{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,2,27]],"date-time":"2026-02-27T06:17:52Z","timestamp":1772173072538,"version":"3.50.1"},"update-to":[{"DOI":"10.1371\/journal.pcbi.1009707","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,1,10]],"date-time":"2022-01-10T00:00:00Z","timestamp":1641772800000}}],"reference-count":87,"publisher":"Public Library of Science (PLoS)","issue":"12","license":[{"start":{"date-parts":[[2021,12,28]],"date-time":"2021-12-28T00:00:00Z","timestamp":1640649600000},"content-version":"vor","delay-in-days":0,"URL":"http:\/\/creativecommons.org\/licenses\/by\/4.0\/"}],"funder":[{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP17H06380"],"award-info":[{"award-number":["JP17H06380"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"publisher","award":["JP19KT0023"],"award-info":[{"award-number":["JP19KT0023"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"publisher"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["JP21H03781"],"award-info":[{"award-number":["JP21H03781"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100001691","name":"Japan Society for the Promotion of Science","doi-asserted-by":"crossref","award":["JP21K17805"],"award-info":[{"award-number":["JP21K17805"]}],"id":[{"id":"10.13039\/501100001691","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100002241","name":"Japan Science and Technology Agency","doi-asserted-by":"crossref","award":["JPMJCR17A4"],"award-info":[{"award-number":["JPMJCR17A4"]}],"id":[{"id":"10.13039\/501100002241","id-type":"DOI","asserted-by":"crossref"}]},{"DOI":"10.13039\/501100002241","name":"Japan Science and Technology Agency","doi-asserted-by":"crossref","award":["ACT-X 21454934 (JPMJAX21AN)"],"award-info":[{"award-number":["ACT-X 21454934 (JPMJAX21AN)"]}],"id":[{"id":"10.13039\/501100002241","id-type":"DOI","asserted-by":"crossref"}]},{"name":"Mitsubishi Foundation Research Grants in the Natural Sciences","award":["202111014"],"award-info":[{"award-number":["202111014"]}]}],"content-domain":{"domain":["www.ploscompbiol.org"],"crossmark-restriction":false},"short-container-title":["PLoS Comput Biol"],"abstract":"<jats:p>Context dependency is a key feature in sequential structures of human language, which requires reference between words far apart in the produced sequence. Assessing how long the past context has an effect on the current status provides crucial information to understand the mechanism for complex sequential behaviors. Birdsongs serve as a representative model for studying the context dependency in sequential signals produced by non-human animals, while previous reports were upper-bounded by methodological limitations. Here, we newly estimated the context dependency in birdsongs in a more scalable way using a modern neural-network-based language model whose accessible context length is sufficiently long. The detected context dependency was beyond the order of traditional Markovian models of birdsong, but was consistent with previous experimental investigations. We also studied the relation between the assumed\/auto-detected vocabulary size of birdsong (i.e., fine- vs. coarse-grained syllable classifications) and the context dependency. It turned out that the larger vocabulary (or the more fine-grained classification) is assumed, the shorter context dependency is detected.