A Discontinuous Tonotopic Organization in the Inferior Colliculus of the Rat

Malmierca, Izquierdo, Cristaudo, Hernández, Pérez-González, Covey, Oliver · Journal of Neuroscience · 2008 · neuroscience · 10.1523/jneurosci.0238-08.2008 · record Malmierca2008-a-discontinuous-tonotopic-organization-in-th
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. Replication status is set to 'replicated' on the paper's own stated grounds, not on outside knowledge: it reports a prior multiunit study finding the same stepwise organisation in cat ('Quotation withheld') and internal convergence across two independently collected datasets ('Our observations are based on two independent datasets that yielded very similar results, one from multiunit mapping and the other from single units.'). If the corpus convention reserves 'replicated' for post hoc independent replication of this paper, downgrade to 'untested'. The facets do not capture the paper's third method strand - anterograde tracer injections in the dorsal cochlear nucleus with Neurolucida 3-D reconstruction - which is anatomical rather than electrophysiological; 'method' had to pick one. The single-unit dataset is partly reused from earlier studies (n = 400 of 604 units), which the paper discloses but does not frame as a limitation. 2026-08-13 schema-0.3 pass: 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
threshold 2 fields · linear regression 2 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 12 concept tags on this paper, none is 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.
tonotopic organization best frequency inferior colliculus CNIC laminae stepwise continuous critical bands frequency response areas multiunit bandpass filters disk-shaped neurons best frequencies are laid out as partially overlapping bandpass filters spaced one third of an octave apart, so near threshold the filter centre frequencies win out over the overlap regions and the measured distribution comes out saw-toothed the discrete physiological frequency steps arise because the map is built from anatomically discrete fibrodendritic laminae, each lamina acting as one frequency-band module disk-shaped isofrequency basilar membrane octave threshold midbrain 3-D reconstruction auditory prosthetic cochlear nucleus width of the plexus BF cross-sectional diameter body temperature depth along the electrode track Electrode steps Mann-Whitney U test Mann–Whitney tests linear regression a linear regression analysis correlation coefficient mean and standard deviation (mean ± SD) histogram (binned frequency distribution) smoothing (fast Fourier transform) fast Fourier transform smoothing curve fitting (sigmoidal function) described by a sigmoidal function logarithmic histogram of best frequencies with peak picking repeated measurement of the same unit over time 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
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

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.0238-08.2008. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 8 · claim_type 1 · concept_edges 3 · content_tags 54 · contribution 1 · controls 1 · limitations 8 · mechanisms 7 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 4 · transfers 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 3 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studythe fine-grained spatial progression of best frequency through the central nucleus of the IC
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object_of_studythe paper's words
research_questionwhether the tonotopic map in the IC is a continuous gradient or a stepwise one, and how it relates to the anatomical laminae
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research_questionthe paper's words
contributionshows the tonotopic progression is stepwise rather than continuous, in both multiunit maps and the distribution of single-unit best frequencies, and matches the anatomical lamina thickness
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contributionthe paper's words
claim_typeexistence
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claim_typethe paper's words
phenomenontonotopic organization in the inferior colliculus
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phenomenonthe paper's words
methodelectrophysiology
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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
mechanismsbest frequencies are modelled as partially overlapping bandpass filters spaced a third of an octave apart, so near threshold the filter centres are favoured and produce a saw-toothed distribution
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mechanisms[2]1.0the tagger's words
mechanismsthe frequency step size matches the axonal lamina thickness, about 140-150 microns
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mechanisms[3]1.0the tagger's words
mechanismstwo to three flat neurons side by side may form the functional unit of a low-frequency lamina
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mechanisms[4]0.9the tagger's words
mechanismselectrode penetrations were angled roughly orthogonal to the isofrequency laminae so the steps would be crossed rather than followed
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mechanisms[5]1.0the tagger's words
mechanismsstep size differs across structures and species and correlates with dendritic field width
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mechanisms[6]0.9the tagger's words
limitationsLaminar width measurements were not corrected for histological shrinkage.
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what this limits: every laminar width figure: MEASUREMENTS ARE PRESENTED WITHOUT CORRECTION FOR SHRINKAGE
limitations[0]the tagger's words
limitationsThe mapping of the finding onto critical bands rests on two unverified assumptions, since the number of critical bands in the rat is unknown.
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what this limits: the mapping onto critical bands: THE NUMBER OF CRITICAL BANDS IN THE RAT IS NOT KNOWN, and the estimate rests on two stated assumptions
limitations[1]the tagger's words
limitationsTwelve penetrations gave no clear or incomplete tonotopic progression and were excluded from analysis.
