Nucleus Taeniae Research Articles

The effect of acute social isolation on neural molecular responses in components of the social decision-making network

Prolonged or chronic social isolation has pronounced effects on animals, ranging from altered stress responses, increased anxiety and aggressive behaviour, and even increased mortality. The effects of shorter periods of isolation are much less well researched; however, short periods of isolation are used routinely for testing animal behaviour and physiology. Here, we studied how a 3 h period of isolation from a cagemate affected neural gene expression in three brain regions that contain important components of the social decision-making network, the hypothalamus, the nucleus taeniae of the amygdala, and the bed nucleus of the stria terminalis, using a gregarious bird as a model (zebra finches). We found evidence suggestive of altered neural activity, synaptic transmission, metabolism, and even potentially pain perception, all of which could create cofounding effects on experimental tests that involve isolating animals. We recommend that the effects of short-term social isolation need to be better understood and propose alternatives to isolating animals for testing.

Noncortical coding of biological motion in newborn chicks' brain.

Biological motion, the typical movement of vertebrates, is perceptually salient for many animal species. Newly hatched domestic chicks and human newborns show a spontaneous preference for simple biological motion stimuli (point-light displays) at birth prior to any visual learning. Despite evidence of such preference at birth, neural studies performed so far have focused on a specialized neural network involving primarily cortical areas. Here, we presented newly hatched visually naïve domestic chicks to either biological or rigid motion stimuli and measured for the first time their brain activation. Immediate Early Gene (c-Fos) expression revealed selective activation in the preoptic area of the hypothalamus and the nucleus taeniae of the amygdala. These results suggest that subpallial/subcortical regions play a crucial role in biological motion perception at hatching, paving the way for future studies on adult animals, including humans.

Varied impacts of social relationships on neuroendocrine state

Social relationships, affiliative social attachments, are important for many species. The best studied types of relationships are monogamous pair bonds. However, it remains unclear how generalizable models of pair bonding are across types of social attachments. Zebra finches are a fascinating system to explore the neurobiology of social relationships because they form various adult bonds with both same- and opposite-sex partners. To test whether different bonds are supported by a single brain network, we quantified individuals' neuroendocrine state after either 24 h or 2 weeks of co-housing with a novel same- or opposite-sex partner. We defined neuroendocrine state by the expression of 22 genes related to 4 major signaling pathways (dopamine, steroid, nonapeptide, and opioid) in six brain regions associated with affiliation or communication [nucleus accumbens (NAc), nucleus taeniae of the amygdala (TnA), medial preoptic area (POM), and periaqueductal gray (PAG), ventral tegmental area, and auditory cortex]. Overall, we found dissociable effects of social contexts (same- or opposite-sex partnerships) and duration of co-housing. Social bonding impacted the neuroendocrine state of four regions in males (NAc, TnA, POM, and PAG) and three regions in females (NAc, TnA, and POM). Monogamous pair bonding specifically appeared to impact male NAc. However, the patterns of gene expression in zebra finches were different than has previously been reported in mammals. Together, our results support the view that there are numerous mechanisms regulating social relationships and highlight the need to further our understanding of how social interactions shape social bonds.

The social environment alters neural responses to a lipopolysaccharide challenge

Sick animals display drastic changes in their behavioral patterns, including decreased activity, decreased food and water intake, and decreased interest in social interactions. These behaviors, collectively called “sickness behaviors”, can be socially modulated. For example, when provided with mating opportunities, males of several species show reduced sickness behaviors. While the behavior is known to change, how the social environment affects neural molecular responses to sickness is not known. Here, we used a species, the zebra finch, Taeniopygia guttata, where males have been shown to decrease sickness behaviors when presented with novel females. Using this paradigm, we obtained samples from three brain regions (the hypothalamus, the bed nucleus of the stria terminalis, and the nucleus taeniae) from lipopolysaccharide (LPS) or control treated males housed under four different social environments. Manipulation of the social environment rapidly changed the strength and co-expression patterns of the neural molecular responses to the immune challenge in all brain regions tested, therefore suggesting that the social environment plays a significant role in determining the neural responses to an infection. In particular, brains of males paired with a novel female showed muted immune responses to LPS, as well as altered synaptic signaling. Neural metabolic activity in response to the LPS challenge was also affected by the social environment. Our results provide new insights into the effects of the social environment on brain responses to an infection, thereby improving our understanding of how the social environment can affect health.

