Unconditioned stimulus pathways to the amygdala: effects of lesions of the posterior intralaminar thalamus on foot-shock-induced c-Fos expression in the subdivisions of the lateral amygdala.

The lateral nucleus of the amygdala (LA) is a site of convergence for auditory (conditioned stimulus) and foot-shock (unconditioned stimulus) inputs during fear conditioning. The auditory pathways to LA are well characterized, but less is known about the pathways through which foot shock is transmitted. Anatomical tracing and physiological recording studies suggest that the posterior intralaminar thalamic nucleus, which projects to LA, receives both auditory and somatosensory inputs. In the present study we examined the expression of the immediate-early gene c-fos in the LA in rats in response to foot-shock stimulation. We then determined the effects of posterior intralaminar thalamic lesions on foot-shock-induced c-Fos expression in the LA. Foot-shock stimulation led to an increase in the density of c-Fos-positive cells in all LA subnuclei in comparison to controls exposed to the conditioning box but not shocked. However, some differences among the dorsolateral, ventrolateral and ventromedial subnuclei were observed. The ventrolateral subnucleus showed a homogeneous activation throughout its antero-posterior extension. In contrast, only the rostral aspect of the ventromedial subnucleus and the central aspect of the dorsolateral subnucleus showed a significant increment in c-Fos expression. The density of c-Fos-labeled cells in all LA subnuclei was also increased in animals placed in the box in comparison to untreated animals. Unilateral electrolytic lesions of the posterior intralaminar thalamic nucleus and the medial division of the medial geniculate body reduced foot-shock-induced c-Fos activation in the LA ipsilateral to the lesion. The number of c-Fos labeled cells on the lesioned side was reduced to the levels observed in the animals exposed only to the box. These results indicate that the LA is involved in processing information about the foot-shock unconditioned stimulus and receives this kind of somatosensory information from the posterior intralaminar thalamic nucleus and the medial division of the medial geniculate body.

Role of the vomeronasal system in intersexual attraction in female mice.

Although it is generally accepted that rodents' sociosexual behavior relies mainly on chemosignals, the specific roles played by the vomeronasal and olfactory systems in detecting these signals are presently unclear. This work reports the results of three experiments aimed at clarifying the role of the vomeronasal system on gender recognition and intersexual attraction, by analyzing the effects of lesions of the accessory olfactory bulbs (AOB) in chemically naïve female mice. The first experiment demonstrates that lesions of the AOB abolish the preference that females show for male-soiled bedding in tests in which the females can contact the bedding, thus having access to both volatile and involatile male chemosignals. The second experiment shows that airborne male-derived chemosignals are not attractive to intact, chemically naïve females but tend to be preferentially explored by females whose AOB has been lesioned. However, repeated exposure to male-soiled bedding has opposite effects in sham-operated and AOB-lesioned female mice. Whereas after this experience sham-operated females show an (acquired) attraction toward male airborne chemosignals, in AOB-lesioned females the same experience makes male-derived volatiles aversive. Finally, in the third experiment we have confirmed that our AOB-lesioned females are able to detect urine-borne male odorants, as well as to discriminate them from the synthetic terpene geraniol. These findings strongly suggest that in mice, the involatile male sexual pheromone that is intrinsically attractive is detected by the vomeronasal system of the females. In addition, the repeated experience of females with male-soiled bedding would probably allow the association of this pheromone, acting as unconditioned stimulus, with olfactory stimuli (odorants) that therefore would become conditioned attractors to the females.

Vomeronasal inputs to the rodent ventral striatum.

