Limbic projections to the ventromedial hypothalamus of the opossum

Limbic projection systems which influence neuronal activity of the ventromedial nucleus of the hypothalamus of the opossum were studied by an evoked potential method. Bipolar concentric recording electrodes were placed in the ventromedial nucleus to monitor activity while exploring the forebrain with a stimulating electrode delivering single rectangular pulses (0.5 msec, 0.4 ma, 0.5–1/sec). Responses in the ventromedial nucleus were consistently evoked by stimulating three forebrain systems. From the dorsal septum, responses with one monophasic component were evoked (latency 3–9 msec). Stimulation more ventrally in the septum evoked, in addition, a second slower component. Stimulation of the corticomedial amygdala produced biphasic responses in the ventromedial nucleus at latencies of 4–9 msec; stimulation of the stria terminalis, the efferent pathway from this division of the amygdaloid complex, yielded almost identical responses at 4- to 6-msec latencies. Responses in the ventromedial nucleus were also recorded to stimulation of the dorsal hippocampus and adjacent fimbria (latencies, 3–7 msec). These responses demonstrate three possibly direct limbic influences on the ventromedial nucleus of a metatherian mammal, and provide new data on the anatomic relationships of this behaviorally significant hypothalamic area.     

Ventromedial nucleus of the hypothalamus: convergent excitatory and inhibitory responses to fimbria and stria terminalis stimulation

Field and extracellular unitary potentials were recorded in the ventromedial nucleus of the hypothalamus (VMH) of urethane-anesthetized rabbits after stimulation of the fimbria and stria terminalis. Stimulation of the lateral portion of the fimbria, which carries fibers from the ventral subiculum of the hippocampal formation, evoked a two-component response. An early excitatory response, with an average latency of 10 msec, predominated along the lateral margins of the VMH. A later inhibitory potential with an average latency of 15 msec was seen predominantly within the central portions of the VMH. Stimulation of the dorsal component of the stria terminalis produced two similar response patterns: an early excitatory response with an average latency of 16 msec, followed by an inhibitory potential with an average latency of 25 msec. The topographical distribution of these two components of the response was nearly identical to that produced by lateral fimbria stimulation. In contrast, stimulation of the ventral component of the stria terminalis evoked a simple excitatory response with an average latency of 10 msec which was maximal within the core of the VMH. Extracellular unitary recordings showed that the early negativity associated with stimulation of each of these three pathways reflects a monosynaptic excitation of VMH cells and that there was convergence of the three excitatory inputs at the single cell level.     

Projections of the septum to the lateral hypothalamus

Evoked potentials recorded in the lateral hypothalamus (LH) of the rat to stimulation of the septum were observed to have response components of 3–6, 10–14, and 18–23 msec. These components were elicited from different regions of the septum; the 3–6 and 10–14 msec components from dorsal and midline regions corresponding to the projection field of the precommissural fornix from the hippocampus and the 18–23 msec component from a ventrolateral region corresponding to projections of the stria terminalis. Stimulation of the hippocampus and stria terminalis evoked responses in the LH of similar configuration but with latencies longer than the 10–14 and 18–23 msec components, respectively. Lesions in the dorsal midline and ventrolateral septum attenuated these responses suggesting that the precommissural fornix and stria terminalis are the pathways mediating the septal evoked components. These data provide a neuroanatomical framework for the dual role of the septum on response patterns elicited from the hypothalamus.     

Amygdaloid and pontine projections to the ventromedial nucleus of the hypothalamus

