Neural pathway from the olfactory bulbs regulating tonic gonadotropin secretion.

Removal of the olfactory bulbs of male golden hamsters results in a marked increase in tonic gonadotropin, prolactin and testosterone secretion which counteracts inhibitory effects of manipulations such as maintenance on short photoperiod, food restriction or treatment with gonadal steroids. The bulbectomy-induced increase in hormone secretion is interpreted to reflect a tonic inhibitory influence of the olfactory bulbs. This inhibition is not dependent upon chemosensory stimulation and may be mediated by olfactory bulb fibers projecting through the lateral olfactory tract to or through the olfactory tubercle. This review will relate these studies conducted on hamsters to results in other species, such as the rat, where the olfactory bulbs enhance serum gonadotropin levels.

Tyrosine hydroxylase neurons in the male hamster chemosensory pathway contain androgen receptors and are influenced by gonadal hormones.

Chemosensory and hormonal signals, both of which are essential for mating in the male Syrian hamster, are relayed through a distinct forebrain circuit. Immunocytochemistry for tyrosine hydroxylase, a catecholamine biosynthetic enzyme, previously revealed immunoreactive neurons in the anterior and posterior medial amygdaloid nucleus, one of the nuclei within this pathway. In addition, dopamine-immunoreactive neurons were located in the posterior, but not the anterior, medial amygdala. In the present study, tyrosine hydroxylase-immunostained neurons were also observed in other areas of the chemosensory pathway, including the posteromedial bed nucleus of the stria terminalis and the posterior, lateral part of the medial preoptic area, while dopamine immunostaining was only seen in the posteromedial bed nucleus of the stria terminalis. The colocalization of tyrosine hydroxylase and androgen receptors was examined in these four tyrosine hydroxylase cell groups by a double immunoperoxidase technique. The percentage of tyrosine hydroxylase-immunolabeled neurons that were also androgen receptor-immunoreactive was highest in the posterior medial amygdaloid nucleus (74%) and the bed nucleus of the stria terminalis (79%). Fewer tyrosine hydroxylase-immunostained neurons in the anterior medial amygdala (33%) and the medial preoptic area (4%) contained androgen receptors. Surprisingly, castration resulted in a significant decrease in the number of tyrosine hydroxylase-immunoreactive neurons only in the anterior medial amygdaloid nucleus, and this effect was transient. Six weeks after castration, the anterior medial amygdala contained 61% fewer tyrosine hydroxylase-immunolabeled neurons, but 12 weeks after gonadectomy, immunostaining returned to intact values. The number of immunostained neurons in testosterone-replaced, castrated hamsters was not significantly different from that of intact or castrated animals at any time. The results of this study indicate that a substantial number of tyrosine hydroxylase-immunostained neurons in the chemosensory pathway are influenced by androgens; the majority of these neurons in the posterior medial amygdala and the posteromedial bed nucleus of the stria terminalis produce androgen receptors, and tyrosine hydroxylase immunoreactivity is altered by castration in the anterior medial amygdala.

Prodynorphin peptide distribution in the forebrain of the Syrian hamster and rat: a comparative study with antisera against dynorphin A, dynorphin B, and the C-terminus of the prodynorphin precursor molecule.