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1009707","type":"journal-article","created":{"date-parts":[[2021,12,28]],"date-time":"2021-12-28T14:50:28Z","timestamp":1640703028000},"page":"e1009707","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":10,"title":["Measuring context dependency in birdsong using artificial neural networks"],"prefix":"10.1371","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-3900-5410","authenticated-orcid":true,"given":"Takashi","family":"Morita","sequence":"first","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0927-3473","authenticated-orcid":true,"given":"Hiroki","family":"Koda","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-7439-9566","authenticated-orcid":true,"given":"Kazuo","family":"Okanoya","sequence":"additional","affiliation":[]},{"ORCID":"https:\/\/orcid.org\/0000-0002-4766-4504","authenticated-orcid":true,"given":"Ryosuke O.","family":"Tachibana","sequence":"additional","affiliation":[]}],"member":"340","published-online":{"date-parts":[[2021,12,28]]},"reference":[{"issue":"9","key":"pcbi.1009707.ref001","doi-asserted-by":"crossref","first-page":"1325","DOI":"10.1016\/j.neunet.2003.06.005","article-title":"Learning and inference in the brain","volume":"16","author":"K Friston","year":"2003","journal-title":"Neural Networks"},{"key":"pcbi.1009707.ref002","doi-asserted-by":"crossref","first-page":"127","DOI":"10.1038\/nrn2787","article-title":"The free-energy principle: a unified brain theory?","volume":"11","author":"K Friston","year":"2010","journal-title":"Nature Reviews Neuroscience"},{"key":"pcbi.1009707.ref003","doi-asserted-by":"crossref","DOI":"10.1515\/9783112316009","volume-title":"Syntactic Structures","author":"N Chomsky","year":"1957"},{"key":"pcbi.1009707.ref004","article-title":"Long Distance Dependencies","author":"B Larson","year":"2017","journal-title":"Oxford Bibliographies"},{"key":"pcbi.1009707.ref005","doi-asserted-by":"crossref","unstructured":"Khandelwal U, He H, Qi P, Jurafsky D. Sharp Nearby, Fuzzy Far Away: How Neural Language Models Use Context. In: Proceedings of the 56th Annual Meeting of the Association for Computational Linguistics (Volume 1: Long Papers). Melbourne, Australia: Association for Computational Linguistics; 2018. p. 284\u2013294.","DOI":"10.18653\/v1\/P18-1027"},{"key":"pcbi.1009707.ref006","doi-asserted-by":"crossref","unstructured":"Dai Z, Yang Z, Yang Y, Carbonell J, Le Q, Salakhutdinov R. Transformer-XL: Attentive Language Models beyond a Fixed-Length Context. In: Proceedings of the 57th Annual Meeting of the Association for Computational Linguistics. Florence, Italy: Association for Computational Linguistics; 2019. p. 2978\u20132988.","DOI":"10.18653\/v1\/P19-1285"},{"issue":"10","key":"pcbi.1009707.ref007","doi-asserted-by":"crossref","first-page":"2091","DOI":"10.1097\/00001756-200007140-00007","article-title":"Lesion of a higher-order song nucleus disrupts phrase level complexity in Bengalese finches","volume":"11","author":"T Hosino","year":"2000","journal-title":"Neuroreport"},{"key":"pcbi.1009707.ref008","doi-asserted-by":"crossref","first-page":"297","DOI":"10.1016\/S0065-3454(04)34008-8","article-title":"Song syntax in Bengalese finches: proximate and ultimate analyses","volume":"34","author":"K Okanoya","year":"2004","journal-title":"Advances in the Study of Behavior"},{"issue":"9","key":"pcbi.1009707.ref009","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1371\/journal.pone.0024516","article-title":"Complex Sequencing Rules of Birdsong Can be Explained by Simple Hidden Markov Processes","volume":"6","author":"K Katahira","year":"2011","journal-title":"PLOS ONE"},{"issue":"44","key":"pcbi.1009707.ref010","doi-asserted-by":"crossref","first-page":"15414","DOI":"10.1523\/JNEUROSCI.1254-12.2012","article-title":"Variable sequencing is actively maintained in a well learned motor skill","volume":"32","author":"TL Warren","year":"2012","journal-title":"Journal of neuroscience"},{"issue":"5","key":"pcbi.1009707.ref011","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1371\/journal.pcbi.1003052","article-title":"Long-range Order in Canary Song","volume":"9","author":"JE Markowitz","year":"2013","journal-title":"PLOS Computational