Efferent projections of the anterior olfactory nucleus

The efferent projection fibers from the rostral and the medial segments of the anterior olfactory nucleus have been studied using the Nauta or the Fink modification of the Nauta method for degenrating axons. Lesions were suitably placed by electrocoagulation. From the rostral segment of the anterior olfactory nucleus efferent fibers by way of the bulbar part of the anterior commissure were traced to the internal granular and the mitral cell layer of the normal side. Preterminal and terminal degeneration were traced by way of the homolateral lateral olfactory tract and these fibers ended in the lateral two-thirds of the olfactory tubercle, the prepiriform, the piriform and the periamygdalar cortices and into the corticomedial segment of the amygdaloid complex. Degenerating fibers were also noted in the dorsomedial nucleus, the lateral habenular nucleus, the lateral hypothalamic area and in the supraoptic area of the same side where the lesion was made. In animals where the lesion was placed in the medial segment of the anterior olfactory nucleus the same fiber distribution was observed as reported when the lesion was placed in the rostral segment of the anterior olfactory nucleus. However, other areas were observed receiving fibers from the medial segment of the nucleus. These areas were the basolateral segment of the amygdaloid complex and the mammillary areas.     

The accessory olfactory bulbs and the lateral telencephalic wall of the rat-fish, Chimaera

The lateral telencephalon of Chimaera possesses several unique features but also has nuclei and fiber systems homologous with those of other sub-mammalian vertebrates. Ventricular ridges, similar to those of reptiles, are quite evident. Accessory olfactory bulbs are associated with the dorsal and ventral parts of each olfactory bulb. These contribute to the lateral olfactory tract. The internal granular layer caudal to the olfactory and the accessory bulbs blends with the anterior olfactory nucleus. Caudal to this nuclear area, the nuclei of the rostral telencephalon are well differentiated. Nuclear areas distinguishable in the lateral hemisphere include: the primordial dorsal pallium, the primordial piriform cortex, the primordial striatal and amygdaloid nuclei, and the lateral zone of the olfactory tubercle. These areas replace dorsal, dorsolateral, ventrolateral and ventral parts of the anterior olfactory nucleus, respectively. The primordial striatum is subdivided into hyperstriatum, neostriatum, paleostriatum augmentatum and paleostriatum primitivum. The amygdaloid area has anterior, corticomedial and basolateral nuclear groups. The basolateral area is best differentiated. The hyperstriatum forms a rostral ventricular eminence; the basolateral amygdaloid nucleus is present in a larger caudal ventricular ridge. Fiber tracts of the lateral wall include the lateral olfactory tract, the lateral corticohabenular tract, the lateral forebrain bundle and the stria terminalis. Nuclei of medial and lateral walls are interrelated through the hippocampal and the anterior commissures.     

The telencephalon of the sea catfish Galeichthys felis.

The sea catfish is a relatively abundant teleost fish. Its placement on the phylogenetic scale remains in question, although its brain resembles grossly those of some of the other teleosts which also possess pedunculated olfactory bulbs. The forebrain of Galeichthys felis consists of two hemispheres which lack lateral ventricles. They are joined at the midline by a thin membrane ventromedially and by the hippocampal and anterior commissures. The telencephalic hemispheres are overlaid by a single, ventrolaterally attached membrane which is continuous rostrally with the roofs of the olfactory ventricles. Six basic nuclear regions are evident in the telencephalon of the sea catfish: a dorsomedial, a ventromedial, a ventral, a lateral, a dorsal and a central. Dorsomedially, the primordial hippocampal formation is divided into an anterior continuation, a primordial dentate gyrus, a primordial cornu ammonis and a primordial subiculum. Ventromedially, the precommissural septum consists of the medial septal nucleus (pars dorsalis and pars ventralis), the lateral septal nucleus and the nuclei of the hippocampal and anterior commissure. Ventrally, the medial and the lateral zones and the medial island constitute the tuberculum olfactorium. The primordial general pallium comprises the dorsal area. Laterally, the primordial piriform cortex and the prepiriform region overlie the relatively large primordial amygdaloid complex, which includes an anterior anygdaloid nucleus, a primordial corticomedial amygdaloid nucleus and a primordial basolateral amygdaloid nucleus. The hyperstriatum, the neostriatum, and paleostriatum augmentatum and the paleostriatum primitivum constitute the central region. Basic fiber systems in Galeichthys felis which are homologous to those of higher vertebrates include the medial and the lateral olfactory tracts, the fornix, the medial and the lateral forebrain bundles, the stria medullaris pathways and the stria terminalis. In spite of the lack of lateral telencephalic ventricles in this form, nuclear areas were defined which, on the basis of topography, cellular morphology and fiber pathways, were homologized with the six basic regions found in the forebrain of higher vertebrates. The basic pattern of the fiber pathways present in the sea catfish corresponds to that found in the other submammalian vertebrates.     

