Learning to Encode Timing: Mechanisms of Plasticity in the Auditory Brainstem

Thanos Tzounopoulos, Nina Kraus · Neuron · 2009 · neuroscience · 10.1016/j.neuron.2009.05.002 · record unknown2009-learning-to-encode-timing-mechanisms-of-plas
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Where this record's text came from. faceted 2026-08 by reading the locally held text (xml); every evidence string verified as an exact substring of that text before merging. This is a Neuron review, so 'method' is review and the limitations quoted are limits of the evidence base it surveys rather than of an experiment the authors ran — the article states no limits on its own procedure. Claim type is existence (plasticity does occur in the brainstem); the same sentences also function as a critique of the traditional hard-wired view, but that view is never attributed to a named source, so no contradicts relation was recorded. Only one claim_relation qualifies: the Reverse Hierarchy Theory is named in the text ("A theory to account for the interactions between sensory input and top-down processes (e.g., attention, language, and memory) is the Reverse Hierarchy Theory (RHT) ( Ahissar and Hochstein, 2004 ).") and the authors explicitly propose extending it subcortically. Authors field is empty in the queue metadata and the text below the line does not name them. 2026-08-12 and 2026-08-15 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
This record’s own method is review, and the tags below follow from that. A review’s findings, quantities and statistics are things stated IN this paper, and some of them report results obtained elsewhere. No tag on this page names a different author. Measured across every review in this corpus, 81% of a review’s tags carry no citation at all — most of them are the review speaking. The corpus does not adjudicate the rest: read an unmarked tag as a claim this paper makes, not necessarily one it made first.
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Other fields define these differently the same word, defined another way in another discipline
plasticity 4 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 23 concept tags on this paper, 1 are grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.
auditory brainstem plasticity intrinsic plasticity spike-timing-dependent plasticity Hebbian anti-Hebbian long-term potentiation long-term depression dorsal cochlear nucleus fusiform cartwheel cells medial nucleus of the trapezoid body auditory brainstem response frequency following response phase locking pitch tracking interaural time differences Reverse Hierarchy Theory corticofugal tinnitus Kv3.1b negative image musicians Synapse-specific plasticity simultaneously disinhibits fusiform cells and strengthens their excitatory input, driving the fusiform hyperactivity that underlies tinnitus Anti-Hebbian STDP reweights parallel-fiber synapses so that inputs predicting incoming sensory activity are weakened, building a 'negative image' that cancels the sensory consequences of the animal's own movements Nitric oxide diffusing out of MNTB principal neurons suppresses postsynaptic Kv3 channels in neighbouring cells, providing intercellular gain control during loud sound Corticofugal (efferent) feedback from higher-order processing levels descends onto brainstem nuclei and is the route by which language and musical experience alter subcortical timing calyx of Held superior olivary complex cochlear nucleus nitric oxide spike-timing-dependent plasticity (STDP) auditory brainstem response (ABR) frequency following response (FFR) Reverse Hierarchy Theory (RHT) relative timing of presynaptic and postsynaptic action potentials sound intensity standard error of the mean fast Fourier transform review

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second readingalternativesan independent re-reading of a facet, recorded whether or not it agrees
relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
conceptconceptsan idea the paper works with, quoted where the paper uses it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
contributioncontributionwhat the paper claims to add
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
object of studyobject_of_studywhat the paper is about — the thing it investigates
phenomenonphenomenonthe effect or regularity under study
research questionresearch_questionthe question the paper sets itself
sensesensesa distinct use of a term, quoted where the paper fixes it
transfertransfersa method or concept carried in from another field

The paper

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Every tag on this page

alternatives 3 · claim_relations 7 · claim_type 1 · concept_edges 6 · concepts 1 · content_tags 48 · contribution 1 · limitations 9 · mechanisms 15 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 6 · transfers 5

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facettagfieldconf.whose words