Active exploration of an environment drives the activation of the hippocampus-amygdala complex of domestic chicks.

In birds, like in mammals, the hippocampus critically mediates spatial navigation through the formation of a spatial map. This study investigates the impact of active exploration of an environment on the hippocampus of young domestic chicks. Chicks that were free to actively explore the environment exhibited a significantly higher neural activation (measured by c-Fos expression) compared with those that passively observed the same environment from a restricted area. The difference was limited to the anterior and the dorsolateral parts of the intermediate hippocampus. Furthermore, the nucleus taeniae of the amygdala showed a higher c-Fos expression in the active exploration group than in the passive observation group. In both brain regions, brain activation was correlated with the number of locations that chicks visited during the test. This suggests that the increase of c-Fos expression in the hippocampus is related to increased firing rates of spatially coding neurons. Furthermore, our study indicates a functional linkage of the hippocampus and nucleus taeniae of the amygdala in processing spatial information. Overall, with the present study, we confirm that in birds, like in mammals, hippocampus and amygdala functions are linked and likely related to spatial representations.

Activation of the Nucleus Taeniae of the Amygdala by Umami Taste in Domestic Chicks (Gallus gallus).

In chickens, the sense of taste plays an important role in detecting nutrients and choosing feed. The molecular mechanisms underlying the taste-sensing system of chickens are well studied, but the neural mechanisms underlying taste reactivity have received less attention. Here we report the short-term taste behaviour of chickens towards umami and bitter (quinine) taste solutions and the associated neural activity in the nucleus taeniae of the amygdala, nucleus accumbens and lateral septum. We found that chickens had more contact with and drank greater volumes of umami than bitter or a water control, and that chicks displayed increased head shaking in response to bitter compared to the other tastes. We found that there was a higher neural activity, measured as c-Fos activation, in response to umami taste in the right hemisphere of the nucleus taeniae of the amygdala. In the left hemisphere, there was a higher c-Fos activation of the nucleus taeniae of the amygdala in response to bitter than in the right hemisphere. Our findings provide clear evidence that chickens respond differently to umami and bitter tastes, that there is a lateralised response to tastes at the neural level, and reveals a new function of the avian nucleus taeniae of the amygdala as a region processing reward information.

Role of aromatase in distinct brain nuclei of the social behaviour network in the expression of sexual behaviour in male Japanese quail.

In male Japanese quail, brain aromatase is crucial for the hormonal activation of sexual behaviour, but the sites producing neuro-oestrogens that are critical for these behaviours have not been completely identified. This study examined the function of aromatase expressed in several nuclei of the social behaviour network on a measure of sexual motivation known as the frequency of rhythmic cloacal sphincter movements (RCSM) and on copulatory behaviour. Sexually experienced castrated males chronically treated with testosterone were stereotaxically implanted with the aromatase inhibitor vorozole (VOR), or cholesterol as control, and tested for sexual behaviour. In experiment 1, males were implanted in the medial preoptic nucleus (POM) with VOR, a manipulation known to reduce the expression of copulatory behaviour. This experiment served as positive control, but also showed that VOR implanted in the dorsomedial or lateral portions of the POM similarly inhibits male copulatory behaviour compared to control implants. In experiments 2 to 4, males received stereotaxic implants of VOR in the periaqueductal gray (PAG), the nucleus taeniae of the amygdala (TnA) and the ventromedial nucleus of the hypothalamus (VMN), respectively. Sexual behaviour was affected only in individuals where VOR was implanted in the PAG: these males displayed significantly lower frequencies of cloacal contact movements, the last step of the copulatory sequence. Inhibition of aromatase in the TnA and VMN did not alter copulatory ability. Overall, RCSM frequency remained unaffected by VOR regardless of implantation site. Together, these results suggest that neuro-oestrogens produced in the POM contribute the most to the control of male copulatory behaviour, while aromatase expressed in the PAG might also participate to premotor aspects of male copulatory behaviour.