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what this limits: the proportion of penetrations showing the organisation: IN 12 PENETRATIONS, NO CLEAR TONOTOPIC ORGANIZATION WAS FOUND ... OR THE TONOTOPIC PROGRESSION WAS INCOMPLETE
limitations[2]the tagger's words
limitationssteps of 50 microns or less were required to see the organisation at all, so the result depends on sampling resolution
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what this limits: any comparison with the earlier mapping literature: ELECTRODE STEPS OF 50 um OR LESS WERE NECESSARY TO OBSERVE THE STEPWISE ORGANIZATION ... PREVIOUS STUDIES USED LARGER STEPS ... AND THUS ARE LIKELY TO HAVE SAMPLED ACROSS THE STEPWISE ORGANIZATION
limitations[3]1.0the tagger's words
limitationsmost single units were collected in earlier studies with unrelated aims, in two laboratories with different equipment
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what this limits: the single-unit half of the evidence: THE SINGLE-UNIT DATA WERE COLLECTED IN PREVIOUS EXPERIMENTS WITH DIFFERENT AIMS UNRELATED TO TONOTOPIC ORGANIZATION
limitations[4]1.0the tagger's words
limitationsthe two recording systems had different frequency ceilings, 25 kHz and 40 kHz, in a study about the distribution of best frequencies
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what this limits: the frequency range every distribution covers: THE HIGHEST FREQUENCY PRODUCED BY THIS SYSTEM WAS LIMITED TO 40 kHz
limitations[5]0.9the tagger's words
limitationsthe number of critical bands in the rat is not known and is estimated from two stated assumptions
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what this limits: the same scope as limitations[1] - the critical-band count is an estimate from assumptions, not a measurement in this species
limitations[6]1.0the tagger's words
limitationsthe cortical comparison rests on reanalysing a figure from another group's paper
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what this limits: the cortical comparison: WE ANALYZED THE BF OF 198 CORTICAL SINGLE UNITS FROM A PREVIOUS STUDY (DORON ET AL., 2002, THEIR FIG. 2) USING THE SAME APPROACH
limitations[7]1.0the tagger's words
transfersthe critical band, imported as the functional interpretation of an anatomical lamina, with the rat count estimated from basilar membrane length
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transfers[0]1.0the tagger's words
transfersthe clinical motivation - midbrain implants in trials need to know the spectral layout they are stimulating
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transfers[1]0.9the tagger's words
controlsthe acoustic output calibrated in situ at each ear with a quarter-inch condenser microphone and a spectrum analyser
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eliminates: a systematic level error at one ear - without in-situ calibration a stated decibel value is a property of the equipment rather than of the sound reaching the animal · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
sensestonotopy here is DISCONTINUOUS by construction of the claim - a stepwise progression with a defined space of tissue per frequency band, not a smooth gradient
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senses[0]1.0the tagger's words
sensesthe continuous view is named as an ASSUMPTION of the field rather than as a finding, which is what makes the paper a critique
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senses[1]1.0the tagger's words
sensesbest frequency is defined as the frequency with the lowest threshold, and threshold as the lowest level eliciting a response - the step structure is a property of BF so defined
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senses[2]1.0the tagger's words
sensesa lamina is simultaneously an anatomical object (axonal band, 140-150 microns) and a functional one (a frequency step) - the paper's claim is that they are the same thing
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senses[3]1.0the 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: multiunit mapping in 25-50 micron steps, single-unit frequency response areas from two laboratories, and 3D reconstruction of anatomically labelled axonal laminae
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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: tonotopy as a discontinuous, laminar organisation
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a second reading DIFFERING from the recorded value. Recorded: tonotopic organization in the inferior colliculus — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsreplicates → Schreiner and Langner, 1997 (stepwise tonotopy in cat CNIC)
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claim_relations[0]the tagger's words
claim_relationscontradicts → Rose et al., 1966; Clopton and Winfield, 1973; Merzenich and Reid, 1974
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claim_relations[1]the tagger's words
claim_relationsextends → Doron et al., 2002 (rat primary auditory cortex single units)
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claim_relations[2]the tagger's words
claim_relationscontradicts → the general view that CNIC tonotopy is a continuous frequency gradient
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claim_relations[3]1.0the tagger's words
claim_relationsexplains-away → the earlier mapping studies that reported continuity, on the grounds that they sampled too coarsely to resolve the steps
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claim_relations[4]1.0the tagger's words
claim_relationsconfirms → Schreiner and Langner's stepwise organisation in cat, at a different step size
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claim_relations[5]1.0the tagger's words
claim_relationsextends → Doron et al.'s cortical single-unit data, reanalysed here and found to show the same periodicity
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claim_relations[6]1.0the tagger's words
claim_relationssupports → the proposal that critical band filters originate at the midbrain
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claim_relations[7]0.9the tagger's words
concept_edgesEVIDENCE CORRECTED (RE-GROUNDED) · stepwise —contrasted-with→ continuous
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EVIDENCE CORRECTED: RE-GROUNDED. The old span argued against a general view without naming either term. This sentence names both and states the contrast in eight words - 'Some appear to be CONTINUOUS, WHEREAS others are STEPWISE' - which is the cleanest contrast span merged tonight. — strength of the relation: 1 · note: the paper's whole claim; and note that the corpus holds both descriptions of IC tonotopy as findings, from this paper and from the studies it refutes
also in: stepwise · continuous
concept_edges[0]0.85the tagger's words
concept_edgesbest frequency —operationalised-as→ frequency response areas
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but best frequency is DEFINED as the frequency with the lowest threshold; 'frequency response areas' is not in the span.
