A Hierarchy of Time Scales for Discriminating and Classifying the Temporal Shape of Sound in Three Auditory Cortical Fields

Osman, Lee, Escabí, Read · The Journal of Neuroscience: The Official Journal of the Society for Neuroscience · 2018 · neuroscience · 10.1523/jneurosci.2871-17.2018 · record Osman2018-a-hierarchy-of-time-scales-for-discriminatin
tag layer · index · concepts · authors · עברית
Where this record's text came from. faceted 2026-08 by reading the locally held text (html); every evidence string verified as an exact substring of that text before merging. Method is single-unit electrophysiology in anaesthetised rats combined with offline decoding (a normalised spike-distance discrimination index and a naive Bayes classifier); I tagged the data-generating method as 'electrophysiology' since the classifiers are analyses of those recordings, not simulations of a model system. Claim type is 'existence' because the contribution is that a graded hierarchy of time scales is present across fields, not a causal mechanism for it. The paper does not describe its anaesthetised, non-behaving preparation as a limitation, so that is not recorded. 2026-08-13, 2026-08-14 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.
How to read this page. The article itself is not reproduced here — this paper is not cleared for redistribution, so what follows is the tag layer alone. The evidence sentences are withheld with the article. The table at the foot lists every tag the record carries; each row names the facet it belongs to, and the whose words column says whether the tag records the paper's own wording or the tagger's sentence about the paper. Every tag links to each other paper in the corpus carrying it. The machine-readable edition of the same thing is the TEI file beside this one, where the tags live in standOff, and the article is at its publisher via the link below. When we have changed our mind. Some entries in the table begin with a word in capitals and the reason for it. EVIDENCE CORRECTED — the finding still stands, but we had quoted the wrong sentence from the paper, and the quotation has been replaced with the right one. RETRACTED — the entry itself was wrong; it stays on the page, marked, with the reason, rather than being deleted. WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below. Each mark carries its own reason, in brackets, because the reasons differ from one entry to the next. An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason. A line reading Audit and a date is a later re-reading of the entry against its whole sentence: DOUBTFUL means it was kept, and the reason says what is loose about it. Quotation marks inside the tagger's sentence. Where a tag is the tagger's own sentence, a phrase in quotation marks inside it is borrowed from the paper — the tagger's wording around it is not. The paper's own sentence is the evidence quote for that row; on this page it is withheld with the article, and the article is at its publisher. Nothing inside quotation marks anywhere on this page is ours: if it is quoted, it is the paper's. Whose words a concept carries. A concept entry records a term the paper itself uses, so the term, the sense given for it, and its evidence are the paper's own words, quoted where the paper fixes them. This is the opposite of a sense entry in the tag table, which is the tagger's sentence about how the paper uses a term. Where a concept entry could not be grounded in the paper's words, the entry says so on its own row.
Other fields define these differently the same word, defined another way in another discipline
population 3 fields · cross-validation 3 fields · anesthesia 2 fields · spike 2 fields · bootstrap 2 fields · discrimination 2 fields · and 3 more →

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 29 concept tags on this paper, 2 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.
A1 VAF cSRAF cortical field onset sustained spike spike rate spike timing discrimination classification classifier time constant time scale response duration sound envelope shape slope duration hierarchy pooling population neuron imprecision noise Poisson spectral anesthesia Longer sustained responses in ventral non-primary cortex require a downstream reader to integrate over a longer time constant, which is what produces the hierarchy of optimal time scales Pooling across neurons averages out the timing jitter of individual spike trains, restoring classification accuracy in temporally imprecise fields Field-specific thalamocortical pathways impose a time-frequency trade-off, buying A1 its temporal resolution at the cost of spectral resolution and vice versa for non-primary fields shape temporal cues encoding time period histogram response delay discrimination index onset spikes sustained spikes sound envelope shape exponential time constant number of neurons Pairwise sound discrimination indices percentage correct classification interaural level difference noise carrier ANOVA two-way ANOVA t-test Student's t test multiple-comparison correction Post hoc Student's t tests with Bonferroni correction Wilcoxon rank-sum test Wilcoxon rank-sum chi-squared test bootstrap cross-validation cross validation approach regression log-linear regression slopes standard error of the mean mean (SEM) effect size / sensitivity index sensitivity index or discrimination index D′ electrophysiology

