Here, we recorded from the premotor nucleus robust nucleus of the Arcopallium (RA) in Bengalese finches and measured whether neural activity covaried with syllable structure across multiple renditions of individual syllables. 
Additionally, prior to sensorimotor learning, we observed elevated TguFmrp expression in the robust nucleus of the Arcopallium (RA) of post-hatch day 30 males, compared with the surrounding telencephalon, suggesting a preparation for this stage of song learning.
Our findings show, remarkably, that the gamma4-subunit transcript is highly enriched in the major nuclei of the song system, including the lateral magnocellular nucleus of the anterior nidopallium (LMAN), the medial magnocellular nucleus of the anterior nidopallium (MMAN), Area X, the robust nucleus of the Arcopallium (RA) and the HVC (used as the proper name), as well as Field L, which innervates the area surrounding HVC.
Among adult songbirds, males have larger brain areas such as the HVC (proper name) and RA (robust nucleus of the Arcopallium) that control the production of learned songs.
Caspase inhibitor infusion near HVC was sufficient to preserve neuron size ipsilaterally in a downstream nucleus, the robust nucleus of the Arcopallium.
Two forebrain regions are involved in distinct aspects of choices; i.e., nucleus accumbens-medial striatum (Ac-MSt) and Arcopallium intermedium (AI), an association area in the lateral forebrain.
Using pharmacological manipulation, electrical stimulation, and extracellular recordings of Uva projection neurons, we study the involvement of Uva in zebra finches for the generation of spontaneous activity and auditory responses in premotor area HVC (used as a proper name) and the downstream robust nucleus of the Arcopallium (RA).
We examined the neuromodulatory effects of noradrenergic and GABA B receptor activation on synaptic inputs to the premotor robust nucleus of the Arcopallium (RA) in zebra finches using whole cell voltage-clamp recording in vitro.
A portion of the Arcopallium also projects to the SNc and VTA and could carry auditory information.
With a species-specific probe for budgerigar androgen receptor mRNA, we found that the androgen receptor was expressed in the vocal areas, such as the central nucleus of the lateral nidopallium, the anterior Arcopallium, the oval nucleus of the mesopallium, the oval nucleus of the anterior nidopallium and the tracheosyringeal hypoglossal nucleus.
The song motor circuit includes premotor and motor cortical analogs, known as HVC (used as a proper name) and RA (the robust nucleus of the Arcopallium), respectively.
On the other hand, cadherin-7 was expressed in the robust nucleus of the Arcopallium (RA) in the sensory learning stage, and its expression was downregulated during the sensorimotor learning stage.
We show that, in the avian vocal control system, activation of the brainstem inspiratory nucleus paraambigualus (PAm), a likely homolog of the mammalian rostral ventral respiratory group, can drive neural activity bilaterally in the forebrain vocal control nuclei HVC (used as a proper name) and the robust nucleus of the Arcopallium (RA).
Proliferating cells were detected immunohistochemically on brain sections by incorporation of pre-training doses of 5-bromodeoxyuridine (BrdU) into DNA; numbers of new cells were counted in the intermediate medial mesopallium, the intermediate Arcopallium, the medial part of the mesopallium and the nidopallium, the dorsocaudal nidopallium, the hippocampus, and the parahippocampal region 24 h and seven days after training.
In the present study, we examined two gaze control centers of the barn owl: the optic tectum (OT) and the Arcopallium gaze fields (AGFs).
Both norepinephrine and dopamine displaced the binding of the radioligand though to a different extent in most of the regions studied (e.g., area X, the lateral part of the magnocellular nucleus of anterior nidopallium, HVC, Arcopallium dorsale, ventral tegmental area and substantia grisea centralis) but not in the robust nucleus of the Arcopallium.
Inside the telencephalon, mesopallium, nidopallium (+ entopallium + Arcopallium) and septum are smaller as well.