The neuroanatomical distribution of the prodynorphin precursor molecule in the forebrain of the male Syrian hamster (Mesocricetus auratus) has been studied with a novel antiserum directed against the C-terminus of the leumorphin [dynorphin B (1-29)] peptide product. C-peptide staining in sections from colchicine-treated hamsters is compared to staining in sections from untreated animals. In addition, the pattern of C-peptide immunostaining in hamster brain is compared to that in the rat brain. Finally, the C-peptide immunolabeling patterns in hamsters and rats are compared to those obtained with antisera to dynorphin A (1-17) and dynorphin B (1-13). Areas of heaviest prodynorphin immunoreactivity in the hamster include the hippocampal formation, lateral septum, bed nucleus of the stria terminalis, medial preoptic area, medial and central amygdaloid nuclei, ventral pallidum, substantia nigra, and numerous hypothalamic nuclei. Although this C-peptide staining pattern is similar to dynorphin staining reported previously in the rat, several species differences are apparent. Whereas moderate dentate gyrus granule cell staining and no CA4 cell staining have been reported in the rat hippocampal formation, intense immunostaining in the dentate gyrus and CA4 cell labeling are observed in the hamster. In addition, the medial preoptic area, bed nucleus of the stria terminalis, and medial nucleus of the amygdala stain lightly for prodynorphin-containing fibers and cells in the rat, compared to heavy cell and fiber staining in the hamster in all three of these regions. In the rat there is no differential staining between tissues processed with the C-peptide, dynorphin A, and dynorphin B antisera, but numerous areas of the hamster brain show striking differences. In most hamster brain areas containing prodynorphin peptides, the C-peptide antiserum immunolabels more cells and fibers than the dynorphin B antiserum, which in turn labels more cells and fibers than dynorphin A antiserum. However, exceptions to this hierarchy of staining intensity are found in the lateral hypothalamus, substantia nigra, arcuate nucleus, and habenula. The differences in staining patterns between rat and hamster are greatest when C-peptide antiserum is used; apparent species differences are present, though less pronounced, in dynorphin B- and dynorphin A-immunostained material.

Differential projections of the anterior and posterior regions of the medial amygdaloid nucleus in the Syrian hamster.

The medial nucleus of the amygdala is important for the neural control of reproductive behavior in the adult male Syrian hamster. Two types of signals are essential for this behavior, chemosensory stimuli and gonadal steroids; these signals appear to be received in different parts of the medial nucleus. The anterior region receives input from olfactory and vomeronasal systems, both of which are required for this behavior, whereas the posterior region receives gonadal hormone inputs. Behavioral studies have also suggested a functional differentiation of these two areas; electrolytic lesions of the anterior, but not the posterior, part eliminates normal sexual behavior. In this study, the efferent projections of the anterior and posterior divisions of the medial nucleus of the amygdala in the Syrian hamster were analyzed throughout the forebrain after injections of the anterograde neuronal tracer, Phaseolus vulgaris-leucoagglutinin. Neurons of the anterior, but not the posterior, medial nucleus, were found to project to numerous olfactory bulb projection areas and to the ventral striatopallidal complex. Within areas of the chemosensory circuitry that control reproductive behavior, the anterior region of the medial nucleus projects to the intermediate part of the posterior bed nucleus of the stria terminalis and the lateral part of the medial preoptic area, whereas the posterior region of the medial nucleus projects to the medial parts of these areas. Differences in targets were also observed in the ventromedial nucleus of the hypothalamus where the anterior region projects to the core while the posterior part projects to the shell of this nucleus. Furthermore, reciprocal projections between the anterior and posterior regions of the medial nucleus were observed. Taken together, these studies support the hypothesis that the anterior and posterior regions of the medial amygdaloid nucleus provide substantially different contributions to the control of reproductive behaviors.

Neurons of origin and fiber trajectory of amygdalofugal projections to the medial preoptic area in Syrian hamsters.

The amygdaloid neurons of origin and the trajectory of amygdaloid fibers to the medial preoptic area of the adult male Syrian hamster were identified by using horseradish peroxidase (HRP) histochemistry. After iontophoresis of HRP into the medial preoptic area, retrogradely labeled amygdaloid neurons were located in the dorsal and caudal parts of the medial amygdaloid nucleus and throughout the amygdalohippocampal area. No amygdaloid neurons were labeled after HRP applications confined to the most rostral portion of the medial preoptic area (anterior to the body of the anterior commissure). Following more caudal medial preoptic area injections (body of the anterior commissure to the suprachiasmatic nucleus) the distribution of retrogradely labeled cells in the medial amygdaloid nucleus and the amygdalohippocampal area revealed no topographic organization of the amygdalopreoptic connections. When amygdaloid neurons were labeled, the amygdalohippocampal area contained two to five times as many HRP-filled cells as the medial amygdaloid nucleus. Retrogradely transported HRP could be followed from the medial preoptic area to the amygdala through fibers in the dorsomedial quadrant of the stria terminalis. In addition, electrolytic lesions of the stria terminalis prior to iontophoresis of HRP into the medial preoptic area prevented retrograde transport to neurons in both the dorsocaudal medial amygdaloid nucleus and the amygdalohippocampal area. These results confirm earlier observations describing the location of autoradiographically labeled efferents from the medial amygdaloid nucleus to the medial preoptic area and provide new information about the restricted region within the medial amygdaloid nucleus from which these projections arise. They also suggest that, unlike the projections from the medial amygdaloid nucleus to the bed nucleus of the stria terminalis, the efferents to the medial preoptic area travel entirely in the stria terminalis.

