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Michael Colombo - One of the best experts on this subject based on the ideXlab platform.
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Neurons in the pigeon Nidopallium caudolaterale, but not the corticoidea dorsolateralis, display value and effort discounting activity
Scientific reports, 2019Co-Authors: Madeline Dykes, Blake Porter, Michael ColomboAbstract:We recorded from single neurons in two areas of the pigeon brain while birds were required to peck a stimulus indicating either a high effort task or a low effort task would follow. Upon completion of the task the birds received the same reward. We found that activity in the Nidopallium caudolaterale, an area equivalent to the mammalian prefrontal cortex, was modulated by the value of the reward that would be received based on how much effort was required to obtain it. Value coding was most prominent during the presentation of the stimulus indicating a high or low effort task, and in the delay period immediately prior to carrying out the effort task. In contrast, activity in the corticoidea dorsolateralis was not modulated by value, however, population firing patterns suggest that it may be involved in associating actions with outcomes. Our findings support the view that activity in the Nidopallium caudolaterale reflects value of reward as a function of effort discounting and as such may serve functions similar to the mammalian anterior cingulate cortex.
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Nidopallium caudolaterale neuronal responses during serial-order behaviour in pigeons.
Behavioural brain research, 2019Co-Authors: Melissa Johnston, Blake Porter, Michael ColomboAbstract:Serial-order behaviour is the ability to complete a sequence of responses in order to obtain a reward. Serial-order tasks can be thought of as either externally-ordered (EO) such that the order of responses is predetermined, or internally-ordered (IO) such that the subject determines the order of responses from trial to trial. Ordinal knowledge (representation of first, second, or third etc.) is a key component of successful serial-order behaviour, and is considered a higher-order cognitive function. The Nidopallium caudolaterale (NCL) is the avian equivalent to the prefrontal cortex, an area of the primate brain important for serial-order behaviour. The importance of the NCL for serial-order behaviour, however, is still unknown. In the current study, we trained pigeons to complete either three-item EO or IO tasks and recorded single-neuron activity from the NCL to determine whether neurons in the NCL code ordinal knowledge. Our results support the view that the NCL is involved in serial-order behaviour by coding ordinal position, at least with respect to the IO task. The absence of any ordinal coding during the EO task could be explained by the different strategies that birds adopt between the EO and IO tasks.
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Delay activity in pigeon Nidopallium caudolaterale during a variable-delay memory task.
Behavioral neuroscience, 2019Co-Authors: Melissa Johnston, Blake Porter, Michael ColomboAbstract:Neurons in the pigeon Nidopallium caudolaterale (NCL) are important for the maintenance of information across delays as long as 3 s. In the current study, we recorded neural activity from the avian NCL of 3 birds trained on a working memory task with three different delay lengths intermixed within a session. We found that when the birds are unable to predict the upcoming delay length there is no evidence that NCL cells engage in temporal coding. Furthermore, delay activity did not differ between correct and incorrect trials. Both findings have implications for the function of delay activity and its role in supporting working memory. (PsycINFO Database Record (c) 2019 APA, all rights reserved).
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Effects of Nidopallium caudolaterale inactivation on serial-order behavior in pigeons ( Columba livia).
Journal of neurophysiology, 2018Co-Authors: Melissa Johnston, Andrew N. Clarkson, Emma K. Gowing, Damian Scarf, Michael ColomboAbstract:We examined the role of the Nidopallium caudolaterale (NCL) in serial-order behavior by training pigeons on a four-item serial-order task and selectively inhibiting the region with TTX. Although TT...
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Neurons in the Pigeon Nidopallium Caudolaterale Display Value-Related Activity.
Scientific reports, 2018Co-Authors: Madeline Dykes, Aylin Klarer, Blake Porter, Jonas Rose, Michael ColomboAbstract:We recorded from neurons in the Nidopallium caudolaterale, the avian equivalent of the mammalian prefrontal cortex, in four birds. The birds were required to peck a stimulus that indicated the amount of reward they would receive (small or large) after a certain delay (short or long). We found that the activity of neurons in the Nidopallium caudolaterale was modulated by the value of the reward that would be received based on the reward amount and the delay to reward. We found that value coding was most prominent during the presentation of the sample period, and less so during the delay period and during the presentation of the reward itself. Our findings support the view that activity in Nidopallium caudolaterale reflects the encoding of the value of reward based on a combination of reward amount and delay to a reward.