The vocal motor pathway (VMP) is a direct connection between HVC (proper name) and the robust nucleus of the Arcopallium (RA), whereas the anterior forebrain pathway (AFP) comprises an indirect circuit from HVC to RA that traverses the basal ganglia.
Neurons in a song premotor nucleus, the robust nucleus of the Arcopallium (RA), show increased intrinsic spontaneous firing rate and soma size when birds are in breeding condition.
In the avian song system the robust nucleus of the Arcopallium (RA) plays a key role in such coordination.
A caudal forebrain area of zebra finches that comprises a part of the caudal nidopallium and a part of the intermediate Arcopallium is highly activated during courtship. Being involved in the integration of external input and previously stored information, as well as in adding motivational factors, the caudal nidopallium and intermediate Arcopallium should be integrative areas receiving input from many other regions of the brain. The intermediate Arcopallium is recipient of input from intermediate and caudal nidopallium, mesopallium and densocellular hyperpallium. Bilateral innervation by ventral intermediate Arcopallium indicates links with sensori-motor pathways, while the projection from the caudal nidopallium to intermediate Arcopallium suggests monosynaptic and disynaptic input to downstream motor pathways. These findings support the idea of an involvement of the caudal nidopallium and the intermediate Arcopallium in the control of courtship behavior..
The volumes of Area X and the robust nucleus of the Arcopallium (RA) were significantly regressed by 7 and 20 days, respectively.
However, recordings in premotor telencephalic nucleus HVC (used as proper name) and RA (robust nucleus of Arcopallium) suggest that song is represented by sparse, fine-grained bursting on the 5-10 ms timescale.
Cell proliferation was examined in the intermediate medial mesopallium (IMM), Arcopallium intermedium (AI), medial part of nidopallium and mesopallium (MNM), nidopallium dorso-caudalis (Ndc), hippocampus (Hp) and area parahippocampalis (APH), as well as in corresponding ventricular zones.
We measured the volumes of the nuclei HVC, robust nucleus of Arcopallium (RA), and area X at 7 and 30 d after exposure to long days plus testosterone in deafened and normally hearing birds.
Although the two song types are similar, the level of neural activity and expression of the immediate early gene egr-1 are higher during undirected than during directed singing in the lateral part of the basal ganglia song nucleus AreaX (LAreaX) and its efferent pallial song nuclei lateral magnocellular nucleus of the anterior nidopallium (LMAN) and the robust nucleus of the Arcopallium (RA).
Considerable GABA labeling was also seen in the shelf area of caudodorsal nidopallium, and the cup area in the Arcopallium, as well as in area X, the lateral magnocellular nucleus of the anterior nidopallium, the robust nucleus of the Arcopallium and nidopallial nucleus HVC.
Testosterone treatment increased the sizes of two SCRs, the HVC and Robust nucleus of the Arcopallium (RA).
Expression was significantly greater in the HVC (used as a proper name) and robust nucleus of the Arcopallium (RA) in males than in females.
One pathway descends directly from HVC to the vocal premotor nucleus RA (the robust nucleus of the Arcopallium) whereas a second pathway descends from HVC into a basal ganglia circuit (the anterior forebrain pathway, AFP) that also terminates in RA.
To study the mechanism, we used expression of the neural activity-induced gene ZENK (or egr-1), which shows singing-regulated expression in a social context-dependent manner: high levels in both pathways when singing undirected and low levels in the lateral part of the loop and in the robust nucleus of the Arcopallium (RA) of the motor pathway when singing directed to another animal.
To address this issue, we first measured four variables for song complexity, i.e., song repertoire size, syllable repertoire size, the mean number of syllables per phrase (MNS) and the number of syllables in the longest phrase (NSLP), and the sizes of three song control nuclei, i.e., HVC, RA (the robust nucleus of the Arcopallium), and Area X in 14 oscine species from eight families.
This nucleus consists of a heterogenous population of inhibitory interneurons (HVC(IN)) and two populations of projection neurons that send axons towards either the robust nucleus of the Arcopallium (HVC(RA)) or the striatal nucleus area X (HVC(X)).