Mating and agonistic behavior produce different patterns of Fos immunolabeling in the male Syrian hamster brain.

Previous work has shown that mating induces the expression of Fos protein within the chemosensory pathways of the male Syrian hamster brain. However, it is not known if this pattern of labeling is specific to mating or the result of social interactions in general. To determine the behavioral specificity of activation within these pathways, Fos immunostaining following mating was compared to that following agonistic behavior. Both mating and agonistic behavior are dependent upon chemosensory cues and gonadal steroids (reviewed in Refs 64, 65) and areas belonging to the olfactory and vomeronasal pathways process chemosensory and hormonal information (reviewed in Ref. 48). The results of this study demonstrate both similarities and differences in brain activation patterns following these two social behaviors. Agonistic behavior increased the number of Fos-immunoreactive neurons within most subdivisions of the medial amygdala, the anteromedial and posterointermediate bed nucleus of the stria terminalis, the ventrolateral septum and the ventral premammillary nucleus of the hypothalamus in a pattern comparable to that observed after mating. This pattern of activation common to mating and agonistic behavior may reflect an increase in an animal's general state of arousal during social interactions. In contrast, although mating and agonistic behavior both activated neurons within the caudal subdivision of the medial nucleus of the amygdala, the anterodorsal level of posteromedial bed nucleus of the stria terminalis and the paraventricular and ventromedial nuclei of the hypothalamus, in these areas either the distribution and/or number of Fos-immunoreactive neurons differed. In addition, agonistic behavior selectively activated neurons within the anterolateral bed nucleus of the stria terminalis, the anterior nucleus of the hypothalamus and the dorsal periaqueductal gray, whereas mating alone activated neurons within the posteroventral level of posteromedial bed nucleus of the stria terminalis and the medial preoptic area. No differences were found between dominant and subordinate males following agonistic behavior. These observations along with results from other laboratories suggest that mating and agonistic behavior activate distinct neural circuits.

6-Hydroxydopamine lesions of the nigrostriatal pathway alter the expression of glutamate decarboxylase messenger RNA in rat globus pallidus projection neurons.

In situ hybridization was used to study the effect of 6-hydroxydopamine-induced damage to the midbrain dopaminergic neurons on the level of glutamate decarboxylase mRNA in globus pallidus neurons in the rat. Some animals received an injection of Fluoro-gold in the entopeduncular nucleus or the substantia nigra prior to the 6-hydroxydopamine lesion in order to identify glutamic acid decarboxylase mRNA levels in pallidal neurons that project to one of these targets. Analysis was carried out on a sample of all pallidal neurons as well as neurons that were identified as projection neurons in control and lesioned groups. The loss of the dopamine-containing neurons in the substantia nigra resulted in significant increases in the percentage of globus pallidus neurons that expressed glutamate decarboxylase mRNA and in the amount of glutamate decarboxylase mRNA per globus pallidus neuron. These increases were noted in a sample of all pallidal neurons, as well as pallidal neurons that were identified as projecting to either the entopeduncular nucleus or the substantia nigra. In control animals, glutamate decarboxylase mRNA was clearly identified in globus pallidus neurons projecting to the entopeduncular nucleus, indicating that this recently reported projection is at least partially GABAergic. The results of this study indicate that substantia nigra dopaminergic neurons regulate globus pallidus neurons in the rat, and that removal of the dopaminergic input to the corpus striatum results in a significant increase in the amount of glutamate decarboxylase mRNA in pallidal neurons. The decreased firing rate of pallidal neurons that is seen following the loss of dopamine input appears to be accompanied by an increase in the level of glutamate decarboxylase mRNA in these neurons.

Tyrosine hydroxylase mRNA-containing neurons in the medial amygdaloid nucleus and the reticular nucleus of the thalamus in the Syrian hamster.

