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The connections of the olfactory bulbs of Podarcis hispanica were studied by tract-tracing of injected horseradish peroxidase. Restricted injections into the main olfactory bulb (MOB) resulted in bilateral terminallike labeling in the medial part of the anterior olfactory nucleus (AON) and in the rostral septum, lateral cortex, nucleus of the lateral olfactory tract, and ventrolateral amygdaloid nucleus. Bilateral retrograde labeling was found in the rostral lateral cortex and in the medial and dorsolateral AON. Ipsilaterally the dorsal cortex, nucleus of the diagonal band, lateral preoptic area, and dorsolateral amygdala showed labeled cell bodies. Retrogradely labeled cells were also found in the midbrain raphe nucleus. Results from injections into the rostral lateral cortex and lateral olfactory tract indicate that the mitral cells are the origin of the centripetal projections of the MOB. Injections in the accessory olfactory bulb (AOB) produced ipsilateral terminallike labeling of the ventral AON, bed nucleus of the accessory olfactory tract, central and ventromedial amygdaloid nuclei, medial part of the bed nucleus of the stria terminalis, and nucleus sphericus. Retrograde labeling of neurons was observed ipsilaterally in the bed nucleus of the accessory olfactory tract and stria terminalis, in the central amygdaloid nucleus, dorsal cortex, and nucleus of the diagonal band. Bilateral labeling of somata was found in the ventral AON, the nucleus sphericus (hilus), and in the mesencephalic raphe nucleus and locus coeruleus. Injections into the dorsal amygdala showed that the mitral neurons are the cells of origin of the AOB centripetal projections. Reciprocal connections are present between AOB and MOB. To our knowledge, this is the first study to address the afferent connections of the olfactory bulbs in a reptile. On the basis of the available data, a discussion is provided of the similarities and differences between the reptilian and mammalian olfactory systems, as well as of the possible functional role of the main olfactory connections in reptiles.

The amygdaloid formation of the lizard Podarcis hispanica can be divided into three main groups of nuclei on the basis of their input from the main and accessory olfactory bulbs: the vomeronasal amygdala, the olfactory amygdala and the dorsal amygdaloid group, the latter group receiving afferents from neither the main (MOB) or the accessory olfactory bulb (AOB). The vomeronasal amygdala has a centrifugal projection to the AOB, an important commissural connection to the contralateral vomeronasal amygdala, a minor projection to nucleus accumbens, and a bilateral projection to the lateral cortex. The olfactory amygdala displays a bilateral afferents from the MOB, receives a contralateral afferent, and is reciprocally connected with the lateral cortex. Moreover, it receives an important input from the vomeronasal amygdala. The dorsal amygdaloid group receives projections from the other two amygdaloid groups, multimodal inputs from the anterior dorsal ventricular ridge and the dorsal cortex, and a putative cholinergic input from the basal telencephalon. Moreover, it is the site of origin of a prominent bilateral amygdalo-striatal projection that extends to the bed nucleus of the stria terminalis and the so-called amygdalo-striatal transition area, through which it may control both visceral and motor activities. The main extratelencephalic output of the amygdala courses through the stria terminalis and terminates in the ventromedial hypothalamic nucleus. The extratelencephalic afferents of the amygdala arise from several hypothalamic and anterior thalamic nuclei, from the mesencephalic and rhombencephalic aminergic cell groups, and from the rhombencephalic parabrachial nucleus.

Efferent projections of the medial cortex of the lizards Podarcis hispanica and Gallotia stehlinii were studied by examining the transport of horseradish peroxidase; results were correlated with those from Timm-stained sections. Two efferent systems were found. The first reaches the distal part of the outer plexiform layer in the medial, dorsomedial, and dorsal cortices, i.e., zones that are negative to Timm staining, and possibly originates from horizontal fusiform neurons. The second reaches the Timm-positive zones in the cortex and septum and is topographically arranged: the vertical portion of the intermediate and caudal medial cortex and the entire rostral medial cortex project to the inner two-thirds of the outer plexiform layer of the dorsomedial cortex and of the medial subfield of the dorsal cortex; to the paraventricular zone of the inner plexiform layer of the medial cortex; and bilaterally to the dorsal part of the dorsal precommissural septum. The dorsal part of the intermediate and caudal medial cortex and the ventralmost folded part of its caudal edge project rostrally to the juxtasomatic zone of the outer plexiform layer and the entire inner plexiform layer of the intermediate and lateral subfields of the dorsal cortex and to the ventral part of the dorsal septum. In its intense Timm reaction and its ultrastructural properties, as reported in earlier studies, the Timm-positive fiber system of the lizard brain shows a close resemblance to the mossy fiber system of the mammalian hippocampus.

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