Biology"},{"issue":"3","key":"pcbi.1009707.ref012","doi-asserted-by":"crossref","first-page":"113","DOI":"10.1016\/j.tics.2011.01.002","article-title":"Songs to syntax: the linguistics of birdsong","volume":"15","author":"RC Berwick","year":"2011","journal-title":"Trends in Cognitive Science"},{"issue":"3","key":"pcbi.1009707.ref013","doi-asserted-by":"crossref","first-page":"364","DOI":"10.1093\/brain\/81.3.364","article-title":"Corticobulbar connexions to the pons and lower brain-stem in man: an anatomical study","volume":"81","author":"HGJM Kuypers","year":"1958","journal-title":"Brain"},{"issue":"3","key":"pcbi.1009707.ref014","doi-asserted-by":"crossref","first-page":"392","DOI":"10.1002\/(SICI)1096-9861(19970120)377:3<392::AID-CNE7>3.0.CO;2-Y","article-title":"Projections of the dorsomedial nucleus of the intercollicular complex (DM) in relation to respiratory-vocal nuclei in the brainstem of pigeon (Columba livia) and zebra finch (Taeniopygia guttata)","volume":"377","author":"JM Wild","year":"1997","journal-title":"Journal of Comparative Neurology"},{"issue":"7176","key":"pcbi.1009707.ref015","doi-asserted-by":"crossref","first-page":"305","DOI":"10.1038\/nature06492","article-title":"Precise auditory\u2013vocal mirroring in neurons for learned vocal communication","volume":"451","author":"JF Prather","year":"2008","journal-title":"Nature"},{"issue":"3","key":"pcbi.1009707.ref016","first-page":"1","article-title":"A Compact Statistical Model of the Song Syntax in Bengalese Finch","volume":"7","author":"DZ Jin","year":"2011","journal-title":"PLOS Computational Biology"},{"issue":"1792","key":"pcbi.1009707.ref017","article-title":"Animal vocal sequences: not the Markov chains we thought they were","volume":"281","author":"A Kershenbaum","year":"2014","journal-title":"Proceedings of the Royal Society of London B: Biological Sciences"},{"issue":"3636","key":"pcbi.1009707.ref018","article-title":"Parallels in the sequential organization of birdsong and human speech","volume":"10","author":"T Sainburg","year":"2019","journal-title":"Nature Communications"},{"key":"pcbi.1009707.ref019","first-page":"5998","volume-title":"Advances in Neural Information Processing Systems 30","author":"A Vaswani","year":"2017"},{"key":"pcbi.1009707.ref020","unstructured":"van den Oord A, Kalchbrenner N, Kavukcuoglu K. Pixel Recurrent Neural Networks; 2016. Available from: https:\/\/arxiv.org\/abs\/1601.06759."},{"key":"pcbi.1009707.ref021","unstructured":"van den Oord A, Dieleman S, Zen H, Simonyan K, Vinyals O, Graves A, et al. WaveNet: A Generative Model for Raw Audio; 2016. Available from: https:\/\/arxiv.org\/abs\/1609.03499."},{"key":"pcbi.1009707.ref022","unstructured":"Dhariwal P, Jun H, Payne C, Kim JW, Radford A, Sutskever I. Jukebox: A Generative Model for Music; 2020. Available from: https:\/\/arxiv.org\/abs\/2005.00341."},{"issue":"2","key":"pcbi.1009707.ref023","doi-asserted-by":"crossref","first-page":"271","DOI":"10.1016\/j.conb.2007.03.011","article-title":"Language evolution and an emergent property","volume":"17","author":"K Okanoya","year":"2007","journal-title":"Current Opinion in Neurobiology"},{"key":"pcbi.1009707.ref024","unstructured":"Kingma DP, Welling M. Auto-Encoding Variational Bayes; 2014. The International Conference on Learning Representations (ICLR) 2014."