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.     

Substance P and neurokinin A are colocalized in the central chemosensory pathway of the male golden hamster

The medial nucleus of the amygdala, bed nucleus of the stria terminalis and medial preoptic area play critical roles in the regulation of mating behavior in the male hamster. Destruction of these nuclei or the pathways that connect them severely disrupt copulation. We have begun identifying the neuropeptides contained in these neurons as a prelude to determining the role of peptide neurotransmitters in the regulation of male copulatory behavior. We have found that substance P is localized within these neurons and is regulated by gonadal steroids. In this study we report 1) that a closely related peptide, NKA (substance K), is also present in the medial nucleus of the amygdala, the bed nucleus of the stria terminalis and the medial preoptic area; 2) that all those neurons which contain SP also contain NKA and 3) testosterone also regulates the production of NKA. Thus, NKA may also play a role in the regulation of male copulatory behavior.     

Paced mating behavior in the female rat following lesions of three regions responsive to vaginocervical stimulation

The present study examines the effects of ibotenic acid lesions of the medial amygdala, the bed nucleus of the stria terminalis and the medial preoptic area on the display of paced mating behavior in female rats. Lesions of either the medial amygdala or the bed nucleus of the stria terminalis have no effect on the display of paced mating behaviors in ovariectomized, hormone-primed rats. In contrast, lesions of the medial preoptic area significantly lengthen contact-return latencies following intromissions and ejaculations and increase withdrawal from the male following intromissions. The present study demonstrates that the medial amygdala and the bed nucleus of the stria terminalis are not involved in the behavioral responses accompanying paced mating behavior, whereas the medial preoptic area is a critical component of the neural circuit mediating paced mating behavior as well as other appetitive aspects of mating.     

Gonadal steroids regulate behavioral responses to pheromones by actions on a subdivision of the medial preoptic nucleus

Mating behavior in male hamsters is initiated by pheromones, detected by two chemosensory systems which converge on the medial nucleus of the amygdala and the bed nucleus of the stria terminalis. Neurons in these areas project to the medial preoptic nucleus. All three of these areas contain androgen receptors. Using Fos as a marker of stimulation we have found that pheromones stimulate neurons in all three areas in intact males but fail to stimulate the magnocellular division of the medial preoptic nucleus of castrates. As this area plays a critical role in the regulation of male mating behavior our results suggest that steroids regulate mating by maintaining the responsiveness of the magnocellular division of the medial preoptic nucleus to pheromones.     

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 hte 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 castratio, 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. © 1993 Wiley-Liss, Inc.     

Sex differences in subregions of the medial nucleus of the amygdala and the bed nucleus of the stria terminalis of the rat

Sex differences are described in subregions of two nuclei of the rat brain: the medical nucleus of the amygdala (MA) and the bed nucleus of the stria terminalis (BNST). The volume of the posterodorsal region of the medial nucleus of the amygdala (MApd) is approximately 85% greater and the volume of the encapsulated region of the bed nucleus of the stria terminalis (BNSTenc) is approximately 97% greater in males than in females. The MApd and BNSTenc are distinct subregions of the MA and BNST. They exhibit intense uptake of gonadal hormones and are anatomically connected to each other and to other sexually dimorphic nuclei. The MA and BNST in general are involved in regulation of several sexually dimorphic functions, including aggression, sexual behavior, gonadotropin secretion and integration of olfactory information. Precise localization of sex differences in subregions of the MA and BNST, such as the MApd and BNSTenc, may facilitate understanding of the neural basis of such functions.     

Photoperiodic regulation of androgen receptor and steroid receptor coactivator-1 in Siberian hamster brain

Seasonal changes in the neuroendocrine actions of gonadal steroid hormones are triggered by fluctuations in daylength. The mechanisms responsible for photoperiodic influences upon the feedback and behavioral effects of testosterone in Siberian hamsters are poorly understood. We hypothesized that daylength regulates the expression of androgen receptor (AR) and/or steroid receptor coactivator-1 (SRC-1) in specific forebrain regions. Hamsters were castrated and implanted with either oil-filled capsules or low doses of testosterone; half of the animals remained in 16L/8D and the rest were kept in 10L/14D for the ensuing 70 days. The number of AR-immunoreactive (AR-ir) cells was regulated by testosterone in medial amygdala and caudal arcuate, and by photoperiod in the medial preoptic nucleus and the posterodorsal medial amygdala. A significant interaction between photoperiod and androgen treatment was found in medial preoptic nucleus and posterodorsal medial amygdala. The molecular weight and distribution of SRC-1 were similar to reports in other rodent species, and short days reduced the number of SRC-1-ir cells in posteromedial bed nucleus of the stria terminalis (BNST) and posterodorsal medial amygdala. A significant interaction between androgen treatment and daylength in regulation of SRC-1-ir was found in anterior medial amygdala. The present results indicate that daylength-induced fluctuations in SRC-1 and AR expression may contribute to seasonally changing effects of testosterone.     