ARC POMC mRNA and PVN alpha-MSH are lower in obese relative to lean zucker rats

The binding sites of nociceptin (also named orphanin FQ), the endogenous ligand of ORL1 (opiate receptor like 1), were localized in rat brain, using an autoradiographic procedure. High levels of binding were observed in the cingulate, retrosplenial, perirhinal, insular and occipital cortex, anterior and posteromedial cortical amygdaloid nuclei, basolateral amygdaloid nucleus, amygdaloid complex, posterior hippocampus, dorsal endopiriform, central medial thalamic, paraventricular, rhomboid thalamic, suprachiasmatic, ventromedial hypothalamic nuclei, mammillary complex, superficial gray layer of the superior colliculus, locus coeruleus, dorsal raphe nucleus. More moderate labelling was observed in the prefrontal, fronto–parietal, temporal, piriform cortex, dentate gyrus, anterior olfactory nucleus, olfactory tubercle, shell of nucleus accumbens, claustrum, lateral septum, laterodorsal thalamic, medial habenular, subthalamic, reuniens thalamic nuclei, subiculum, periaqueductal grey matter and pons. A lower binding site density was observed in the anterior and medial hippocampus, olfactory bulb, caudate putamen, the core of the nucleus accumbens, medial septum, ventrolateral, ventroposterolateral and mediodorsal thalamic nuclei, lateral and medial geniculate nuclei, hypothalamic area, substantia nigra, ventral tegmentum area and interpedoncular nucleus. A moderate and similar labelling was found in the dorsal and ventral horn of the spinal cord. No labelling was apparent in the corpus callosum. Thus, it appears that the ORL1 receptor is particularly abundant in the cerebral cortex, limbic system of the rat brain and some areas involved in pain perception.     

Amygdalopetal projections in the cat. I. Cortical afferent connections. A study with retrograde and anterograde tracing techniques

The cortical afferent connections of the amygdaloid complex of the cat have been studied by means of retrograde tracing of horseradish peroxidase and the fluorescent substances bisbenzimid and nuclear yellow. Subsequently, anterograde tracing experiments were carried out in order to define more precisely the termination areas of the corticoamygdaloid fibers. The results of the present study indicate that the main and accessory olfactory bulbs, the anterior olfactory nucleus, the prepiriform cortex and discrete regions of the medial frontal lobe, the insular and temporal cortices, as well as the perirhinal and entorhinal cortices and the ventral subiculum project to the amygdaloid complex. The main termination sites of these projections are the central, basolateral, and lateral amygdaloid nuclei. Neocortical regions project to the lateral nucleus and the lateral division of the lateral central nucleus. The mesocortical regions project predominantly to the basolateral nucleus and a medial division of the lateral central nucleus. In addition, area 35 distributes fibers to the lateral nucleus and the entorhinal cortex projects to the cortical nuclei of the amygdaloid complex. Fibers from the infralimbic area only reach the region of the medial central nucleus. Of the allocortical regions the prepiriform cortex distributes its fibers to the lateral, basolateral, and cortical nuclei, whereas the ventral subiculum projects to the medial division of the lateral central nucleus and the cortical nuclei. In the neocortical and most of the mesocortical regions the cells which project to the lateral and basolateral amygdaloid nuclei lie in layer III, whereas the cells which project to the central nucleus are located in layer V.     

Differential projections of the main and accessory olfactory bulb in the frog.

