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Olfactory bulb connections with basal rhinencephalon in the ferret: an evoked potential and neuroanatomical study.

The lateral olfactory tract (LOT) of the ferret has been shown to project to most of the pyriform lobe, as in the cat. Only a small medio-posterior region of the pyriform cortex (the angular cortex), which has a distinctly different appearance in Nissl stained sections, proves to be devoid of olfactory connections. Despite the fact that sub-areas can be recognized within olfactory cortex, there is an underlying constancy in design throughout. Latency measurements indicate that fine collateral branches of theLOT reach the posterior olfactory cortex, whereas mainly larger diameter (faster conducting) fibres comprise the contribution to anterior olfactory cortex. Thus in the present context it is sufficient to recognise just an anterior and posterior subdivisionof olfactory cortex, contingent on the above criterion and correlated with cytoarchitectural features, chiefly variations in cortical layers II and III of the regions concerned. Pyriform cortex directly in receipt of LOT fibres, and associated deeper cortical zones, generated potentials in the olfactory bulbs which can only be attributed to centrifugal input, thus further substantiating an olfactory role for this cortex. In addition, non-cortical regions in receipt of LOT fibres, namely the anterior olfactory nucleus, olfactory tubercle, cortical amygdaloid nucleus and nucleus of the lateral olfactory tract, also contribute centrifugal input to the bulbs. All these regions are thereby capable of providing a rather direct feedback on olfactory bulb activity. The exact pathways concerned have yet to be determined.

Animals

Telencephalic efferents of the tiger salamander Ambystoma tigrinum tigrinum (Green).

The efferent projections of the telencephalon in the tiger salamander were examined by the Nauta and Fink-Heimer methods following unilateral hemispherectomies, rostral hemispheric ablations and pallial lesions. The cerebral hemisphere connects with most areas of the contralateral hemisphere via the pallial, anterior and habenular commissures. The descending fibers travel in the medial and lateral forebrain bundles and in the tracts comprising the stria medullaris. Degenerating fibers and terminals were present throughout the diencephalon but were more abundant ipsilaterally. Fibers reach the pretectum and optic tectum via dorsal and ventral pathways. There is a heavy projection to the midbrain tegmentum and a sparse projection to the tectum via the ipsilateral lateral forebrain bundle. This tract continues into the medulla oblongata and the cervical spinal cord. Rostral and dorsal hemispheric ablations revealed that the majority of fibers forming the olfacto-peduncular tract originate in the ventral, rostral one-third of the hemisphere. It was also determined that the majority of the descending efferent fibers located in the lateral forebrain bundle originate from the caudal lateral hemispheric wall, and that these fibers form connections characteristic of mammalian corticofugal and striatofugal systems. The cytoarchitecture and connections of the caudal lateral hemispheric wall suggest that it is homologous to parts of motor isocortex and amygdala of amniotes.

Ambystoma

Olfactory bulb projections in the bullfrog Rana catesbeiana.

The projections of the accessory and main olfactory bulbs of the bullfrog are described as part of a long term analysis of the morphological differences in amphibian and reptilian telancephalons. Unilateral aspiration of the accessory olfactory bulb results in an ipsilateral projection to the pars lateralis of the amygdala via the accessory olfactory tract. Degenerating fibers from the accessory olfactory bulb are tracable into the cell-free zone between the dorsal striatum and the lateral pallium, and projections to these neural populations may also exist. Unilateral lesions of the main olfactory bulb reveal two major secondary pathways: an ipsilateral medial olfactory tract that projects to the rostral ventromedial portion of the medial pallium, the postolfactory eminence and the rostal, lateral and medial septal nuclei; and an ipsilateral lateral olfactory tract that projects to the dorsal striatum, the lateral pallium and the ventral half of the dorsal pallium. Two crossed secondary olfactory pathways to the contralateral telencephalon decussate via the habenular commissure after entering the ipsilateral stria medullaris. A crossed lateral pathway terminates in the dorsal striatum, the caudal, lateral pallium and the ventral portion of the dorsal pallium. A crossed medial pathway terminates in the internal granule layer of the main olfactory bulb.

Amygdala

The efferent connections of the suprachiasmatic nucleus of the hypothalamus.