},{"issue":"5","key":"pcbi.1009707.ref025","doi-asserted-by":"crossref","first-page":"859","DOI":"10.1038\/s41386-018-0303-6","article-title":"DeepSqueak: a deep learning-based system for detection and analysis of ultrasonic vocalizations","volume":"44","author":"KR Coffey","year":"2019","journal-title":"Neuropsychopharmacology"},{"key":"pcbi.1009707.ref026","article-title":"Inferring low-dimensional latent descriptions of animal vocalizations","author":"J Goffinet","year":"2019","journal-title":"bioRxiv"},{"key":"pcbi.1009707.ref027","doi-asserted-by":"crossref","unstructured":"Morita T, Koda H. Exploring TTS without T Using Biologically\/Psychologically Motivated Neural Network Modules (ZeroSpeech 2020). In: Proceedings of Interspeech 2020; 2020. p. 4856\u20134860.","DOI":"10.21437\/Interspeech.2020-3127"},{"key":"pcbi.1009707.ref028","first-page":"6306","volume-title":"Advances in Neural Information Processing Systems 30","author":"A van den Oord","year":"2017"},{"issue":"12","key":"pcbi.1009707.ref029","doi-asserted-by":"crossref","first-page":"2041","DOI":"10.1109\/TASLP.2019.2938863","article-title":"Unsupervised Speech Representation Learning Using WaveNet Autoencoders","volume":"27","author":"J Chorowski","year":"2019","journal-title":"IEEE\/ACM Transactions on Audio, Speech, and Language Processing"},{"issue":"3","key":"pcbi.1009707.ref030","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1371\/journal.pone.0092584","article-title":"Semi-Automatic Classification of Birdsong Elements Using a Linear Support Vector Machine","volume":"9","author":"RO Tachibana","year":"2014","journal-title":"PLOS ONE"},{"issue":"12","key":"pcbi.1009707.ref031","doi-asserted-by":"crossref","first-page":"1157","DOI":"10.1007\/s00359-015-1046-z","article-title":"Variability in the temporal parameters in the song of the Bengalese finch (Lonchura striata var. domestica)","volume":"201","author":"RO Tachibana","year":"2015","journal-title":"Journal of Comparative Physiology A"},{"key":"pcbi.1009707.ref032","volume-title":"Information science and statistics","author":"CM Bishop","year":"2006"},{"issue":"10","key":"pcbi.1009707.ref033","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1371\/journal.pcbi.1008228","article-title":"Finding, visualizing, and quantifying latent structure across diverse animal vocal repertoires","volume":"16","author":"T Sainburg","year":"2020","journal-title":"PLOS Computational Biology"},{"issue":"1","key":"pcbi.1009707.ref034","doi-asserted-by":"crossref","first-page":"37","DOI":"10.1177\/001316446002000104","article-title":"A Coefficient of Agreement for Nominal Scales","volume":"20","author":"J Cohen","year":"1960","journal-title":"Educational and Psychological Measurement"},{"key":"pcbi.1009707.ref035","unstructured":"Roseberg A, Hirschberg J. V-Measure: A conditional entropy-based external cluster evaluation measure. In: Proceedings of the 2007 Joint Conference on Empirical Methods in Natural Language Processing and Computational Natural Language Learning (EMNLP-CoNLL). Association for Computational Linguistics; 2007. p. 410\u2013420."},{"key":"pcbi.1009707.ref036","volume-title":"Studies in cognition","author":"JR Anderson","year":"1990"},{"issue":"4","key":"pcbi.1009707.ref037","doi-asserted-by":"crossref","first-page":"751","DOI":"10.1037\/a0034245","article-title":"A Role for the Developing Lexicon in Phonetic Category Acquisition","volume":"120","author":"NH Feldman","year":"2013","journal-title":"Psychological Review"},{"issue":"1","key":"pcbi.1009707.ref038","doi-asserted-by":"crossref","first-page":"159","DOI":"10.2307\/2529310","article-title":"The Measurement of Observer Agreement for Categorical Data","volume":"33","author":"JR Landis","year":"1977","journal-title":"Biometrics"},{"key":"pcbi.1009707.ref039","first-page":"237","volume-title":"A Field Guide to Dynamical Recurrent Networks","author":"S Hochreiter","year":"2001"},{"key":"pcbi.1009707.ref040","doi-asserted-by":"crossref","unstructured":"Abnar S, Zuidema W. Quantifying Attention Flow in Transformers. In: Proceedings of the 58th Annual Meeting of the Association for Computational Linguistics. Online: Association for Computational Linguistics; 2020. p. 4190\u20134197.","DOI":"10.18653\/v1\/2020.acl-main.385"},{"issue":"48","key":"pcbi.1009707.ref041","doi-asserted-by":"crossref","first-page":"30046","DOI":"10.1073\/pnas.1907367117","article-title":"Emergent