The central projections of the main olfactory bulb and the accessory olfactory bulb of the adult leopard frog (Rana pipiens) were reexamined, by using a horseradish peroxidase anterograde tracing method that fills axons with a continuous deposit of reaction product. The fine morphology preserved by this method allowed the terminal fields of the projection tracts to be delineated reliably, and for the first time. Herrick's amygdala has been newly subdivided into cortical and medial nuclei on the basis of cytoarchitecture, dendritic morphology, and the differential projections of the main and accessory olfactory tracts. The main olfactory bulb projects through the medial and lateral olfactory tracts to the postolfactory eminence, the rostral end of the medial cortex, the rostral end of the medial septal nucleus, the cortical amygdaloid nucleus, the nucleus of the hemispheric sulcus, and both the dorsal and ventral divisions of the lateral cortex, including its retrobulbar fringe. The lateral olfactory tract overlaps the dorsal edge of the striatal plate along the ventral border of the lateral cortex, but it is not certain whether any striatal cells are postsynaptic to the tract fibers. The lateral cortex is the largest of these territories, and receives the terminals of the main olfactory projection throughout its extent. It extends from the olfactory bulb to the posterior pole, and from the striatum to the summit of the hemisphere, where it borders the dorsal cortex. The medial and lateral olfactory tracts combine in the region of the amygdala to form a part of the stria medullaris thalami. These fibers cross in the habenular commissure and terminate in the contralateral cortical amygdaloid nucleus and periamygdaloid part of the lateral cortex. Cells projecting to the main olfactory bulb are found in the diagonal band and adjacent cell groups, but there is no evidence of an interbulbar projection arising from either the olfactory bulb proper or a putative anterior olfactory nucleus. The accessory olfactory bulb projects through the accessory olfactory tract to the medial and cortical amygdaloid nuclei. A fascicle of the tract crosses in the anterior commissure to terminate in the contralateral amygdala. While the main and accessory olfactory projections may converge in the cortical amygdaloid nucleus, the medial amygdaloid nucleus is connected exclusively with the accessory olfactory bulb.     

Afferent connections to the amygdaloid complex of the rat and cat: II. Afferents from the hypothalamus and the basal telencephalon

The projections from the basal telencephalon and hypothalamus to each nucleus of the amygdaloid complex of the rat, and to the central amygdala of the cat, were investigated by the use of retrograde transport of horseradish peroxidase (HRP). The enzyme was injected stereotaxically by microiontophoresis, using three different approaches. The ventral pallidum (Heimer, '78) and ventral part of the globus pallidus were found to project to the lateral and basolateral nuclei of the amygdala. The substantia innominata projects diffusely to the entire amygdaloid complex, except to the lateral nucleus and the caudal part of the medial nucleus. The anterior amygdaloid area shows a similar projection field, the only difference being that this structure does not project to any parts of the medial nucleus. The dorsal subdivision of the nucleus of the lateral olfactory tract sends fibers to the ipsilateral as well as the contralateral basolateral nucleus, and possibly to the ipsilateral basomedial and cortical amygdala. The ventral subdivision of the nucleus of the lateral olfactory tract was massively labeled after an injection in the ipsilateral central nucleus, but this injection affected the commissural component of the stria terminalis. The nucleus of the horizontal limb of the diagonal band of Broca connects with the medial, central, and anterior cortical nuclei, whereas the bed nucleus of stria terminalis and medial preoptic area are related to the medial nucleus predominantly. The lateral preoptic area is only weakly labeled after intra-amygdaloid HRP injections. The hypothalamo-amygdaloid projections terminate preponderantly in the medial part of the amygdaloid complex. Thus, axons from neurons in the area dorsal and medial to the paraventricular nucleus of the hypothalamus distribute to the medial nucleus and intra-amygdaloid part of the bed nucleus of stria terminalis. Most of the amygdalopetal fibers from the ventromedial, ventral premammillary, and arcuate nuclei of the hypothalamus end in the medial nucleus, but some extend into the central nucleus. A few fibers from the ventromedial nucleus of the hypothalamus reach the basolateral nucleus. The lateral hypothalamic area projects heavily to the central nucleus, and more sparsely to the medial and basolateral nuclei. The dorsal hypothalamic area and supramammillary nucleus show restricted projections to the central and basolateral nuclei, respectively. There are only a modest number of crossed hypothalamo-amygdaloid fibers. Most of these originate in the ventromedial nucleus of the hypothalamus and terminate in the contralateral medial nucleus. The projections from the basal telencephalon and hypothalamus to the central nucleus of the amygdala of the cat are similar to the corresponding projections in the rat.     