Vertebrates sense chemical signals through the olfactory and vomeronasal systems. In squamate reptiles, which possess the largest vomeronasal system of all vertebrates, the accessory olfactory bulb projects to the nucleus sphericus, which in turn projects to a portion of the ventral striatum known as olfactostriatum. Characteristically, the olfactostriatum is innervated by neuropeptide Y, tyrosine hydroxylase and serotonin immunoreactive fibers. In this study, the possibility that a structure similar to the reptilian olfactostriatum might be present in the mammalian brain has been investigated. Injections of dextran-amines have been aimed at the posteromedial cortical amygdaloid nucleus (the putative mammalian homologue of the reptilian nucleus sphericus) of rats and mice. The resulting anterograde labeling includes the olfactory tubercle, the islands of Calleja and sparse terminal fields in the shell of the nucleus accumbens and ventral pallidum. This projection has been confirmed by injections of retrograde tracers into the ventral striato-pallidum that render retrograde labeling in the posteromedial cortical amygdaloid nucleus. The analysis of the distribution of neuropeptide Y, tyrosine hydroxylase, serotonin and substance P in the ventral striato-pallidum of rats, and the anterograde tracing of the vomeronasal amygdaloid input in the same material confirm that, similar to reptiles, the ventral striatum of mammals includes a specialized vomeronasal structure (olfactory tubercle and islands of Calleja) displaying dense neuropeptide Y-, tyrosine hydroxylase- and serotonin-immunoreactive innervations. The possibility that parts of the accumbens shell and/or ventral pallidum could be included in the mammalian olfactostriatum cannot be discarded.

Unconditioned stimulus pathways to the amygdala: effects of posterior thalamic and cortical lesions on fear conditioning.

Plasticity in the lateral nucleus of the amygdala is thought to be critical for the acquisition of Pavlovian fear conditioning. The pathways that transmit auditory conditioned stimulus information originate in auditory processing regions of the thalamus and cortex, but the pathways mediating transmission of unconditioned stimuli to the amygdala are poorly understood. Recent studies suggest that somatosensory (footshock) unconditioned stimulus information is also relayed in parallel to the lateral nucleus of the amygdala from the thalamus (the posterior intralaminar thalamic complex, PIT) and the cortex (parietal insular cortex). In the present study we reexamined this issue. Our results showed that bilateral electrolytic lesions of the PIT alone blocked fear conditioning, whereas bilateral excitotoxic PIT lesions had no effect. These electrolytic PIT lesions did not affect fear conditioning using a loud noise as unconditioned stimulus, defining the effects of PIT lesions as a disruption of somatosensory as opposed to auditory processing. Finally, we performed combined bilateral excitotoxic lesions of the PIT nuclei and electrolytic lesions of the parietal insular cortex. These, like excitotoxic lesions of PIT alone, had no effect on the acquisition of fear conditioning. Thus, somatosensory regions of the thalamus and cortex may well be important routes of unconditioned stimulus transmission to the amygdala in fear conditioning, but information about the unconditioned somatosensory stimulus is also transmitted from other sources that send fibers through, but do not form essential synapses in, the thalamus en route to the amygdala.

Distribution of PSA-NCAM expression in the amygdala of the adult rat.

Synaptic plasticity in the amygdala appears to be necessary for the generation of emotional memories. However, the molecular bases of this plasticity are not fully understood. Because the polysialylated form of the neural cell adhesion molecule (PSA-NCAM) has been implicated in memory consolidation in the hippocampus and temporal cortex, we have studied in detail the expression of this molecule in the adult rat amygdala with an antibody against PSA-NCAM. Our results demonstrate for the first time the presence of PSA-NCAM in the adult rat amygdala. Immunoreactive somata and processes are abundant in the amygdalo-hippocampal transition area, central nucleus, intra-amygdaloid bed nucleus of the stria terminalis, anterior and posterior cortical nuclei, periamygdaloid cortex and medial nucleus of the amygdala. In addition PSA-NCAM immunoreactive neuronal somata and processes exist in the lateral, basal and accessory basal nuclei, anterior amygdaloid area and amygdalo-striatal area. The presence of this molecule in areas that receive olfactory or vomeronasal input could reflect the intrinsic plasticity of these chemosensory systems. PSA-NCAM expression in the lateral amygdala could indicate its participation in the plastic events that lead to the generation of emotional memories such as those related to fear conditioning.

Striato-amygdaloid transition area lesions reduce the duration of tonic immobility in the lizard Podarcis hispanica.

