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The standard nomenclature that has been used for many telencephalic and related brainstem structures in birds is based on flawed assumptions of homology to mammals. In particular, the outdated terminology implies that most of the avian telencephalon is a hypertrophied basal ganglia, when it is now clear that most of the avian telencephalon is neuro...
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... example, the striatum is rich in dopaminergic fibers and acetylcholinesterase activity, whereas the pallidum is poor in both (Fig. 2B,E). The various subpallial structures, the new terms, and the evidence for homology accepted by the Nomenclature Forum are detailed below on a structure-by-structure basis and summarized in Table 2. Note that the Forum concluded that sufficient evidence existed to recognize a ventrocaudal part of the subpallium as part of the amygdala, as addressed in the section on the archistriatum. ...
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... Smith-Fernandez et al., 1998;Puelles et al., 2000). For these reasons, and additional ones summarized in Reiner et al. (1998a), the Forum replaced the arcane name lobus parolfactorius (meaning lobe next to the olfactory bulb) with the term medial striatum and recognized it as (to- gether with the paleostriatum augmentatum) forming the avian dorsal striatum (Table 2 ...
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... these reasons, as well as reasons summarized in Reiner et al. (1998a), the Forum concluded that the name paleostriatum augmentatum (with its inaccurate evolu- tionary and cellular implications of being a pallidal deriv- ative) should be abandoned and replaced with the term lateral striatum (Table 2, Figs. 5A-F, 6A-C). ...
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... primitivum (PP) 3 Globus pallidus (GP). The Forum recommended that the paleostriatum primitivum henceforth be called the globus pallidus (Table 2, Figs. 5A-F, 6B,C). ...
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... the data are thus beginning to suggest a largely striatal nature for the INP, the Forum concluded that it was still premature to conclude unequivocally that the INP is a striatal territory. Thus, the Forum decided to leave the name for the INP unaltered (Table 2, Figs. 5D, 6B). ...
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... these reasons, the Forum recognized and recom- mended that the rostral ventromedial part of the former LPO of birds be called the nucleus accumbens and that the term medial striatum only be used to refer to the remain- der of the LPO (Table 2, Fig. 6A,D). As in mammals, however, a precise cytoarchitectonic border between the dorsal striatum and nucleus accumbens is not evident, and a neurochemical criterion by which to distinguish the two fields unambiguously has not been identified. ...
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... bulb input and resembles the olfactory tubercle of mammals in its neurochemistry and connectivity (Heimer et al., 1985(Heimer et al., , 1997Reiner and Karten, 1985; Roberts et al., 2002). The Forum thus endorsed the previously recognized homology of this cell group to the similarly named mammalian cell group and recommended no name change (Table 2, Figs. 6A,D, 8B). The Forum recommends a slight modification of the ab- breviation for the olfactory tubercle (i.e., TuO) so it is not in conflict with the common abbreviation for the optic tract (i.e., ...
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... comparable cell group is present in turtles, crocodilians, and lizards Brauth, 1984;Reiner, 1987;Reiner and Carraway, 1987;Russchen et al., 1987;Russchen and Jonker, 1988). The Forum thus recognized this cell group, which has some- times been called the ventral paleostriatum ( Kuenzel and Masson, 1988), and recommended it be referred to as the ventral pallidum (Table 2 , Figs. 5C,D, 6B). The word ven- tral is used because this provides the VP with a name that is positionally appropriate with respect to its more dorsal counterpart, the GP, which has also been termed the dor- sal pallidum. ...
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... extension occupies a small but conspicuous bulge into the lateral edge of the inferior aspect of the telencephalic ventricle, and it may be surrounded by the true nucleus accumbens at very rostral levels (Reiner et al., , 1984bAste et al., 1998a,b). The Forum thus recommended that the term nucleus accumbens be discon- tinued as the name for the region identified as accumbens in Karten and Hodos (1967) and that the lateral part of the bed nucleus of the stria terminalis be employed in- stead (Table 2, Figs. 5C,D,F, 6B,C). ...
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... region in birds lies within the path of the apparent avian counterpart of the stria terminalis ( Zeier and Karten, 1971). The Forum thus recommended that the term medial part of the bed nucleus of the stria terminalis be employed for this region in birds (Table 2, Fig. 6C). By in situ hybridization histochemistry and im- munocytochemistry for arginine vasotocin, a single BSTM nucleus has been identified in Japanese quail ( Aste et al., 1998a,b), whereas two BSTM subnuclei have been identi- Fig. 5. Images of transverse sections of pigeon brain immunola- beled for various markers to show the location of several subpallial cell groups affected by the nomenclature revision. ...