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.     

Differential Localization of Estrogen Receptors in Various Vasopressin Synthesizing Nuclei of the Rat Brain

Vasopressin (VP) cells in the bed nucleus of the stria terminalis, medial amygdaloid nucleus and supraoptic and paraventricular nuclei are influenced by gonadal steroids. The present paper examined whether VP cells in the bed nucleus of the stria terminalis, medial amygdaloid nucleus, and supraoptic and paraventricular nuclei contain estrogen receptors. Brains from adult short-term castrated, colchicine-treated male rats were fixed with 4% paraformaldehyde and 0.5% glutaraldehyde. In the immunocytochemical double-staining procedure Vibratome sections were first incubated with an estrogen receptor antibody (#H222) and stained with diaminobenzidine-Ni⁺. Following methanol-hydrogen peroxide washes, sections were incubated with anti-neurophysin and stained with diaminobenzidine. Parvocellular cells in the bed nucleus of the stria terminalis and medial amygdaloid nucleus were double-stained with a blue-black nucleus (indicating the estrogen receptors) surrounded by brown cytoplasm (resulting from VP-neurophysin-immunoreactivity). Our results provide the first direct anatomical evidence supporting the hypothesis that gonadal steroids' influence of parvocellular VP cells in the bed nucleus of the stria terminalis and medial amygdaloid nucleus is mediated directly via estrogen receptors localized in nuclei of VP neurons. We were unable to co-localize any estrogen receptors in VP and oxytocin cells of magnocellular size in the supraoptic, paraventricular and anterior commissural nuclei, suggesting that estrogen indirectly affects these magnocellular hypothalamic cells.     

Projections from Ventrolateral Hypothalamic Neurons Containing Progestin Receptor- and Substance P-Immunoreactivity to Specific Forebrain and Midbrain Areas in Female Guinea Pigs

Many neurons within the ventrolateral hypothalamus in guinea pigs contain estrogen-induced progestin receptors as well as substance P. Retrograde tracing combined with immunocytochemistry was used to determine the specific projections of this subset of steroid- sensitive cells. Unilateral Fluoro-Gold injections into the dorsal midbrain, including the central gray, labeled a large proportion of the ventrolateral hypothalamic neurons immunoreactive for both progestin receptors and substance P (approximately 30%); substantially fewer of these neurons were labeled by unilateral Fluoro-Gold injections into the preoptic area (approximately 6%), medial amygdala (approximately 10%), or the bed nucleus of the stria terminalis (approximately 11 %). The projections of progestin receptor-immunoreactive neurons in the ventrolateral hypothalamus were similar to those of progestin receptor/substance P double-labeled neurons, while a slightly lower percentage of the ventrolateral hypothalamic, substance P-immunoreactive neurons tended to project to each of these areas. These pathways may prove to be components of the neural circuitry underlying a variety of functions influenced by gonadal steroid hormones and substance P, such as female sexual behavior, salt intake, nociception and aggression.     

Intraneuronal Convergence of Tactile and Hormonal Stimuli Associated with Female Reproduction in Rats

Stimulation of the vagina and cervix, by mating or manual probing, elicits many behavioral and endocrine changes associated with female reproduction in rats. We and others have identified neurons in the medial preoptic area, medial division of the bed nucleus of the stria terminalis, posterodorsal portion of the medial amygdala, ventromedial hypothalamus, dorsomedial hypothalamus and midbrain central gray that increase Fos expression in response to vaginal-cervical stimulation (VCS). In the present study, we used a double-label immunofluorescent technique to determine if any of these VCS-responsive neurons also contained estrogen receptor-immunoreactivity. We found that over 80% of the VCS-induced Fos-IR neurons in the medial division of the bed nucleus of the stria terminalis also contained estrogen receptor-immunoreactivity. Furthermore, high percentages of VCS-responsive neurons in the medial preoptic area, posterodorsal medial amygdala, ventromedial hypothalamus and midbrain central gray contained estrogen receptor-immunoreactivity as well. These results suggest that sensory and hormonal information associated with female reproduction converge on specific populations of neurons and may be integrated at the molecular level within these neurons.     

Androgen and estrogen concentrating neurons in chemosensory pathways of the male Syrian hamster brain.