To confirm previous immunocytochemical findings in colchicine-treated Syrian hamsters, in situ hybridization was used to investigate the distribution of TH mRNA-containing cells in the medial amygdaloid nucleus (Me) and the thalamic reticular nucleus (Rt) of untreated hamsters. TH mRNA-producing neurons were observed in anterior and posterior Me and throughout Rt, similar to the distribution of TH-immunostained cells in these areas of animals receiving colchicine. These data confirm that TH is normally produced in amygdaloid and thalamic cell groups which lie outside the classical catecholamine systems.

Axon-sparing lesions of the medial nucleus of the amygdala decrease affiliative behaviors in the prairie vole (Microtus ochrogaster): behavioral and anatomical specificity.

The neural basis of affiliative behavior was examined in the prairie vole, a rodent that exhibits high levels of social contact and paternal behavior. In the first study, the axon-sparing excitotoxin N-methyl-D,L-aspartic acid (NMA) produced lesions in the basolateral nucleus of the amygdala or the corticomedial amygdala. Males with corticomedial lesions showed significantly less contact with a familiar adult female and a pup when compared with males with lesions of the basolateral nucleus or controls. This behavioral change was not associated with changes in exploratory behavior, motor function, performance in an olfactory task, fearfulness, physical well-being, or body temperature. In a second study, NMA lesions restricted to the medial nucleus also decreased paternal behavior. Neurons in the medial nucleus of the amygdala appear to be essential for the normal expression of paternal care in this species.

The medial extended amygdala in male reproductive behavior. A node in the mammalian social behavior network.

Hormonal and chemosensory signals regulate social behaviors in a wide variety of mammals. In the male Syrian hamster, these signals are integrated in nuclei of the medial extended amygdala, where olfactory and vomeronasal system transmission is modulated by populations of androgen- and estrogen-sensitive neurons. Evidence from behavioral changes following lesions and from immediate early gene expression supports the hypothesis that the medial extended amygdala and medial preoptic area belong to a circuit that functions selectively in male sexual behavior. However, accumulated behavioral, neuroanatomical, and neuroendocrine data in hamsters, other rodents, and other mammals indicate that this circuit is embedded in a larger integrated network that controls not only male mating behavior, but female sexual behavior, parental behavior, and various forms of aggression. In this context, perhaps an individual animal's social responses can be more easily understood as a repertoire of closely interrelated, hormone-regulated behaviors, shaped by development and experience and modulated acutely by the environmental signals and the hormonal milieu of the brain.

Non invasive in vivo anatomical studies of the oscine brain by high resolution MRI microscopy.

We describe in this paper an in vivo Magnetic Resonance Imaging (MRI) procedure that allows one to obtain three-dimensional high quality images of the entire brain of small passerine birds such as the canary with a slice thickness of 58 micron and an image resolution of 78 microns. This imaging procedure was completed in 70 min on anaesthetised birds that later recovered uneventfully and could be reused for subsequent additional imaging. To illustrate the high resolution and anatomical detail that can be achieved, examples of coronal images through the entire hypothalamus are provided in the same sectioning plane as the previously published canary brain atlas. The data set can be used to create sections in any desired plane and the entire data set can be viewed from any point of view in a volume rendered image. This provides a useful tool in understanding the three-dimensional organisation of the brain. Similar procedures can also be applied on fixed brains and might allow an even better anatomical resolution of images because time constrains no longer limit the duration of image acquisition. The non-invasive MRI technique enables to study neuroanatomical features with a high resolution and without killing the animal subjects so that measures can be obtained in a same individual both before and after an experimental treatment.

MPOA and BNST lesions in male Syrian hamsters: differential effects on copulatory and chemoinvestigatory behaviors.

Electrolytic lesions were made in the medial preoptic area (MPOA) and bed nucleus of the stria terminalis (BNST) to evaluate their participation in the neural regulation of copulatory and chemoinvestigatory behaviors in male hamsters. Damage to either the MPOA or the BNST caused severe deficits in copulatory performance in a subset of the animals in each group. In the MPOA group all males displaying severe deficits had lesions which included a small central region of the caudal MPOA. In the BNST group, animals with severe copulatory deficits all had large lesions which covered most of both the medial and lateral parts of the nucleus. In contrast, MPOA and BNST lesions differentially affected chemoinvestigatory behaviors. MPOA lesions did not affect any of the males' anogenital investigation rates or attraction to female odors, even though some of these hamsters had stopped mating completely. Males with BNST lesions, on the other hand, all displayed significant reductions in their chemoinvestigatory responding even though the majority of them continued to mate normally. We suggest that the MPOA and BNST may in part regulate male sexual behavior by differentially responding to 'attractant' and 'mounting' substances within female hamster vaginal secretion.