linguistic structure in artificial neural networks trained by self-supervision","volume":"117","author":"CD Manning","year":"2020","journal-title":"Proceedings of the National Academy of Sciences"},{"issue":"8","key":"pcbi.1009707.ref042","doi-asserted-by":"crossref","first-page":"2176","DOI":"10.1523\/JNEUROSCI.2383-16.2017","article-title":"Dynamic Encoding of Acoustic Features in Neural Responses to Continuous Speech","volume":"37","author":"B Khalighinejad","year":"2017","journal-title":"Journal of Neuroscience"},{"issue":"3997","key":"pcbi.1009707.ref043","doi-asserted-by":"crossref","first-page":"585","DOI":"10.1126\/science.173.3997.585","article-title":"Songs of Humpback Whales","volume":"173","author":"RS Payne","year":"1971","journal-title":"Science"},{"issue":"4471","key":"pcbi.1009707.ref044","doi-asserted-by":"crossref","first-page":"801","DOI":"10.1126\/science.7433999","article-title":"Monkey responses to three different alarm calls: evidence of predator classification and semantic communication","volume":"210","author":"R Seyfarth","year":"1980","journal-title":"Science"},{"issue":"1","key":"pcbi.1009707.ref045","doi-asserted-by":"crossref","first-page":"13","DOI":"10.1111\/brv.12160","article-title":"Acoustic sequences in non-human animals: a tutorial review and prospectus","volume":"91","author":"A Kershenbaum","year":"2016","journal-title":"Biological Reviews"},{"issue":"1","key":"pcbi.1009707.ref046","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1006\/anbe.1998.0923","article-title":"Pitfalls in the categorization of behaviour: a comparison of dolphin whistle classification methods","volume":"57","author":"VM Janik","year":"1999","journal-title":"Animal Behaviour"},{"key":"pcbi.1009707.ref047","unstructured":"Jang E, Gu S, Poole B. Categorical Reparameterization with Gumbel-Softmax. In: Proceedings of the 5th International Conference on Learning Representations (ICLR); 2017."},{"key":"pcbi.1009707.ref048","unstructured":"Louizos C, Swersky K, Li Y, Welling M, Zemel RS. The Variational Fair Autoencoder. In: Bengio Y, LeCun Y, editors. Proceedings of the 4th International Conference on Learning Representations (ICLR); 2016."},{"issue":"59","key":"pcbi.1009707.ref049","first-page":"1","article-title":"Domain-Adversarial Training of Neural Networks","volume":"17","author":"Y Ganin","year":"2016","journal-title":"Journal of Machine Learning Research"},{"issue":"8","key":"pcbi.1009707.ref050","doi-asserted-by":"crossref","first-page":"1735","DOI":"10.1162\/neco.1997.9.8.1735","article-title":"Long short-term memory","volume":"9","author":"S Hochreiter","year":"1997","journal-title":"Neural Computation"},{"issue":"11","key":"pcbi.1009707.ref051","doi-asserted-by":"crossref","first-page":"2531","DOI":"10.1162\/089976602760407955","article-title":"Real-Time Computing Without Stable States: A New Framework for Neural Computation Based on Perturbations","volume":"14","author":"W Maass","year":"2002","journal-title":"Neural Computation"},{"issue":"5667","key":"pcbi.1009707.ref052","doi-asserted-by":"crossref","first-page":"78","DOI":"10.1126\/science.1091277","article-title":"Harnessing Nonlinearity: Predicting Chaotic Systems and Saving Energy in Wireless Communication","volume":"304","author":"H Jaeger","year":"2004","journal-title":"Science"},{"issue":"9","key":"pcbi.1009707.ref053","doi-asserted-by":"crossref","first-page":"2727","DOI":"10.1073\/pnas.84.9.2727","article-title":"Neural networks that learn temporal sequences by selection","volume":"84","author":"S Dehaene","year":"1987","journal-title":"Proceedings of the National Academy of Sciences"},{"key":"pcbi.1009707.ref054","doi-asserted-by":"crossref","first-page":"402","DOI":"10.1109\/TASLP.2019.2956145","article-title":"Neural