object_of_studywhether the auditory brainstem, long treated as a hard-wired relay optimised for timing, is a site of experience-dependent plasticity
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object_of_studythe paper's words
research_questiondoes the mammalian auditory brainstem express the cellular mechanisms that could support the learning effects seen in humans
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research_questionthe paper's words
contributionassembles slice physiology and human evoked-potential evidence to overturn the view that early auditory processing is non-plastic, and identifies cell-specific synaptic and intrinsic plasticity in brainstem nuclei
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contributionthe paper's words
claim_typeexistence
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claim_typethe paper's words
phenomenonauditory brainstem plasticity
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phenomenonthe paper's words
methodreview
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also in: review
methodthe paper's words
mechanismsmechanisms
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mechanisms[0]the tagger's words
mechanismsmechanisms
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mechanisms[1]the tagger's words
mechanismsmechanisms
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mechanisms[2]the tagger's words
mechanismsmechanisms
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mechanisms[3]the tagger's words
mechanismsthe field inferred incapacity from function - a structure optimised for fast reliable signalling was assumed to be ill-suited to plasticity, and the test was not run
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mechanisms[4]1.0the tagger's words
mechanismsand effort was directed to the cortex instead, which is where the plasticity literature accumulated
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mechanisms[5]1.0the tagger's words
mechanismsthe reversal is built from two independent lines that meet in the middle - slice physiology in animals and non-invasive evoked potentials in humans
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mechanisms[6]1.0the tagger's words
mechanismsadding an in-vivo-like feature to the standard slice preparation reversed a standing conclusion about the same nucleus
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mechanisms[7]1.0the tagger's words
mechanismsand the authors generalise the warning rather than confining it to that result
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mechanisms[8]1.0the tagger's words
mechanismsplasticity is shown to be feature-specific rather than a gain change, which is what makes it evidence of learning rather than of arousal
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mechanisms[9]1.0the tagger's words
mechanismsthe functional interpretation of the dorsal cochlear nucleus is imported wholesale from the electrosensory lobe of a fish, where anti-Hebbian plasticity cancels self-generated input
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mechanisms[10]1.0the tagger's words
mechanismsthe argument that the same job exists in audition is made by matching the problem, not the anatomy - both systems must separate external signals from self-generated ones
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mechanisms[11]1.0the tagger's words
mechanismshuman timescales are stacked deliberately: lifelong language and musical experience, then training over weeks, then attention within a session
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mechanisms[12]0.95the tagger's words
mechanismsthe same plasticity mechanisms are used to explain a pathology, which converts the review's claim into something testable in tinnitus
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mechanisms[13]0.95the tagger's words
mechanismsa theory built for visual cortex is extended below the cortex, and the extension is flagged as the authors' view rather than an established result
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mechanisms[14]0.95the tagger's words
limitationsThe synaptic-plasticity evidence base comes overwhelmingly from brain slices, which differ substantially from the intact brain.
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what this limits: every plasticity claim in the review: MOST STUDIES OF SYNAPTIC PLASTICITY IN THE AUDITORY BRAINSTEM HAVE BEEN PERFORMED IN BRAIN SLICES
limitations[0]the tagger's words
limitationsHow naturally occurring noise exposure would induce the plasticity proposed to underlie tinnitus is not established.
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what this limits: the tinnitus account specifically: it is obvious that ELECTRICAL STIMULATION OF PARALLEL FIBERS CAN INDUCE SYNAPTIC PLASTICITY, but how naturally occurring noise exposure would do so is not established
limitations[1]the tagger's words
limitationsHow brainstem pathways adapt to ongoing sensory activity and contribute to perception of environmental sound remains largely unknown.