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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
controlcontrolsan experiment run to eliminate a rival explanation
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
unrecordable relationunhoused_relationsa relation the paper asserts that this vocabulary cannot express, with the reason

The paper

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1523/jneurosci.2871-17.2018. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 10 · claim_type 1 · concept_edges 7 · concepts 2 · content_tags 77 · contribution 1 · controls 5 · limitations 7 · mechanisms 6 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 21 · transfers 6 · unhoused_relations 2

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 7 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studywhether primary and non-primary auditory cortex encode the temporal shape of the sound envelope on different time scales
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
object_of_studythe paper's words
research_questionwhether a functional hierarchy of the kind established in vision for receptive field size exists in audition along the time axis instead
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research_questionthe paper's words
contributionresponse duration and the integration time constant that best discriminates sound shape both increase in rank order A1 < VAF < cSRAF, so the non-primary fields carry the same cue on a longer time scale
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contributionthe paper's words
claim_typeexistence
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claim_typethe paper's words
phenomenontemporal coding of sound envelope shape across auditory cortical fields
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[0]the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[1]the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[2]the tagger's words
mechanismslonger response durations in the ventral fields give a downstream reader more time to integrate, so the time constant that best discriminates sound shape is longer there
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mechanisms[3]1.0the tagger's words
mechanismsonset spikes carry most of the discrimination in every field, and what is left after they are removed is what separates the fields - the sustained spikes work better in VAF and cSRAF
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mechanisms[4]1.0the tagger's words
mechanismsand pooling compensates for imprecision - performance rises 2 to 3 fold over the average single neuron, with half maximal performance reached from fewer than 10 neurons
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mechanisms[5]1.0the tagger's words
limitationswe CANNOT DISAMBIGUATE potential neural codes for sound burst slope versus duration AS THESE CUES COVARY IN OUR SOUNDS
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what this limits: any code attributed to slope or to duration separately: WE CANNOT DISAMBIGUATE POTENTIAL NEURAL CODES FOR SOUND BURST SLOPE VERSUS DURATION AS THESE CUES COVARY IN OUR SOUNDS
limitations[0]the tagger's words
limitationswe HOLD SPECTRAL AND SPATIAL CUES CONSTANT to examine neural coding of temporal cues in the sound envelope independently
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what this limits: the scope of the temporal-coding claim: WE HOLD SPECTRAL AND SPATIAL CUES CONSTANT TO EXAMINE NEURAL CODING OF TEMPORAL CUES IN THE SOUND ENVELOPE INDEPENDENTLY
limitations[1]the tagger's words
limitationsTWO CANDIDATE CODES THAT THIS DESIGN CANNOT SEPARATE, SAID OUT LOUD: we CANNOT DISAMBIGUATE potential neural codes for sound burst slope versus duration AS THESE CUES COVARY IN OUR SOUNDS
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
what this limits: distinguishing a code for the slope of the envelope from a code for the duration of the sound burst
limitations[2]1.0the tagger's words
limitationsthe spike-distance metric is biased at long time constants, and the fix is SYNTHETIC DATA - a periodic spike train with matched temporal statistics, extended to 100 s, standing in for the recording
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what this limits: the spike-distance results at long time constants: AS tau APPROACHES THE TOTAL LENGTH OF THE SPIKE TRAIN (2 s), THE SPIKE DISTANCE IS BIASED AND YIELDS LARGE VALUES
limitations[3]1.0the tagger's words
limitationsthe classifier assumes response bins from different neurons are INDEPENDENT, which the paper states as a design property rather than as a fact about the brain
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what this limits: the population classification figures: BY DESIGN, THE NAIVE BAYES CLASSIFIER TREATS RESPONSES BINS FROM DIFFERENT NEURONS AS IF THEY ARE INDEPENDENT
limitations[4]1.0the tagger's words
limitationsthe first 500 ms of every spike train is discarded, so every number here is about the ADAPTED STEADY-STATE segment and not about the response to a sound that has just begun