The role of posterior pallial amygdala in mediating motor behaviors in pigeons

The posterior pallial amygdala (PoA) is located on the basolateral caudal telencephalon, including the basal division of PoA (PoAb) and the compact division of PoA (PoAc). PoA plays a vital role in emotion regulation and is considered a part of the amygdala in birds. However, the regulatory functions responsible for motor behaviors and emotions between PoAb and PoAc are poorly understood. Therefore, we studied the structure and function of PoA by tract-tracing methods, constant current electrical stimulation, and different dopamine receptor drug injections in pigeons (Columba livia domestica). PoAb connects reciprocally with two nuclear groups in the cerebrum: 1) a continuum comprising the temporo–parieto–occipitalis, corticoidea dorsolateralis, hippocampus, and parahippocampalis areas and 2) rostral areas of the hemisphere, including the nucleus septalis lateralis and nucleus taeniae amygdalae. Extratelencephalic projections of PoAb terminate in the lateral hypothalamic nucleus and are scattered in many limbic midbrain regions. PoAb and PoAc mainly mediated the turning movement. In the ‘open-field’ test, D1 agonist and D2 antagonist could significantly reduce the latency period for entering into the central area and increase the residence time in the central area, whereas D1 antagonist and D2 agonist had the opposite effect. PoAb and PoAc are important brain areas that mediate turning behavior.

American Crow Brain Activity in Response to Conspecific Vocalizations Changes When Food Is Present.

Social interaction among animals can occur under many contexts, such as during foraging. Our knowledge of the regions within an avian brain associated with social interaction is limited to the regions activated by a single context or sensory modality. We used 18F-fluorodeoxyglucose positron emission tomography (FDG-PET) to examine American crow (Corvus brachyrhynchos) brain activity in response to conditions associated with communal feeding. Using a paired approach, we exposed crows to either a visual stimulus (the sight of food), an audio stimulus (the sound of conspecifics vocalizing while foraging) or both audio/visual stimuli presented simultaneously and compared to their brain activity in response to a control stimulus (an empty stage). We found two regions, the nucleus taenia of the amygdala (TnA) and a medial portion of the caudal nidopallium, that showed increased activity in response to the multimodal combination of stimuli but not in response to either stimulus when presented unimodally. We also found significantly increased activity in the lateral septum and medially within the nidopallium in response to both the audio-only and the combined audio/visual stimuli. We did not find any differences in activation in response to the visual stimulus by itself. We discuss how these regions may be involved in the processing of multimodal stimuli in the context of social interaction.

Neurotranscriptomic changes associated with chick-directed parental care in adult non-reproductive Japanese quail

For many species, parental care critically affects offspring survival. But what drives animals to display parental behaviours towards young? In mammals, pregnancy-induced physiological transformations seem key in preparing the neural circuits that lead towards attraction (and reduced-aggression) to young. Beyond mammalian maternal behaviour, knowledge of the neural mechanisms that underlie young-directed parental care is severely lacking. We took advantage of a domesticated bird species, the Japanese quail, for which parental behaviour towards chicks can be induced in virgin non-reproductive adults through a sensitization procedure, a process that is not effective in all animals. We used the variation in parental responses to study neural transcriptomic changes associated with the sensitization procedure itself and with the outcome of the procedure (i.e., presence of parental behaviours). We found differences in gene expression in the hypothalamus and bed nucleus of the stria terminalis, but not the nucleus taeniae. Two genes identified are of particular interest. One is neurotensin, previously only demonstrated to be causally associated with maternal care in mammals. The other one is urocortin 3, causally demonstrated to affect young-directed neglect and aggression in mammals. Because our studies were conducted in animals that were reproductively quiescent, our results reflect core neural changes that may be associated with avian young-directed care independently of extensive hormonal stimulation. Our work opens new avenues of research into understanding the neural basis of parental care in non-placental species.

Current Stimulation of the Midbrain Nucleus in Pigeons for Avian Flight Control.

A number of research attempts to understand and modulate sensory and motor skills that are beyond the capability of humans have been underway. They have mainly been expounded in rodent models, where numerous reports of controlling movement to reach target locations by brain stimulation have been achieved. However, in the case of birds, although basic research on movement control has been conducted, the brain nuclei that are triggering these movements have yet to be established. In order to fully control flight navigation in birds, the basic central nervous system involved in flight behavior should be understood comprehensively, and functional maps of the birds’ brains to study the possibility of flight control need to be clarified. Here, we established a stable stereotactic surgery to implant multi-wire electrode arrays and electrically stimulated several nuclei of the pigeon’s brain. A multi-channel electrode array and a wireless stimulation system were implanted in thirteen pigeons. The pigeons’ flight trajectories on electrical stimulation of the cerebral nuclei were monitored and analyzed by a 3D motion tracking program to evaluate the behavioral change, and the exact stimulation site in the brain was confirmed by the postmortem histological examination. Among them, five pigeons were able to induce right and left body turns by stimulating the nuclei of the tractus occipito-mesencephalicus (OM), nucleus taeniae (TN), or nucleus rotundus (RT); the nuclei of tractus septo-mesencephalicus (TSM) or archistriatum ventrale (AV) were stimulated to induce flight aviation for flapping and take-off with five pigeons.