Onur Güntürkün - One of the best experts on this subject based on the ideXlab platform.
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fMRI Reveals a Novel Region for Evaluating Acoustic Information for Mate Choice in a Female Songbird.
Current biology : CB, 2018Co-Authors: Lisbeth Van Ruijssevelt, Onur Güntürkün, Yining Chen, Kaya Von Eugen, Julie Hamaide, Geert De Groof, Marleen Verhoye, Sarah C. Woolley, Annemie Van Der LindenAbstract:Summary Selection of sexual partners is among the most critical decisions that individuals make and is therefore strongly shaped by evolution. In social species, where communication signals can convey substantial information about the identity, state, or quality of the signaler, accurate interpretation of communication signals for mate choice is crucial. Despite the importance of social information processing, to date, relatively little is known about the neurobiological mechanisms that contribute to sexual decision making and preferences. In this study, we used a combination of whole-brain functional magnetic resonance imaging (fMRI), immediate early gene expression, and behavior tests to identify the circuits that are important for the perception and evaluation of courtship songs in a female songbird, the zebra finch ( Taeniopygia guttata ). Female zebra finches are sensitive to subtle differences in male song performance and strongly prefer the longer, faster, and more stereotyped courtship songs to non-courtship renditions. Using BOLD fMRI and EGR1 expression assays, we uncovered a novel region involved in auditory perceptual decision making located in a sensory integrative region of the avian central Nidopallium outside the traditionally studied auditory forebrain pathways. Changes in activity in this region in response to acoustically similar but categorically divergent stimuli showed stronger parallels to behavioral responses than an auditory sensory region. These data highlight a potential role for the caudocentral Nidopallium (NCC) as a novel node in the avian circuitry underlying the evaluation of acoustic signals and their use in mate choice.
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Blocking NMDA-receptors in the pigeon's "prefrontal" caudal Nidopallium impairs appetitive extinction learning in a sign-tracking paradigm.
Frontiers in behavioral neuroscience, 2015Co-Authors: Daniel Lengersdorf, David Marks, Metin Uengoer, Maik C. Stüttgen, Onur GüntürkünAbstract:Extinction learning provides the ability to flexibly adapt to new contingencies by learning to inhibit previously acquired associations in a context-dependent manner. The neural networks underlying extinction learning were mostly studied in rodents using fear extinction paradigms. To uncover invariant properties of the neural basis of extinction learning, we employ pigeons as a model system. Since the prefrontal cortex of mammals is a key structure for extinction learning, we assessed the role of N-methyl-D-aspartate receptors (NMDARs) in the Nidopallium caudolaterale, the avian functional equivalent of mammalian prefrontal cortex. Since NMDARs in prefrontal cortex have been shown to be relevant for extinction learning, we locally antagonized NMDARs through 2-Amino-5-phosphonovalerianacid (APV) during extinction learning in a within-subject sign-tracking ABA-renewal paradigm. APV-injection slowed down extinction learning and in addition also caused a disinhibition of responding to a continuously reinforced control stimulus.. In subsequent retrieval sessions, spontaneous recovery was increased while ABA renewal was unaffected. The effect of APV resembles that observed in studies of fear extinction with rodents, suggesting common neural substrates of extinction under both appetitive and aversive conditions and highlighting the similarity of mammalian prefrontal cortex and the avian caudal Nidopallium despite 300 million years of independent evolution.