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... com- parable cell group is present in members of all reptilian orders ( Brauth and Kitt, 1980;Brauth, 1984;Reiner and Carraway, 1987;Medina et al., 1993;Smeets, 1994). The Forum thus recommended that the name for the medial septal nucleus remain unchanged (Table 2, Fig. 6C). ...
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... com- parable cell group is present in all reptilian orders (Brauth, 1984;Reiner, 1987;Smeets, 1994). The Forum thus recommended that the name for the lateral septal nucleus remain unchanged (Table 2, Figs. 5C,D, 6B,C). ...
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... basalis (Bas) 3 Nucleus basorostralis pal- lii (Bas). Although the term nucleus basalis as it has been used in birds to refer to a sensory structure of the pallium does not possess any root words implying an as- sociation with the basal ganglia, the name used for this sensory cell group needed to be changed because the Fo- rum had already reserved that same name for the avian homologue of the basal forebrain cholinergic cell field in mammals ( Table 2). The structure that has been called nucleus basalis in birds is not located in the subpallium and is not a cholinergic cell group, but rather is a trigemi- norecipient pallial sensory cell group ( Witkovsky et al., 1973;Wild et al., 1985) and, in some species, also a general somatosensory recipient nucleus ( Wild et al., 1997Wild et al., , 2001). ...
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... Besides NC, we found that MSNs and larger neurons (likely pallidal neurons and interneurons) were activated in different parts of the basal ganglia. Levels of activation were significantly higher in LSt and GP [important for motor functions in birds (Veenman et al., 1995;Reiner et al., 2004)] of Trained, No-Association, and Undertrained birds which attempted to obtain the reward vs. Baseline controls. Furthermore, Area X and MSt, which are components of the anterior forebrain pathway in zebra finches (Brainard and Doupe, 2002) were activated in crows after training on the visual discrimination task. ...
The caudolateral nidopallium (NCL, an analog of the prefrontal cortex) is known to be involved in learning, memory, and discrimination in corvids (a songbird), whereas the involvement of other brain regions in these phenomena is not well explored. We used house crows (Corvus splendens) to explore the neural correlates of learning and decision-making by initially training them on a shape discrimination task followed by immunohistochemistry to study the immediate early gene expression (Arc), a dopaminoceptive neuronal marker (DARPP-32, Dopamine- and cAMP-regulated phosphoprotein, Mr 32 kDa) to understand the involvement of the reward pathway and an immature neuronal marker (DCX, doublecortin) to detect learning-induced changes in adult neurogenesis. We performed neuronal counts and neuronal tracing, followed by morphometric analyses. Our present results have demonstrated that besides NCL, other parts of the caudal nidopallium (NC), avian basal ganglia, and intriguingly, vocal control regions in house crows are involved in visual discrimination. We have also found that training on the visual discrimination task can be correlated with neurite pruning in mature dopaminoceptive neurons and immature DCX-positive neurons in the NC of house crows. Furthermore, there is an increase in the incorporation of new neurons throughout NC and the medial striatum which can also be linked to learning. For the first time, our results demonstrate that a combination of structural changes in mature and immature neurons and adult neurogenesis are linked to learning in corvids.
... One study found a significant decrease in 5-HIAA but no differences in the 5-HT levels in the amygdala of birds from an HFP line [16]. The amygdala, which contains the arcopallium, leads to a pathway to the brainstem, is responsible for motor function, and may be involved in anxiety and fear in avian species [59,60]. Serotonin is also important for the regulation of this area in the brain [31]. ...