concept_edges[3]1.0the tagger's words
concept_edgestonotopic organization —operationalised-as→ multiunit
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strength of the relation: 0.9 · note: RE-GROUNDS another row of this record: the old span named NEITHER endpoint. Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span reports stepwise tonotopy in multiunit recordings; operationalised-as is loose.
concept_edges[6]0.95the tagger's words
content_tagsMann-Whitney U test
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content_tags.statistics[0]the tagger's words
content_tagslinear regression
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content_tags.statistics[1]the tagger's words
content_tagscorrelation coefficient
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content_tags.statistics[2]the tagger's words
content_tagsmean and standard deviation
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content_tags.statistics[3]the tagger's words
content_tagshistogram (binned frequency distribution)
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content_tags.statistics[4]the tagger's words
content_tagssmoothing (fast Fourier transform)
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content_tags.statistics[5]the tagger's words
content_tagscurve fitting (sigmoidal function)
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content_tags.statistics[6]the tagger's words
content_tagslogarithmic histogram of best frequencies with peak picking
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content_tags.statistics[7]0.85the tagger's words
content_tagsrepeated measurement of the same unit over time
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content_tags.statistics[8]1.0the tagger's words
content_tagsSampling best frequency every 25-50 micrometres revealed a stair-step rather than smooth progression in every penetration through the central nucleus of the inferior colliculus.
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content_tags.findings[0]the tagger's words
content_tagsBest frequency stayed within 0.1 octaves for about 151 micrometres of electrode travel and then jumped by about a third of an octave.
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content_tags.findings[1]the tagger's words
content_tagsBest frequencies of 604 independently recorded single units were not uniformly distributed but clustered into 16 peaks spaced about a third of an octave apart.
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content_tags.findings[2]the tagger's words
content_tagsStep thickness and step size were statistically indistinguishable between low-frequency and high-frequency regions, so laminae are similar throughout the central nucleus.
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content_tags.findings[3]the tagger's words
content_tagsThe thinnest anatomically labelled axonal laminae, once corrected for histological shrinkage, are the same size as the physiological frequency steps, tying the stair-step map to the fibrodendritic laminae.
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content_tags.findings[4]the tagger's words
content_tagsReanalysed cortical single units showed a similar multipeaked best-frequency distribution, with peaks about 0.31 octaves apart.
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content_tags.findings[5]the tagger's words
content_tagsTaken together the multiunit and single-unit data contradict the standard assumption that the inferior colliculus contains a continuous low-to-high frequency gradient.