Amygdaloid and pontine projections to the feline ventromedial nucleus of the hypothalamus (HVM) were studied with retrograde transport of horseradish peroxidase (HRP) and anterograde transport of tritiated amino acids. Following injections of HRP into HVM, amygdaloid neurons were labeled in the ipsilateral cortical and medial nuclei and the ventral portion of the parvocellular part of the basal nucleus. In experiments in which HRP was injected into the tuberal hypothalamus following stria terminalis lesions, it was determined that amygdaloid neurons projecting to HVM by way of the stria terminalis were located in the cortical and medial nuclei while those projecting through another route, presumably the ventral amygdalofugal pathway, were found in the rostral part of the medial nucleus and the parvocellular basal nucleus. Following HRP injection into lateral hypothalamus at the level of HVM, labeled neurons were seen in the magnocellular basal nucleus. After preoptic injections, neurons containing the HRP reaction product were in cortical and medial nuclei and magnocellular and parvocellular parts of the basal nucleus. In addition to cells in the amygdala, rostral pontine neurons were labeled after HRP injections into HVM. The cells were located ipsilateral to the injection, mostly in the dorsal nucleus of the lateral lemniscus, lateral and dorsolateral to the brachium conjunctivum. The pontine cells labeled following HVM injections of HRP were different from those labeled following lateral hypothalamic and preoptic region injections. The pontine projection to HVM was confirmed using axoplasmic transport autoradiography. A mixture of tritiated leucine and tritiated proline was injected into the lateral pontine region labeled after HRP injections into HVM. Labeled axons ascending in the medial forebrain bundle terminated throughout the rostro-caudal extent of HVM.     

Effect of fastigial stimulation on neuronal activity in the red nucleus

Discharges from cells recorded with microelectrodes in the red nucleus of anesthetized cats could be increased or decreased by stimulation of contralateral or ipsilateral cerebellar fastigial nucleus. Contralateral stimulation altered the activity in 10% of the cells studied, whereas ipsilateral stimulation altered 22% of the cells. Furthermore, ipsilateral stimulation evoked responses with latencies of 2–10 msec (mean = 6.4). We concluded that there are polysynaptic (but no monosynaptic) pathways from both fastigial nuclei to the red nucleus.     

Heterogeneous synaptic inputs from the ventrolateral periaqueductal gray matter to neurons responding to somatosensory stimuli in the rostral ventromedial medulla of rats

The ventrolateral cell column of the midbrain periaqueductal gray matter (vl-PAG) plays a major role in the attenuation of pain behaviour. It is established that this effect is exerted via modulation of neuronal activities in the rostral ventromedial medulla (RVM). Until recently it has been generally accepted that the vl-PAG exerts its modulatory effects upon RVM neurons through a direct monosynaptic pathway. However, recent data suggest that an additional indirect, di- or polysynaptic pathway may also exist. Using in vivo intracellular recordings we tested this hypothesis, by studying synaptic responses of somatosensory receptive RVM neurons evoked by electric stimulation of the vl-PAG in rats. RVM neurons were regarded as somatosensory receptive if they responded to electrical stimulation of the sciatic nerve. Most of the recorded RVM cells were excited by vl-PAG stimulation. Some of them responded with a short onset latency (3.6+/-0.9 ms) corresponding to monosynaptic excitation. All of these neurons were also excited by sciatic nerve stimulation at nociceptive intensities. In contrast to this, another proportion of the recorded RVM neurons responded with a four times longer (14.8+/-3 ms) onset latency to the vl-PAG stimulation, corresponding to polysynaptic modulation. All of these neurons were inhibited by sciatic nerve stimulation. The findings show that RVM neurons receive heterogeneous monosynaptic and polysynaptic inputs from the vl-PAG. The results also suggest that the monosynaptic and polysynaptic pathways modulate the activity of functionally distinct groups of RVM neurons.     

Electrophysiological study of the response of medial prefrontal cortex neurons to stimulation of the basolateral nucleus of the amygdala in the rat.

The neural connections from the basolateral nucleus of the amygdala (BLA) to the medial prefrontal cortex (MPC) in urethane-anesthetized rats were investigated. Extracellular recordings were made from 200 neurons with spontaneous firing in the MPC, and the BLA was electrically stimulated. The most frequent response to BLA stimulation was inhibition (63.5%). Excitatory responses were found in 17 units (8.5%), while 56 neurons (28%) did not change their spontaneous firing after BLA stimulation. Inhibitory responses showed a wide range of latencies, suggesting the coexistence of mono- and polysynaptic pathways. On the contrary, the excitatory responses seem to be mediated by a monosynaptic pathway. BLA projections to the MPC play a predominantly inhibitory role in the spontaneous activity of prefrontal neurons. This inhibition may modulate central motor systems and motivated behaviors.     