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what this limits: what the brainstem literature covers: MUCH LESS IS KNOWN ABOUT HOW THESE PATHWAYS ADAPT TO ONGOING SENSORY ACTIVITY AND HOW THEY CONTRIBUTE TO THE PERCEPTION AND INTERPRETATION OF ENVIRONMENTAL SOUNDS, INCLUDING SPEECH
limitations[2]the tagger's words
limitationsmost of the plasticity evidence comes from slices, which differ from the intact brain in ways that have already changed one conclusion
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what this limits: the plasticity evidence base as a whole: Quotation withheld THIS EXPERIMENTAL APPROACH HAS SUBSTANTIAL DIFFERENCES FROM STUDYING INTACT BRAINS IN VIVO
limitations[3]1.0the tagger's words
limitationshow noise exposure would induce the plasticity proposed to cause tinnitus is not established
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duplicate of: limitations[1] · what this limits: the same scope as limitations[1] - how naturally occurring noise exposure would induce the plasticity proposed to underlie tinnitus is not established
limitations[4]1.0the tagger's words
limitationsthe pathophysiology of tinnitus remains poorly understood despite its prevalence
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what this limits: any mechanistic claim about the disorder: DESPITE THE WIDE PREVALENCE OF TINNITUS, THE PATHOPHYSIOLOGY OF THE DISORDER IS POORLY UNDERSTOOD
limitations[5]1.0the tagger's words
limitationswhether the fish-derived cancellation function applies to the dorsal cochlear nucleus is posed as an open question, not answered
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what this limits: the transfer of the cancellation function from the fish circuit: A KEY QUESTION IS WHETHER ANTI-HEBBIAN PLASTICITY OBSERVED IN THE DCN CAN SERVE A SIMILAR FUNCTION
limitations[6]1.0the tagger's words
limitationswhether brainstem plasticity is driven top-down, locally, or by both is unresolved
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what this limits: the locus of control for brainstem plasticity: whether changes are INITIATED/MODULATED IN A TOP-DOWN FASHION ... OR THROUGH LOCAL MECHANISMS OF ADAPTATION is unresolved
limitations[7]1.0the tagger's words
limitationshow these pathways adapt and contribute to perception is much less understood than how they transmit timing
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duplicate of: limitations[2] · what this limits: the same scope as limitations[2] - transmission of timing is understood far better than adaptation or perception
limitations[8]1.0the tagger's words
transfersThe Reverse Hierarchy Theory, first formulated to describe top-down influence of higher visual cortical areas on lower ones, is taken over to explain top-down modulation of auditory processing and extended below cortex to the brainstem.
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transfers[0]the tagger's words
transfersThe adaptive-sensory-processor / 'negative image' account of anti-Hebbian STDP in the cerebellum-like electrosensory lobe is imported as a candidate functional model for the mammalian dorsal cochlear nucleus.
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transfers[1]the tagger's words
transfersanti-Hebbian STDP building a negative image that cancels the sensory consequences of the animal's own movements, imported as the candidate function of the dorsal cochlear nucleus
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transfers[2]1.0the tagger's words
transfersthe Reverse Hierarchy Theory, built for higher-to-lower visual cortical areas and extended here to corticofugal control of the brainstem
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transfers[3]0.95the tagger's words
transfersthe parallel-fiber and climbing-fiber arrangement, used both as the structural analogy for the DCN and as the model for how auditory nerve activity might induce plasticity
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transfers[4]0.95the tagger's words
conceptsauditory brainstem response
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concepts[12]0.85the paper's words
sensesSTDP - Hebbian at fusiform cells and anti-Hebbian at cartwheel cells; ONE induction protocol producing opposite signs, decided by cell type
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senses[0]1.0the tagger's words
sensesthe auditory brainstem - as a hard-wired relay optimised for timing versus as a plastic, experience-modified structure; the paper is the transition between the two readings
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senses[1]1.0the tagger's words
sensesplasticity - synaptic (changed connection strength) versus intrinsic (changed excitability through ion channels); the review insists a circuit needs one or the other
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senses[2]1.0the tagger's words
sensesexperience effects - as a generalised gain change versus as feature-specific alteration; only the second counts as learning
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senses[3]1.0the tagger's words
sensesthe MNTB - as a simple faithful relay versus as a nucleus that fails under realistic activity; the reading depends on what the slice preparation includes
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senses[4]1.0the tagger's words
sensesbrainstem plasticity - as adaptive learning versus as the cause of a disease; the same mechanism read with opposite valence in tinnitus
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senses[5]0.95the tagger's words
alternativesclaim_type: critique
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a second reading DIFFERING from the recorded value. Recorded: existence — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.claim_typethe tagger's words