Source-Filter Waveform Models for Statistical Parametric Speech Synthesis","volume":"28","author":"X Wang","year":"2020","journal-title":"IEEE\/ACM Transactions on Audio, Speech, and Language Processing"},{"key":"pcbi.1009707.ref055","article-title":"Latent space visualization, characterization, and generation of diverse vocal communication signals","author":"T Sainburg","year":"2019","journal-title":"bioRxiv"},{"issue":"1","key":"pcbi.1009707.ref056","doi-asserted-by":"crossref","first-page":"3","DOI":"10.1037\/0735-7036.104.1.3","article-title":"Experimental determination of a unit of song production in the zebra finch (Taeniopygia guttata)","volume":"104","author":"J Cynx","year":"1990","journal-title":"Journal of Comparative Psychology"},{"issue":"4","key":"pcbi.1009707.ref057","doi-asserted-by":"crossref","first-page":"745","DOI":"10.1016\/S0003-3472(05)80386-0","article-title":"Models for song learning in the zebra finch: fathers or others?","volume":"39","author":"H Williams","year":"1990","journal-title":"Animal Behaviour"},{"issue":"3","key":"pcbi.1009707.ref058","doi-asserted-by":"crossref","first-page":"278","DOI":"10.1037\/0735-7036.106.3.278","article-title":"Syllable chunking in zebra finch (Taeniopygia guttata) song","volume":"106","author":"H Williams","year":"1992","journal-title":"Journal of Comparative Psychology"},{"key":"pcbi.1009707.ref059","doi-asserted-by":"crossref","first-page":"980","DOI":"10.3389\/fpsyg.2016.00980","article-title":"Zebra Finch Song Phonology and Syntactical Structure across Populations and Continents\u2014A Computational Comparison","volume":"7","author":"RF Lachlan","year":"2016","journal-title":"Frontiers in Psychology"},{"key":"pcbi.1009707.ref060","doi-asserted-by":"crossref","unstructured":"Chiu CC, Sainath TN, Wu Y, Prabhavalkar R, Nguyen P, Chen Z, et al. State-of-the-Art Speech Recognition with Sequence-to-Sequence Models. In: 2018 IEEE International Conference on Acoustics, Speech and Signal Processing (ICASSP); 2018. p. 4774\u20134778.","DOI":"10.1109\/ICASSP.2018.8462105"},{"key":"pcbi.1009707.ref061","doi-asserted-by":"crossref","unstructured":"Dunbar E, Cao XN, Benjumea J, Karadayi J, Bernard M, Besacier L, et al. The Zero Resource Speech Challenge 2017. In: 2017 IEEE Automatic Speech Recognition and Understanding Workshop (ASRU); 2017. p. 323\u2013330.","DOI":"10.1109\/ASRU.2017.8268953"},{"key":"pcbi.1009707.ref062","doi-asserted-by":"crossref","unstructured":"van Niekerk B, Nortje L, Kamper H. Vector-quantized neural networks for acoustic unit discovery in the ZeroSpeech 2020 challenge. In: Proceedings of Interspeech 2020; 2020. p. 4836\u20134840.","DOI":"10.21437\/Interspeech.2020-1693"},{"key":"pcbi.1009707.ref063","unstructured":"Baevski A, Hsu WN, Conneau A, Auli M. Unsupervised Speech Recognition. In: Advances in Neural Information Processing Systems 34 (NeurIPS 2021); 2021. Available from: https:\/\/papers.nips.cc\/paper\/2021\/hash\/ea159dc9788ffac311592613b7f71fbb-Abstract.html."},{"issue":"3","key":"pcbi.1009707.ref064","doi-asserted-by":"crossref","first-page":"400","DOI":"10.1109\/TASSP.1987.1165125","article-title":"Estimation of Probabilities from Sparse Data for the Language Model Component of a Speech Recognizer","volume":"35","author":"SM Katz","year":"1987","journal-title":"IEEE Transactions on Acoustics, Speech, and Signal Processing"},{"key":"pcbi.1009707.ref065","doi-asserted-by":"crossref","unstructured":"Kneser R, Ney H. Improved Backing-off for N-gram Language Modeling. In: Proceedings of the IEEE International Conference on Acoustics, Speech and Signal. vol. 1; 1995. p. 181\u2013184.","DOI":"10.1109\/ICASSP.1995.479394"},{"issue":"45","key":"pcbi.1009707.ref066","doi-asserted-by":"crossref","first-page":"17710","DOI":"10.1523\/JNEUROSCI.2181-13.2013","article-title":"Neural