{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,10,20]],"date-time":"2026-10-20T03:45:01Z","timestamp":1792467901374,"version":"4.5.0"},"reference-count":50,"publisher":"American Physiological Society","issue":"3","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":["Journal of Neurophysiology"],"published-print":{"date-parts":[[2006,9]]},"abstract":"Trial-by-trial variability is important in feedback-based motor learning. Variation in motor output enables evaluation mechanisms to differentially reinforce patterns of motor activity that produce desired behaviors. Here, we studied neural substrates of variability in the performance of adult birdsong, a complex, learned motor skill used for courtship. Song performance is more variable when male birds sing alone (undirected) than when they sing to females (directed). We test the role of the anterior forebrain pathway (AFP), an avian basal ganglia\u2013forebrain circuit, in this socially driven modulation of song variability. We show that lesions of the lateral magnocellular nucleus of the anterior nidopallium (LMAN), the output nucleus of the AFP, cause a reduction in the moment-by-moment variability in syllable structure during undirected song to the level present during directed song. This elimination of song modulation is immediate and long-lasting. We further show that the degree of syllable variability and its modulation are both attenuated in older birds, in concert with decreased variability of LMAN activity in these birds. In contrast to the requirement of LMAN for social modulation of syllable structure, we find that LMAN is not required for modulation of other features of song, such as the number of introductory elements and motif repetitions and the ordering of syllables or for other motor and motivational aspects of courtship. Our findings suggest that a key function of avian basal ganglia circuitry is to regulate vocal performance and plasticity by specifically modulating moment-by-moment variability in the structure of individual song elements.<\/jats:p>","DOI":"10.1152\/jn.01138.2005","type":"journal-article","created":{"date-parts":[[2006,5,24]],"date-time":"2006-05-24T20:44:41Z","timestamp":1148503481000},"page":"1441-1455","source":"Crossref","is-referenced-by-count":244,"title":["Lesions of an Avian Basal Ganglia Circuit Prevent Context-Dependent Changes to Song Variability"],"prefix":"10.1152","volume":"96","author":[{"given":"Mimi H.","family":"Kao","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Michael S.","family":"Brainard","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"24","reference":[{"key":"R1","doi-asserted-by":"publisher","DOI":"10.1073\/pnas.91.26.12413"},{"key":"R2","doi-asserted-by":"publisher","DOI":"10.1007\/BF01939388"},{"key":"R3","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1668-05.2005"},{"key":"R4","unstructured":"Bottjer SW\n .\n Developmental regulation of basal ganglia circuitry during the sensitive period for vocal learning in songbirds. In:\n Behavioral Neurobiology of Birdsong\n , edited by Zeigler HP and Marler P. New York: New York Academy of Sciences, 2004, p. 395\u2013415."},{"key":"R5","doi-asserted-by":"publisher","DOI":"10.1126\/science.6719123"},{"key":"R6","doi-asserted-by":"publisher","DOI":"10.1159\/000113342"},{"key":"R7","unstructured":"Brainard MS\n .\n Contributions of the anterior forebrain pathway to vocal plasticity. In:\n Behavioral Neurobiology of Birdsong\n , edited by Zeigler HP and Marler P. New York: New York Academy of Sciences, 2004, p. 377\u2013394."},{"key":"R8","doi-asserted-by":"publisher","DOI":"10.1038\/35008083"},{"key":"R9","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.21-07-02501.2001"},{"key":"R10","doi-asserted-by":"crossref","unstructured":"Cooper BG\n and Goller F.\n Physiological insights into the social-context-dependent changes in the rhythm of the song motor program.\n J Neurophysiol\n 95: 3798\u20133809, 2006.","DOI":"10.1152\/jn.01123.2005"},{"key":"R11","doi-asserted-by":"publisher","DOI":"10.1037\/0735-7036.104.1.3"},{"key":"R12","doi-asserted-by":"publisher","DOI":"10.1006\/anbe.2001.1744"},{"key":"R13","doi-asserted-by":"publisher","DOI":"10.1121\/1.1458024"},{"key":"R14","doi-asserted-by":"publisher","DOI":"10.1146\/annurev.neuro.22.1.567"},{"key":"R15","unstructured":"Doya K\n and Sejnowski TJ.\n A computational model of avian song learning. In:\n The New Cognitive Neurosciences\n , edited by Gazzaniga MS. Cambridge, MA: MIT Press, 2000, p. 469\u2013482."},{"key":"R16","unstructured":"Dunn AM\n and Zann RA.