The medial preoptic area (MPOA), bed nucleus of the stria terminalis (BNST), and medial amygdaloid nucleus (Me) are essential for male sexual behavior in the Syrian hamster. These nuclei received chemosensory stimuli and gonadal steroid signals, both of which are required for mating behavior. The objective of this study was to compare the distribution of androgen- and estrogen-concentrating neurons in MPOA, BNST, and Me in the adult male hamster using steroid autoradiography for estradiol (E2), testosterone (T) and dihydrotestosterone (DHT). Adult males (n = 4 per group) received two i.p. injections of tritiated steroid 4-7 days after castration. Six-microns frozen sections through the brain were mounted onto emulsion-coated slides, and exposed for 11-16 months. In MPOA, BNST, and Me, neurons were more abundant and heavily labelled after [3H]E2 treatment than after either [3H]T or [3H]DHT. Tritiated estradiol- and DHT-labeled cells were found throughout the rostrocaudal extent of Me, with a high concentration in posterodorsal Me. Tritiated testosterone treatment labelled cells largely within posterodorsal Me. In MPOA, the majority of E2-, T-, and DHT-labelled neurons were in the medial preoptic nucleus (MPN) and the preoptic continuation of the posteromedial bed nucleus of the stria terminalis (BNSTpm). Few T-labelled cells were present outside these subdivisions. In the BNST, E2- and DHT-labelled neurons were present in all subdivisions, whereas T labelling was confined to the antero- and posteromedial subdivisions of BNST. These results suggest that the distribution of androgen- and estrogen receptor-containing neurons overlap considerably in nuclei which transmit chemosensory signals in the control of mating behavior.     

Adrenalectomy alters the response of neurons in the bed nucleus of the stria terminalis to electrical stimulation of the medial amygdala

This study was performed to characterize the effects of adrenalectomy (AD)Q on electrical activity and synaptic responses of bed nucleus of the stria terminalis (BNST) and preoptic area (PDA) neurons, which are involved in the control of limbic-hypothalamo-pituitary-adrenocortical (LHPA) activity. Adrenalectomy altered the response of BNST neurons to medial amygdala (AME) stimulation, increasing the proportion of excitatory responses and reducing the number of cells inhibited. No such effects were found for neurons within the PDA. The basal activity of neurons recorded within the BNST and PDA, as well as the latencies and duration of responses, was not af2fected. The specificity of the effects upon BNST, but not POA, neurons suggests that the response of BNST neurons to AME stimulation is corticosteroid dependent, whereas the response of preoptic neurons is not.     

Experience facilitates vomeronasal and olfactory influence on Fos expression in medial preoptic area during pheromone exposure or mating in male hamsters

Chemosensory stimuli are essential for mating in male hamsters but either main olfactory or vomeronasal input is sufficient in sexually experienced males. Activation in central chemosensory pathways and medial preoptic area, after stimulation with female chemosignals or after mating, was estimated by counting neurons expressing Fos protein in experienced and naive males, with or without vomeronasal organ lesions. Regions counted included main and accessory olfactory bulbs, corticomedial amygdala, bed nucleus stria terminalis and medial preoptic area. Chemosensory stimulation was more effective in activating medial preoptic area in experienced than in naive males. In experienced males with vomeronasal organs removed, main olfactory input was as effective in activating medial preoptic area as was the combination of main and accessory input available to intact animals. As previously reported, the main olfactory input remaining after vomeronasal lesions in naive males was poorly effective in activating medial preoptic area, and these animals had impaired mating behavior. The change in access of chemosensory input to medial preoptic area after experience suggests that an experience-dependent synaptic modulation in this pathway, perhaps in the amygdala, may underlie some changes in mating behavior with experience.     

Alteration in [11C]vorozole binding to aromatase in neuronal cells of rat brain induced by anabolic androgenic steroids and flutamide

In a previous study, we demonstrated that androgenic-anabolic steroids increased aromatase expression in the bed nucleus of stria terminalis and preoptic area in rat brain, as evaluated using autoradiography with [11C]vorozole, a potential positron emission tomography tracer for aromatase. In this study, we explored whether the increase in aromatase binding is mediated via androgen receptors and whether this increase occurs in neurons or glial cells. Rats were given nandrolone decanoate (15 mg/kg body weight once every 3 days) and flutamide (20 mg/kg/day) alone or in combination for 20 days. Results indicated a significant increase of [11C]vorozole binding by nandrolone decanoate in the bed nucleus of the stria terminalis and preoptic area, as in our previous study. Flutamide treatment, on the other hand, decreased [11C]vorozole binding in the bed nucleus of the stria terminalis, preoptic area, and medial amygdala. Immunohistochemical examination demonstrated that upregulation of aromatase expression occurred in neurons. Our findings suggest that aromatase is regulated through an androgen receptor-mediated system. This aromatase-specific tracer and the positron emission tomography technique could be useful for exploring the role of aromatase in anabolic androgenic steroids abusers.     