Mating activates androgen receptor-containing neurons in chemosensory pathways of the male Syrian hamster brain.

Fos-immunoreactivity is induced during mating in the male Syrian hamster in limbic areas that relay chemosensory information and contain receptors for gonadal steroid hormones. The induction of Fos is an index of neuronal activation. After mating, c-fos expression is greatest in subnuclei of the medial amygdaloid nucleus (Me), bed nucleus of the stria terminalis (BNST), and medial preoptic area (MPOA). The present study determined if individual neurons in these activated subnuclei contain androgen receptors. We aim to understand how essential chemosensory and hormonal signals are integrated to control copulation. Adult male hamsters (n = 6) were allowed to mate with a sexually receptive female for 30 min. They were perfused 1 h later with 4% paraformaldehyde and 40 microns frozen sections were processed for immunocytochemistry using antisera against Fos (Cambridge Research Biochemicals) and the androgen receptor (G.S. Prins). The brains of three non-mated males were also processed for Fos immunocytochemistry. Mating significantly increased the number of Fos-immunoreactive neurons within subnuclei of Me, BNST, and MPOA relative to non-mated males (P < 0.05). These nuclei contained abundant androgen receptors. In the corticomedial amygdala, 20-40% of Fos-immunoreactive neurons in mated hamsters expressed androgen receptors. Although few androgen receptors are found in the anteromedial and postero-intermediate subdivisions of the BNST, these areas exhibited 26% and 47% co-localization, respectively. In posteromedial BNST, which contains large numbers of steroid receptor-containing neurons, androgen receptors were identified in 48% of Fos-immunoreactive neurons. In the MPOA, 54% of Fos-immunoreactive neurons expressed the androgen receptor throughout the rostrocaudal extent of the medial preoptic nucleus (MPN).(ABSTRACT TRUNCATED AT 250 WORDS)

Testosterone regulates substance P within neurons of the medial nucleus of the amygdala, the bed nucleus of the stria terminalis and the medial preoptic area of the male golden hamster.

The medial nucleus of the amygdala, bed nucleus of the stria terminalis, and medial preoptic area appear to mediate steroidal regulation of mating behavior in male rodents. The mechanism of action has not been determined. One way testosterone could enhance neuronal function is by increasing neurotransmitter levels, thus altering neuronal transmission. To assess this hypothesis, we examined the effect of castration and testosterone treatment on substance P levels in the neurons of these three brain regions. Brains from male Syrian hamsters that were (1) gonadally intact, (2) castrated for 13 weeks, or (3) castrated for 9 weeks and treated with testosterone for 4 weeks, were processed for substance P, and the numbers of substance P immunoreactive neurons in the medial nucleus of the amygdala, bed nucleus of the stria terminalis, and medial preoptic area were determined. Castration reduced the number of substance P neurons in the bed nucleus of the stria terminalis and medial preoptic area relative to those in intact hamsters; the number of substance P neurons in these regions was restored by testosterone treatment. Castration did not reduce the number of substance P neurons in the medial nucleus of the amygdala; however, testosterone treatment increased the numbers of these neurons when compared to intacts. Thus, testosterone regulates substance P levels in areas that regulate mating behavior. As substance P enhances male copulatory behavior our results suggest that testosterone may regulate copulatory behavior by enhancing substance P levels in medial nucleus of the amygdala, bed nucleus of the stria terminalis and medial preoptic area.

Prodynorphin- and substance P-containing neurons project to the medial preoptic area in the male Syrian hamster brain.