\n Undirected song in wild finch flocks: context and effects of mate removal.\n Ethology\n 102\n : 529\u2013539, 1996."},{"key":"R17","doi-asserted-by":"publisher","DOI":"10.1002\/1097-4695(20010205)46:2<142::AID-NEU60>3.0.CO;2-R"},{"key":"R18","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.3387-05.2006"},{"key":"R19","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.19-23-10461.1999"},{"key":"R20","doi-asserted-by":"publisher","DOI":"10.1038\/6306"},{"key":"R21","unstructured":"Immelmann K\n .\n Song development in zebra finch and other Estrildid finches. In:\n Bird Vocalisations\n , edited by Hinde RA. London: Cambridge, 1969, p. 61\u201374."},{"key":"R22","doi-asserted-by":"publisher","DOI":"10.1016\/S0896-6273(00)80594-2"},{"key":"R23","doi-asserted-by":"crossref","unstructured":"Johnson F\n and Bottjer SW.\n Afferent influences on cell death and birth during development of a cortical nucleus necessary for learned vocal behavior in zebra finches.\n Development\n 120\n : 13\u201324, 1994.","DOI":"10.1242\/dev.120.1.13"},{"key":"R24","doi-asserted-by":"publisher","DOI":"10.1038\/nature03127"},{"key":"R25","doi-asserted-by":"publisher","DOI":"10.1002\/neu.20109"},{"key":"R26","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.19-21-09385.1999"},{"key":"R27","doi-asserted-by":"publisher","DOI":"10.1038\/20933"},{"key":"R28","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.20-13-05054.2000"},{"key":"R29","doi-asserted-by":"publisher","DOI":"10.1007\/BF02476084"},{"key":"R30","doi-asserted-by":"publisher","DOI":"10.1002\/neu.480240805"},{"key":"R31","doi-asserted-by":"publisher","DOI":"10.1038\/334149a0"},{"key":"R32","doi-asserted-by":"publisher","DOI":"10.1016\/0163-1047(92)90757-U"},{"key":"R33","doi-asserted-by":"publisher","DOI":"10.1016\/0163-1047(93)91215-9"},{"key":"R34","doi-asserted-by":"publisher","DOI":"10.1002\/cne.901650405"},{"key":"R35","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pbio.0030153"},{"key":"R36","unstructured":"Perkel DJ\n .\n Origin of the anterior forebrain pathway. In:\n Behavioral Neurobiology of Birdsong\n , edited by Zeigler HP and Marler P. New York: New York Academy of Sciences, 2004, p. 736\u2013748."},{"key":"R36A","doi-asserted-by":"publisher","DOI":"10.1002\/cne.20118"},{"key":"R37","doi-asserted-by":"publisher","DOI":"10.1016\/S0959-4388(00)00155-0"},{"key":"R38","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.11-09-02896.1991"},{"key":"R39","doi-asserted-by":"publisher","DOI":"10.1111\/j.1439-0310.1980.tb01044.x"},{"key":"R40","unstructured":"Sutton RS\n and Barto AG.\n Reinforcement Learning: An Introduction.\n Cambridge, MA: MIT Press, 1998."},{"key":"R41","doi-asserted-by":"publisher","DOI":"10.1126\/science.1058522"},{"key":"R42","doi-asserted-by":"publisher","DOI":"10.1152\/jn.2000.84.3.1224"},{"key":"R43","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.14-11-06924.1994"},{"key":"R44","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.18-21-09088.1998"},{"key":"R45","doi-asserted-by":"crossref","unstructured":"Williams H\n .\n Choreography of song, dance and beak movements in the zebra finch (\n Taeniopygia guttata\n ).\n J Exp Biol\n 204\n : 3497\u20133506, 2001.","DOI":"10.1242\/jeb.204.20.3497"},{"key":"R46","doi-asserted-by":"publisher","DOI":"10.1002\/(SICI)1097-4695(199904)39:1<14::AID-NEU2>3.0.CO;2-X"},{"key":"R47","doi-asserted-by":"publisher","DOI":"10.1126\/science.273.5283.1871"},{"key":"R48","doi-asserted-by":"crossref","unstructured":"Zann RA\n .\n The Zebra Finch: A Synthesis of Field and Laboratory Studies.\n Oxford, UK: Oxford, 1996.","DOI":"10.1093\/oso\/9780198540793.001.0001"},{"key":"R49","doi-asserted-by":"publisher","DOI":"10.1523\/JNEUROSCI.1891-04.2004"}],"container-title":["Journal of Neurophysiology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.physiology.org\/doi\/pdf\/10.1152\/jn.01138.2005","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,2,4]],"date-time":"2024-02-04T09:31:21Z","timestamp":1707039081000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.physiology.org\/doi\/10.1152\/jn.01138.2005"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2006,9]]},"references-count":50,"journal-issue":{"issue":"3","published-print":{"date-parts":[[2006,9]]}},"alternative-id":["10.1152\/jn.01138.2005"],"URL":"https:\/\/doi.org\/10.1152\/jn.01138.2005","relation":{},"ISSN":["0022-3077","1522-1598"],"issn-type":[{"value":"0022-3077","type":"print"},{"value":"1522-1598","type":"electronic"}],"subject":[],"published":{"date-parts":[[2006,9]]}}}