The efferent connections of the suprachiasmatic nucleus of the hypothalamus have been studied in the rat by the injection of 3H-proline into the nucleus and the surrounding regions of the rostral hypothalamus, and by the injection of the enzyme marker, horseradish peroxidase, into the region of the ventromedial hypothalamic nucleus. After an injection of 3H-proline confined to the ventral portion of the suprachiasmatic nucleus, transported label can be followed, in the autoradiographs, dorsally and caudally in the periventricular area as far as the caudal end of the ventromedial nucleus, into the triangular area between this nucleus and the arcuate nucleus, and along the ventral aspect of the tuberal region, just lateral to the ventromedial nucleus. A small number of silver grains are also seen over the internal lamina of the median eminence. No label can be followed rostrally or immediately lateral to the nucleus. Comparable injections into adjoining regions of the hypothalamus (especially the anterior hypothalamic area, the medial preoptic area, and the retrochiasmatic region) show transported label over the same regions, but with a somewhat different pattern of grain distribution; in addition, the anterior hypothalamic area shows an extensive projection through the medial forebrain bundle to the mammillary and supramammillary nuclei, the midbrain tegmentum, and certain of the midline thalamic nuclei. Although it is difficult in our autoradiographs to distinguish between the course of the efferent fibers from the suprachiasmatic nucleus and the zones in which they terminate, our evidence favors a termination among the cells of the periventricular area, and upon dendrites of the cells in the ventromedial, dorsomedial and arcuate nuclei, which extend beyond the limits of the nuclei into the periventricular area and to the area beneath the ventromedial nucleus.

Amygdala

The differential projections of the olfactory bulb and accessory olfactory bulb in mammals.

Three species were studied, the rabbit, opossum and rat. Lesions of the main olfactory bulb caused terminal degeneration, assayed by the Fink-Heimer method, to occur in the ipsilateral olfactory tubercle, prepyriform cortex (including its periamygdaloid part), ventrolateral entorhinal area, and in anterior and posterolateral divisions of the cortical amygdaloid nucleus. The various parts of the ipsilateral anterior olfactory nucleus and the rostroventral end of the anterior continuation of the hippocampus (hippocampal rudiment) also received this projection. Lesions of the accessory olfactory bulb, which receives its sensory input from the vomeronasal (Jacobson's) organ, caused terminal degeneration to occur in the medial amygdaloid nucleus and in a posteromedial part of the cortical amygdaloid nucleus. This projection was conveyed by an accessory olfactory tract, which is accompanied in part of its course by a small nucleus, the bed nucleus of the accessory olfactory tract. The accessory olfactory tract is initially a part of the lateral olfactory tract but becomes increasingly indivuated at more posterior levels. It parts company with the lateral olfactory tract at the rostral end of the amygdaloid region, and, in addition to distributing to the medio-cortical amygdaloid region, it enters the stria terminalis to terminate in the bed nucleus of the stria terminalis in a small region bearing cytoarchitectonic resemblance to the medial amygdaloid nucleus. The topographic segregation of the areas of termination of the olfactory and accessory olfactory (vomeronasal) projections is suggestive of a functional dichotomy in the organization of the olfactory system...

Amygdala

The distribution of the olfactory tracts in the bullhead catfish, Ictalurus nebulosus.

Using degeneration staining methods, central projections of the olfactory tracts in the bullhead catfish, Ictalurus nebulosus were studied. Seporate lesions were made of the lateral and medial olfactory tracts in an attempt to separate their zones of projection. Three major terminal fields were found: lateral, medial and central-posterior complex. Two additional minor projections were seen: interbulbar and hypothalamic. The lateral division of the olfactory tract terminates in lateral, central and hypothalamic terminal fields. The medial tract projects to lateral, meedial, posterior and hypothalamic fields. All terminal fields were found bilaterally; the lateral tract partly crossing in both the anterior commissure. A small intellbulbar commissure was also found to contain medial tract fibers. Evidence is given to show that the anterior olfactory nucleus, if present in catfish, may be located within the olfactory bulb itself. Similarities between mammalian and teleost olfactory systems are also discussed.

Animals

Association and commissural fiber systems of the olfactory cortex of the rat.

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) 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 predominatnly 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 contralteral 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. Projection to olfactory areas from the hypothalamus, thalamus, diagonal band, and biogenic amine cell groups have been briefly described.

Animals

Anatomical and experimental study of certain association fascicles in the cortex of the rabbit (Oryctalagus cuniculus).

1. All lesions resulted in degeneration of the short intracortical association fibers in cortical layer I and of the short subcortical fibers which extended to the corona radiata before ending in the deeper layers of the overlying neopallium. 2. From all the lesions fibers were traced through the corona radiata to the subcallosal or the so-called superior fronto-occipital association bundle. This bundle had projection fibers to the orbitofrontal cortex. 3. From the lesion in the orbitofrontal neopallium, the orbitofrontal-pyriform connections were established. Such fibers coursed on the dorsal edge of the lateral olfactory tract and distributed to the pyriform cortex and to the nucleus of the lateral olfactory tract. 4. The uncinate fasciculus of man derived its name from its arching course from the base of the frontal lobe to the temporal lobe. Because of the more caudal position of the amygdala in the rabbit, the comparable fasciculus passed directly caudally and exhibited only slight arching. This fasciculus in the rabbit had the typical dorsal and ventral parts. The dorsal part arose from the orbitofrontal cortex to distribute to the pyriform and the temporal lobe cortices. The ventral portion extended into the olfactory tuberculum and the anterior amygdaloid area. 5. The paraventricular component of the transverse frontal fasciculus interconnected the neopallium with the medial part of the olfactory tuberculum. It had origins in the frontal and possibly in other neocortical areas. 6. The cingulum interconnected the medial portion of the olfactory tubercle, the septum, the various cingulate areas and areas of the neopallium with each other. 7. Therefore, the New Zealand white rabbit had short association fibers which were mainly neopallial in origin and termination and long association fibers which had both a neopallial and a limbic component.