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what this limits: every number in the paper, which describes the adapted response and not the onset: FOR ALL DATA ANALYSES, THE FIRST 500 ms WAS REMOVED FROM EACH SPIKE TRAIN TO RESTRICT ANALYSIS TO THE ADAPTED STEADY-STATE SEGMENTS
limitations[5]1.0the tagger's words
limitationsfuture exploration will be needed to determine whether classification is even more robust when spectral, temporal and spatial features are COMBINED, and WHEN ANIMALS ARE ATTENDING during a behavioral task, AS HAPPENS IN NATURAL SETTINGS
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what this limits: the generality to combined cues: FUTURE EXPLORATION WILL BE NEEDED TO DETERMINE WHETHER NEURONAL CLASSIFICATION OF SOUND'S TEMPORAL SHAPE IS EVEN MORE ROBUST ... WHEN SPECTRAL, TEMPORAL AND SPATIAL FEATURES ARE COMBINED
limitations[6]0.9the tagger's words
transfersa Bernoulli Naive Bayes classifier of McCallum and Nigam 1998, used to READ OUT THE SPIKE TRAINS from individual neurons and CATEGORIZE THE SOUND SHAPE
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transfers[0]the tagger's words
transfersthe sensitivity index d-prime, a measure of behavioural discriminability, is repurposed to normalize a spike-train distance so that neural discrimination is expressed on a behavioural scale
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transfers[1]the tagger's words
transfersthe functional hierarchy of successive stages with progressively larger receptive fields, carried from space into the time axis
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
transfers[2]1.0the tagger's words
transfersan exponential-kernel spike distance, normalized by trial variance so that it reads as a sensitivity index
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transfers[3]1.0the tagger's words
transfersa D-prime of 1 corresponds to about 70% accuracy, WHICH IS TYPICALLY CONSIDERED A CRITICAL LIMIT FOR DISCRIMINATING A SIGNAL BEHAVIORALLY
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transfers[4]1.0the tagger's words
transfersa Bernoulli naive Bayes classifier reading spike trains as bit patterns, with its independence assumption declared and used deliberately
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transfers[5]1.0the tagger's words
controlsthe noise carrier varied across trials and across bursts, unfrozen
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eliminates: spectral cues and repeated fine structure - THIS GUARANTEES THAT, ON AVERAGE, THE ONLY PHYSICAL CUES AVAILABLE TO EACH NEURON ARISE FROM THE TEMPORAL ENVELOPE · outcome (stated by the tagger, no quotation): held
controls[0]1.0the tagger's words
controlsspectral and spatial cues held constant across every condition
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eliminates: frequency and location differences contributing to what is read as temporal coding · outcome (stated by the tagger, no quotation): held
controls[1]1.0the tagger's words
controlseach sound discriminated against itself
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eliminates: a metric that returns a positive value for identical inputs - the diagonal of every matrix is zero · outcome (stated by the tagger, no quotation): held
controls[2]1.0the tagger's words
controlsthe spike count distribution tested against a Poisson model before the rate classifier was built on it
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eliminates: a rate code scored under a distribution the data do not follow · outcome (stated by the tagger, no quotation): held
controls[3]1.0the tagger's words
controlsrecording depth restricted to 400 to 650 micrometers
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eliminates: differences between fields being differences between cortical layers - this is the depth corresponding to layer 4 in each field · outcome (stated by the tagger, no quotation): held
controls[4]1.0the tagger's words
conceptsdiscrimination
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concepts[9]the paper's words
conceptsduration
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: duration
concepts[19]the paper's words
unhoused_relationsslope → duration
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THE TWO CANNOT BE SEPARATED IN THIS STUDY - the DESIGN form of the negative relation / No edge type can express it: all eight assert that a relation HOLDS. A confounding statement bounds what the paper's own results can mean, and it lives where nothing downstream will find it. PRESERVED HERE BECAUSE THE EDGE IT DEFEATED WAS WITHDRAWN. 'slope --distinct-from--> duration' was retracted in an earlier note of ours for asserting what this sentence denies; without this record the finding would have left the corpus with the edge. 'these cues COVARY in our sounds.'
unhoused_relations[0]the tagger's words
unhoused_relationssound classification performance → temporal imprecision
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INVERSE ASSOCIATION, ordered across three cortical fields (A1 > VAF > cSRAF): the better the spike-timing classification, the LESS temporally imprecise the responses. No edge type carries polarity, so recording this as 'causes' loses the sign that is the whole finding.