Encoding and Integration of Learned Probabilistic Sequences in Avian Sensory-Motor Circuitry","volume":"33","author":"KE Bouchard","year":"2013","journal-title":"Journal of Neuroscience"},{"issue":"39","key":"pcbi.1009707.ref067","doi-asserted-by":"crossref","first-page":"12936","DOI":"10.1523\/JNEUROSCI.2690-10.2010","article-title":"Linked Control of Syllable Sequence and Phonology in Birdsong","volume":"30","author":"MJ Wohlgemuth","year":"2010","journal-title":"Journal of Neuroscience"},{"issue":"2","key":"pcbi.1009707.ref068","doi-asserted-by":"crossref","first-page":"192069","DOI":"10.1098\/rsos.192069","article-title":"Difficulties in analysing animal song under formal language theory framework: comparison with metric-based model evaluation","volume":"7","author":"T Morita","year":"2020","journal-title":"Royal Society Open Science"},{"issue":"2","key":"pcbi.1009707.ref069","doi-asserted-by":"crossref","first-page":"257","DOI":"10.1109\/5.18626","article-title":"A tutorial on hidden Markov models and selected applications in speech recognition","volume":"77","author":"LR Rabiner","year":"1989","journal-title":"Proceedings of the IEEE"},{"issue":"3","key":"pcbi.1009707.ref070","doi-asserted-by":"crossref","first-page":"306","DOI":"10.1016\/j.cognition.2010.11.001","article-title":"The learnability of abstract syntactic principles","volume":"118","author":"A Perfors","year":"2011","journal-title":"Cognition"},{"issue":"7","key":"pcbi.1009707.ref071","doi-asserted-by":"crossref","first-page":"190139","DOI":"10.1098\/rsos.190139","article-title":"Superregular grammars do not provide additional explanatory power but allow for a compact analysis of animal song","volume":"6","author":"T Morita","year":"2019","journal-title":"Royal Society Open Science"},{"issue":"27","key":"pcbi.1009707.ref072","doi-asserted-by":"crossref","first-page":"10023","DOI":"10.1523\/JNEUROSCI.1606-11.2011","article-title":"Neural Coding of Syntactic Structure in Learned Vocalizations in the Songbird","volume":"31","author":"H Fujimoto","year":"2011","journal-title":"Journal of Neuroscience"},{"issue":"12","key":"pcbi.1009707.ref073","doi-asserted-by":"crossref","first-page":"3273","DOI":"10.1111\/j.1460-9568.2008.06291.x","article-title":"Population coding of song element sequence in the Bengalese finch HVC","volume":"27","author":"J Nishikawa","year":"2008","journal-title":"European Journal of Neuroscience"},{"issue":"1","key":"pcbi.1009707.ref074","doi-asserted-by":"crossref","first-page":"95","DOI":"10.2326\/osj.5.95","article-title":"Dynamical neural representation of song syntax in Bengalese Finch: a model study","volume":"5","author":"J Nishikawa","year":"2006","journal-title":"Ornithological Science"},{"issue":"4","key":"pcbi.1009707.ref075","doi-asserted-by":"crossref","first-page":"303","DOI":"10.1007\/BF02551274","article-title":"Approximation by superpositions of a sigmoidal function","volume":"2","author":"G Cybenko","year":"1989","journal-title":"Mathematics of Control, Signals and Systems"},{"key":"pcbi.1009707.ref076","doi-asserted-by":"crossref","unstructured":"Jin L, Gupta MM, Nikiforuk PN. Universal approximation using dynamic recurrent neural networks: discrete-time version. In: Proceedings of ICNN\u201995\u2014International Conference on Neural Networks. vol. 1; 1995. p. 403\u2013408.","DOI":"10.1109\/ICNN.1995.488134"},{"key":"pcbi.1009707.ref077","volume-title":"Bird Families of the World","author":"CB Frith","year":"1998"},{"issue":"4","key":"pcbi.1009707.ref078","first-page":"967","article-title":"Courtship Ethology of Carola\u2019s Parotia (Parotia Carolae)","volume":"123","author":"EI Scholes","year":"2006","journal-title":"The Auk"},{"key":"pcbi.1009707.ref079","doi-asserted-by":"crossref","first-page":"161","DOI":"10.1016\/S0066-1856(68)80003-2","article-title":"The