Gonadal steroid control of preprocholecystokinin mRNA expression in the limbic-hypothalamic circuit: Comparison of adult with neonatal steroid treatments

The neuropeptide cholecystokinin (CCK) is involved in the regulation of female, but not male, reproductive behavior. In both sexes, estrogen regulates the expression of CCK in adulthood within the bed nucleus of the stria terminalis and medial amygdaloid nucleus. These areas are parts of an interconnected limbic system-hypothalamic circuit, the development of which is influenced by estrogen during the early postnatal period. This is the same period during which central nervous system (CNS) expression of CCK is dramatically increased, suggesting that the male and female patterns of CCK expression may be the result of early postnatal exposure to estrogen. In the present experiment, the expression of preprocholecystokinin (pCCK) mRNA was determined by in situ hybridization with an isotopically labeled pCCK complementary RNA and emulsion autoradiography in animals whose neonatal and adult gonadal steroid levels had been manipulated. The number of pCCK-expressing cells in animals that were gonadectomized as adults was determined by neonatal estrogen, but stimulation with steroids in adulthood induced a similar number of pCCK-expressing cells in both sexes in the medial amygdala and bed nucleus of the stria terminalis. Neonatal treatment of females with estrogen or testosterone, followed by ovariectomy in adulthood, eliminated the sex difference in pCCK mRNA expression. Males treated neonatally with the aromatase inhibitor androstenedione (to block metabolism of testosterone to estrogen) and orchidectomized in adulthood had a level of pCCK mRNA expression that was similar to that of ovariectomized females. These data suggest that, during neonatal development, estrogen determines the constitutive expression of pCCK mRNA in the medial amygdala and bed nucleus of the stria terminalis, resulting in higher levels of pCCK mRNA expression in males than in females. However, exogenous gonadal steroids induce the same levels of pCCK mRNA expression in adult females, indicating that the levels of gonadal steroids and the patterns of their secretion are the predominant influences on the sexually dimorphic adult levels of pCCK mRNA expression. © 1994 Wiley-Liss, Inc.     

Evidence for a ventral non-strial pathway from the amygdala to the bed nucleus of the stria terminalis in the male golden hamster

Male hamsters in which the stria terminalis (ST) had been interrupted either by electrolytic lesions or knife cuts, or normal control males, received iontophoretic injections of horseradish peroxidase in either the bed nucleus of the stria terminalis (BNST) or the medial preoptic-anterior hypothalamic area (MPOAH). Comparison of intact and ST-lesioned brains revealed the existence of a ventral non-strial pathway, from cells in the medial amygdaloid nucleus (M) to the preoptic portion of the BNST but not to the MPOAH. Since bilateral lesions of M completely eliminate male hamster mating behavior, but ST lesions do not, we suggest that the ventral pathway to the BNST may be an important route by which M influences male copulatory behavior.     

Intrinsic GABAergic neurons in the rat central extended amygdala

The present study examined the distribution, morphology, and connections of gamma-aminobutyric acid-immunoreactive (GABA-IR) neuros in the three principal components of the central extended amygdala: the central amygdaloid nucleus, the bed nucleus of the stria terminalis (BNST) and the sublenticular substantia innominata. In the central nucleus, large numbers of GABA-IR neurons were identified in the lateral, lateral capsular, and ventral subdivisions, though in the medial subdivision, GABA-IR neurons were only present at very caudal levels. Combined immunocytochemistry-Golgi impregnation revealed that GABA-IR neurons in the lateral central nucleus were medium-sized spiny neurons that were morphologically similar to GABAergic neurons in the striatum. Injections of horseradish peroxidase into the bed nucleus of the stria terminalis labeled a major proportion of the GABA-IR neurons in the central nucleus. In the bed nucleus, the majority of GABA-IR neurons were located in the anterolateral subdivision, ventral part of the posterolateral subdivision and the parastrial subdivision. GABA-IR neurons in the anterolateral bed nucleus were of the typical mediumsized spiny type. Injections of horseradish peroxidase into the central nucleus labeled a few GABA-IR neurons in the posterior part of the anterolateral bed nucleus. GABA-IR neurons were identified in the sublenticular substantia innominata and medial shell of the nucleus accumbens and contributed to the continuum of GABA-IR extending from the central nucleus to the bed nucleus. Injections of horseradish peroxidase (HRP) into the central nucleus, but not the BNST, labeled a few GABA-IR neurons in the substantia innominata. The data point to GABA-IR neurons being a characteristic feature of the central extended amygdala and that GABA-IR neurons participate in the long intrinsic connections linking the major components of this structure. Since lesions of the stria terminalis and basolateral amygdaloid nucleus failed to deplete GABA-IR terminals in the central nucleus, the role of GABA in local and short intrinsic connections in the central extended amygdala is discussed. Further, physiological findings implicating the intrinsic GABAergic system of the central extended amygdala in the tonic inhibition of brainstem efferents are reviewed.     

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.