unhoused_relations[1]the tagger's words
sensesA HIERARCHY OF TIME SCALES: response durations and optimal time constants for discriminating sound envelope shape INCREASE IN RANK ORDER WITH A1 < VAF < cSRAF, so sustained spiking is more prominent and discrimination more robust in NON-PRIMARY cortex
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senses[0]1.0the tagger's words
sensesAND THE RANK ORDER REVERSES WITH THE MEASURE: A1 and VAF have significantly HIGHER mean discrimination than cSRAF across all 45 pairs - CONVERSELY cSRAF is better across 30 pairwise comparisons when the sound is slower than 11 Hz. The same three fields, opposite orders, one figure apart
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senses[1]1.0the tagger's words
sensesand reverses again by task: A1 AND VAF OUTPERFORM cSRAF in 10 alternative forced choice classification, where the ranking A1 > VAF > cSRAF is the INVERSE of the time-scale ranking
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senses[2]1.0the tagger's words
sensesand once more by stimulus, inside the pooling analysis: the effect of pooling was GREATER ON AVERAGE IN A1 than in cSRAF, HOWEVER the benefit was GREATER IN cSRAF for two of the longest duration sound bursts - THE ADVANTAGES OF POOLING VARY WITH THE TIME SCALE OF THE SOUND
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senses[3]1.0the tagger's words
senseswith the inverse relation stated as the finding rather than as a caveat: the ability to classify 10 sound shapes IS Quotation withheld, A1 > VAF > cSRAF - the field that discriminates on the longest time scale classifies worst
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senses[4]1.0the tagger's words
sensesTHE OPERATIONAL DEFINITION OF `ONSET SPIKE` IS A WINDOW THAT DIFFERS EIGHTFOLD BETWEEN THE FIELDS BEING COMPARED: the onset response time window is 29.6 ms for A1 and 180.7 ms for cSRAF at the slowest sound, and a single window is measured PER CONDITION AND PER REGION
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about this quote: A TABLE IS JOINED INTO THE 'evidence' QUOTE: after the table's title ('Onset response time windows vary…') it runs on through the table's column heads and cells ('f c 2 4 6 8 11 16 …'). The numbers this row states are table cells; read the quote as a title, then a table.
senses[5]1.0the tagger's words
sensesand the paper says so plainly: onset response time windows VARY WITH SOUND BURST HALF-DURATION AND SLOPE as determined by the sound shaping filter, and are given FOR EACH SOUND BURST VARIATION AND CORTICAL FIELD
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senses[6]1.0the tagger's words
sensesA TERM REDEFINED MID-PAPER, EXPLICITLY: onset spikes are a MIXTURE of stimulus-evoked and spontaneous activity, a subset is categorised as spontaneous BY RANDOM SAMPLING against a Poisson criterion, and IN THE REMAINDER OF THIS MANUSCRIPT WE USE THE TERM ONSET SPIKES TO DENOTE THE EVOKED ONSET SPIKES ONLY
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senses[7]1.0the tagger's words
sensesSYNTHETIC DATA INSIDE THE MEASUREMENT CHAIN, DECLARED: as the time constant approaches the length of the spike train THE SPIKE DISTANCE IS BIASED, so the spike train was EXTENDED BY SYNTHESIZING a periodic spike train WITH MATCHED TEMPORAL STATISTICS, sampling rate and times from the original, out to 100 s
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senses[8]1.0the tagger's words
sensesTWO CANDIDATE CODES THIS DESIGN CANNOT SEPARATE: we CANNOT DISAMBIGUATE potential neural codes for sound burst slope versus duration AS THESE CUES COVARY IN OUR SOUNDS - and the untestable possibility is still stated, that non-primary cortices are more sensitive to duration
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senses[9]1.0the tagger's words
sensesan imported threshold that gives the index its meaning: a D-prime of 1 corresponds to about 70% accuracy, WHICH IS TYPICALLY CONSIDERED A CRITICAL LIMIT for discriminating a signal behaviorally - a behavioral criterion borrowed to read a number computed from spikes
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senses[10]1.0the tagger's words
sensesan assumption declared and then used as a feature: the naive Bayes classifier TREATS RESPONSE BINS FROM DIFFERENT NEURONS AS IF THEY ARE INDEPENDENT, which ALLOWS THE CLASSIFIER TO TAKE ADVANTAGE OF any independent and nonredundant responses
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senses[11]1.0the tagger's words
sensesand a scope restriction that is easy to miss: THE FIRST 500 ms WAS REMOVED FROM EACH SPIKE TRAIN to restrict analysis to the ADAPTED STEADY-STATE segments
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senses[12]1.0the tagger's words
sensesTHE PARAMETER IS THE FINDING: the exponential time constant DETERMINES THE TEMPORAL RESOLUTION OR INTEGRATION TIME of the metric, large values leave mostly slow fluctuations in spike rate and small values make it sensitive to precise timing - the range tested is what lets one number stand for two rival codes
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senses[13]1.0the tagger's words
sensesSPIKE RATE IS NEAR CHANCE AND SAID TO BE SO: all three fields classify with LOW PERFORMANCE ACCURACY based on spike rate alone, and spike count per 2 s IS NOT AN OPTIMAL NEURAL SIGNAL for coding sound shape
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senses[14]1.0the tagger's words
sensesand pooling does not rescue it - pooled spike-rate classification is CLOSE TO CHANCE LEVELS in all cortical fields, while the timing-based classifier reaches 80 to 100%