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content_tags.findings[6]the tagger's words
content_tagsmultiunit maps show a distinct stepwise progression of best frequency
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content_tags.findings[7]1.0the tagger's words
content_tagssingle-unit best frequencies cluster at intervals of about one third of an octave
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content_tags.findings[8]1.0the tagger's words
content_tagsthe two lines of evidence argue against a continuous gradient
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content_tags.findings[9]1.0the tagger's words
content_tagsearlier studies reporting continuity used larger electrode steps and would have sampled across the structure
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content_tags.findings[10]1.0the tagger's words
content_tagsbest frequency of a unit stays constant over 23 to 80 minutes, varying 50 times less than the step size
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content_tags.findings[11]1.0the tagger's words
content_tagsaxonal lamina thickness and frequency step size are both about 140-150 microns
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content_tags.findings[12]1.0the tagger's words
content_tagsreanalysing another group's cortical single units shows the same periodicity, r = 0.998
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content_tags.findings[13]1.0the tagger's words
content_tagsthe data suggest 8-12 laminae of 0.29-0.36 octaves each, similar to the 0.333-0.375-octave critical bands suggested for the mouse
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content_tags.findings[14]1.0the tagger's words
content_tagsmidbrain filters appear roughly twice as broad as peripheral auditory filters
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content_tags.findings[15]1.0the tagger's words
content_tagstonotopic organization
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content_tags.definitions[0]the tagger's words
content_tagswidth of the plexus
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content_tags.definitions[2]the tagger's words
content_tagsbandpass filters
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content_tags.definitions[4]1.0the tagger's words
content_tagsthreshold
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also in: threshold
content_tags.definitions[5]the tagger's words
content_tagsBF
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also in: BF
content_tags.definitions[6]the tagger's words
content_tagsBF
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also in: BF
content_tags.variables[2]the tagger's words
content_tagscross-sectional diameter
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content_tags.variables[3]the tagger's words
content_tagsbody temperature
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content_tags.variables[4]the tagger's words
content_tagsbest frequency
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content_tags.variables[6]1.0the tagger's words
content_tagsdepth along the electrode track
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content_tags.variables[7]1.0the tagger's words
content_tagsElectrode steps
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content_tags.variables[8]1.0the tagger's words
content_tags137 — pigmented rats from which extracellular activity was recorded
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content_tags.quantities[0]the tagger's words
content_tags604 — single-unit frequency response areas obtained, from 125 rats
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content_tags.quantities[1]the tagger's words
content_tags0.34 ± 0.11 octaves — mean jump in best frequency at each step of the multiunit staircase (460 steps)
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content_tags.quantities[2]the tagger's words
content_tags14 ± 3.7 — average number of frequency steps per electrode track
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content_tags.quantities[3]the tagger's words
content_tagsr = 0.99855 — correlation between the 12 matched best-frequency peaks of the single-unit and multiunit datasets
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content_tags.quantities[4]the tagger's words
content_tags104 ± 21 μm — mean cross-sectional diameter of the seven thinnest anatomically labelled axonal laminae
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content_tags.quantities[5]the tagger's words
content_tags0.006 ± 0.0002 octaves — drift of best frequency at a fixed recording site over 23-80 minutes, about 50 times smaller than a staircase step
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content_tags.quantities[6]the tagger's words
content_tags142 ± 69 μm — mean thickness of a best-frequency step in low-frequency regions (below 4 kHz)
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content_tags.quantities[7]the tagger's words
content_tags2180 ± 460 μm — average length of the recorded dorsoventral electrode tracks
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content_tags.quantities[8]the tagger's words
content_tags∼8–12 — estimated number of critical bands in the rat, based on an 8 mm basilar membrane
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[9]the tagger's words
content_tagsabout 0.29 octaves per step in rat, against 0.278 in cat and about 1 octave in rabbit MGV — step size across structures and species
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
value source note: THE VALUE DRAWS ON THE PAPER BEYOND THIS QUOTE: part of it is printed elsewhere in the paper, not in this quote ('or rat [∼ 0.29 octaves (present data)]…').
content_tags.quantities[10]0.9the tagger's words
content_tags140-150 microns — both the axonal lamina thickness and the spatial period of the frequency steps
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[11]1.0the tagger's words
content_tags0.006 octaves mean change, variance 0.0002, 50 times smaller than the 0.34 octave jumps — stability of best frequency over 23 to 80 minutes
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[12]1.0the tagger's words
content_tags8-12 critical bands from an 8 mm basilar membrane, and 8-12 laminae observed — two independent estimates agreeing
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[13]1.0the tagger's words
content_tags604 frequency response areas from 125 rats, 400 of them from earlier studies — the single-unit dataset and its provenance
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[14]1.0the tagger's words
content_tags137 rats of both sexes — the full sample
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[15]1.0the tagger's words
content_tags25 kHz in one system and 40 kHz in the other — the differing stimulus ceilings of the two laboratories contributing single units
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
value source note: THE VALUE DRAWS ON THE PAPER BEYOND THIS QUOTE: part of it is printed elsewhere in the paper, not in this quote ('frequency produced by this system was limited to 25 kHz.').
content_tags.quantities[16]0.9the tagger's words
content_tagsr = 0.99841 — correlation between IC and cortical best-frequency distributions
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[17]1.0the tagger's words

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