The development of the human amygdala during early embryonic life,

The development of the human amygdaloid complex is described beginning at the time that it first appears, when the cerebral hemispheres begin to evaginate (approximately 8--9 mm), and including a fetus of 27.4 mm CR length Both Nissl and protargol silver series, transversely and sagittally sectioned, were used. The earliest cell migration representing the striatal complex is from the germinal epithelium in the region of the interventricular foramen, lateral to the primordial hippocampal formation. This is the characteristic topographic location of the amygdala, which, therefore, is the first portion of the human striatal complex to appear embryologically. All of the amygdaloid nuclei identified develop by the migration of neuroblasts from the germinal epithelium. The three main subdivisions of the amygdaloid complex (anterior amygdaloid area, corticomedial complex and basolateral nuclear group) are identifiable almost immediately after the primordial amygdala appears (9.5 mm in the material studied). In this embryo the cortical and medial nuclei are identifiable, but the central nucleus was not seen until 22.2 mm and the nucleus of the lateral olfactory tract was not identified even in the oldest fetuses. The basolateral complex differentiates much later than the corticomedial complex and all of the nuclear components became identifiable at the same time (20.7 mm embryo). However the basal nucleus is best represented, the accessory basal next in size, and the lateral only just appearing. In the oldest fetus the subdivisions of the basal and accessory basal nuclei seen in the adult brain are recognizable. Even before the individual nuclei appear, the basolateral complex is much larger than the corticomedial complex. The lateral to medial rotation of the amygdaloid complex has not yet begun in the oldest fetuses, but the posterior end of the lateral ventricle is beginning to turn anteriorly. During early development, all cells of the amygdala are derived from the lateral striatal ridge, the only one present until 14.0 mm. At 20.7 mm and later neuroblasts are contributed from the medial striatal ridge also. Although the greater part of the basolateral amygdaloid complex is derived from the lateral ridge, some cells are contributed from the medial ridge, particularly to the accessory basal nucleus. By contrast, the corticomedial complex is derived in part from the medial striatal ridge, after it appears, even though the early development is from the lateral ridge. Comparisons between the amygdala at different developmental stages and the amygdala in fishes, in amphibians, in reptiles and in lower mammals are made. The amygdala in fishes, in amphibians, in reptiles and in lower mammals are made. The topographic relationships between the amygdala and the hippocampal formation, the piriform cortex and the caudate, the putamen and the globus pallidus are considered.     

Projections of the nucleus and tracts of the stria terminalis following lesions at the level of the anterior commissure.

The projections of the stria terminalis were traced with the Fink-Heimer stain following lesions at the level of the anterior commissure. The pre-commissural stria terminalis is amygdalofugal only, and projects to the nucleus of the anterior commissure, the medial preoptic area, the ventral portion of the capsule surrounding the ventromedial nucleus, and to the area closely adjacent to the periventricular nucleus by way of the medial corticohypothalamic tract. The postcommissural stria terminalis is both amygdalofugal and amygdalopetal. Its hypothalamic projection is to the lateral preoptic area and the bed nucleus of the stria terminalis, and to the lateral hypothalamus by way of the lateral preoptic area. The amygdaloid projection is mainly to the basolateral nucleus, with fewer terminations to the basomedial nucleus and the area surrounding the central nucleus. The projections of the bed nucleus of the stria terminalis are quite similar to the postcommissural stria, except for an additional projection to the magnocellular paraventricular and dorsal periventricular nuclei by way of the lateral filiform tract. The commissural stria terminalis projects contralaterally to cells within its fiber bundle and the posterior limb of the anterior commissure.     

Quantitative 3H-thymidine radiographic analyses of neurogenesis on the rat amygdala