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senses[15]1.0the tagger's words
senseswith the number of neurons needed stated per field: HALF MAXIMAL PERFORMANCE from fewer than 10 neurons everywhere, but A1 and VAF require 3 and 5 against 16 for cSRAF
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senses[16]1.0the tagger's words
sensesand a resolution limit read off the point where performance drops - a HALF-RESOLUTION LIMIT ON THE ORDER OF 20 ms
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senses[17]1.0the tagger's words
sensesTHE HIERARCHY IS CLAIMED AS DIRECTIONAL: non-primary cortex COULD CERTAINLY POOL RESPONSES FROM A1 to classify sounds BUT THE REVERSE IS LESS LIKELY, and structures reading cSRAF would need longer integration times than structures reading A1
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senses[18]1.0the tagger's words
sensesA CONCEPT MOVED FROM SPACE TO TIME, AGAIN: functional hierarchies IN THE VISUAL CORTICES support the concept that classifying objects requires successive stages with a PROGRESSIVE INCREASE IN RECEPTIVE FIELD SIZE, and this study supports the idea that A SIMILAR PROGRESSION EXISTS IN AUDITORY CORTICES IN THE TIME DOMAIN
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senses[19]1.0the tagger's words
sensesand the specialisation is framed as a trade-off rather than a ranking: A1 has SUPERIOR TEMPORAL RESOLUTION AT THE EXPENSE OF SPECTRAL RESOLUTION, and the non-primary fields the reverse
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senses[20]1.0the tagger's words
alternativesclaim_type: existence
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a second reading reaching the same value. Recorded: existence — note: a second reading reaching the same value from a different sentence. Not applied - a person decides.
alternatives.claim_typethe tagger's words
alternativesmethod: sweep an exponential smoothing time constant from 1 to 256 ms, take the value that maximises a normalised spike-distance discrimination index, and read the same spike trains with a naive Bayes classifier in a 10-alternative task
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a second reading DIFFERING from the recorded value. Recorded: electrophysiology — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.methodthe tagger's words
alternativesphenomenon: temporally precise spiking that follows the rise, fall and duration of the sound envelope rather than its spectrum
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a second reading DIFFERING from the recorded value. Recorded: temporal coding of sound envelope shape across auditory cortical fields — 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 → Phillips et al., 2002; Heil, 2003; Zheng and Escabí, 2013; Lee et al., 2016
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claim_relations[0]the tagger's words
claim_relationsreplicates → Malone et al., 2015 (A1 response duration versus sound burst duration)
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claim_relations[1]the tagger's words
claim_relationsbuilds-on → Larson et al., 2009 (robustness of pairwise discrimination to response jitter)
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claim_relations[2]the tagger's words
claim_relationsbuilds-on → Van Rossum (2001) spike-distance metric
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claim_relations[3]the tagger's words
claim_relationsextends → prior work showing that response duration tracks sound envelope shape in A1, by measuring the same thing in two non-primary fields
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claim_relations[4]1.0the tagger's words
claim_relationscontradicts → the reading that a longer time scale means better sound coding - the field with the longest time scale classifies worst
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differs in: the task: pairwise discrimination against 10-alternative classification
claim_relations[5]1.0the tagger's words
claim_relationssupports → a spike-timing code over a spike-rate code, since rate classification stays near chance even when pooled
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claim_relations[6]1.0the tagger's words
claim_relationssupports → pooling as compensation for single-neuron imprecision, extended to a field where responses are sluggish and imprecise
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claim_relations[7]1.0the tagger's words
claim_relationspredicts → that brain structures reading cSRAF need longer integration times than structures reading A1, and that pooling runs from A1 to non-primary rather than the reverse
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claim_relations[8]0.9the tagger's words
claim_relationsqualifies → its own hierarchy claim, which holds for time constants and durations but reverses for discrimination and for classification
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differs in: THE AXIS IS WHICH COMPARISON SET, AND THE ORDERING REVERSES BETWEEN THEM. 'A1 and VAF have significantly higher mean discrimination than cSRAF averaged across all 45 pairwise discriminations. Conversely, cSRAF neurons are marginally better at discriminating across 30 pairwise sound comparisons [at low rates].' The same three fields rank one way over one set of comparisons and the other way over a subset - so a hierarchy claim is a claim about the averaging window, and this record's [32] gives the time-constant ordering that does hold.