Convergence of projections from the rat hippocampal formation, medial geniculate and basal forebrain onto single amygdaloid neurons: an in vivo extra- and intracellular electrophysiological study.

We recorded extra- and intracellular responses from rat amygdaloid neurons in vivo after electrical stimulation of the hippocampal formation (dentate gyrus, hippocampal fields CA3 and CA4, entorhinal cortex, subicular complex); medial geniculate; and basal forebrain (diagonal band, ventral pallidum, olfactory tubercle, nucleus accumbens, bed nucleus of stria terminalis, lateral preoptic area, substantia innominata). Stimulation of hippocampal formation structures evoked IPSPs or EPSP-IPSP sequences in which the IPSP had a lower threshold than the EPSP. Recordings from candidate inhibitory neurons in the amygdala indicated that excitatory afferents from the hippocampal formation contact both amygdaloid inhibitory and principal neurons (feedforward inhibition), and that the inhibitory neurons have a lower threshold of activation. Medial geniculate stimulation also evoked EPSP-IPSP sequences. In marked contrast to these results, stimulation of basal forebrain structures evoked short latency IPSPs in amygdaloid neurons. This provides the first physiological evidence for direct inhibition of the amygdala by the basal forebrain. Basal forebrain stimulation also evoked EPSP-IPSP sequences in amygdaloid neurons. Individual amygdaloid neurons could show responses to stimulation of the hippocampal formation, basal forebrain, and medial geniculate, indicating that synaptic input from these areas converges onto single amygdaloid cells. The findings provide further information about the synaptic organization of afferents to the amygdala, and indicate that single amygdaloid neurons play a role in the synaptic integration of input from these diverse sources.     

Postsynaptic potentials in neurons of the pigeon''s optic tectum in response to afferent stimulation from the retina and other visual structures: an intracellular study

The responses of the cells in the pigeon's optic tectum to electrical stimulation of the contralateral optic nerve, the ipsilateral visual Wulst and the opposite optic tectum were intracellularly recorded. Optic nerve or visual Wulst stimulation elicited 3 types of responses: (1) a pure EPSP which gave rise to one or two action potentials; (2) an EPSP which sometimes gave rise to a spike, followed by an IPSP; and (3) a pure IPSP. Opposite tectum stimulation evoked in the tectal cells either a pure IPSP or a pure EPSP. The mono- or polysynaptic nature of the pathways involved in the excitatory and inhibitory responses of the tectal cells was assessed by increasing the frequency of the optic nerve stimulation. At low stimulus rates (2-6 Hz), all the excitatory events showing latencies longer than 5 ms were blocked suggesting that they were polysynaptic. Excitatory events having latencies shorter than 5 ms were generally able to follow high rate frequencies of optic nerve stimulation (40, 50 or 90 Hz) and we considered them to be monosynaptic. All but 3 IPSPs evoked by optic nerve stimulation, were blocked by stimulus rates beyond 5 Hz. Thus, although most IPSPs are generated through polysynaptic paths, direct retino-tectal inhibitory paths may also exist. The latency of the responses of individual cells to optic nerve, visual Wulst and opposite tectum stimulation show that the polysynaptic IPSPs to optic nerve stimulation did not involve relays in the visual Wulst or the opposite tectum.     