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Serotonin release in the caudal Nidopallium of adult laying hens genetically selected for high and low feather pecking behavior: An in vivo microdialysis study
Behavioural brain research, 2014Co-Authors: Marjolein S. Kops, Joergen B. Kjaer, Onur Güntürkün, Koen G.c. Westphal, Gerdien A.h. Korte-bouws, Berend Olivier, J. Elizabeth Bolhuis, S. Mechiel KorteAbstract:Severe feather pecking (FP) is a detrimental behavior causing welfare problems in laying hens. Divergent genetic selection for FP in White Leghorns resulted in strong differences in FP incidences between lines. More recently, it was shown that the high FP (HFP) birds have increased locomotor activity as compared to hens of the low FP (LFP) line, but whether these lines differ in central serotonin (5-hydroxytryptamine, 5-HT) release is unknown. We compared baseline release levels of central 5-HT, and the metabolite 5-HIAA in the limbic and prefrontal subcomponents of the caudal Nidopallium by in vivo microdialysis in adult HFP and LFP laying hens from the ninth generation of selection. A single subcutaneous d-fenfluramine injection (0.5 mg/kg) was given to release neuronal serotonin in order to investigate presynaptic storage capacity. The present study shows that HFP hens had higher baseline levels of 5-HT in the caudal Nidopallium as compared to LFP laying hens. Remarkably, no differences in plasma tryptophan levels (precursor of 5-HT) between the lines were observed. d-fenfluramine increased 5-HT levels in both lines similarly indirectly suggesting that presynaptic storage capacity was the same. The present study shows that HFP hens release more 5-HT under baseline conditions in the caudal Nidopallium as compared to the LFP birds. This suggests that HFP hens are characterized by a higher tonic 5-HT release.
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The putative pigeon homologue to song bird LMAN does not modulate behavioral variability.
Behavioural brain research, 2014Co-Authors: Sascha Helduser, Maren Westkott, Klaus Pawelzik, Onur GüntürkünAbstract:The active generation of behavioral variability is thought to be a pivotal element in reinforcement based learning. One example for this principle is song learning in oscine birds. Oscines possess a highly specialized set of brain areas that compose the song system. It is yet unclear how the song system evolved. One important hypothesis assumes a motor origin of the song system, i.e. the song system may have developed from motor pathways that were present in an early ancestor of extant birds. Indeed, in pigeons neural pathways are present that parallel the song system. We examined whether one component of these pathways, a forebrain area termed Nidopallium intermedium medialis pars laterale (NIML), is functionally comparable to its putative homologue, the lateral magnocellular nucleus of the anterior Nidopallium (LMAN) of the song system. LMAN conveys variability into the motor output during singing; a function crucial for song learning and maintenance. We tested if NIML is likewise associated with the generation of variability. We used a behavioral paradigm in which pigeons had to find hidden target areas on a touch screen to gain food rewards. Alterations in pecking variability would result in changes of performance levels in this search paradigm. We found that pharmacological inactivation of NIML did not reduce pecking variability contrasting increases of song stereotypy observed after LMAN inactivation.
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Stimulus-response-outcome coding in the pigeon Nidopallium caudolaterale.
PloS one, 2013Co-Authors: Sarah Starosta, Onur Güntürkün, Maik C. StüttgenAbstract:A prerequisite for adaptive goal-directed behavior is that animals constantly evaluate action outcomes and relate them to both their antecedent behavior and to stimuli predictive of reward or non-reward. Here, we investigate whether single neurons in the avian Nidopallium caudolaterale (NCL), a multimodal associative forebrain structure and a presumed analogue of mammalian prefrontal cortex, represent information useful for goal-directed behavior. We subjected pigeons to a go-nogo task, in which responding to one visual stimulus (S+) was partially reinforced, responding to another stimulus (S-) was punished, and responding to test stimuli from the same physical dimension (spatial frequency) was inconsequential. The birds responded most intensely to S+, and their response rates decreased monotonically as stimuli became progressively dissimilar to S+; thereby, response rates provided a behavioral index of reward expectancy. We found that many NCL neurons' responses were modulated in the stimulus discrimination phase, the outcome phase, or both. A substantial fraction of neurons increased firing for cues predicting non-reward or decreased firing for cues predicting reward. Interestingly, the same neurons also responded when reward was expected but not delivered, and could thus provide a negative reward prediction error or, alternatively, signal negative value. In addition, many cells showed motor-related response modulation. In summary, NCL neurons represent information about the reward value of specific stimuli, instrumental actions as well as action outcomes, and therefore provide signals useful for adaptive behavior in dynamically changing environments.
