A Discontinuous Tonotopic Organization in the Inferior Colliculus of the Rat — TEI-tagged view

Malmierca, Izquierdo, Cristaudo, Hernández, Pérez-González, Covey, Oliver · 2008 · neuroscience · doi:10.1523/jneurosci.0238-08.2008 · full text at the source · TEI xml · tagged edition · index · concepts · authors
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 ('A single previous study with multiunit recordings has shown a similar stepwise tonotopic organization in the CNIC of the cat ( Schreiner and Langner, 1997 ).') 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.
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, whether or not the words occur in the text below — a number is how many papers in the corpus carry that tag; a tag with no number is unique to this paper
tonotopic organization4 best frequency7 inferior colliculus7 CNIC4 laminae stepwise continuous3 critical bands frequency response areas multiunit4 bandpass filters disk-shaped neurons electrophysiology110

Our tag layer for this paper

Cross-disciplinary analysis
content-tags & facets: phenomenon · claim-type · cross-domain-transfer · mechanism · claim-relation · replication-status · limitation — grounded in the full text
phenomenontonotopic organization in the inferior colliculus— “The present study shows a discontinuous tonotopic organization in the CNIC.”
claim-typeexistence— “Here, physiological and anatomical data show that best frequency is represented in a discontinuous manner in the inferior colliculus, the major auditory structure of the midbrain.”
method — evidence— “Extracellular single-unit or multiunit activity was recorded from the IC in 137 pigmented rats ( Rattus norvergicus , Rj:Long–Evans; body weight, 109–420 g) of both sexes.”
object of study — evidence— “It is not known how this tonotopic pattern corresponds with the BF of single neurons, nor is it known how the stepwise progression seen by Schreiner and Langner (1997) relates to the anatomical laminae within the IC.”
research question — evidence— “The tonotopic organization in the CNIC is often assumed to be a continuous progression from low to high frequency ( Rose et al., 1966 ; Clopton and Winfield, 1973 ; Merzenich and Reid, 1974 ). However, multiunit recordings of best frequency (BF) at threshold from the CNIC in the cat have described a stepwise progression in the tonotopic…
contribution — evidence— “Multiunit maps demonstrate a distinct stepwise organization in the order of best frequency progression. Furthermore, independent data from single neurons show that best frequencies at octave intervals of approximately one-third are more prevalent than others.”
transferfrom human psychoacoustics: the critical band, imported as the functional interpretation of an anatomical lamina, with the rat count estimated from basilar membrane length— “The number of critical bands in the rat is not known, but a rough estimate can be based on two assumptions: (1) the basilar membranes of mammals are scale models of each other and critical bands cover equal distances on the basilar membrane ( Greenwood, 1990 ); and (2) one critical band is thought to cover ∼1 mm (0.7–1.3 mm) on the basilar…transfers[0]
transferfrom auditory prosthetics: the clinical motivation - midbrain implants in trials need to know the spectral layout they are stimulating— “Furthermore, understanding this spectral coding will help to further develop promising new auditory prosthetic devices that use electrodes implanted in the IC and are now in clinical trials”transfers[1]
mechanismbest 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— “Thus, near threshold, frequencies at the center of the filters are favored over those at the overlapping regions, and it results in the multi-modal, saw-toothed pattern of frequency distribution.”mechanisms[0]
mechanismthe discrete physiological frequency steps arise because the map is built from anatomically discrete fibrodendritic laminae, each lamina acting as one frequency-band module— “Thus, our anatomical and physiological data suggest that the tonotopic map in the CNIC in the rat is composed of discrete physiological laminae that correspond to the anatomical fibrodendritic laminae of the CNIC.”mechanisms[1]
mechanismbest 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— “At the IC level, best frequencies may be organized as partially overlapping bandpass filters with a center-to-center separation of one-third octave. Thus, near threshold, frequencies at the center of the filters are favored over those at the overlapping regions, and it results in the multi-modal, saw-toothed pattern of frequency…mechanisms[2]
mechanismthe frequency step size matches the axonal lamina thickness, about 140-150 microns— “Our data in the rat show that the axonal laminar thickness and the frequency step changes are similar across frequency (140–150 μm).”mechanisms[3]
mechanismtwo to three flat neurons side by side may form the functional unit of a low-frequency lamina— “The dendritic arbors of flat cells are ∼50 μm thick and less flat cells are ∼100 μm thick on average ( Malmierca et al., 1993 , 1995 ), suggesting that two to three flat neurons organized side-by-side”mechanisms[4]
mechanismelectrode penetrations were angled roughly orthogonal to the isofrequency laminae so the steps would be crossed rather than followed— “Penetrations were from anterodorsal to posteroventral through the IC at an angle 10° from the frontal plane (see Fig. 8 C ). This angle is approximately orthogonal to the predominant orientation of isofrequency laminae”mechanisms[5]
mechanismstep size differs across structures and species and correlates with dendritic field width— “A comparison of dendritic fields suggests that the larger step size is positively correlated with a larger dendritic field width”mechanisms[6]