Behaviour of Free-living Chimpanzees in the Gombe Stream Reserve","volume":"1","author":"J van Lawick-Goodall","year":"1968","journal-title":"Animal Behaviour Monographs"},{"issue":"1","key":"pcbi.1009707.ref080","doi-asserted-by":"crossref","first-page":"162","DOI":"10.1086\/204484","article-title":"Representation of Action Through Iconic Gesture in a Captive Lowland Gorilla","volume":"37","author":"JE Tanner","year":"1996","journal-title":"Current Anthropology"},{"issue":"3","key":"pcbi.1009707.ref081","doi-asserted-by":"crossref","first-page":"878","DOI":"10.1523\/JNEUROSCI.1804-14.2015","article-title":"Audition-Independent Vocal Crystallization Associated with Intrinsic Developmental Gene Expression Dynamics","volume":"35","author":"C Mori","year":"2015","journal-title":"Journal of Neuroscience"},{"issue":"2","key":"pcbi.1009707.ref082","doi-asserted-by":"crossref","first-page":"1728","DOI":"10.1111\/ejn.14057","article-title":"Singing activity-driven Arc expression associated with vocal acoustic plasticity in juvenile songbird","volume":"48","author":"S Hayase","year":"2018","journal-title":"European Journal of Neuroscience"},{"issue":"2","key":"pcbi.1009707.ref083","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1109\/TAC.1959.1104847","article-title":"On adaptive control processes","volume":"4","author":"R Bellman","year":"1959","journal-title":"IRE Transactions on Automatic Control"},{"issue":"8","key":"pcbi.1009707.ref084","first-page":"707","article-title":"Binary Codes Capable of Correcting Deletions, Insertions and Reversals","volume":"10","author":"VI Levenshtein","year":"1966","journal-title":"Soviet Physics Doklady"},{"key":"pcbi.1009707.ref085","doi-asserted-by":"crossref","unstructured":"Bowman SR, Vilnis L, Vinyals O, Dai A, Jozefowicz R, Bengio S. Generating Sentences from a Continuous Space. In: Proceedings of The 20th SIGNLL Conference on Computational Natural Language Learning; 2016. p. 10\u201321.","DOI":"10.18653\/v1\/K16-1002"},{"key":"pcbi.1009707.ref086","doi-asserted-by":"crossref","unstructured":"Zhao T, Zhao R, Eskenazi M. Learning Discourse-level Diversity for Neural Dialog Models using Conditional Variational Autoencoders. In: Proceedings of the 55th Annual Meeting of the Association for Computational Linguistics (Volume 1: Long Papers). Association for Computational Linguistics; 2017. p. 654\u2013664.","DOI":"10.18653\/v1\/P17-1061"},{"key":"pcbi.1009707.ref087","first-page":"2825","article-title":"Scikit-learn: Machine Learning in Python","volume":"12","author":"F Pedregosa","year":"2011","journal-title":"Journal of Machine Learning Research"}],"updated-by":[{"DOI":"10.1371\/journal.pcbi.1009707","type":"new_version","label":"New version","source":"publisher","updated":{"date-parts":[[2022,1,10]],"date-time":"2022-01-10T00:00:00Z","timestamp":1641772800000}}],"container-title":["PLOS Computational Biology"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1009707","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2024,9,15]],"date-time":"2024-09-15T04:41:58Z","timestamp":1726375318000},"score":1,"resource":{"primary":{"URL":"https:\/\/dx.plos.org\/10.1371\/journal.pcbi.1009707"}},"subtitle":[],"editor":[{"given":"Fr\u00e9d\u00e9ric E.","family":"Theunissen","sequence":"first","affiliation":[]}],"short-title":[],"issued":{"date-parts":[[2021,12,28]]},"references-count":87,"journal-issue":{"issue":"12","published-online":{"date-parts":[[2021,12,28]]}},"URL":"https:\/\/doi.org\/10.1371\/journal.pcbi.1009707","relation":{"has-preprint":[{"id-type":"doi","id":"10.1101\/2020.05.09.083907","asserted-by":"object"}]},"ISSN":["1553-7358"],"issn-type":[{"value":"1553-7358","type":"electronic"}],"subject":[],"published":{"date-parts":[[2021,12,28]]}}}