Additionally, we found novel inputs to area X from the nidopallium and Arcopallium, the mesencephalic central gray, and the dorsolateralis anterior (DLL) and posterior (DLP) lateralis in the thalamus.
In addition, our experiments indicate that the HVC projection neurons that project into nucleus robust nucleus of the Arcopallium (RA) are born locally from the ventricular region immediately dorsal to HVC.
Additional weaker signals were detected in the rostral forebrain, Arcopallium and mesencephalic regions.
We first showed that the participation of NMDA receptor NR2B subunits in synaptic currents in the robust nucleus of the Arcopallium (RA), a critical location for integration of signals during song learning by young birds, decreases from young birds to adults.
One pathway descends directly from HVC to the vocal premotor nucleus RA (the robust nucleus of the Arcopallium) whereas a second pathway descends from HVC into a basal ganglia circuit (the anterior forebrain pathway, AFP) that also terminates in RA.
Here we show that the expression of the gene encoding the middle-weight neurofilament (NF-M), an important component of the neuronal cytoskeleton and a useful tool for studying the cytarchitectonic organization of mammalian cortical areas, is highly enriched in large neurons within pallial song control nuclei (nucleus HVC, robustus nucleus of the Arcopallium, and lateral magnocellular nucleus of the nidopallium) of male zebra finches (Taeniopygia guttata).
Testing with single units recorded from individual electrodes within the robust nucleus of the Arcopallium of zebra finches and with recordings from an array placed within the motor cortex of macaque monkeys demonstrates that the approach can identify occurrences of specified patterns with good time precision in a broad range of neurophysiological data..
Retrogradely labeled cells were located within the Arcopallium, the hyperpallium apicale (HA) and the temporo-parieto-occipital area (TPO). Descending projections from HA, Arcopallium, and TPO were mainly or exclusively ipsilateral with the contralateral projection being extremely small.
Excitotoxic lesion of the bilateral Arcopallium intermedium also selectively reduced the choice of the six grains, while leaving actual cost investment (number of pecks and handling time) unaltered. Operant peck latencies somewhat depended on food rewards, but were not affected by lesions of the Arcopallium or the ventral striatum. The Arcopallium could contribute to foraging behaviors by enabling chicks to overcome the handling cost, thus gaining more beneficial food. Furthermore, the present results indicate doubly dissociated functional roles of the ventral striatum and the Arcopallium, the former in the cost of traveling for food and the latter in the cost of handling food, respectively..
Similarly, brain mass and high vocal center (HVC), robust nucleus of the Arcopallium (RA), and lateral magnocellular nucleus of the anterior nidopallium (LMAN) volumes did not covary with nestling condition and growth measurements.
Spontaneous activity patterns in the nucleus robustus of the Arcopallium (RA) and in HVC (high vocal center) of the sleeping songbird resemble premotor patterns in these areas observed during singing.
In Experiment 1, males exhibited increased expression of both RPL17 and RPL37 compared to females in Area X, the robust nucleus of the Arcopallium (RA), and the ventral ventricular zone (VVZ), which may provide neurons to Area X.
L-aspartate- and L-glutamate-immunoreactive neurons in the Arcopallium and posterior amygdaloid pallium were identified and counted by using fluorescence microscopy and confocal laser scanning microscopy. Most labeled neurons of Arcopallium were enriched in glutamate as well as aspartate. However, the Arcopallium and posterior amygdaloid pallium differed from a neighboring telencephalic region (nidopallium; formerly neostriatum) by containing a substantial proportion of cells singly labeled for L-aspartate (15%, vs.
Our experiments indeed show that the lateral nidopallium receives input from a variety of telencephalic regions including the primary and secondary areas of both visual pathways, the globus pallidus, the caudolateral nidopallium functionally comparable to the prefrontal cortex, the caudomedial nidopallium involved in song perception and storage of song-related memories, and some parts of the Arcopallium.