Neuroanatomical data suggest that the lizard striato-amygdaloid transition area is homologous with the mammalian central amygdala. In order to investigate possible functional similarities, tonic immobility was induced in adult lizards and its duration recorded. Each lizard was then randomly assigned to one of three treatments: (1) bilateral striato-amygdaloid transition area lesions, (2) bilateral dorsal cortex lesions or (3) untreated controls. Three days after trial 1, each lizard was subjected to a second trial and the tonic immobility duration recorded. The mean tonic immobility duration in lizards with striato-amygdaloid transition area lesions was significantly shorter (80.5%; p < 0.0033) in trial 2 than in trial 1. There were no inter-trial differences within dorsal cortex-lesioned lizards or untreated controls. There was a significant treatment effect on tonic immobility duration in trial 2 (p < 0.0001). The mean tonic immobility duration of lizards with striato-amygdaloid transition area lesions was significantly shorter than that of dorsal cortex-lesioned lizards (72.2%; p < 0.01) or untreated controls (78.2%; p < 0.01). There was no significant difference in mean tonic immobility duration between dorsal cortex-lesioned lizards and untreated controls. Tonic immobility is considered to be an anti-predator behaviour that reflects the underlying state of fear. Therefore, the reduced tonic immobility duration in lizards with striato-amygdaloid transition area lesions reflects a reduction of fear. These results provide the first data to indicate a functional similarity between the lizard striato-amygdaloid transition area and the mammalian central amygdala.

Colecistitis aguda litiásica asociada a brucella melitensis. Presentación de un caso y revisión de la bibliografía / Acute lithiasic cholecystitis associated with Brucella melitensis. Case report and review of the literature

Aportamos el segundo caso de colecistitis aguda litiásica durante el curso de una brucelosis sistémica, según la revisión de la bibliografía mundial. Se describe su semiología clínica y tratamiento. Se resalta la importancia de considerar dicha enfermedad, como concomitante a otras, en zonas geográficas consideradas endémicas para la brucelosis (AU)

Distribution of CGRP-like immunoreactivity in the chick and quail brain.

Calcitonin gene-related peptide (CGRP)-containing neurones have been implicated in the transmission of visceral sensory information to the cortex and in the control of arterial blood pressure in mammals. However, little is known about its function in other vertebrates. As a first step toward investigating the function of CGRP in birds, its distribution was studied in the domestic chick and quail brain by means of immunocytochemistry, by using antibodies against rat CGRP. The distribution of CGRP immunoreactivity in the chick and quail central nervous system was found to be similar. CGRP-immunoreactive (CGRPi) perikarya were not present in the telencephalon. In the diencephalon, CGRPi perikarya were present mainly in the shell of the thalamic nucleus ovoidalis, the nucleus semilunaris paraovoidalis, the nucleus dorsolateralis posterior thalami, and in the hypothalamic nucleus of the ansa lenticularis. In the brainstem, CGRPi perikarya were present in the nucleus mesencephalicus nervi trigemini, the nucleus tegmenti ventralis, the locus coeruleus, the nucleus linearis caudalis and in the parabrachial region. In addition CGRPi perikarya were found in the motor nuclei of the III, IV, V, VI, VII, IX, X, and XII cranial nerves. The telencephalon contained CGRPi fibres within the paleostriatal complex (mainly in the ventral paleostriatum), parts of the neostriatum and ventral hyperstriatum, parts of the archistriatum, and the septum. In the diencephalon, the densest plexus of CGRPi fibres was observed in the dorsal reticular thalamus. A less dense CGRPi innervation was present in some dorsal thalamic nuclei and in the medial and periventricular hypothalamus. The pretectum and midbrain tegmentum also contained CGRPi fibres, whereas the optic tectum was virtually devoid of immunolabelling. Scattered CGRPi fibres were observed in the central grey and neighbouring pontine areas. Some of the sensory fibres of the trigeminal, vagal, glossopharyngeal, and spinal nerves were also CGRPi. The results of comparative studies indicate that the presence of CGRP in some thalamo-telencephalic projections is a primitive feature of the forebrain of amniotes. Therefore, the brain areas giving rise to and receiving such a projection in different vertebrates, are likely to be homologous.