Prenatal testosterone triggers long-term behavioral changes in male zebra finches: unravelling the neurogenomic mechanisms

BackgroundMaternal hormones, like testosterone, can strongly influence developing offspring, even generating long-term organizational effects on adult behavior; yet, the mechanisms facilitating these effects are still unclear. Here, we experimentally elevated prenatal testosterone in the eggs of zebra finches (Taeniopygia guttata) and measured male aggression in adulthood along with patterns of neural gene expression (RNA-seq) and DNA methylation (MethylC-Seq) in two socially relevant brain regions (hypothalamus and nucleus taenia of the amygdala). We used enrichment analyses and protein-protein interaction networks to find candidate processes and hub genes potentially affected by the treatment. We additionally identified differentially expressed genes that contained differentially methylated regions.ResultsWe found that males from testosterone-injected eggs displayed more aggressive behaviors compared to males from control eggs. Hundreds of genes were differentially expressed, particularly in the hypothalamus, including potential aggression-related hub genes (e.g., brain derived neurotrophic factor). There were also enriched processes with well-established links to aggressive phenotypes (e.g., somatostatin and glutamate signaling). Furthermore, several highly connected genes identified in protein-protein interaction networks also showed differential methylation, including adenylate cyclase 2 and proprotein convertase 2.ConclusionsThese results highlight genes and processes that may play an important role in mediating the effects of prenatal testosterone on long-term phenotypic outcomes, thereby providing insights into the molecular mechanisms that facilitate hormone-mediated maternal effects.

Neural basis of unfamiliar conspecific recognition in domestic chicks (Gallus Gallus domesticus)

Domestic chickens are able to distinguish familiar from unfamiliar conspecifics, however the neuronal mechanisms mediating this behaviour are almost unknown. Moreover, the lateralisation of chicks’ social recognition has only been investigated at the behavioural level, but not at the neural level. The aim of the present study was to test the hypothesis that exposure to unfamiliar conspecifics will selectively activate septum, hippocampus or nucleus taeniae of the amygdala of young domestic chicks. Moreover we also wanted to test the lateralisation of this response. For this purpose, we used the immediate early gene product c-Fos to map neural activity. Chicks were housed in pairs for one week. At test, either one of the two chicks was exchanged by an unfamiliar individual (experimental ‘unfamiliar’ group) or the familiar individual was briefly removed and then placed back in its original cage (control ‘familiar’ group). Analyses of chicks’ interactions with the familiar/unfamiliar social companion revealed a higher number of social pecks directed towards unfamiliar individuals, compared to familiar controls. Moreover, in the group exposed to the unfamiliar individual a significantly higher activation was present in the dorsal and ventral septum of the left hemisphere and in the ventral hippocampus of the right hemisphere, compared to the control condition. These effects were neither present in other subareas of hippocampus or septum, nor in the nucleus taeniae of the amygdala. Our study thus indicates selective lateralised involvement of domestic chicks’ septal and hippocampal subregions in responses to unfamiliar conspecific.

Selective response of the nucleus taeniae of the amygdala to a naturalistic social stimulus in visually naive domestic chicks

The detection of animate beings at the onset of life is important for phylogenetically distant species, such as birds and primates. Naïve chicks preferentially approach a stimulus resembling a conspecific (a stuffed fowl) over a less naturalistic one (a scrambled version of the stuffed fowl, presenting the same low-level visual features as the fowl in an unnatural configuration). The neuronal mechanisms underlying this behavior are mostly unknown. However, it has been hypothesized that innate social predispositions may involve subpallial brain areas including the amygdala. Here we asked whether a stuffed hen would activate areas of the arcopallium/amygdala complex, in particular the nucleus taeniae of the amygdala (TnA) or septum. We measured brain activity by visualizing the immediate early gene product c-Fos. After exposure to the hen, TnA showed higher density of c-Fos expressing neurons, compared to chicks that were exposed to the scrambled stimulus. A similar trend was present in the lower portion of the arcopallium, but not in the upper portion of the arcopallium or in the septum. This demonstrates that at birth the TnA is already engaged in responses to social visual stimuli, suggesting an important role for this nucleus in the early ontogenetic development of social behavior.