replicates Schreiner and Langner, 1997 (stepwise tonotopy in cat CNIC)— “A single previous study with multiunit recordings has shown a similar stepwise tonotopic organization in the CNIC of the cat ( Schreiner and Langner, 1997 ).”claim_relations[0]
contradicts Rose et al., 1966; Clopton and Winfield, 1973; Merzenich and Reid, 1974— “Previous studies ( Rose et al., 1966 ; Clopton and Winfield, 1973 ; Merzenich and Reid, 1974 ) used larger steps in their electrode mapping and thus are likely to have sampled across the stepwise organization.”claim_relations[1]
extends Doron et al., 2002 (rat primary auditory cortex single units)— “To determine whether the BFs of single units in the primary auditory cortex of the rat follow a stepwise progression, we analyzed the BF of 198 cortical single units from a previous study ( Doron et al., 2002 , their Fig. 2 ) using the same approach that we used for our 604 IC single units.”claim_relations[2]
contradicts the general view that CNIC tonotopy is a continuous frequency gradient— “Together, both types of data argue against the general view that the tonotopic organization in the CNIC forms a continuous frequency gradient. Rather, it seems that there is a…claim_relations[3]
explains-away the earlier mapping studies that reported continuity, on the grounds that they sampled too coarsely to resolve the stepsclaim_relations[4]
confirms Schreiner and Langner's stepwise organisation in cat, at a different step size
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claim_relations[5]
extends Doron et al.'s cortical single-unit data, reanalysed here and found to show the same periodicity
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claim_relations[6]
supports the proposal that critical band filters originate at the midbrain
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claim_relations[7]
sensetonotopy 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]
sensethe 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]
sensebest 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]
sensea 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]
controlthe acoustic output calibrated in situ at each ear with a quarter-inch condenser microphone and a spectrum analyser — held — rules out 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
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controls[0]
noteEVIDENCE CORRECTED (RE-GROUNDED) · stepwise —contrasted-with→ continuous — 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
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concept_edges[0]
retractedEVIDENCE CORRECTED (RE-GROUNDED) · stepwise —contrasted-with→ continuous — 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.
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concept_edges[0]
auditbest frequency —operationalised-as→ frequency response areas — 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.
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concept_edges[3]
audittonotopic organization —operationalised-as→ multiunit — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span reports stepwise tonotopy in multiunit recordings; operationalised-as is loose.
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concept_edges[6]
notetonotopic organization —operationalised-as→ multiunit — RE-GROUNDS another row of this record: the old span named NEITHER endpoint.
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concept_edges[6]
weighttonotopic organization —operationalised-as→ multiunit — 0.9
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concept_edges[6]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
for which uselimitation — every laminar width figure: MEASUREMENTS ARE PRESENTED WITHOUT CORRECTION FOR SHRINKAGE
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limitations[0]
for which uselimitation — the mapping onto critical bands: THE NUMBER OF CRITICAL BANDS IN THE RAT IS NOT KNOWN, and the estimate rests on two stated assumptions
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limitations[1]
for which uselimitation — the proportion of penetrations showing the organisation: IN 12 PENETRATIONS, NO CLEAR TONOTOPIC ORGANIZATION WAS FOUND ... OR THE TONOTOPIC PROGRESSION WAS INCOMPLETE
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limitations[2]
for which uselimitation — 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
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limitations[3]
for which uselimitation — the single-unit half of the evidence: THE SINGLE-UNIT DATA WERE COLLECTED IN PREVIOUS EXPERIMENTS WITH DIFFERENT AIMS UNRELATED TO TONOTOPIC ORGANIZATION
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limitations[4]
for which uselimitation — the frequency range every distribution covers: THE HIGHEST FREQUENCY PRODUCED BY THIS SYSTEM WAS LIMITED TO 40 kHz
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limitations[5]
for which uselimitation — the same scope as limitations[1] - the critical-band count is an estimate from assumptions, not a measurement in this species
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limitations[6]
for which uselimitation — 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
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limitations[7]
replicationreplicated
limitationLaminar width measurements were not corrected for histological shrinkage.
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limitations[0]
limitationThe 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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limitations[1]
limitationTwelve penetrations gave no clear or incomplete tonotopic progression and were excluded from analysis.