claim_relations[9]1.0the tagger's words
concept_edgesA1 —contrasted-with→ cSRAF
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
strength of the relation: 1
also in: A1 · cSRAF
concept_edges[1]1.0the tagger's words
concept_edgesA1 —causes→ discrimination
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but A1 and VAF showed HIGHER mean discrimination than cSRAF; causes is loose.
also in: A1 · discrimination
concept_edges[6]1.0the tagger's words
concept_edgesdiscrimination —operationalised-as→ time constant
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strength of the relation: 1 · note: THE PARAMETER IS THE FINDING - large values leave mostly slow fluctuations in spike rate, small values make the metric sensitive to precise timing, and the sweep is what lets one number stand for two rival codes Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but discrimination was computed FOR EACH time constant to find the optimal one; operationalised-as is loose.
concept_edges[17]1.0the tagger's words
concept_edgesnoise —contrasted-with→ envelope
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strength of the relation: 1 · note: the carrier varied unfrozen across trials and bursts, which GUARANTEES THAT ON AVERAGE THE ONLY PHYSICAL CUES AVAILABLE TO EACH NEURON ARISE FROM THE TEMPORAL ENVELOPE Audit 2026-09-28: DOUBTFUL - read against its span and kept, but noise carriers varied while envelopes were the object of study; contrasted-with is loose.
also in: noise · envelope
concept_edges[19]1.0the tagger's words
concept_edgesspectral —contrasted-with→ envelope
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its span and kept, but spectral cues were held constant to study envelope cues; contrasted-with is loose.
also in: spectral · envelope
concept_edges[20]1.0the tagger's words
concept_edgesspike rate —causes→ classification
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strength of the relation: 1 · note: spike count or equivalent spike rate per 2 s IS NOT AN OPTIMAL NEURAL SIGNAL for coding sound shape in primary or non-primary cortex Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but classification BASED ON spike rate performed near chance; causes with no direction is loose.
concept_edges[22]1.0the tagger's words
concept_edgesVAF —contrasted-with→ cSRAF
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differs in: THE AXIS IS THE OPTIMAL TIME CONSTANT, AND THE THREE FIELDS ARE ORDERED ON IT. 'Quotation withheld.' The comparison is made FOR THESE SAME SOUNDS, so the stimulus is held constant and the difference is a property of the fields - and it is an ORDERING, which a binary edge cannot hold: the fact that VAF sits between the other two is the content, and it needs two edges to say. · strength of the relation: 1
also in: VAF · cSRAF
concept_edges[32]1.0the tagger's words
content_tagsANOVA
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also in: ANOVA
content_tags.statistics[0]the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[1]the tagger's words
content_tagsmultiple-comparison correction
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content_tags.statistics[2]the tagger's words
content_tagsWilcoxon rank-sum test
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content_tags.statistics[3]the tagger's words
content_tagschi-squared test
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content_tags.statistics[4]the tagger's words
content_tagsbootstrap
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also in: bootstrap
content_tags.statistics[5]the tagger's words
content_tagscross-validation
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content_tags.statistics[6]the tagger's words
content_tagsregression
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also in: regression
content_tags.statistics[7]the tagger's words
content_tagsstandard error of the mean
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content_tags.statistics[8]the tagger's words
content_tagseffect size / sensitivity index
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content_tags.statistics[9]the tagger's words
content_tagstwo-way ANOVA
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also in: two-way ANOVA
content_tags.statistics[10]1.0the tagger's words
content_tagsWilcoxon rank-sum
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.statistics[11]1.0the tagger's words
content_tagsPoisson
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also in: Poisson
content_tags.statistics[12]1.0the tagger's words
content_tagsD′
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: D′
content_tags.statistics[13]1.0the tagger's words
content_tagsAverage response duration to the sound bursts grows from primary to ventral non-primary cortex in the order A1, VAF, cSRAF.
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content_tags.findings[0]the tagger's words
content_tagsThe integration time constant that gives best pairwise sound-shape discrimination also grows in the same order A1, VAF, cSRAF, establishing a hierarchy of time scales.
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content_tags.findings[1]the tagger's words
content_tagsOnset spikes carry most of the discrimination in all three fields, while the later sustained spikes discriminate better in the two non-primary fields than in A1.
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content_tags.findings[2]the tagger's words
content_tagsSpike timing outperforms spike rate as a code for identifying which of ten envelope shapes was played, in primary and non-primary cortex alike.