Neurogenesis in the rat amygdala was examined with ³H-thymidine radiography. The animals were the offspring of pregnant females given two injections of ³H-thymidine on consecutive days in an overlapping series: Embryonic day (E) 12 + E13, + E14. E21 + E22. On 60 days of age, the percentage of labelled cells and the proportion of cells added during each day of formation were determined at several anatomical levels within the following components of the amygdala: anterior amygdaloid area, bed nuclei of the lateral and accessory olfactory tracts, central, medial, anterior cortical, posterolateral cortical, posteromedial cortical, basomedial, basolateral, and lateral nuclei, the amygdalo-hippocampal area, and the intercalated masses. All large and many small neurons originate in most nuclei between E13 and E17, those in the intercalated masses between E15 and E19, those in the amygdalo-hippocampal area between E16 and E19. The anterior amygdaloid area, intercalated masses, central, medial, posterolateral cortical, posteromedial cortical, basomedial, basolateral, and lateral nuclei have strong rostral-to-caudal intranuclear gradients. There are five additional intranuclear gradients: 1) medial to lateral in the central nucleus, anterior amygdaloid area, and anterior intercalated masses; 2) lateral to medial in the bed nucleus of the lateral olfactory tract and basolateral nucleus; 3) superficial to deep in the amygdalo-hippocampal area, posterolateral, and posteromedial cortical nuclei; 4) ventral to dorsal in the medial nucleus; and 5) dorsal to ventral between the small and large-celled parts of the lateral nucleus. Only the bed nucleus of the accessory olfactory tract and the anterior cortical nucleus do not have intranuclear gradients. Between 10 and 15% of the total cell population in most nuclei are very small neurons and/or glia which originate simultaneously between E18 and E20. This population is absent in the ventral part of the medial, anterior cortical, and anterior basomedial nuclei; these contiguous areas may form a distinct subunit in the amygdala. In contrast to the pronounced intranuclear gradient, internuclear gradients are weak. Neurogenesis in the central nucleus and corticomedial and basolateral complexes appears to take place both concurrently and independently. There are groups of early-originating neurons in the central, medial, and basolateral nuclei located near the periphery of the amygdala. Each of these groups is surrounded by younger neurons farther within the interior. The youngest calls are in the centrally placed intercalated masses. These settling patterns suggest that cells in the amygdala arise simultaneously from more than one neuroepithelial source during morphogenesis. The chronology of neurogenesis in the amygdala can be related to some of its anatomical connections. Rostral-to-caudal gradients in the corticomedial complex may be timed to coincide with early vs. late arrival of olfactory fibers. The subdivisions bases on intranuclear gradients in the central, medial, and basolateral nuclei match the subdivisions based on patterns of anatomical connections.     

Connections of the olfactory bulb and nucleus olfactorius anterior in the hedgehog (Erinaceus europaeus): fluorescent tracers and HRP study

The projections of the main olfactory bulbs (MOBs) and the dorsal part of the anterior olfactory nucleus (NOA) in the hedgehog (Erinaceus europaeus) have been studied by fluorescent tracers and the horseradish peroxidase method (HRP), respectively, to reveal the pattern of labeling from these structures. After different dye injections in both MOBs, labeled cells were present in the following structures: tenia tecta, vertical limb of the diagonal band of Broca, and medial septal nucleus in the ipsilateral injection site; and the NOA, piriform cortex, nucleus of the lateral olfactory tract, horizontal limb of the diagonal band of Broca, posterolateral cortical amygdaloid nucleus, anterior amygdaloid area, and dorsal raphe nucleus in both hemispheres. Structures showing double-labeled cells were the NOA, horizontal limb of the diagonal band of Broca, nucleus of the lateral olfactory tract, anterior amygdaloid area, and posterolateral cortical amygdaloid nucleus. After HRP injections in the dorsal part of the NOA, labeled cells were distributed in the NOA, nucleus of the lateral olfactory tract, posterolateral cortical amygdaloid nucleus, piriform cortex, horizontal and vertical limbs of the diagonal band of Broca, mitral cell layer of the MOB, tenia tecta, anterior amygdaloid area, and the contralateral NOA. We suggest that the contralateral projection nuclei to the MOB of the hedgehog, unusual in other mammals, and the large number of cells with axonal collaterals projecting to both hemispheres, may be a strategy in these animals to bilaterally integrate brain functions at the expense of its reduced corpus callosum.     

Efferent connections of the nucleus of the lateral olfactory tract in the rat

The efferent connections of the nucleus of the lateral olfactory tract (LOT) were examined in the rat with the Phaseolus vulgaris leucoagglutinin (PHA-L) technique. Our observations reveal that layers II and III of LOT have largely segregated outputs. Layer II projects chiefly ipsilaterally to the olfactory bulb and anterior olfactory nucleus, bilaterally to the anterior piriform cortex, dwarf cell cap regions of the olfactory tubercle and lateral shell of the accumbens, and contralaterally to the lateral part of the interstitial nucleus of the posterior limb of the anterior commissure. Layer III sends strong bilateral projections to the rostral basolateral amygdaloid complex, which are topographically organized, and provides bilateral inputs to the core of the accumbens, caudate-putamen, and agranular insular cortex (dorsal and posterior divisions). Layer II projects also to itself and to layers I and II of the contralateral LOT, whereas layer III projects to itself, to ipsilateral layer II, and to contralateral layer III of LOT. In double retrograde labeling experiments using Fluorogold and cholera toxin subunit b tracers, LOT neurons from layers II and III were found to provide collateral projections to homonymous structures on both sides of the brain. Unlike other parts of the olfactory amygdala, LOT neither projects directly to the extended amygdala nor to the hypothalamus. Thus, LOT seemingly influences nonpheromonal olfactory-guided behaviors, especially feeding, by acting on the olfactory bulb and on ventral striatal and basolateral amygdaloid districts that are tightly linked to lateral prefrontal cortical operations.     