A complex system of neural pathways, collectively known as the microbiota–gut–brain (MGB) axis, interconnects the gut microbiota, the gastrointestinal system, and the brain along with its periphery. Previous studies have demonstrated that modulation of the MGB axis can influence stress-related behaviors such as anxiety. This connection becomes apparent in scenarios like agonistic behavior in laying hens, which is characterized by aggressive head and feather pecks, that can ultimately result in cannibalism and death. The objective was to examine the effects of a dietary synbiotic on agonistic behavior, plasma and brain monoamines, stress parameters, and cecal microbiota counts via modulation of the MGB axis. A total of 396 W36 Hy-Line laying hens were provided at random with a control (CON: basal diet) or treatment (SYN: basal diet supplemented with synbiotic) diet from 50 to 60 weeks old (nine pens/treatment, 22 birds/pen). Blood samples and video recordings (three consecutive days/week) were taken at 50 and 60 weeks. At 60 weeks, three hens/pen were euthanized for brain and cecal microbiota collection. Threatening, fighting, head, body, and feather pecking all occurred less frequently at 60 weeks in the SYN group (p < 0.05). Plasma corticosterone, adrenocorticotropic hormone, dopamine, and serotonin were significantly lower while tryptophan and 5-hydroxyindoleacetic acid were significantly higher in birds from the SYN group (p < 0.05). Significant differences in serotonin, 5-hydroxyindoleacetic acid, dopamine, homovanillic acid, and 3,4-dihydroxyphenylacetic acid were observed in the hypothalamus, hippocampus, and amygdala of the brain. Serotonin and dopamine turnover rates were significantly different in all three regions of the brain (p < 0.05). Cecal counts of Lactobacillus and Bifidobacterium were significantly higher in the SYN group (p < 0.05). Synbiotic supplementation resulted in many significant differences, indicating activation of the serotonergic systems and modulation of both the MGB axis and HPA axis with positive effects on welfare and stress.
... The Wulst is a laminated forebrain structure organized into two different functional regions, a smaller, anterior somatosensory region and a larger, posterior region processing visual information [23]. The visual Wulst is composed of four laminae: running from dorsal to ventral the hyperpallium apicale (HA), interstitial nucleus of the hyperpallium apicale (IHA), hyperpallium intercalatum (HI), and densocellular part of the hyperpallium (HD) [24,25]. It is worth noting that HA and HD have connections with the HF [23,26,27]. ...
... The hyperpallium (H) or Wulst is composed of four layers in pigeons (Atoji and Wild, 2019). However, only the most ventral layer, the densocellular part of the hyperpallium (HD), and the dorsal part of the hyperpallium (HA) are readily identified in the crow and other songbirds (Reiner et al., 2004a;Jarvis et al., 2013;Lovell et al., 2020;Kersten et al., 2022). The HD has been renamed to MD and the HA is referred to as H in songbirds, based on cell clustering patterns (Jarvis et al., 2013;Kersten et al., 2022). ...
The avian telencephalic structure nidopallium caudolaterale (NCL) functions as an analog to the mammalian prefrontal cortex. In crows, corvid songbirds, it plays a crucial role in higher cognitive and executive functions. These functions rely on the NCL's extensive telencephalic connections. However, systematic investigations into the brain-wide connectivity of the NCL in crows or other songbirds are lacking. Here, we studied its input and output connections by injecting retrograde and anterograde tracers into the carrion crow NCL. Our results, mapped onto a published carrion crow brain atlas, confirm NCL multisensory connections and extend prior pigeon findings by identifying a novel input from the hippocampal formation. Furthermore, we analyze crow NCL efferent projections to the arcopallium and report newly identified arcopallial neurons projecting bilaterally to the NCL. These findings help to clarify the role of the NCL as central executive hub in the corvid songbird brain.
... For thalamofugal structures we primarily used the terminology from the atlas of Kuenzel and Masson 39 . For many of the forebrain regions, we adopted the nomenclature of Reiner et al. (2004) 75 . Furthermore, we used recent literature 76 for nomenclature of several thalamofugal structures. ...
... For thalamofugal structures we primarily used the terminology from the atlas of Kuenzel and Masson 39 . For many of the forebrain regions, we adopted the nomenclature of Reiner et al. (2004) 75 . Furthermore, we used recent literature 76 for nomenclature of several thalamofugal structures. ...
Amniotes feature two principal visual processing systems: the tectofugal and thalamofugal pathways. In most mammals, the thalamofugal pathway predominates, routing retinal afferents through the dorsolateral geniculate complex to the visual cortex. In most birds, the thalamofugal pathway often plays the lesser role with retinal afferents projecting to the principal optic thalami, a complex of several nuclei that resides in the dorsal thalamus. This thalamic complex sends projections to a forebrain structure called the Wulst, the terminus of the thalamofugal visual system. The thalamofugal pathway in birds serves many functions such as pattern discrimination, spatial memory, and navigation/migration. A comprehensive analysis of avian species has unveiled diverse subdivisions within the thalamic and forebrain structures, contingent on species, age, and techniques utilized. In this study, we documented the thalamofugal system in three dimensions by integrating histological and contrast-enhanced computed tomography imaging of the avian brain. Sections of two-week-old chick brains were cut in either coronal, sagittal, or horizontal planes and stained with Nissl and either Gallyas silver or Luxol Fast Blue. The thalamic principal optic complex and pallial Wulst were subdivided on the basis of cell and fiber density. Additionally, we utilized the technique of diffusible iodine-based contrast-enhanced computed tomography (diceCT) on a 5-week-old chick brain, and right eyeball. By merging diceCT data, stained histological sections, and information from the existing literature, a comprehensive three-dimensional model of the avian thalamofugal pathway was constructed. The use of a 3D model provides a clearer understanding of the structural and spatial organization of the thalamofugal system. The ability to integrate histochemical sections with diceCT 3D modeling is critical to better understanding the anatomical and physiologic organization of complex pathways such as the thalamofugal visual system.