Further studies on peripeduncular-hypothalamic pathways involved in sexual behavior in the female rat

A series of preliminary experiments demonstrated that injection of 22 mM sodium pentobarbital in the brain of the rat blocked synaptic transmission at the site of injection; the same concentration of pentobarbital did not block fiber conduction. Based on the latter information, 22 mM pentobarbital was applied to different parts of the peripeduncular-hypothalamic pathways responsible for the conduction and generation of potentials evoked in the ventromedial nucleus (VMN) by stimuli applied to the peripeduncular nucleus (PPN), to determine whether participation of the amygdala and bed nucleus of the stria terminalis involves the transynaptic activation of neuron somas at these places or the operation of passing fibers only. We determined that potentials evoked in the VMN by PPN stimulation involves synaptic activity in both the lateral amygdaloid nucleus and the bed nucleus of the stria terminalis. Both structures receive PPN-originated activity independently, and both structures contribute to the generation of PPN-VMN evoked responses, presumably through temporal or spatial summation of inputs in the VMN. We also showed that activity in the lateral amygdaloid nucleus is conducted toward the VMN along fibers in the stria terminalis. We propose that the synaptic interactions thus demonstrated serve as integrating relays for different sensory modalities and hormone actions regulating sexual behavior.     

Correlation between intrinsic firing patterns and thalamocortical synaptic responses of neurons in mouse barrel cortex.

We used a thalamocortical slice preparation to record both spike trains and synaptically evoked responses from neurons of mouse barrel cortex. Cells were classified as regular spiking (RS), intrinsically bursting (IB), or fast spiking (FS) according to their temporal firing patterns when injected with current. RS cells were further separated into two subtypes, RS1 and RS2 cells, the latter encountered only in the infragranular layers. Synaptic responses were elicited by focal electrical stimuli in the ventrobasal nucleus of the thalamus (VB) while holding the cells at different membrane potentials. Postsynaptic potentials were classified as excitatory (EPSPs) or inhibitory (IPSPs), and their latencies were measured from the onset of the extracellularly recorded fiber volley in layer IV. EPSPs fell into three groups, according to latency. Those in the early cluster had latencies shorter than 1 msec and were coincident with the postsynaptic layer IV population response; they were considered monosynaptic. A second group, with latencies between 1.3 and 2.5 msec, were coincident with all IPSPs and were classified as disynaptic. The rest had latencies longer than 5 msec and were considered polysynaptic. The synaptic order of a cell was correlated with its laminar position and its electrophysiological class. Specifically, monosynaptic responses were restricted to infragranular RS cells and to FS cells, while disynaptic EPSPs were found in supragranular RS cells and in IB cells. Disynaptic IPSPs were found in both deep and superficial layers; in the deep layers they nearly always followed monosynaptic EPSPs, while in the superficial layers they were mostly found in isolation. We conclude that the intrinsic spiking characteristics of a neuron are an important determinant of its position in the cortical circuit and may have a substantial role in determining its response properties.     

Mechanism of inhibition by the amygdala in the lateral hypothalamic area of rats

OOMURA, Y. AND T. ONO. Mechanism of inhibition by the amygdala in the lateral hypothalamic area of rats. BRAIN RES. BULL. 8(6) 653–666, 1982.—The inhibition of neuronal activity in the lateral hypothalamus (LHA) of the rat by the basolateral nucleus of the amygdala (AL) was investigated by analyzing evoked potentials, single unit discharges and intracellular synaptic potentials. A single volley to the AL induced a negative-positive-wave in the LHA. The negative-wave threshold was lower than that of the positive wave. Analysis of depth profiles showed that the negative- and positive-waves appeared first at the dorsal margin of the LHA, peaked within the LHA, and were clearly different from each other. The effects of acute lesions showed the negative-wave to be conducted through the direct amygdalo-hypothalamic pathway. The positive-wave; through the stria terminalis. Stimulation of the stria terminalis produced positive evoked potentials with latencies shorter than those of the positive-waves. When conditioning and test stimuli were delivered to the AL, the negative-wave was inhibited for about 90 msec by the evoked positive-wave. Single AL stimuli evoked single unit discharges followed by inhibition of spontaneous firing for about 100 msec. Single stria terminalis stimuli inhibited spontaneous firing for the duration of the positive evoked potential. Intracellular LHA recording during single AL stimuli showed the presence of an EPSP followed by a 100 msec long lasting IPSP. The negative and positive extracellular potentials corresponded to these synaptic potentials. Inward current injection of 1 to 1.4 nA reversed the IPSP's indicating a -15 mV hyperpolarization difference between the IPSP reversal potential and the resting potential in LHA cells. The ionic mechanism of the IPSP is also discussed.     