alternativesmethod: narrative review pairing rodent brain-slice recordings of synaptic and intrinsic plasticity with non-invasive human auditory brainstem responses
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a second reading DIFFERING from the recorded value. Recorded: review — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.methodthe tagger's words
alternativesphenomenon: experience-dependent plasticity in subcortical auditory processing
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a second reading DIFFERING from the recorded value. Recorded: auditory brainstem plasticity — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsextends → Reverse Hierarchy Theory (Ahissar and Hochstein, 2004)
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claim_relations[0]the tagger's words
claim_relationscontradicts → the traditional view that early sensory processing stages are hard-wired and non-plastic
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claim_relations[1]1.0the tagger's words
claim_relationscritiques → the inference from specialisation for precise timing to incapacity for plasticity
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claim_relations[2]1.0the tagger's words
claim_relationscontradicts → the treatment of the MNTB as a simple and faithful relay nucleus
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claim_relations[3]1.0the tagger's words
claim_relationsbuilds-on → Krishnan and colleagues, who first showed that language experience alters brainstem pitch encoding
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claim_relations[4]1.0the tagger's words
claim_relationsextends → the Reverse Hierarchy Theory, from intracortical feedback to subcortical processing
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claim_relations[5]0.95the tagger's words
claim_relationssupports → the account of tinnitus as centrally maintained rather than purely cochlear
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claim_relations[6]0.9the tagger's words
concept_edgesHebbian —contrasted-with→ anti-Hebbian
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strength of the relation: 1
also in: Hebbian · anti-Hebbian
concept_edges[0]the tagger's words
concept_edgesfusiform —contrasted-with→ cartwheel cells
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strength of the relation: 1
concept_edges[1]the tagger's words
concept_edgesRETRACTED (WRONG TYPE: a KIND, not a PART) · intrinsic plasticity —is-a→ plasticity
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RETRACTED: WRONG TYPE: a KIND, not a PART. Intrinsic plasticity is a sort of plasticity, not a component of it. — strength of the relation: 0.95
concept_edges[2]the tagger's words
concept_edgesfrequency following response —part-of→ auditory brainstem response
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'ABR'; endpoint abbreviated.
concept_edges[3]the tagger's words
concept_edgesplasticity —causes→ tinnitus
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strength of the relation: 0.75 Audit 2026-09-28: DOUBTFUL - read against its span and kept, but plasticity-driven disinhibition is linked to tinnitus; causes is loose.
also in: plasticity · tinnitus
concept_edges[4]the tagger's words
concept_edgesdorsal cochlear nucleus —part-of→ auditory brainstem
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strength of the relation: 0.95
concept_edges[7]the tagger's words
content_tagsstandard error of the mean
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content_tags.statistics[0]the tagger's words
content_tagsfast Fourier transform
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content_tags.statistics[1]0.75the tagger's words
content_tagsphase locking
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also in: phase locking
content_tags.statistics[2]0.95the tagger's words
content_tagsContrary to the traditional view that early sensory stages are not plastic, the auditory brainstem is dynamic.
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content_tags.findings[0]the tagger's words
content_tagsIn the dorsal cochlear nucleus one protocol produces LTP in fusiform cells and LTD in cartwheel cells.
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content_tags.findings[1]the tagger's words
content_tagsLanguage experience reshapes brainstem coding: speakers of tonal languages show enhanced neural representation of pitch.
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content_tags.findings[2]the tagger's words
content_tagsExperience-driven changes in the brainstem are selective to particular stimulus features rather than an across-the-board amplification of all responses.
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content_tags.findings[3]the tagger's words
content_tagsAcoustic environment tunes intrinsic excitability: in quiet conditions PKC phosphorylation of Kv3.1b potassium channels lowers the potassium current so that neurons fire only at low frequencies.
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content_tags.findings[4]the tagger's words
content_tagsThe medial nucleus of the trapezoid body is not a faithful relay: under in-vivo-like spontaneous activity it fails to entrain reliably to prolonged high-frequency stimuli because of short-term plasticity, even though it still encodes stimulus onset precisely.
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content_tags.findings[5]the tagger's words
content_tagsThe authors argue that the Reverse Hierarchy Theory, formulated for cortex, extends to subcortical processing and may itself initiate or modulate brainstem plasticity.