... No pixilation adjustments or manipulations of the captured images were undertaken, except for the adjustment of contrast, brightness, and levels using Adobe Photoshop. The neuroanatomical demarcations and nomenclature used in the current study were derived fromReiner et al. (2004), Izawa and Watanabe(2007), andBrauth et al. (2011). ...
The orexinergic system, while having several roles, appears to be a key link in the balance between arousal and satiety. In birds, to date, this system has only been examined anatomically in four species, all with brains smaller than 3.5 g and of a limited phylogenetic range. Here, using orexin-A immunohistochemistry, we describe the distribution, morphology and nuclear parcellation of orexinergic neurons within the hypothalami of a Congo grey and a Timneh grey parrot, a pied crow, an emu, and a common ostrich. These birds represent a broad phylogeny, with brains ranging in size from 7.85–26.5 g. Within the hypothalami of the species studied, the orexinergic neurons were organized in two clusters, a densely packed paraventricular hypothalamic nucleus cluster located within the medial hypothalamus, but not contacting the ventricle, and a more loosely packed lateral hypothalamic cluster in the lateral hypothalamus. Stereological analysis revealed a strong correlation, using phylogenetic generalized least squares regression analyses, between brain mass and the total number of orexinergic neurons, as well as soma parameters such as volume and area. Orexinergic axonal terminals evinced two types of boutons, larger and the smaller en passant boutons. Unlike the orexinergic system in mammals, that has several variances in cluster organization, that of the birds studied, in the present and previous studies, shows organization invariance, despite the differences in brain and body mass, phylogeny and life-history of the species studied. This indicates that the orexinergic system has a specific and non-variant function in birds as compared to mammals.
... We showed abundant astrocytes in like white matter regions, such as the pallidum and mesencephalon, corresponding to the FPL and FRL, respectively. The lateral prosencephalic fascicle (FPL), also known as the medial telencephalic fascicle, is a bundle of nerve fibers that connects the hypothalamus with the limbic system, which is involved in the reward system and basal ganglia (Reiner et al., 2004a;Hernandez et al., 2006;Bradbury and Vehrencamp, 2011;Coenen et al., 2018) It is believed that the FPL plays similar roles in birds, coordinating song production in response to territorial defense and sexual selection, seeking a final reward such as deterring intruders or reproducing. In juvenile zebra finches, Taeniopygia guttata, the accuracy of song imitation correlates with brain areas distinct from those in the song motor control system. ...
The house wren shows complex song, and the rufous-tailed hummingbird has a simple song. The location of vocal brain areas supports the song’s complexity; however, these still need to be studied. The astrocytic population in songbirds appears to be associated with change in vocal control nuclei; however, astrocytic distribution and morphology have not been described in these species. Consequently, we compared the distribution and volume of the vocal brain areas: HVC, RA, Area X, and LMAN, cell density, and the morphology of astrocytes in the house wren and the rufous-tailed hummingbird. Individuals of the two species were collected, and their brains were analyzed using serial Nissl- NeuN- and MAP2-stained tissue scanner imaging, followed by 3D reconstructions of the vocal areas; and GFAP and S100β astrocytes were analyzed in both species. We found that vocal areas were located close to the cerebral midline in the house wren and a more lateralized position in the rufous-tailed hummingbird. The LMAN occupied a larger volume in the rufous-tailed hummingbird, while the RA and HVC were larger in the house wren. While Area X showed higher cell density in the house wren than the rufous-tailed hummingbird, the LMAN showed a higher density in the rufous-tailed hummingbird. In the house wren, GFAP astrocytes in the same bregma where the vocal areas were located were observed at the laminar edge of the pallium (LEP) and in the vascular region, as well as in vocal motor relay regions in the pallidum and mesencephalon. In contrast, GFAP astrocytes were found in LEP, but not in the pallidum and mesencephalon in hummingbirds. Finally, when comparing GFAP astrocytes in the LEP region of both species, house wren astrocytes exhibited significantly more complex morphology than those of the rufous-tailed hummingbird. These findings suggest a difference in the location and cellular density of vocal circuits, as well as morphology of GFAP astrocytes between the house wren and the rufous-tailed hummingbird.