Melissa Johnston - One of the best experts on this subject based on the ideXlab platform.
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Nidopallium caudolaterale neuronal responses during serial-order behaviour in pigeons.
Behavioural brain research, 2019Co-Authors: Melissa Johnston, Blake Porter, Michael ColomboAbstract:Serial-order behaviour is the ability to complete a sequence of responses in order to obtain a reward. Serial-order tasks can be thought of as either externally-ordered (EO) such that the order of responses is predetermined, or internally-ordered (IO) such that the subject determines the order of responses from trial to trial. Ordinal knowledge (representation of first, second, or third etc.) is a key component of successful serial-order behaviour, and is considered a higher-order cognitive function. The Nidopallium caudolaterale (NCL) is the avian equivalent to the prefrontal cortex, an area of the primate brain important for serial-order behaviour. The importance of the NCL for serial-order behaviour, however, is still unknown. In the current study, we trained pigeons to complete either three-item EO or IO tasks and recorded single-neuron activity from the NCL to determine whether neurons in the NCL code ordinal knowledge. Our results support the view that the NCL is involved in serial-order behaviour by coding ordinal position, at least with respect to the IO task. The absence of any ordinal coding during the EO task could be explained by the different strategies that birds adopt between the EO and IO tasks.
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Delay activity in pigeon Nidopallium caudolaterale during a variable-delay memory task.
Behavioral neuroscience, 2019Co-Authors: Melissa Johnston, Blake Porter, Michael ColomboAbstract:Neurons in the pigeon Nidopallium caudolaterale (NCL) are important for the maintenance of information across delays as long as 3 s. In the current study, we recorded neural activity from the avian NCL of 3 birds trained on a working memory task with three different delay lengths intermixed within a session. We found that when the birds are unable to predict the upcoming delay length there is no evidence that NCL cells engage in temporal coding. Furthermore, delay activity did not differ between correct and incorrect trials. Both findings have implications for the function of delay activity and its role in supporting working memory. (PsycINFO Database Record (c) 2019 APA, all rights reserved).
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Effects of Nidopallium caudolaterale inactivation on serial-order behavior in pigeons ( Columba livia).
Journal of neurophysiology, 2018Co-Authors: Melissa Johnston, Andrew N. Clarkson, Emma K. Gowing, Damian Scarf, Michael ColomboAbstract:We examined the role of the Nidopallium caudolaterale (NCL) in serial-order behavior by training pigeons on a four-item serial-order task and selectively inhibiting the region with TTX. Although TT...
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Pigeon NCL and NFL neuronal activity represents neural correlates of the sample.
Behavioral neuroscience, 2017Co-Authors: Melissa Johnston, Catrona Anderson, Michael ColomboAbstract:Four birds were trained on a delayed matching-to-sample task with common outcomes where correct responses during both red and green trials yielded reward. We recorded neuronal activity from the avian Nidopallium caudolaterale, the avian equivalent of the mammalian prefrontal cortex, and the avian Nidopallium frontolaterale, a higher-order visual processing region. In both regions we found sustained activity during the delay period of both red and green trials. These findings provide the first evidence that delay activity in the pigeon's Nidopallium caudolaterale and Nidopallium frontolaterale represent a neural correlate for the to-be-remembered sample stimulus. (PsycINFO Database Record
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Neural correlates of sample-coding and reward-coding in the delay activity of neurons in the entopallium and Nidopallium caudolaterale of pigeons (Columba livia)
Behavioural brain research, 2016Co-Authors: Melissa Johnston, Catrona Anderson, Michael ColomboAbstract:We recorded neuronal activity from the Nidopallium caudolaterale, the avian equivalent of mammalian prefrontal cortex, and the entopallium, the avian equivalent of the mammalian visual cortex, in four birds trained on a differential outcomes delayed matching-to-sample procedure in which one sample stimulus was followed by reward and the other was not. Despite similar incidence of reward-specific and reward-unspecific delay cell types across the two areas, overall entopallium delay activity occurred following both rewarded and non-rewarded stimuli, whereas Nidopallium caudolaterale delay activity tended to occur following the rewarded stimulus but not the non-rewarded stimulus. These findings are consistent with the view that delay activity in entopallium represents a code of the sample stimulus whereas delay activity in Nidopallium caudolaterale represents a code of the possibility of an upcoming reward. However, based on the types of delay cells encountered, cells in NCL also code the sample stimulus and cells in ENTO are influenced by reward. We conclude that both areas support the retention of information, but that the activity in each area is differentially modulated by factors such as reward and attentional mechanisms.