To determine if substance P- or prodynorphin-containing neurons of the medial nucleus of the amygdala and medial bed nucleus of the stria terminalis send projections to the medial preoptic area in the male Syrian hamster, we placed a fluorescent retrograde tract tracer (either Fluoro-gold, or rhodamine- or fluorescein-impregnated latex microspheres) into the medial preoptic area. Five to seven days later, the animals were treated with colchicine, allowed to survive for 48 h and the brains were processed for immunofluorescence histochemistry. Tissue sections were incubated in either rat anti-substance P or rabbit anti-C-peptide (the C-terminal sequence of dynorphin B) antiserum followed by incubation in either fluorescein- or rhodamine-conjugated anti-rabbit or anti-rat antiserum. When the injection site of retrograde tracer was centered within the caudal one-third of the medial preoptic area, labeled cell bodies were observed caudally in the medial part of the bed nucleus of the stria terminalis. Retrogradely labeled cell bodies were also observed in the posterodorsal subdivision of the medial nucleus of the amygdala. Both prodynorphin and substance P immunolabeling were observed in retrogradely labeled neurons in these two areas but fewer of these projection neurons were immunolabeled with substance P antiserum than with C-peptide antiserum. These projections may play a role in the peptidergic modulation of reproductive behavior in this species.

The colocalization of substance P and prodynorphin immunoreactivity in neurons of the medial preoptic area, bed nucleus of the stria terminalis and medial nucleus of the amygdala of the Syrian hamster.

To determine the extent of colocalization of substance P (SP) and prodynorphin peptides within neurons of the medial nucleus of the amygdala (AMe), medial bed nucleus of the stria terminalis (BNSTm) and medial preoptic area (MPOA), we incubated colchicine-treated Syrian hamster brain tissue in an antiserum mixture containing rat anti-SP antibody combined with 1 of 3 rabbit antibodies against prodynorphin peptides: anti-dynorphin A(1-17), anti-dynorphin B(1-13) or anti-C-peptide. This was followed by incubation in a secondary antiserum mixture containing fluorescein-labelled anti-rabbit and rhodamine-labelled anti-rat antibodies. Sections were viewed with an epifluorescence microscope using blue light excitation for fluorescein and green light excitation for rhodamine. Colocalization of SP and prodynorphin labelling was observed in neurons of the caudal parts of AMe, BNSTm and MPOA, areas which are essential for male mating behavior. The colocalization was most extensive in the dorsolateral part of the caudal MPOA, the caudodorsal part of the BNSTm, and in the posterodorsal subdivision of AMe. Although all 3 dynorphin peptides coexisted with SP in these areas, dynorphin B did so less than C-peptide, and dynorphin A less than dynorphin B.

A species-specific population of tyrosine hydroxylase-immunoreactive neurons in the medial amygdaloid nucleus of the Syrian hamster.

The medial amygdaloid nucleus (Me) is part of a neural pathway that regulates sexual behavior in the male Syrian hamster. To characterize the neurochemical content of neurons in this nucleus, brains from colchicine-treated adult male and female hamsters were immunocytochemically labeled using antibodies that recognize the catecholamine-synthesizing enzymes, tyrosine hydroxylase (TH), dopamine-beta-hydroxylase (DBH) and phenylethanolamine-N-methyltransferase (PNMT), as well as dopamine. A large population of TH-immunoreactive (TH-IR) neurons was observed throughout Me of male and female hamsters, primarily concentrated in the midrostral and caudal portions of the nucleus. The somata were generally small to medium in size and bipolar. Brains from animals that did not receive colchicine contained a limited number of TH-IR neurons in Me as reported previously. The DBH and PNMT antisera did not label any cells in Me of colchicine-treated animals, and the dopamine antiserum labeled neurons in the same location as the caudal group of TH-IR cells. Therefore, these caudal TH-IR neurons are interpreted to be dopaminergic. The rostral group of TH-IR neurons, on the other hand, may be producing only the immediate precursor of dopamine, L-3,4-dihydroxyphenylalanine (L-DOPA). The TH-synthesizing neurons in Me of the Syrian hamster appear to be a species-specific group of cells located outside of the previously described catecholaminergic cell groups.

Lesions of the ventral striatum mimic the effect of olfactory bulbectomy to prevent short photoperiod-induced testicular regression in golden hamsters.