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content_tags.findings[6]the tagger's words
content_tagsone induction protocol produces LTP in fusiform cells and LTD in cartwheel cells
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content_tags.findings[7]1.0the tagger's words
content_tagsspeakers of tonal languages have enhanced neural representation of pitch in the brainstem
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content_tags.findings[8]1.0the tagger's words
content_tagsmusical experience produces more robust brainstem encoding of speech, linguistic pitch patterns and vocal emotion
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content_tags.findings[9]1.0the tagger's words
content_tagsbrainstem encoding is disrupted in subsets of poor readers and children with autism
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content_tags.findings[10]1.0the tagger's words
content_tagsbrainstem responses are affected by attention and are larger to a speech syllable than to its time-reversed version
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content_tags.findings[11]1.0the tagger's words
content_tagsexperience does not produce a generalised gain change but alters specific stimulus features
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content_tags.findings[12]1.0the tagger's words
content_tagsbrainstem processing can also be changed by auditory training over weeks, not only by lifelong experience
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content_tags.findings[13]1.0the tagger's words
content_tagsadding in-vivo-like spontaneous activity to MNTB slices revealed synaptic failures at high frequency that earlier studies had not seen
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content_tags.findings[14]1.0the tagger's words
content_tagsso the MNTB is not a simple faithful relay and may not entrain reliably to sustained high-frequency input
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content_tags.findings[15]1.0the tagger's words
content_tagsphosphorylation state of Kv3.1b channels sets whether neurons fire at low or high frequency, tracking the acoustic environment
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content_tags.findings[16]1.0the tagger's words
content_tagsactivity in one MNTB neuron can modulate the excitability of its neighbours, a candidate gain reduction in loud noise
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content_tags.findings[17]1.0the tagger's words
content_tagsretrograde GABA signalling in the lateral superior olive adjusts the excitation-inhibition balance underlying sound localisation
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content_tags.findings[18]1.0the tagger's words
content_tagssynapse-specific plasticity that disinhibits and excites fusiform cells can produce hyperactivity like that seen in animals with behavioural signs of tinnitus
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content_tags.findings[19]0.85the tagger's words
content_tagscortical activation shapes the tuning of cochlear nucleus neurons, giving a route for top-down control
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content_tags.findings[20]1.0the tagger's words
content_tagsintrinsic plasticity
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content_tags.definitions[0]the tagger's words
content_tagsspike-timing-dependent plasticity (STDP)
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content_tags.definitions[1]the tagger's words
content_tagsauditory brainstem response (ABR)
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content_tags.definitions[2]the tagger's words
content_tagsfrequency following response (FFR)
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content_tags.definitions[3]the tagger's words
content_tagstinnitus
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also in: tinnitus
content_tags.definitions[4]the tagger's words
content_tagspitch tracking
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content_tags.definitions[5]the tagger's words
content_tagsReverse Hierarchy Theory (RHT)
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content_tags.definitions[6]the tagger's words
content_tagsauditory brainstem response
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content_tags.definitions[7]1.0the tagger's words
content_tagsfrequency following response
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content_tags.definitions[8]1.0the tagger's words
content_tagsspike-timing-dependent plasticity
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content_tags.definitions[9]1.0the tagger's words
content_tagsintrinsic plasticity
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content_tags.definitions[10]1.0the tagger's words
content_tagsTinnitus
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also in: Tinnitus
content_tags.definitions[11]1.0the tagger's words
content_tagsReverse Hierarchy Theory
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content_tags.definitions[12]1.0the tagger's words
content_tags<4000 Hz — Upper frequency limit of the 'low-frequency' sounds that the binaural timing circuitry of the mammalian brainstem is specialised to localise
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content_tags.quantities[0]the tagger's words
content_tags4–12 µm — Diameter of the myelinated globular-bushy-cell axons giving rise to the calyx of Held, large enough to give fast conduction velocity
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content_tags.quantities[1]the tagger's words
content_tags200 µs — Delay with which inhibitory input reaches the cat lateral superior olive after the excitatory input
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content_tags.quantities[2]the tagger's words
content_tags40 ms time bins, 1 ms interval — Window length and step of the running short-term Fourier analysis used to extract pitch contours from the brainstem response
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content_tags.quantities[3]the tagger's words
content_tags0.5 ms — Scale of the x-axis tick marks in Figure 1, indicating that the experience-related timing disruptions and enhancements are of the order of tenths of a millisecond
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content_tags.quantities[4]the tagger's words
content_tags<4000 Hz — the low-frequency range localised by interaural timing in mammals
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content_tags.quantities[5]0.85the tagger's words
content_tagsmicrosecond range — the interaural phase differences the brainstem resolves
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content_tags.quantities[6]1.0the tagger's words
content_tagslast 10 years — the period over which brainstem timing mechanisms were worked out
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content_tags.quantities[7]0.8the tagger's words
content_tagsrelative timing of presynaptic and postsynaptic action potentials
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content_tags.variables[0]1.0the tagger's words
content_tagssound intensity
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content_tags.variables[1]0.9the tagger's words
content_tagspitch tracking
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content_tags.variables[2]0.8the tagger's words

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