Monika Sadananda - One of the best experts on this subject based on the ideXlab platform.
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Afferentation of a caudal forebrain area activated during courtship behavior: a tracing study in the zebra finch (Taeniopygia guttata).
Brain research, 2007Co-Authors: Monika Sadananda, Stefan Korte, Hans-joachim BischofAbstract: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. This activation is thought to reflect the processing of information that is necessary for the choice of an appropriate mate. In addition to the information on the potential mate, control of courtship behavior includes motivational aspects. 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. Our results indeed show that the caudal Nidopallium receives input from a variety of telencephalic regions including the secondary visual and auditory areas. The intermediate arcopallium is recipient of input from intermediate and caudal Nidopallium, mesopallium and densocellular hyperpallium. Regions closely associated with the song control nuclei also innervate both regions. There are also specific visual and auditory thalamic inputs, while specific motivating catecholaminergic mesencephalic afferents include the ventral tegmental area, the substantia nigra and the locus coeruleus. In addition, non-specific activation reaches these areas from the mesencephalic reticular formation. 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.
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Afferentation of the lateral Nidopallium: A tracing study of a brain area involved in sexual imprinting in the zebra finch (Taeniopygia guttata).
Brain research, 2006Co-Authors: Monika Sadananda, Hans-joachim BischofAbstract:The lateral forebrain of zebra finches that comprises parts of the lateral Nidopallium and parts of the lateral mesopallium is supposed to be involved in the storage and processing of visual information acquired by an early learning process called sexual imprinting. This information is later used to select an appropriate sexual partner for courtship behavior. Being involved in such a complicated behavioral task, the lateral Nidopallium should be an integrative area receiving input from many other regions of the brain. 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. There are also a number of thalamic, mesencephalic, and brainstem efferents including the catecholaminergic locus coeruleus and the unspecific activating reticular formation. The spatial distribution of afferents suggests a compartmentalization of the lateral Nidopallium into several subdivisions. Based on its connections, the lateral Nidopallium should be considered as an area of higher order processing of visual information coming from the tectofugal and the thalamofugal visual pathways. Other sensory modalities and also motivational factors from a variety of brain areas are also integrated here. These findings support the idea of an involvement of the lateral Nidopallium in imprinting and the control of courtship behavior.
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Acetylcholinesterase in central vocal control nuclei of the zebra finch (Taeniopygia guttata)
Journal of Biosciences, 2004Co-Authors: Monika SadanandaAbstract:The distribution of acetylcholinesterase (AChE) in the central vocal control nuclei of the zebra finch was studied using enzyme histochemistry. 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). The identified sites of cholinergic and/or cholinoceptive neurons are similar to the cholinergic presence in vocal control regions of other songbirds such as the song sparrow, starling and another genus of the zebra finch ( Poephila guttata ), and to a certain extent in parallel vocal control regions in vocalizing birds such as the budgerigar. AChE presence in the vocal control system suggests innervation by either afferent projecting cholinergic systems and/or local circuit cholinergic neurons. Co-occurrence with choline acetyltransferase (ChAT) indicates efferent cholinergic projections. The cholinergic presence in parts of the zebra finch vocal control system, such as the area X, that is also intricately wired with parts of the basal ganglia, the descending fibre tracts and brain stem nuclei could underlie this circuitry’s involvement in sensory processing and motor control of song.
Sarah W Bottjer - One of the best experts on this subject based on the ideXlab platform.