The expression at posthatch day 9 (P9) was heavy in almost the entire telencephalon and showed heavier expression in SVZ and song regions such as the high vocal center (HVC) and the robust nucleus of Arcopallium (RA).
Then, we examined the distribution of SP and ENK in four control nuclei, two in the motor pathway, i.e., HVC and the robust nucleus of Arcopallium (RA), and the other two in the forebrain pathway, i.e., Area X and the lateral magnocellular nucleus of the anterior nidopallium (LMAN).
The synaptic connection from high vocal center (HVC) to robust nucleus of the Arcopallium (RA) is a pivotal part of vocal motor pathway in songbirds.
However, the neural regions underlying these vocalizations, such as HVC, area X, and RA (robust nucleus of Arcopallium), remain understudied.
In parallel, the expression of PKC beta1 increases transiently 2 weeks after deafening, and then decreases gradually in the robust nucleus of the Arcopallium (RA) of Bengalese finches, similar to the pattern observed during developmental song learning.
The highest net-stimulated [ (35)S]GTPgammaS binding values induced by the selective CB(1) receptor agonist WIN55,212-2 were observed in the nucleus paramedianus internus thalami, and high values of [ (35)S]GTPgammaS binding were observed in the TnA, Mol, Arcopallium dorsale and Arcopallium intermedium.
Later establishment and maintenance in adulthood of small somata and neuropil staining within regions of rostral telencephalon [ HVC and robust nucleus of the Arcopallium (RA)] are consistent with a vocal motor role for cannabinoid signaling.
During singing, neurons in premotor nucleus RA (robust nucleus of the Arcopallium) of the zebra finch produce complex temporal sequences of bursts that are recapitulated during sleep.
Retrograde tracer injections into the VLT revealed an ipsilateral forebrain input from the visual Wulst, from subregions of the Arcopallium, and bilateral afferents from the optic tectum.
A potential site of NE action is the robust nucleus of the Arcopallium (RA): RA receives noradrenergic inputs and has adrenergic receptors, and it is a sensorimotor area instrumental to song production.
The effects of bilateral chemical lesions of the ventral striatum (nucleus accumbens and the surrounding areas in the medial striatum) and Arcopallium (major descending area of the avian telencephalon) were examined in 1-2-weeks old domestic chicks. In task 1, bilateral lesions of the ventral striatum (but not the Arcopallium) enhanced the impulsiveness of the chicks' choices, suggesting that choices based on the anticipated proximity were selectively changed. Neural correlates of anticipated food rewards in the ventral striatum (but not those in the Arcopallium) could allow chicks to invest appropriate amount of work-cost in approaching distant food resources..
Nissl staining was used to measure the volumes of four telencephalic song nuclei: Area X, HVC, the robust nucleus of the Arcopallium (RA), and the lateral portion of the magnocellular nucleus of the anterior nidopallium (LMAN).
We show that, in the context of song auditory stimulation, Arc expression is induced in several telencephalic auditory areas, most prominently the caudomedial nidopallium and mesopallium, whereas in the context of singing, Arc is also induced in song control areas, namely nucleus HVC, used as a proper name, the robust nucleus of the Arcopallium and the interface nucleus of the nidopallium.
Behavioral studies in barn owls indicate that both the optic tectum (OT) and the auditory Arcopallium (AAr) mediate sound localization through the presence of neurons that respond only when sound comes from a circumscribed direction in space.
We found that brief stimulation in the forebrain nuclei HVC (used as a proper name) and RA (robust nucleus of the Arcopallium) caused a short-latency truncation of ongoing song syllables, which ultimately led to a cessation of the ongoing motor sequence.
In the neural song circuit, strong expression was found in high vocal center (HVC), para-HVC, and at a very low level in the robust nucleus of the Arcopallium (RA).
