been described previously. Following injections of 3H-leucine into the PPCa transported labeling is present over the entire expanse of the entorhinal cortex and is located over layer Ib with the greatest density in its superficial part. Injections of 3H-leucine into the PPCp give rise to transported labeling over much of the entorhinal cortex. No labeling was found over the most medial parts of the medial subdivision (VMEA) of the lateral entorhinal area and the medial entorhinal area. Labeling occupies layer Ib, especially its middle part, and layers II and III. Both PPCa and PPCp appear to project most heavily to the dorsal (DLEA) and ventral (VLEA) subdivisions of the lateral entorhinal area. From the retrograde experiments it can be inferred that cells of layers II and III of the PPCa project predominantly to the DLEA, whereas those of the PPCp project predominantly to the VLEA. The MEA receives its heaviest projection from layer II of both PPCa and PPCp.(ABSTRACT TRUNCATED AT 250 WORDS)