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content_tags.findings[3]the tagger's words
content_tagsA single neuron's spike timing classifies the ten envelope shapes at roughly 25-40% correct, well above the 10% chance level.
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content_tags.findings[4]the tagger's words
content_tagsPooling neuron output raises classification accuracy to a maximum of 80-100%.
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content_tags.findings[5]the tagger's words
content_tagsPeak response delay lengthens with the envelope shape parameter in all three cortical fields alike.
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.findings[6]the tagger's words
content_tagsResponse duration tracks envelope shape four times more steeply in VAF and seventeen times more steeply in cSRAF than in A1.
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.findings[7]the tagger's words
content_tagsA1 and VAF reach half-maximal classification by pooling only 3 and 5 neurons respectively, whereas cSRAF needs 16.
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content_tags.findings[8]the tagger's words
content_tagsA rate code built from spike counts leaves A1 neurons below 15% correct, close to the 10% chance level.
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.findings[9]the tagger's words
content_tagsresponse durations and optimal time constants for discriminating sound envelope shape INCREASE IN RANK ORDER WITH A1 < VAF < cSRAF
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content_tags.findings[10]1.0the tagger's words
content_tagsA1 and VAF have significantly higher mean discrimination than cSRAF across all 45 pairwise discriminations, CONVERSELY cSRAF neurons are marginally better across 30 pairwise comparisons when sound burst f c is <11 Hz
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content_tags.findings[11]1.0the tagger's words
content_tagsA1 AND VAF OUTPERFORM cSRAF in 10 alternative forced choice classification scenarios
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content_tags.findings[12]1.0the tagger's words
content_tagsthe effect of pooling was GREATER ON AVERAGE IN A1 than in cSRAF, HOWEVER the benefit of pooling was GREATER IN cSRAF for two of the longest duration sound bursts
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content_tags.findings[13]1.0the tagger's words
content_tagsthe ability to classify 10 sound shapes with spike-timing pattern is Quotation withheld, A1 > VAF > cSRAF
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content_tags.findings[14]1.0the tagger's words
content_tagsonset response time windows VARY WITH SOUND SHAPE AND ACROSS CORTICAL FIELDS - A1 29.6 where cSRAF is 180.7 at f c of 2
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about this quote: A TABLE IS JOINED INTO THE 'evidence' QUOTE: after the table's title ('Onset response time windows vary…') it runs on through the table's column heads and cells ('f c 2 4 6 8 11 16 …'). The numbers this row states are table cells; read the quote as a title, then a table.
content_tags.findings[15]1.0the tagger's words
content_tagsonset response time windows VARY WITH SOUND BURST HALF-DURATION AND SLOPE as determined by the sound shaping filter
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content_tags.findings[16]1.0the tagger's words
content_tagsIN THE REMAINDER OF THIS MANUSCRIPT WE USE THE TERM ONSET SPIKES TO DENOTE THE EVOKED ONSET SPIKES ONLY, after a subset was categorised as spontaneous by random sampling
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content_tags.findings[17]1.0the tagger's words
content_tagsthe spike distance IS BIASED as the time constant approaches the length of the spike train, so the train was extended BY SYNTHESIZING a periodic spike train WITH MATCHED TEMPORAL STATISTICS out to 100 s
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content_tags.findings[18]1.0the tagger's words
content_tagswe CANNOT DISAMBIGUATE potential neural codes for sound burst slope versus duration AS THESE CUES COVARY IN OUR SOUNDS
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content_tags.findings[19]1.0the tagger's words
content_tagsthe naive Bayes classifier TREATS RESPONSE BINS FROM DIFFERENT NEURONS AS IF THEY ARE INDEPENDENT, by design
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content_tags.findings[20]1.0the tagger's words
content_tagsTHE FIRST 500 ms WAS REMOVED FROM EACH SPIKE TRAIN to restrict analysis to the adapted steady-state segments
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content_tags.findings[21]1.0the tagger's words
content_tagsspike count or equivalent spike rate per 2 s IS NOT AN OPTIMAL NEURAL SIGNAL for coding sound shape in primary and non-primary auditory cortices
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.findings[22]1.0the tagger's words
content_tagspooled spike-rate classification is CLOSE TO CHANCE LEVELS in all cortical fields while the timing classifier reaches 80 to 100% accuracy
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content_tags.findings[23]1.0the tagger's words
content_tagsA1 and VAF require fewer neurons, 3 and 5, than cSRAF at 16, to reach half maximal classification
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.findings[24]1.0the tagger's words
content_tagsonset spikes alone discriminate sound shape well in all three fields, WHEREAS SUSTAINED RESPONSES PROVIDE MORE EFFECTIVE DISCRIMINATION IN NON-PRIMARY CORTICES
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content_tags.findings[25]1.0the tagger's words
content_tagsnon-primary cortex COULD CERTAINLY POOL RESPONSES FROM A1 to classify sounds BUT THE REVERSE IS LESS LIKELY
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content_tags.findings[26]1.0the tagger's words