Organization of the ophidian amygdala: chemosensory pathways to the hypothalamus.

Although recent studies in squamate reptiles have importantly clarified how chemical information is processed in the reptilian brain, how the amygdala relays chemosensory inputs to the hypothalamus to influence chemically guided behaviors is still poorly documented. To identify these chemosensory pathways, the amygdalo-hypothalamic projections, intra-amygdaloid circuitry and afferents from the lateral cortex (LC) to the amygdala were investigated by injecting conjugated dextran-amines into the hypothalamus, amygdala, and LC of garter snakes. The amygdala was divided into olfactory recipient (ventral anterior and external amygdalae), vomeronasal recipient (nucleus sphericus, NS, and medial amygdala, MA), and nonchemosensory (e.g., posterior dorsal ventricular ridge, PDVR, and dorsolateral amygdaloid nucleus, DLA) subdivisions. Rostroventral (LCrv) and dorsocaudal subdivisions of the LC were distinguished. In addition to receiving afferents from the main olfactory bulb, the olfactory amygdala receives afferents from NS and projects to the NS, PDVR, and dorsal hypothalamus. The NS has only a minor projection to the lateral hypothalamus, whereas the MA, which receives afferents from the LCrv and NS, has projections to the ventromedial hypothalamic (VMH) and lateral posterior hypothalamic nuclei. Among the nonchemosensory amygdaloid structures, the PDVR receives afferents from the LCrv and the olfactory amygdala and projects to the VMH, whereas DLA receives afferents from the LCrv and NS, and projects to the periventricular hypothalamus. These results substantially clarify the olfactory and vomeronasal tertiary connections and demonstrate that parts of the nonchemosensory amygdala play a major role in relaying chemosensory information to the hypothalamus.

Afferents to the red nucleus in the lizard Podarcis hispanica: putative pathways for visuomotor integration.

The afferents to the red nucleus from visual and nonvisual forebrain centers have been investigated in the lizard Podarcis hispanica by using both retrograde and anterograde transport of tracers. Because the red nucleus constitutes a key structure in the limb premotor system, these sensory afferents probably are involved in visuomotor and other forms of sensorimotor integration. After tracer injections aimed at the red nucleus, retrograde labeling was found in the reticular thalamus, the subthalamus, the nucleus of the posterior commissure, as well as in two retinorecipient nuclei, namely, the ventral lateral and pretectal geniculate nuclei, where labeled cells are especially abundant. These geniculorubral projections have been confirmed by means of anterograde tracing with dextranamine injections. On the other hand, small injections of tracers in the retina demonstrated that its projections to the ventral lateral and pretectal geniculate nuclei are organized in a point-to-point fashion. Moreover, small tracer injections into the optic tectum of Podarcis indicated that the ventral lateral geniculate nucleus also receives a precisely organized tectal afferent. Taken together, these results strongly suggest that geniculorubral projections might constitute the neuroanatomical substrate for the generation of quick locomotor responses to appropriate visual stimuli. Additional ventral thalamic, subthalamic, and pretectal afferents to the red nucleus are likely to subserve other kinds of sensorimotor integration. These results help to clarify the organization of the reptilian motor system, including the telencephalic control of motor responses, and to unravel some of the major trends in the evolution of the limb premotor network of tetrapodian vertebrates.

Septal complex of the telencephalon of lizards: III. Efferent connections and general discussion.