... Four current intensities (0.3, 0.4, 0.5, 0.6 mA) and four frequencies (12, 16, Table 1 Micro-stimulation electrode implantation site. NO 20, 24 Hz) were tested in this work. The interval between different stimulation intensities was 3-5 minutes. ...
... The terminology used in the present study is based on the Avian Brain Nomenclature Forum [20] and the atlas of Karten and Hodos (1967) [21]. The 3D coordinate values of LHy were determined based on the Nissl staining results and the MRI brain imaging data [17]. ...
... This suggests that a small pre-existing bias to process song with the left NCM becomes more pronounced through experience; a process through which hemispheric dominance arises 27 . We hypothesize that the same experience-dependent lateralization pattern may be present in the HVC (acronym used as a proper name 33 ), which is a premotor nucleus in the songbird brain that is part of the primary motor pathway and the anterior forebrain pathway (Fig. 1c) [34][35][36] . The HVC also receives input from auditory regions ( Fig. 1a-b), and thus it is critical in both song learning and song production 37,38 . ...
Juvenile male zebra finches (Taeniopygia guttata) must be exposed to an adult tutor during a sensitive period to develop normal adult song. The pre-motor nucleus HVC (acronym used as a proper name), plays a critical role in song learning and production (cf. Broca’s area in humans). In the human brain, left-side hemispheric dominance in some language regions is positively correlated with proficiency in linguistic skills. However, it is unclear whether this pattern depends upon language learning, develops with normal maturation of the brain, or is the result of pre-existing functional asymmetries. In juvenile zebra finches, even though both left and right HVC contribute to song production, baseline molecular activity in HVC is left-dominant. To test if HVC exhibits hemispheric dominance prior to song learning, we raised juvenile males in isolation from adult song and measured neuronal activity in the left and right HVC upon first exposure to an auditory stimulus. Activity in the HVC was measured using the immediate early gene (IEG) zenk (acronym for zif-268, egr-1, NGFI-a, and krox-24) as a marker for neuronal activity. We found that neuronal activity in the HVC of juvenile male zebra finches is not lateralized when raised in the absence of adult song, while normally-reared juvenile birds are left-dominant. These findings show that there is no pre-existing asymmetry in the HVC prior to song exposure, suggesting that lateralization of the song system depends on learning through early exposure to adult song and subsequent song-imitation practice.
... No pixelation adjustments or manipulation of the captured images were undertaken, except for the adjustment of contrast, brightness, and levels using Adobe Photoshop. The neuroanatomical demarcations and nomenclature used in the current study were derived from Kuenzel and van Tienhoven (1982), Medina and Reiner (1994), Reiner et al. (2004), Izawa and Watanabe (2007), and Brauth et al. (2011). ...
We examined the presence/absence and parcellation of cholinergic neurons in the hypothalami of five birds: a Congo grey parrot ( Psittacus erithacus ), a Timneh grey parrot ( P. timneh ), a pied crow ( Corvus albus ), a common ostrich ( Struthio camelus ), and an emu ( Dromaius novaehollandiae ). Using immunohistochemistry to an antibody raised against the enzyme choline acetyltransferase, hypothalamic cholinergic neurons were observed in six distinct clusters in the medial, lateral, and ventral hypothalamus in the parrots and crow, similar to prior observations made in the pigeon. The expression of cholinergic nuclei was most prominent in the Congo grey parrot, both in the medial and lateral hypothalamus. In contrast, no evidence of cholinergic neurons in the hypothalami of either the ostrich or emu was found. It is known that the expression of sleep states in the ostrich is unusual and resembles that observed in the monotremes that also lack hypothalamic cholinergic neurons. It has been proposed that the cholinergic system acts globally to produce and maintain brain states, such as those of arousal and rapid‐eye‐movement sleep. The hiatus in the cholinergic system of the ostrich, due to the lack of hypothalamic cholinergic neurons, may explain, in part, the unusual expression of sleep states in this species. These comparative anatomical and sleep studies provide supportive evidence for global cholinergic actions and may provide an important framework for our understanding of one broad function of the cholinergic system and possible dysfunctions associated with global cholinergic neural activity.