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Cortical inter-hemispheric circuits for multimodal vocal learning in songbirds
The Journal of comparative neurology, 2017Co-Authors: Amy K. Paterson, Sarah W BottjerAbstract:Vocal learning in songbirds and humans is strongly influenced by social interactions based on sensory inputs from several modalities. Songbird vocal learning is mediated by cortico-basal ganglia circuits that include the SHELL region of lateral magnocellular nucleus of the anterior Nidopallium (LMAN), but little is known concerning neural pathways that could integrate multimodal sensory information with SHELL circuitry. In addition, cortical pathways that mediate the precise coordination between hemispheres required for song production have been little studied. In order to identify candidate mechanisms for multimodal sensory integration and bilateral coordination for vocal learning in zebra finches, we investigated the anatomical organization of two regions that receive input from SHELL: the dorsal caudolateral Nidopallium (dNCLSHELL) and a region within the ventral arcopallium (Av). Anterograde and retrograde tracing experiments revealed a topographically organized inter-hemispheric circuit: SHELL and dNCLSHELL, as well as adjacent nidopallial areas, send axonal projections to ipsilateral Av; Av in turn projects to contralateral SHELL, dNCLSHELL, and regions of Nidopallium adjacent to each. Av on each side also projects directly to contralateral Av. dNCLSHELL and Av each integrate inputs from ipsilateral SHELL with inputs from sensory regions in surrounding Nidopallium, suggesting that they function to integrate multimodal sensory information with song-related responses within LMAN-SHELL during vocal learning. Av projections share this integrated information from the ipsilateral hemisphere with contralateral sensory and song-learning regions. Our results suggest that the inter-hemispheric pathway through Av may function to integrate multimodal sensory feedback with vocal-learning circuitry and coordinate bilateral vocal behavior.
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conjunction of vocal production and perception regulates expression of the immediate early gene zenk in a novel cortical region of songbirds
Journal of Neurophysiology, 2010Co-Authors: Sarah W Bottjer, Tanya L Alderete, Daniel ChangAbstract:The cortical nucleus LMAN (lateral magnocellular nucleus of the anterior Nidopallium) provides the output of a basal ganglia pathway that is necessary for acquisition of learned vocal behavior during development in songbirds. LMAN is composed of two subregions, a core and a surrounding shell, that give rise to independent pathways that traverse the forebrain in parallel. The LMANshell pathway forms a recurrent loop that includes a cortical region, the dorsal region of the caudolateral Nidopallium (dNCL), hitherto unknown to be involved with learned vocal behavior. Here we show that vocal production strongly induces the IEG product ZENK in dNCL of zebra finches. Hearing tutor song while singing is more effective at inducing expression in dNCL of juvenile birds during the auditory–motor integration stage of vocal learning than is hearing conspecific song. In contrast, hearing conspecific song is relatively more effective at inducing expression in adult birds, regardless of whether they are producing song. Furthermore, ZENK+ neurons in dNCL include projection neurons that are part of the LMANshell recurrent loop and a high proportion of dNCL projection neurons express ZENK in singing juvenile birds that hear tutor song. Thus juvenile birds that are actively refining their vocal pattern to imitate a tutor song show high levels of ZENK induction in dNCL neurons when they are singing while hearing the song of their tutor and low levels when they hear a novel conspecific. This pattern indicates that dNCL is a novel brain region involved with vocal learning and that its function is developmentally regulated.
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Parallel pathways for vocal learning in basal ganglia of songbirds
Nature Neuroscience, 2010Co-Authors: Sarah W Bottjer, Brie AltenauAbstract:The cortical nucleus LMAN (lateral magnocellular nucleus of the anterior Nidopallium) provides the output of a basal ganglia pathway that is necessary for vocal learning in juvenile songbirds. The shell subregion of LMAN (LMAN_shell) gives rise to recurrent loops that may subserve specific learning-related functions. We found that lesions in the LMAN_shell pathway caused no immediate disruption of vocal behavior, but prevented the development of stable vocal sequences and the ability to imitate vocal sounds. The shell subregion of nucleus LMAN is an output for the basal ganglia in song birds. The authors report that lesions of the this region do not immediately disrupt vocal behavior but do prevent the development of stable vocal sequences and the ability to imitate vocal sounds.