Neurophysiology of limbic system pathways in the rat: Projections from the amygdala to the entorhinal cortex

We studied the responses of rat entorhinal neurons to electrical stimulation of the amygdala. Four main results were obtained: (1) excitatory postsynaptic potentials were recorded in entorhinal neurons in response to electrical stimulation of the amygdala. Cells in layers II, III and V of the entorhinal cortex were responsive. (2) Excitatory responses were followed by inhibitory postsynaptic potentials. (3) Frequency potentiation of both excitatory and inhibitory responses was observed when 10/s stimulation was used. (4) Three amygdala neurons were antidromically activated by entorhinal stimulation; and two layer II entorhinal cells that were excited by amygdala stimulation were also antidromically activated by dentate gyrus stimulation. These results provide evidence for a monosynaptic, excitatory projection from the amygdala to the entorhinal cortex. In addition, the data indicate that amygdala neurons are only one synapse removed from the excitation of dentate gyrus granule cells.     

Anesthetic-dependent excitability changes in the hypothalamic ventromedial nucleus of the rat

In acute experiments in rats anesthetized with urethane, the field potentials, population spike, and unit activity evoked in the hypothalamic ventromedial nucleus (HVM) by amygdaloid stimulation are significantly increased with respect to control when preceded by a conditioning volley at 20- to 100-ms intervals. Under pentobarbital anesthesia, in contrast, the evoked responses were inhibited by the conditioning stimulus for similar interstimulus intervals. In unanesthetized animals chronically implanted with stimulating and recording electrodes, a facilitation of responses by a conditioning stimulus was observed when they were awake or anesthetized with urethane. When the same animals were anesthetized with pentobarbital the HVM evoked response was inhibited by a conditioning pulse. Frequency facilitation and post-tetanic potentiation of HVM responses were markedly enhanced under urethane, whereas in pentobarbital-anesthetized animals inhibition predominated. Picrotoxin reversed the inhibition under pentobarbital to facilitation. These results suggest that the HVM neuron population is under both excitatory and inhibitory influences from the amygdala, the former being predominant in awake and urethane-anesthetized animals and the latter being expressed under pentobarbital anesthesia and is probably mediated by γ-aminobutyric acid.     

Pathways conducting amygdaloid influence on feminine sexual behavior in the rat

In ovariectomized female rats primed with estradiol benzoate and progesterone, electrochemical stimulation of medial amygdaloid nucleus, anterior part (AMEa), increased lordotic frequency, but this effect was not found in rats with transection of the stria terminalis (ST) or the ventral amygdalofugal pathways (VAF). On the other hand, the decrease in lordotic frequency observed after stimulation of the posterior part of the lateral amygdaloid nucleus (ALp), persisted in animals with transection of the VAF, but was eliminated in animals with section of the ST. The effect of transecting these pathways on the multiunit and field responses evoked in the hypothalamic ventromedial nucleus (VMN) by stimulation of both amygdaloid nuclei was also studied. Responses evoked by AMEa stimulation were smaller or disappeared after lesion of either the ST or the VAF, but responses evoked by ALp stimulation were affected by lesion of the ST only. Findings are interpreted as suggesting that the ST conducts activity originating in AMEa and ALp which affects sexual receptivity in a complementary fashion, whereas VAF contains AMEa originated fibers whose activation increase receptivity. These behavioral effects may be the consequence of modulatory action on VMN neuronal activity.     