The projections of the septum of the lizard Podarcis hispanica (Lacertidae) were studied by combining retrograde and anterograde neuroanatomical tracing. The results confirm the classification of septal nuclei into three main divisions. The nuclei composing the central septal division (anterior, lateral, medial, dorsolateral, and ventrolateral nuclei) displayed differential projections to the basal telencephalon, preoptic and anterior hypothalamus, lateral hypothalamic area, dorsal hypothalamus, mammillary complex, dorsomedial anterior thalamus, ventral tegmental area, interpeduncular nucleus, raphe nucleus, torus semicircularis pars laminaris, reptilian A8 nucleus/substantia nigra and central gray. For instance, only the medial septal nucleus projected substantially to the thalamus whereas the anterior septum was the only nucleus projecting to the caudal midbrain including the central gray. The anterior and lateral septal nuclei also differ in the way in which their projection to the preoptic hypothalamus terminated. The midline septal division is composed of the dorsal septal nucleus, nucleus septalis impar and nucleus of the posterior pallial commissure. The latter two nuclei projected to the lateral habenula and, at least the nucleus of the posterior pallial commissure, to the mammillary complex. The dorsal septal nucleus projected to the preoptic and periventricular hypothalamus and the anterior thalamus, but its central part seemed to project to the caudal midbrain (up to the midbrain central gray). Finally, the ventromedial septal division (ventromedial septal nucleus) showed a massive projection to the anterior and the lateral tuberomammillary hypothalamus. Data on the connections of the septum of P. hispanica and Gecko gekko are discussed from a comparative point of view and used for better understanding of the functional anatomy of the tetrapodian septum.

Identification of the reptilian basolateral amygdala: an anatomical investigation of the afferents to the posterior dorsal ventricular ridge of the lizard Podarcis hispanica.

The presence of multimodal association in the telencephalon of reptiles has been investigated by tracing the afferent connections to the posterior dorsal ventricular ridge (PDVR) of the lizard Podarcis hispanica. The PDVR receives telencephalic afferents from the lateral (olfactory) and dorsal cortices, and from the three unimodal areas of the anterior dorsal ventricular ridge, in a convergent manner. From the diencephalon, it receives afferents from the dorsomedial anterior and medial posterior thalamic nuclei, and from several hypothalamic nuclei. Brainstem afferents to the PDVR originate in the dorsal interpeduncular nucleus, the nucleus of the lateral lemniscus and parabrachial nucleus. The afferents to the thalamic nuclei that project to the PDVR have also been studied. The dorsomedial anterior thalamic nucleus receives projections mainly from limbic structures, whereas the medial posterior thalamic nucleus is the target of projections from structures with a clear sensory significance (optic tectum, torus semicircularis, nuclei of the lateral and spinal lemniscus, superior olive and trigeminal complex). As a result, the PDVR appears as an associative centre that receives visual, auditory, somatosensory and olfactory information from several telencephalic and non-telencephalic centres, and a multimodal projection from the medial posterior thalamic nucleus. This pattern of afferents of the PDVR is similar to that of the caudal neostriatum in birds and the basolateral division of the mammalian amygdala. These results indicate that a multimodal amygdala is already present in reptiles, and has probably played a key role in the evolution of the vertebrate brain.

Ascending projections from the optic tectum in the lizard Podarcis hispanica.

The ascending projections of the optic tectum, including their cells of origin, have been studied in the lizard Podarcis hispanica by means of a two-step experimental procedure. First, tracers were injected in the tectum to study the anterograde labeling in the forebrain. Second, the cells of origin of these projections have been identified by analyzing the retrograde labeling after tracer injections in the thalamus, hypothalamus, and pretectum. Three main tectal ascending pathways have been described: the dorsal tecto-thalamic tract (dtt), the medial tecto-thalamic tract (mtt), and the ventral tecto-thalamic tract (vtt). The dtt originates in radial cells of layers 5 and 7 and bipolar cells of layers 8 and 10 that project to the lateral neuropile of the dorsal lateral geniculate nucleus (GLD), to the intergeniculate leaflet (IGL), and to the ventral lateral geniculate nucleus (GLV). The mtt arises from radial neurons of layers 3 and 5 and bilaterally reaches the putative reticular thalamus and its boundary with the hypothalamus, the rostral IGL, and the area triangularis (AT). The vtt is composed of fibers from ganglion and multipolar cells of the layer 7 that project bilaterally to the nucleus of the vtt, the ventrolateral thalamic nucleus, the medial posterior thalamic nucleus (MP), the nucleus rotundus (Rot), the IGL, and the cell plate of the GLD. Therefore, the GLD receives not only direct retinal afferents but also two different tectal inputs, thus constituting a convergence point in the two visual pathways to the telencephalon. Moreover, different tectal cells specifically project to the ventrolateral thalamus and to pretectal nuclei. These results are discussed from comparative and functional viewpoints.

