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Theta (\u223c3-12 Hz) rhythms play a key role by phase-locking to recurring acoustic features marking syllable boundaries. Reliable synchronization to quasi-rhythmic inputs, whose variable frequency can dip below cortical theta frequencies (down to \u223c1 Hz), requires \u201cflexible\u201d theta oscillators whose underlying neuronal mechanisms remain unknown. Using biophysical computational models, we found that the flexibility of phase-locking in neural oscillators depended on the types of hyperpolarizing currents that paced them. Simulated cortical theta oscillators flexibly phase-locked to slow inputs when these inputs caused both (i) spiking and (ii) the subsequent buildup of outward current sufficient to delay further spiking until the next input. The greatest flexibility in phase-locking arose from a synergistic interaction between intrinsic currents that was not replicated by synaptic currents at similar timescales. Flexibility in phase-locking enabled improved entrainment to speech input, optimal at mid-vocalic channels, which in turn supported syllabic-timescale segmentation through identification of vocalic nuclei. Our results suggest that synaptic and intrinsic inhibition contribute to frequency-restricted and -flexible phase-locking in neural oscillators, respectively. Their differential deployment may enable neural oscillators to play diverse roles, from reliable internal clocking to adaptive segmentation of quasi-regular sensory inputs like speech.<\/jats:p>","DOI":"10.1371\/journal.pcbi.1008783","type":"journal-article","created":{"date-parts":[[2021,4,14]],"date-time":"2021-04-14T19:05:10Z","timestamp":1618427110000},"page":"e1008783","update-policy":"https:\/\/doi.org\/10.1371\/journal.pcbi.corrections_policy","source":"Crossref","is-referenced-by-count":16,"title":["Differential contributions of synaptic and intrinsic inhibitory currents to speech segmentation via flexible phase-locking in neural oscillators"],"prefix":"10.1371","volume":"17","author":[{"ORCID":"https:\/\/orcid.org\/0000-0002-6798-7191","authenticated-orcid":true,"given":"Benjamin 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Shamir","year":"2009","journal-title":"PLoS computational biology"},{"key":"pcbi.1008783.ref045","doi-asserted-by":"crossref","DOI":"10.7554\/eLife.06213","article-title":"Speech encoding by coupled cortical theta and gamma oscillations","volume":"4","author":"A Hyafil","year":"2015","journal-title":"Elife"},{"key":"pcbi.1008783.ref046","doi-asserted-by":"crossref","first-page":"130","DOI":"10.1016\/j.cognition.2017.11.003","article-title":"Pre-linguistic segmentation of speech into syllable-like units","volume":"171","author":"O R\u00e4s\u00e4nen","year":"2018","journal-title":"Cognition"},{"issue":"1","key":"pcbi.1008783.ref047","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1038\/s41467-020-16956-5","article-title":"Combining predictive coding and neural oscillations enables online syllable recognition in natural speech","volume":"11","author":"S Hovsepyan","year":"2020","journal-title":"Nature 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Adams","year":"2019","journal-title":"Journal of neurophysiology"},{"key":"pcbi.1008783.ref063","doi-asserted-by":"crossref","unstructured":"Garofolo JS, Lamel LF, Fisher WM, Fiscus JG, Pallett DS. DARPA TIMIT acoustic-phonetic continous speech corpus CD-ROM. NIST speech disc 1-1.1. 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Neural entrainment determines the words we hear. 2017;.","DOI":"10.1101\/175000"},{"key":"pcbi.1008783.ref087","unstructured":"Brown M, Tanenhaus MK, Dilley L. Syllable inference as a mechanism for spoken language understanding. Topics in Cognitive Science. 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Hasselmo","year":"2004","journal-title":"Progress in brain research"},{"issue":"6","key":"pcbi.1008783.ref115","doi-asserted-by":"crossref","first-page":"710","DOI":"10.1016\/j.conb.2006.09.002","article-title":"The role of acetylcholine in learning and memory","volume":"16","author":"ME Hasselmo","year":"2006","journal-title":"Current opinion in neurobiology"},{"issue":"4","key":"pcbi.1008783.ref116","doi-asserted-by":"crossref","first-page":"339","DOI":"10.1162\/NETN_a_00024","article-title":"Switching between internal and external modes: a multiscale learning principle","volume":"1","author":"CJ Honey","year":"2017","journal-title":"Network Neuroscience"},{"issue":"1","key":"pcbi.1008783.ref117","doi-asserted-by":"crossref","first-page":"324","DOI":"10.1037\/h0076177","article-title":"Relationship between hippocampal theta activity and running speed in the rat","volume":"88","author":"WL McFarland","year":"1975","journal-title":"Journal of comparative and physiological 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Program TSYLB (version 2 revision 1.1); 1996."},{"key":"pcbi.1008783.ref127","unstructured":"Kahn D. Syllable-based generalizations in English. 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