Social odour activates the hippocampal formation in zebra finches (Taeniopygia guttata).

Experiments from our research group have demonstrated that the olfactory sense of birds, which has been considered as unimportant for a long time, plays a prominent role as communication channel in social behaviour. Odour cues are used e.g. by zebra finch chicks to recognize the mother, by adult birds to distinguish their own eggs from others, or to recognize kin. While there is quite a lot of evidence for the importance of odour for social behaviour, it is not known as yet which brain areas may be involved in the processing of socially relevant odours. We therefore compared the brain activation pattern of zebra finch males exposed to their own offspring odour with that induced by a neutral odour stimulus. By measuring head saccade changes as behavioural reaction and using the expression of the immediate early gene product c-Fos as brain activity marker, we show here that the activation pattern, namely the activity difference between the left and the right hemisphere, of several hippocampal areas in zebra finch males is altered by the presentation of the odour of their own nestlings. In contrast, the nucleus taeniae of the amygdala (TnA) exhibits a tendency of a reduction of c-Fos activation in both hemispheres as a consequence of exposure to the nestling odour. We conclude that the hippocampus is involved in odour based processing of social information, while the role of TnA remains unclear.

Simulated viral infection in early-life alters brain morphology, activity and behavior in zebra finches (Taeniopygia guttata)

Early-life immune challenges (ELIC) have long-term effects on adult behavior and brain development. ELIC studies on birds are still few, but they are epidemiologically crucial since birds are important hosts of many mosquito-borne viruses. In this study, we administered a viral infection mimicking agent, Polyinosinic: polycytidylic acid (Poly I:C), to nestling zebra finches on post-hatch day 14. When birds became sexually mature, their general activity (i.e., hopping, feeding behavior) and mosquito defense behaviors (i.e., hops, head movements, pecks, wing movements, foot movements, and scratches) were measured. Following behavioral trials, brains of male birds were collected for anatomical and histochemical analyses. Poly I:C challenge had sex-dependent effects on general activity and mosquito defense behaviors. When compared to control females, Poly I:C challenged females hopped and fed less often in their general activities, but hopped more often in the presence of mosquitoes. Poly I:C challenged males did not differ from control males in any behaviors. Brain analysis revealed that the nucleus taeniae of the amygdala (TnA) of Poly I:C challenged males were smaller in volume yet had more neurons expressing immediate-early gene proteins compared with controls, suggesting a more active TnA. These results suggest that immune challenges early in the life could have long-term effects on behaviors and brains of zebra finches, which may influence disease spread and fitness of individual birds.

Rapid effects of estradiol on aggression depend on genotype in a species with an estrogen receptor polymorphism

The white-throated sparrow (Zonotrichia albicollis) represents a powerful model in behavioral neuroendocrinology because it occurs in two plumage morphs that differ with respect to steroid-dependent social behaviors. Birds of the white-striped (WS) morph engage in more territorial aggression than do birds of the tan-striped (TS) morph, and the TS birds engage in more parenting behavior. This behavioral polymorphism is caused by a chromosomal inversion that has captured many genes, including estrogen receptor alpha (ERα). In this study, we tested the hypothesis that morph differences in aggression might be explained by differential sensitivity to estradiol (E2). We administered E2 non-invasively to non-breeding white-throated sparrows and quantified aggression toward a conspecific 10 min later. E2 administration rapidly increased aggression in WS birds but not TS birds, consistent with our hypothesis that differential sensitivity to E2 may at least partially explain morph differences in aggression. To query the site of E2 action in the brain, we administered E2 and quantified Egr-1 expression in brain regions in which expression of ERα is known to differ between the morphs. E2 treatment decreased Egr-1 immunoreactivity in nucleus taeniae of the amygdala, but this effect did not depend on morph. Overall, our results support a role for differential effects of E2 on aggression in the two morphs, but more research will be needed to determine the neuroanatomical site of action.

Neural activation following offensive aggression in Japanese quail.