Portal hypertensive colopathy: histologic appearance of the colonic mucosa.

BACKGROUND/AIMS: The aim of this study was to evaluate macroscopic and histologic disorders of the colon in rats with portal hypertension. METHODOLOGY: The animals were divided into two groups: 10 rats in the experimental group and 10 controls. Portal hypertension was induced by a partial ligation of the portal vein. We assessed the histologic appearance and macroscopic alterations of the colonic mucosa 12 weeks after the induction of portal hypertension. RESULTS: The macroscopic results showed the presence of the typical signs of portal hypertension, with dilatation and tortuosity of the mesenteric veins and hyperemia of the rectum and colon. Histologic study revealed a marked dilatation in the microcirculation at the level of the ascending colon (20 +/- 2 micra in diameter) and rectum (30 +/- 4), with a tendency to group the lesions. Submucosal edema without inflammatory signs was observed. CONCLUSION: The existence of these lesions in the colon and rectum can be the cause of hemorrhage, as in the gastropathy of portal hypertension and should be considered in cases of undiagnosed hemorrhage in patients with portal hypertension.

Efferents and centrifugal afferents of the main and accessory olfactory bulbs in the snake Thamnophis sirtalis.

This study reinvestigates the efferent projections of the main and accessory olfactory bulbs and describes for the first time the centrifugal projections to the main and accessory olfactory bulbs in the snake Thamnophis sirtalis, using the intraaxonal transport of the anterograde and retrograde tracer biotinylated dextran amine and the retrograde tracer horseradish peroxidase. The olfactory projection consists of three tracts: the lateral olfactory tract, which projects bilaterally to the lateral cortex and the rostral amygdala, crossing the midline through the stria medullaris-habenular commissure system; the intermediate olfactory tract, which projects ipsilaterally to the olfactory tubercle and contributes to the contralateral projection; and the medial olfactory tract, which projects ipsilaterally to the dorsomedial retrobulbar formation. The vomeronasal projection is formed by a single tract, the accessory olfactory tract, that projects ipsilaterally to the nucleus of the accessory olfactory tract, the medial amygdala and the nucleus sphericus. The centrifugal projections to the main and accessory olfactory bulb are composed of two components: one that arises in areas that receive the olfactory or vomeronasal input (neurons in the olfactory tubercle, retrobulbar formation and lateral cortex project to the main olfactory bulb; and neurons in the nucleus of the accessory olfactory tract, the medial amygdala and the nucleus sphericus project to the accessory olfactory bulb), and another that arises in areas not directly implicated in processing the chemosensory information (the nucleus of the diagonal band of Broca and the dorsal cortex). These data allow the recognition of the general pattern of organization of the reptilian olfactory and vomeronasal systems.

Afferent and efferent connections of the nucleus sphericus in the snake Thamnophis sirtalis: convergence of olfactory and vomeronasal information in the lateral cortex and the amygdala.