Efferent projections of the infralimbic (area 25) region of the medial prefrontal cortex in the rat: an anterograde tracer PHA-L study

The efferent projections of the infralimbic region (IL) of the medial prefrontal cortex of the rat were examined by using the anterograde transport of Phaseolus vulgaris leucoagglutinin (PHA-L). Major targets of the IL were found to include the agranular insular cortex, olfactory tubercle, perirhinal cortex, the whole amygdaloid complex, caudate putamen, accumbens nucleus, bed nucleus of the stria terminalis, midline thalamic nuclei, the lateral preoptic nucleus, paraventricular nucleus, supramammillary nucleus, medial mammillary nucleus, dorsal and posterior areas of the hypothalamus, ventral tegmental area, central gray, interpeduncular nucleus, dorsal raphe, lateral parabrachial nucleus and locus coeruleus. Previously unreported projections of the IL to the anterior olfactory nucleus, piriform cortex, anterior hypothalamic area and lateroanterior hypothalamic nucleus were observed. The density of labeled terminals was especially high in the agranular insular cortex, olfactory tubercle, medial division of the mediodorsal nucleus of the thalamus, dorsal hypothalamic area and the lateral division of the central amygdaloid nucleus. Several physiological and pharmacological studies have suggested that the IL functions as the 'visceral motor' cortex, involved in autonomic integration with behavioral and emotional events. The present investigation is the first comprehensive study of the IL efferent projections to support this concept.     

Projections from the amygdaloid complex to the piriform cortex: A PHA-L study in the rat

Projections from the amygdala to the piriform cortex are proposed to provide a pathway via which the emotional system can modulate the processing of olfactory information as well as mediate the spread of seizure activity in epilepsy. To understand the details of the distribution and topography of these projections, we injected the anterograde tracer Phaseolus vulgaris-leucoagglutinin into different nuclear divisions of the amygdaloid complex in 101 rats and analyzed the distribution and density of projections in immunohistochemically processed preparations. The heaviest projections from the amygdala to the piriform cortex originated in the medial division of the lateral nucleus, the periamygdaloid and sulcal subfields of the periamygdaloid cortex, and the posterior cortical nucleus. The heaviest terminal labeling was observed in layers Ib and III of the medial aspect of the posterior piriform cortex. Lighter projections to the posterior piriform cortex originated in the dorsolateral division of the lateral nucleus, the magnocellular and parvicellular divisions of the basal and accessory basal nuclei, and the anterior cortical nucleus. The projections to the anterior piriform cortex were light and originated in the dorsolateral and medial divisions of the lateral nucleus, the magnocellular division of the basal and accessory basal nuclei, the anterior and posterior cortical nuclei, and the periamygdaloid subfield of the periamygdaloid cortex. The results indicate that only selective amygdaloid nuclei or their subdivisions project to the piriform cortex. In addition, substantial projections from several amygdaloid nuclei converge in the medial aspect of the posterior piriform cortex. Via these projections, the amygdaloid complex can modulate the processing of olfactory information in the piriform cortex. In pathologic conditions such as epilepsy, these connections might provide pathways for the spread of seizure activity from the amygdala to extra-amygdaloid regions.     