Amygdala

The luteinizing hormone-releasing hormone (LH-RH) neuronal networks of the guinea pig brain. I. Intra- and extra-hypothalamic projections.

In the guinea pig brain, LH-RH-containing cell bodies are located not only within the classical hypophysiotrophic area but also in the medial preoptic area, septum and olfactory tubercle. LH-RH fiber tracts project not only to the primary portal plexus in the median eminence but also throughout the limbic forebrain and limbic midbrain regions. Using radiofrequency lesions in different brain regions, the projections of LH-RH cell bodies were determined. Cells in the medial preoptic area project ot the organum vasculosum of the lamina terminalis (OVLT), the suprachiasmatic nucleus, the mammillary body complex and the ventral tegmental area. LH-RH neurons in both the medial septal nucleus and medial preoptic area project via the stria medullaris to the medial habenular nucleus and from there via the fasciculus retroflexus to the interpeduncular nucleus of the midbrain. Other LH-RH neurons in the medial septal nucleus, nucleus of the diagonal band of Broca and olfactory tubercle are congregated in small clusters around large blood vessels which penetrate into this area, and they do not appear to send axons outside their immediate vicinity. The types of LH-RH axonal terminations and the roles of these peptide-containing neurons are discussed.

Animals

Morphine and naloxone effects on olfactory evoked electrographic activity in the amygdala.

This research tested morphine and naloxone effects on evoked EEG and unit activity in 3 opiate-relevant brain areas in response to electrical stimulation of the olfactory bulb in acute, unanesthesized rat preparations. Stimulation evoked clear EEG responses in the amygdala (Amyg) and sometimes in the other areas (caudate and central grey); morphine (15 mg/kg) depressed the Amyg response in some rats, but enhanced it in others, and naloxone usually reversed both kinds of effect. Stimulation caused excitatory unit impulse reponses in the Amyg, and morphine unexpectedly increased the magnitude of the stimulus-evoked excitation; naloxone reversed this enhancement. In control rats, naloxone often decreased the Amyg evoked response. Stimulus-evoked increases in unit activity in the caudate and central grey, when they did occur, were depressed by morphine, but naxoxone had no consistent reversing effect. Both the EEG and unit data indicate that morphine excites, or disinhibits, certain neurons associated with the olfactory-Amyg pathway. There was also some evidence that this pathway contains endorphinergic elements.

Amygdala

An autoradiographic study of the efferent connections of the preoptic region in the rat.

The normal morphology of the rat preoptic region has been briefly described on the basis of Nissl- and silver-stained preparations and its efferent connections have been studied autoradiographically in over 50 rat brains with single small injections of 3H-proline, or various mixtures of 3H-proline, 3H-leucine, and 3H-lysine. Injections in the anteroventral part of the lateral preoptic area labeled fibers projecting through, and perhaps to, the anterior and lateral hypothalamic areas and ending in the supramammillary region, and ventral fiber lamina of the mammillary complex; other labeled fibers ended in the periventricular hypothalamic gray and the internal lamina of the median eminence. The posteromedial lateral preoptic area projects to the same regions, as well as to the medial septal-diagonal band complex, and to the lateral habenula through the stria medullaris. Injections of the posterolateral lateral preoptic area labeled each of the above fiber systems as well as fibers to the main olfactory bulb, anterior olfactory nucleus and taenia tecta. Other fibers coursed over the genu of the corpus callosum, through the stria terminalis and ansa peduncularis to the medial, cortical and basal amygdaloid nuclei and the anterior amygdaloid area, and through the medial forebrain bundle to the substantia nigra. The transition region between the lateral preoptic and lateral hypothalamic areas at the level of the supraoptic nucleus has widespread connections as a whole (a) with the medial septal-diagonal band complex, lateral septum and bed nucleus of the stria terminalis, (b) through or to most of the hypothalamus, the substantia nigra, central tegmental field, central gray, superior central nucleus, and the locus coeruleus, (c) through the stria medullaris to the lateral habenula (bilaterally), parataenial, paraventricular, and mediodorsal nuclei of the thalamus, (d) through the stria terminalis and ansa peduncularis to the central, medial and cortical nuclei of the amygdala, and (e) to the main olfactory bulb, anterior olfactory nucleus, cingulate bundle, olfactory tubercle, medial septal-diagonal band complex and the lateral septum.

Amygdala