This paper is an account of the afferent and efferent projections of the nucleus sphericus (NS), which is the major secondary vomeronasal structure in the brain of the snake Thamnophis sirtalis. There are four major efferent pathways from the NS: 1) a bilateral projection that courses, surrounding the accessory olfactory tract, and innervates several amygdaloid nuclei (nucleus of the accessory olfactory tract, dorsolateral amygdala, external amygdala, and ventral anterior amygdala), the rostral parts of the dorsal and lateral cortices, and the accessory olfactory bulb; 2) a bilateral projection that courses through the medial forebrain bundle and innervates the olfactostriatum (rostral and ventral striatum); 3) a commissural projection that courses through the anterior commissure and innervates mainly the contralateral NS; and 4) a meager bilateral projection to the lateral hypothalamus. On the other hand, important afferent projections to the NS arise solely in the accessory olfactory bulb, the nucleus of the accessory olfactory tract, and the contralateral NS. This pattern of connections has three important implications: first, the lateral cortex probably integrates olfactory and vomeronasal information. Second, because the NS projection to the hypothalamus is meager and does not reach the ventromedial hypothalamic nucleus, vomeronasal information from the NS is not relayed directly to that nucleus, as previously reported. Finally, a structure located in the rostral and ventral telencephalon, the olfactostriatum, stands as the major tertiary vomeronasal center in the snake brain. These three conclusions change to an important extent our previous picture of how vomeronasal information is processed in the brain of reptiles.

Amygdalo-hypothalamic projections in the lizard Podarcis hispanica: a combined anterograde and retrograde tracing study.

The cells of origin and terminal fields of the amygdalo-hypothalamic projections in the lizard Podarcis hispanica were determined by using the anterograde and retrograde transport of the tracers, biotinylated dextran amine and horseradish peroxidase. The resulting labeling indicated that there was a small projection to the preoptic hypothalamus, that arose from the vomeronasal amygdaloid nuclei (nucleus sphericus and nucleus of the accessory olfactory tract), and an important projection to the rest of the hypothalamus, that was formed by three components: medial, lateral, and ventral. The medial projection originated mainly in the dorsal amygdaloid division (posterior dorsal ventricular ridge and lateral amygdala) and also in the centromedial amygdaloid division (medial amygdala and bed nucleus of the stria terminalis). It coursed through the stria terminalis and reached mainly the retrochiasmatic area and the ventromedial hypothalamic nucleus. The lateral projection originated in the cortical amygdaloid division (ventral anterior and ventral posterior amygdala). It coursed via the lateral amygdalofugal tract and terminated in the lateral hypothalamic area and the lateral tuberomammillary area. The ventral projection originated in the centromedial amygdaloid division (in the striato-amygdaloid transition area), coursed through the ventral peduncle of the lateral forebrain bundle, and reached the lateral posterior hypothalamic nucleus, continuing caudally to the hindbrain. Such a pattern of the amygdalo-hypothalamic projections has not been described before, and its functional implications in the transfer of multisensory information to the hypothalamus are discussed. The possible homologies with the amygdalo-hypothalamic projections in mammals and other vertebrates are also considered.

Septal complex of the telencephalon of the lizard Podarcis hispanica. II. Afferent connections.

The afferent connections to the septal complex were studied in the lizard Podarcis hispanica (Lacertidae) by means of a combination of retrograde and anterograde tracing. The results of these experiments allow us to classify the septal nuclei into three main divisions. The central septal division (anterior, lateral, dorsolateral, ventrolateral, and medial septal nuclei plus the nucleus of the posterior pallial commissure) receives a massive, topographically organized, cortical projection (medial, dorsal, and ventral areas) and widespread afferents from the tuberomammillary hypothalamus and the basal telencephalon. Moreover, it receives discrete projections from the dorsomedial anterior thalamus, the ventral tegmentum, the midbrain raphe, and the locus coeruleus. The ventromedial septal division (ventromedial septal nucleus) receives a massive projection from the anterior hypothalamus, dense serotonergic innervation, and a faint amygdalohypothalamic projection, but it is devoid of direct cortical input. The midline septal division (nucleus septalis impar and dorsal septal nucleus) receives a nontopographic cortical projection (dorsomedial and dorsal cortices) and afferents from the preoptic hypothalamus, the dorsomedial anterior thalamus, the midbrain central gray, and the reptilian A8 nucleus/substantia nigra. Our results indicate that the cortex provides a physiologically complex, massive input to the septum that terminates over the whole dendritic tree of septal cells. In contrast, most of the ascending afferents make axosomatic contacts by means of pericellular nests. The chemical nature of the main septal afferents and the comparative implications of the available hodological data on the organization of the septal complex of tetrapod vertebrates are discussed.