The topographic organization of associational fibers of the olfactory system in the rat, including centrifugal fibers to the olfactory bulb

This study analyzed the topographic organization of the associational fibers within the olfactory cortex of the rat, by using the autoradiographic method. Small injections of 3H-leucine were placed in all of the subdivisions of the olfactory cortex, to label selectively the fibers arising in each area. Intracortical fibers were identified from all of the olfactory cortical areas except the olfactory tubercle and were classified into two major systems (the layer Ib system and the layer II-deep Ib system) on the basis of their laminar pattern of termination (see Luskin and Price, '83). The layer Ib fiber system arises in the anterior olfactory nucleus, piriform cortex, and lateral entorhinal area, and is broadly organized in relation to the lateral olfactory tract. Cortical areas deep to or near the lateral olfactory tract are preferentially interconnected with areas near the tract, while parts of the cortex lateral and caudal to the lateral olfactory tract are most heavily interconnected with areas lateral, caudal, and medial to the tract. Commissural projections from the anterior olfactory nucleus and the anterior piriform cortex match some (but not all) components of the ipsilateral layer Ib fiber system. The layer II-deep Ib fiber system arises in three small areas--the ventral tenia tecta, the dorsal peduncular cortex, and the periamygdaloid cortex. The fibers from the ventral tenia tecta terminate in layer II of the anterior olfactory nucleus and are topographically organized. The fibers from the dorsal peduncular cortex and the periamygdaloid cortex are more widely distributed, especially in the lateral and caudal parts of the cortex. Two other intracortical projections do not fit into either of these fiber systems. The nucleus of the lateral olfactory tract projects bilaterally to the islands of Calleja and the medial edge of the anterior piriform cortex. The anterior cortical nucleus projects to many parts of the olfactory cortex, but the fibers end in both superficial and deep parts of layer I (layer Ia and Ib). There are projections from several of the olfactory cortical areas to the cortical areas surrounding the olfactory cortex. Virtually all of the olfactory areas also project to the ventral and dorsal endopiriform nuclei deep to the piriform cortex and/or to the polymorph zone deep to the olfactory tubercle. In addition, projections have been demonstrated to the deep amygdaloid nuclei, especially from the more ventromedial and caudal parts of the olfactory cortex.     

Afferent and efferent connections of the olfactory bulbs in the lizard Podarcis hispanica.

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.     

Efferent projections of the basolateral amygdala in the opossum, Didelphis virginiana

The autoradiographic anterograde axonal transport technique was used to study efferent projections of the opossum basolateral amygdala. All nuclei of the basolateral amygdala send topographically organized fibers to the bed nucleus of the stria terminalis (BST) via the stria terminalis (ST). Injections into rostrolateral portions of the basal nuclei label fibers that surround the commissural bundle of the ST, cross the midline by passing along the outer aspect of the anterior commissure, and terminate primarily in the contralateral BST, anterior subdivision of the basolateral nucleus (BLa), ventral putamen, and olfactory cortex. Each of the basal nuclei project ipsilaterally to the anterior amygdaloid area, substantia innominata and topographically to the ventral part of the striatum and adjacent olfactory tubercle. The posterior subdivision of the basolateral nucleus (BLp), but not the basomedial nucleus (BM), projects to the ventromedial hypothalamic nucleus. BLa and BLp have projections to the nucleus of the lateral olfactory tract and also send fibers to the central nucleus, as does the lateral nucleus (L). The lateral nucleus also has a strong projection to BM and both nuclei project to the amygdalo-hippocampal area. BLa and BLp send axons to the ventral subiculum and ventral lateral entorhinal area whereas L projects only to the latter area. The lateral nucleus and BLp project to the perirhinal cortex and the posterior agranular insular area. The BLa sends efferents to the anterior agranular insular area. Rostrally this projection is continuous with a projection to the entire frontal cortex located rostral and medial to the orbital sulcus. All of the nuclei of the basolateral amygdala project to areas on the medial wall of the frontal lobe that appear to correspond to the prelimbic and infralimbic areas of other mammals. Despite the great phylogenetic distance separating the opossum from placental mammals, the projections of the opossum basolateral amygdala are very similar to those seen in other mammals. The unique frontal projections of the opossum BLa to the dorsolateral prefrontal cortex appear to be related to the distinctive organization of the mediodorsal thalamic nucleus and prefrontal cortex in this species.     