Connections of the hypothalamic paraventricular necleus with the neurohypophysis, median eminence, amygdala, lateral septum and midbrain periaqueductal gray: An electrophysiological study in the rat

Extracellular recordings were obtained from 555 paraventricular (PVN) nucleus neurons in pentobarbital-anesthetized male rats. Cells were examined for their spontaneous activity patterns and response to single 1-Hz electrical stimulation of the neurohypophysis, median eminence, amygdala, lateral septum (LS) and midbrain periaqueductal gray (PAG).Neurohypophyseal stimulation evokedantidromic activation from 109 neurons. Among spontaneouly active neurohypophyseal neurons, evidence of a recurrent inhibitory pathway usually required pituitary stimulus intensities twice threshold for antidromic activation. Orthodromic excitatory or inhibitory responses followed amygdala and LS stimulation, but not PAG stimulation. The amygdala influence was predominantly inhibitory to ‘phasic’ (putative vasopressin-secreting) PVN neurohypophyseal neurons. Neurohypophyseal stimulation evokedorthodromic responses from 124 PVN cells; some of these neurons were also responsive to stimulation in other sites.Median eminence stimulation evoked antidromic responses from 37 PVN neurons; some of these cells also displayed phasic activity but not evidence for recurrent inhibition. Twelve cells in this group were also activated antidromically from both the median eminence and the neurohypophysis; collision tests suggest that the median eminence innervation may be an axon collateral of a neurohypophyseal pathway. Amygdala stimulation was inhibitory to some cells in this category.Amygdala, LS and PAG stimulation evoked antidromic activation from a small number of PVN cells, but none of these cells appeared to innervate more than one area, including the neurohypophysis, and none displayed phasic activity. Orthodromic responses were recorded among other PVN neurons after stimulation in these sites; however, PAG stimulation was the least effective stimulation area.These observations provide additional electrophysiological data that confirm efferent PVN connections to all areas tested, afferent connections from amygdala and LS but not PAG, and the possibility for coordinated activity among PVN neurons through local recurrent or common afferent connections.     

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.     

A comparison of the effects of electrical stimulation of the lateral and ventromedial hypothalamus on the activity of neurons in ventral tegmental area and substantia nigra

Extracellular unit recordings were obtained from neurons in the ventral tegmental area (VTA), the substantia nigra, zona compacta (SNC) and zona reticulata (SNR) of adult female albino rats anaesthetized with urethane and chloral hydrate. Neurons were divided into two types based on their electrophysiological characteristics; Type I neurons had long duration action potentials (>2.6 msec) and slow discharge rates and Type II neurons had shorter duration action potentials and a wider range of discharge rates. Both types of neurons were found in the VTA and SNC, but there were only Type II neurons in the SNR. The effects of single pulse stimuli delivered to the ipsilateral ventromedial (VMH) or lateral (LH) hypothalamic areas on activities of the two types of neurons were investigated. Only a small portion of neurons in the VTA and SNC responded to VMH stimulation, but in contrast a majority of the two types of neurons in the VTA and SNC responded to LH stimulation. Most neurons in the SNR did not respond to VMH or to LH stimulation. Type II neurons in the VTA and SNC were predominantly suppressed by LH stimulation with short onset latencies (<6 msec), indicating the possibility of monosynaptic mediation. However Type I neurons in the VTA and SNC were activated and suppressed and the onset latencies of these responses were relatively longer. The high proportion of neurons of VTA and SNC responding to electrical stimulation of LH is consistent with anatomical evidence. Suppression and activation of Type I neurons in VTA and SNC suggest that the LH exerts modulatory influences on these neurons of the midbrain.     

Inhibitory action of the ventral noradrenergic bundle on the lateral hypothalamic neurons through alpha-noradrenergic mechanisms in the rat

Electrical stimulation of the ventral noradrenergic bundle (V-NA bundle) produced 3 types of responses in lateral hypothalamic neurons: IPSPs, a polysynaptic EPSP-IPSP sequence and antidromic spikes. The IPSPs were considered to be monosynaptic due to the fixed latencies seen at stimulus intensities. Iontophoretic application of an alpha-NA antagonist blocked only the presumed monosynaptic inhibition. Most of the glucose-sensitive neurons were inhibited by stimulation of the V-NA bundle. These results may account for the hyperphagia and obesity produced by selective lesions of the V-NA bundle.     

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.