content_tagsshape temporal cues
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content_tags.definitions[0]the tagger's words
content_tagsencoding time
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: encoding time
content_tags.definitions[1]the tagger's words
content_tagsperiod histogram
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[2]the tagger's words
content_tagsresponse delay
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[3]the tagger's words
content_tagsresponse duration
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[4]the tagger's words
content_tagsdiscrimination index
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[5]the tagger's words
content_tagsonset spikes
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: onset spikes
content_tags.definitions[6]the tagger's words
content_tagssustained spikes
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.definitions[7]the tagger's words
content_tagsonset
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: onset
content_tags.definitions[9]1.0the tagger's words
content_tagstime constant
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: time constant
content_tags.definitions[10]1.0the tagger's words
content_tagsenvelope
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: envelope
content_tags.definitions[11]0.9the tagger's words
content_tagssound envelope shape
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[0]the tagger's words
content_tagsexponential time constant
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content_tags.variables[1]the tagger's words
content_tagsnumber of neurons
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[2]the tagger's words
content_tagsPairwise sound discrimination indices
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[3]the tagger's words
content_tagspercentage correct classification
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[4]the tagger's words
content_tagscortical field
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content_tags.variables[6]the tagger's words
content_tagsinteraural level difference
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content_tags.variables[8]the tagger's words
content_tagsnoise carrier
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: noise carrier
content_tags.variables[10]the tagger's words
content_tagstime constant
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
also in: time constant
content_tags.variables[11]1.0the tagger's words
content_tagsresponse duration
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[12]1.0the tagger's words
content_tagsdiscrimination
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[13]1.0the tagger's words
content_tagsclassification
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content_tags.variables[14]1.0the tagger's words
content_tags223 single neurons from 16 male Brown Norway rats — total sample of recorded units and animals
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content_tags.quantities[0]the tagger's words
content_tags54, 104, and 65 — number of single neurons analysed in A1, VAF and cSRAF respectively
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[1]the tagger's words
content_tags19.6 (±0.1), 28.9 (±0.4), 53.6 (±1.9) — mean response duration in ms for A1, VAF and cSRAF averaged over all sound conditions
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content_tags.quantities[2]the tagger's words
content_tags0.97 (± 0.01), 0.98 (±0.01), and 0.92 (±0.01) — mean pairwise discrimination index across all 45 sound pairs in A1, VAF and cSRAF
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content_tags.quantities[3]the tagger's words
content_tags2 to threefold — increase in classification performance from pooling the neural population relative to the average single neuron
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content_tags.quantities[4]the tagger's words
content_tags−122, −113, and −80, respectively; r 2 = 0.82, 0.83 and 0.89 — log-linear regression slopes and fit quality for peak response delay versus envelope shape in A1, VAF and SRAF
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content_tags.quantities[5]the tagger's words
content_tagsF (9,1323) = 29.6, p = 0 — effect of sound shape on temporal-classifier performance
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content_tags.quantities[6]the tagger's words
content_tags10 different variations — number of envelope shape conditions, spanning fc from 2 to 64 Hz
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content_tags.quantities[7]the tagger's words
content_tags29.6 — onset response time window for A1 at f c of 2, where the cSRAF value in the same row of the same table is 180.7
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content_tags.quantities[8]1.0the tagger's words
content_tags100 s — the length of the SYNTHESIZED periodic spike train that stands in for the recording so the distance metric is not biased at long time constants
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content_tags.quantities[9]1.0the tagger's words
content_tags500 ms — the segment removed from the start of every spike train before any analysis
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content_tags.quantities[10]1.0the tagger's words
content_tags20 ms — the half-resolution limit read off the point where discrimination performance drops
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content_tags.quantities[11]1.0the tagger's words
content_tags3 and 5 — neurons needed by A1 and VAF for half maximal classification, against 16 for cSRAF
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content_tags.quantities[12]1.0the tagger's words

Interpretive choices

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© 2026 Shir Sivroni · ORCID 0009-0000-2597-2824