Association and commissural fiber systems of the olfactory cortex of the rat. I. Systems originating in the piriform cortex and adjacent areas

The association and commissural fiber systems arising in the olfactory cortical areas caudal to the olfactory peduncle (the piriform cortex, nucleus of the lateral olfactory tract, anterior cortical nucleus of the amygdala, periamygdaloid cortex and entorhinal cortex) have been studied utilizing horseradish peroxidase as both an anterograde and a retrograde axonal tracer. In the piriform cortex two sublaminae within layer II (IIa and IIb) and layer III have been found to give rise to distinctly different projections. Retrograde cell labeling experiments indicate that the association fiber projection from layer IIb is predominantly caudally directed, while the projection from layer III is predominantly rostrally directed. Cells in layer IIa project heavily to areas both caudal and rostral to the piriform cortex. The commissural fibers from the piriform cortex are largely restricted in their origin to layer IIb of the anterior part of the piriform cortex and in their termination on the contralateral side to the posterior part of the piriform cortex and adjacent olfactory cortical areas. A projection to the olfactory bulb has also been found to arise from cells in layers IIb and III of the ipsilateral piriform cortex, but not in layer IIa. In addition to those from the piriform cortex, association projections have also been found from other olfactory cortical areas. The nucleus of the lateral olfactory tract has a heavy bilateral projection to the medial part of the anterior piriform cortex and the lateral part of the olfactory tubercle (as well as a lighter projection to the olfactory bulb); both the anterior cortical nucleus of the amygdala and the periamygdaloid cortex project ipsilaterally to several olfactory cortical areas. The entorhinal cortex has been found to project to the medial parts of the olfactory tubercle and the olfactory peduncle. The olfactory tubercle is the only olfactory cortical area from which no association fiber systems (instrinsic or extrinsic) have been found to originate. A broad topographic organization exists in the distribution of the fibers from several of the olfactory areas. This is most obvious in the anterior part of the olfactory cortex, in which fibers from the more rostral areas (the anterior olfactory nucleus and the anterior piriform cortex) terminate in regions near the lateral olfactory tract, while those from more caudal areas (the posterior piriform cortex and the entorhinal cortex) terminate in areas further removed, both laterally and medially, from the tract. Projections to olfactory areas from the hypothalamus, thalamus, diagonal band, and biogenic amine cell groups have been briefly described.     

Thalamoamygdaloid projections in the rat: a test of the amygdala's role in sensory processing.

We used the autoradiographic tract-tracing method to define the amygdaloid projection fields after injecting 3H-amino acids into individual thalamic nuclei in the rat. The parvicellular division of the ventroposterior nucleus, the thalamic taste relay, projected lightly to the central and lateral amygdaloid nuclei. The central medial, interanteromedial, and paraventricular thalamic nuclei, viscerosensory relays of the thorax and abdomen, projected heavily to the amygdala. All projected to the basolateral amygdaloid nucleus, the paraventricular nucleus in addition having terminations in the central nucleus, the amygdaloid portion of the nucleus of the stria terminalis, and the amygdalohippocampal transition area. The magnocellular division of the medial geniculate, a thalamic auditory (and, to a moderate degree, a spinothalamic) relay, sent heavy projections to the central, accessory basal, lateral, and anterior cortical nuclei, and to the anterior amygdaloid area and the nucleus of the accessory olfactory tract. Other thalamic nuclei projecting to the amygdala, for which functions could not be associated, were the paratenial and subparafascicular nuclei. The former projected to the lateral, basal, and posterolateral cortical nuclei; the latter projected very lightly to the central, medial, and basal accessory nuclei. These results show that, like the cortical amygdaloid nuclei, which are sensory (olfactory) in nature, the subcortical amygdaloid nuclei must have major sensory functions. These thalamic afferents, when correlated with cortical and brainstem data from the literature, suggested that the amygdala is in receipt of sensory information from many modalities. To uncover the manner by which such information is processed by the amygdala and relayed to effector areas of the brain, six hypothetical mechanisms relating to modality specificity and convergence were posited. By charting sensory-related afferents to all subdivisions of the amygdala, each nucleus was characterized as to its mechanism of information processing. Four proposed amygdaloid systems emerged from this analysis. A unimodal corticomedial amygdaloid system relays pheromonal information from the accessory olfactory bulb to medial basal forebrain and hypothalamic areas. A second system--the lateral-basomedial--collects and combines input from a number of sensory modalities and distributes it to the same basal forebrain and hypothalamic areas as the corticomedial. The central system appears to concentrate the effect of viscerosensory information arriving from multiple brainstem, thalamic, cortical, and amygdaloid sources; this information is combined with significant auditory and spinothalamic inputs from the thalamus and cortex. The central system projects to lateral nuclei in the basal forebrain, hypothalamus, and brainstem.(ABSTRACT TRUNCATED AT 400 WORDS)