ABSTRACTAggression is a fundamental part of animal social behaviour. In avian species, little is known about its neural representation. In particular, neural activity following offensive aggression has not been studied in detail. Here, we investigated the patterns of brain activation using immediate-early gene (IEG) expression in male Japanese quail that showed pronounced aggressive behaviours during a 30 min male–male interaction and compared them to those of males that did not interact with a conspecific. In aggressive males, we found a massive induction of the IEG ZENK in pallial brain structures such as the intermediate medial mesopallium, the caudomedial mesopallium and the intermediate medial nidopallium. To a lesser extent, activation was observed in subpallial areas such as the nucleus taeniae of the amygdala and in the medial portion of the bed nucleus of the stria terminalis. Our data suggest that the modulation of aggressive behaviour involves the integration of multisensory information.

ZENK induction in the zebra finch brain by song: Relationship to hemisphere, rhythm, oestradiol and sex.

Oestradiol is abundant in the zebra finch auditory forebrain and has the capacity to modulate neural responses to auditory stimuli with specificity as a result of both hemisphere and sex. Arrhythmic song induces greater ZENK expression than rhythmic song in the caudomedial nidopallium (NCM), caudomedial mesopallium (CMM) and nucleus taeniae (Tn) of adult zebra finches. The increases in the auditory regions (i.e. NCM and CMM) may result from detection of errors in the arrhythmic song relative to the learned template. In the present study, zebra finches were treated with oestradiol, the aromatase inhibitor fadrozole or a control and then exposed to rhythmic or arrhythmic song to assess the effect of oestradiol availability on neural responses to auditory rhythms. ZENK mRNA was significantly greater in the left hemisphere within the NCM, CMM and Tn. Main effects of sex were detected in both auditory regions, with increased ZENK in males in the NCM and in females in the CMM. In the CMM, an effect of hormone treatment also existed. Although no pairwise comparison was statistically significant, the pattern suggested greater ZENK expression in control compared to both fadrozole- and oestradiol-treated birds. In the NCM, an interaction between sex and hormone treatment suggested that the sex effect was restricted to control animals. An additional interaction in the NCM among sex, stimulus rhythmicity and hemisphere indicated that the strongest effect of laterality was present in males exposed to arrhythmic song. The hormone effects suggest that an optimal level of oestradiol may exist for processing rhythmicity of auditory stimuli. The overall pattern for left lateralisation parallels the left lateralisation of language processing in humans and may suggest that this hemisphere is specialised for processing conspecific vocalisations. The reversed sex differences in the NCM and CMM suggest that males and females differentially rely on components of the auditory forebrain for processing conspecific song.

Transmitter receptors reveal segregation of the arcopallium/amygdala complex in pigeons (Columba livia).

At the beginning of the 20th century it was suggested that a complex group of nuclei in the avian posterior ventral telencephalon is comparable to the mammalian amygdala. Subsequent findings, however, revealed that most of these structures share premotor characteristics, while some indeed constitute the avian amygdala. These developments resulted in 2004 in a change of nomenclature of these nuclei, which from then on were named arcopallial or amygdala nuclei and referred to as the arcopallium/amygdala complex. The structural basis for the similarities between avian and mammalian arcopallial and amygdala subregions is poorly understood. Therefore, we analyzed binding site densities for glutamatergic AMPA, NMDA and kainate, GABAergic GABAA , muscarinic M1 , M2 and nicotinic acetylcholine (nACh; α4 β2 subtype), noradrenergic α1 and α2 , serotonergic 5-HT1A and dopaminergic D1/5 receptors using quantitative in vitro receptor autoradiography combined with a detailed analysis of the cyto- and myelo-architecture. Our approach supports a segregation of the pigeon's arcopallium/amygdala complex into the following subregions: the arcopallium anterius (AA), the arcopallium ventrale (AV), the arcopallium dorsale (AD), the arcopallium intermedium (AI), the arcopallium mediale (AM), the arcopallium posterius (AP), the nucleus posterioris amygdalopallii pars basalis (PoAb) and pars compacta (PoAc), the nucleus taeniae amgygdalae (TnA) and the area subpallialis amygdalae (SpA). Some of these subregions showed further subnuclei and each region of the arcopallium/amygdala complex are characterized by a distinct multi-receptor density expression. Here we provide a new detailed map of the pigeon's arcopallium/amygdala complex and compare the receptor architecture of the subregions to their possible mammalian counterparts.