The present study was designed to explore whether these regions, as well as three song control nuclei [ area X, the high vocal center (HVC), and the robust nucleus of the Arcopallium (RA)], might be involved differentially in song produced within compared to outside of a breeding context.
This dissociation between ZENK mRNA and ZENK protein was regionally specific to the robust nucleus of the Arcopallium (RA), a region that is well known for its control of vocal-motor behavior in birds.
Prominent labeling was also evident in the nucleus taeniae and subpallial amygdala, but not in the Arcopallium in film autoradiograms.
These regions include the hippocampal complex, the medial nidopallium, and the ventromedial Arcopallium.
Here we induced seasonal-like changes in the song systems of adult white-crowned sparrows and used extracellular recording in acute brain slices from those individuals to study physiological properties of neurons in the robust nucleus of the Arcopallium (RA), a pre-motor nucleus necessary for song production.
However, at this time, none of the steroids had any effects on the volumes of two other song control nuclei, Area X of the medial striatum and the robust nucleus of the Arcopallium (RA), that are efferent targets of HVC, known to be regulated by androgen in canaries and also to play a role in the control of adult song.
the preoptic area, bed nucleus striae terminalis, Arcopallium, nucleus intercollicularis, periaqueductal gray and the ventral tegmental area.
To investigate how the sparse HVC code is transformed into continuous vocal patterns, we recorded in the singing zebra finch from populations of neurons in the robust nucleus of Arcopallium (RA), a premotor area intermediate between HVC and the motor neurons.
In conclusion, social experience during an early sensitive period of song learning in the zebra finch is essential for the normal development of PKC expression in the robust nucleus of the Arcopallium (RA), a premotor nucleus related to vocal plasticity..
Here, we have tested the hypothesis that neurotrophin expression in the robust nucleus of the Arcopallium (RA), the telencephalic output controlling song, regulates song variability.
We measured gonadal activity and the size of song nuclei (high vocal center, robust nucleus of the Arcopallium, and area X) during each population's breeding and nonbreeding periods.
Development and use of an anti-zebra finch cannabinoid receptor type 1 (CB1) antibody demonstrates distinct, dense cannabinoid receptor expression within song regions including Area X, lMAN (lateral magnocellular nucleus of anterior nidopallium), HVC, RA (robust nucleus of Arcopallium), and L2.
Changes in the volumes of song control nuclei, such as HVC and the robust nucleus of the Arcopallium (RA), are observed seasonally.
AChE fibres and cells are intensely labelled in the forebrain nucleus area X, strongly labelled in high vocal centre (HVC) perikarya, and moderately to lightly labelled in the somata and neuropil of vocal control nuclei robust nucleus of Arcopallium (RA), medial magnocellular nucleus of the anterior nidopallium (MMAN) and lateral magnocellular nucleus of the anterior nidopallium (LMAN).
This included relatively lower levels of all four AMPA subunits in lMAN, strikingly higher levels of the kainite subunit GluR5 in the robust nucleus of the Arcopallium (RA), higher and lower levels respectively of the NMDA subunits NR2A and NR2B in most vocal nuclei and lower levels of the metabotropic group I subtypes (mGluR1 and -5) in most vocal nuclei and the group II subtype (mGluR2), showing a unique expression pattern of very low levels in RA and very high levels in HVC.
DM had reciprocal fiber connections with the V-shaped layer, Ma, and DL as well as with several subdivisions of the Arcopallium.
The sectors of the hyperstriatum composing the Wulst (i.e., the hyperstriatum accessorium intermedium, and dorsale), the hyperstriatum ventrale, the neostriatum, and the archistriatum have been renamed (respectively) the hyperpallium (hypertrophied pallium), the mesopallium (middle pallium), the nidopallium (nest pallium), and the Arcopallium (arched pallium).
Telencephalic areas devoid of chBRS-3.5 signals were the entopallium, Arcopallium anterius, globus pallidus, nucleus intrapeduncularis, tuberculum olfactorius, nucleus septalis lateralis, hypothalamic and thalamic areas.
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