Bilateral olfactory bulbectomy (BX) or bilateral transection of the rostral lateral olfactory tract (LOT) at the level of the anterior olfactory nucleus markedly increases gonadotropin secretion and prevents the testicular regression associated with maintenance on short photoperiod in golden hamsters. In an effort to further elucidate the neural tracts involved in this influence on gonadotropin secretion, lesions were placed in several potential pathways. Hamsters underwent sham surgery (SH), bilateral BX, or electrolytic or radiofrequency lesions of the: medial nucleus of the amygdala (MeX) caudal LOT just rostral to the medial nucleus of the amygdala (LOTX); or ventral striatum (VSX). Lesions were either bilateral or unilateral with contralateral olfactory bulbectomy. All animals were then placed on short photoperiod (LD 10:14) for 10 weeks and testicular size and body weight were assessed at weekly intervals. Lesion placement was assessed in brain sections stained with cresyl violet and animals with misplaced lesions were excluded. The following represent the number of animals in each group undergoing testicular regression in response to short photoperiod: SH: 32/35; BX: 8/31 (P < 0.01 vs. SH); MeX: 5/5; caudal LOTX: 8/9 and VSX: 3/8 (P < 0.05 vs. SH). Serum LH, FSH and testosterone at the end of the study correlated with the testicular regression data. These results suggest that the tonic inhibitory effect of the olfactory bulbs on gonadotropin secretion is mediated by fibers that exit the LOT rostral to the amygdala and project medially, either passing through or synapsing in the ventral striatum.

Role of the hypothalamic paraventricular nucleus in neuroendocrine responses to daylength in the golden hamster.

Daylength regulates reproduction in golden hamsters through a mechanism which involves the pineal indoleamine, melatonin. Retinal input to the suprachiasmatic nucleus of the hypothalamus (SCN) and sympathetic innervation of the pineal are critical to the inhibition of reproduction by short photoperiods. Since the hypothalamic paraventricular nucleus (PVN) receives extensive input from the SCN in the rat, and may influence autonomic function via its brainstem and spinal cord projections, we studied the role of this nucleus in photoperiodically induced gonadal regression in the hamster. Bilateral electrolytic destruction of either the paraventricular nucleus (PVN) or suprachiasmatic nucleus (SCN) of the hypothalamus completely blocked testicular regression induced by either blinding or exposure to short days (10L:14D). Lesions in the retrochiasmatic hypothalamus (RCA) which may have interrupted the pathway of previously identified efferents from the SCN to the PVN were also effective in preventing short day-induced gonadal regression. Pineal melatonin content was measured in intact and lesioned hamsters sacrificed 3-5 h before lights on, at the time of the expected nocturnal peak. While SCN and RCA lesions significantly reduced pineal melatonin content, PVN lesions were still more effective in this regard. We conclude that the hamster's neuroendocrine response to photoperiod is mediated by neural pathways which include retinohypothalamic input to the SCN and efferents from this nucleus to the PVN which travel dorsocaudally through the retrochiasmatic area of the hypothalamus. Effectiveness of lesions restricted to the PVN suggests that direct projections from the PVN to spinal autonomic centers convey photoperiodic information which regulates pineal, and hence gonadal, function.

Bilateral transection of the lateral olfactory tract but not removal of the vomeronasal organs inhibits short-photoperiod-induced testicular regression in golden hamsters.

It is now known that removal of the olfactory bulbs increases basal gonadotropin secretion and prevents short-photoperiod-induced testicular regression in Syrian hamsters. The experiments described in the present paper were an attempt to determine which neuronal systems associated with the olfactory bulbs are responsible for this influence on the reproductive neuroendocrine axis. In the first experiment, removal of the vomeronasal organ failed to influence gonadotropin secretion or testes weight in hamsters on long or short photoperiod, suggesting that the vomeronasal-accessory olfactory pathway is not individually responsible for the effect of the olfactory bulbs on gonadotropin secretion. In the second experiment, bilateral transection of the lateral olfactory tracts (LOT) did prevent short-photoperiod-induced testicular regression and the associated decrease in gonadotropin secretion. Since the nervus terminalis is confined to the surface of the medical olfactory bulb pathway, the results of LOT transection indicate that the nervus terminalis, which itself contains gonadotropin releasing hormone, does not mediate the influence of the olfactory bulbs on gonadotropin secretion. These results further suggest that the olfactory bulb influence on gonadotropin secretion is due to neural connections to the pyriform cortex, entorhinal cortex or amygdala.