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limitations[2]
limitationsteps of 50 microns or less were required to see the organisation at all, so the result depends on sampling resolution
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limitations[3]
limitationmost single units were collected in earlier studies with unrelated aims, in two laboratories with different equipment
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limitations[4]
limitationthe 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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limitations[5]
limitationthe number of critical bands in the rat is not known and is estimated from two stated assumptions
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limitations[6]
limitationthe cortical comparison rests on reanalysing a figure from another group's paper
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limitations[7]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingSampling 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]
findingBest 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]
findingBest 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]
findingStep 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]
findingThe 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]
findingReanalysed 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]
findingTaken 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]
findingmultiunit maps show a distinct stepwise progression of best frequency
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content_tags.findings[7]
findingsingle-unit best frequencies cluster at intervals of about one third of an octave
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content_tags.findings[8]
findingthe two lines of evidence argue against a continuous gradient
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content_tags.findings[9]
findingearlier studies reporting continuity used larger electrode steps and would have sampled across the structure
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content_tags.findings[10]
findingbest 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]
findingaxonal lamina thickness and frequency step size are both about 140-150 microns
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content_tags.findings[12]
findingreanalysing another group's cortical single units shows the same periodicity, r = 0.998
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content_tags.findings[13]
findingthe 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]
findingmidbrain filters appear roughly twice as broad as peripheral auditory filters
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content_tags.findings[15]
quantity137 — pigmented rats from which extracellular activity was recorded
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content_tags.quantities[0]
quantity604 — single-unit frequency response areas obtained, from 125 rats
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content_tags.quantities[1]
quantity0.34 ± 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]
quantity14 ± 3.7 — average number of frequency steps per electrode track
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content_tags.quantities[3]
quantityr = 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]
quantity104 ± 21 μm — mean cross-sectional diameter of the seven thinnest anatomically labelled axonal laminae
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content_tags.quantities[5]
quantity0.006 ± 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]
quantity142 ± 69 μm — mean thickness of a best-frequency step in low-frequency regions (below 4 kHz)
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content_tags.quantities[7]
quantity2180 ± 460 μm — average length of the recorded dorsoventral electrode tracks
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content_tags.quantities[8]
quantity∼8–12 — estimated number of critical bands in the rat, based on an 8 mm basilar membrane
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content_tags.quantities[9]
quantityabout 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
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content_tags.quantities[10]
value source notequantity — 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]
quantity140-150 microns — both the axonal lamina thickness and the spatial period of the frequency steps
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content_tags.quantities[11]
quantity0.006 octaves mean change, variance 0.0002, 50 times smaller than the 0.34 octave jumps — stability of best frequency over 23 to 80 minutes
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content_tags.quantities[12]
quantity8-12 critical bands from an 8 mm basilar membrane, and 8-12 laminae observed — two independent estimates agreeing
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content_tags.quantities[13]
quantity604 frequency response areas from 125 rats, 400 of them from earlier studies — the single-unit dataset and its provenance
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content_tags.quantities[14]
quantity137 rats of both sexes — the full sample
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content_tags.quantities[15]
quantity25 kHz in one system and 40 kHz in the other — the differing stimulus ceilings of the two laboratories contributing single units
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content_tags.quantities[16]
value source notequantity — 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]
quantityr = 0.99841 — correlation between IC and cortical best-frequency distributions
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content_tags.quantities[17]
variableBF (dependent)
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content_tags.variables[2]
variablecross-sectional diameter (dependent)
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content_tags.variables[3]
variablebody temperature (control)
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content_tags.variables[4]
variablebest frequency
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[6]
variabledepth along the electrode track
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[7]
variableElectrode steps
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[8]
definitiontonotopic organization
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[0]
definitionwidth of the plexus
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[2]
definitionbandpass filters
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[4]
definitionthreshold
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[5]
definitionBF
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[6]
key-termstonotopic organization, best frequency, inferior colliculus, CNIC, laminae, stepwise, continuous, critical bands, frequency response areas, multiunit, bandpass filters, disk-shaped, isofrequency, basilar membrane, octave, threshold, midbrain, 3-D reconstruction, auditory prosthetic, cochlear nucleus
Statistics — cross-field join keys
field-invariant statistical terms, used in their statistical sense (gated on a number / symbol / test). A p-value or a standard deviation means the SAME thing in physics, biology, psychology and sociology — so these are the strongest cross-field join keys. Grounded verbatim.
Mann-Whitney U test p as written: Mann–Whitney tests
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[0]
linear regression as written: a linear regression analysis
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[1]
correlation coefficient r as written: strongly correlated ( r = 0.99855)
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[2]
mean and standard deviation ± as written: (mean ± SD)
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[3]
histogram (binned frequency distribution) as written: displayed as a histogram with 0.1 octave bins
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[4]
smoothing (fast Fourier transform) as written: fast Fourier transform smoothing
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[5]
curve fitting (sigmoidal function) as written: described by a sigmoidal function
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[6]
logarithmic histogram of best frequencies with peak picking
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[7]
repeated measurement of the same unit over time
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[8]
Second readings
a facet read a second time, independently, from its own sentence — and whether the two readings agree
claim typecritique — a second reading DIFFERING from the recorded value; recorded: existence
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.claim_type
methodmultiunit mapping in 25-50 micron steps, single-unit frequency response areas from two laboratories, and 3D reconstruction of anatomically labelled axonal laminae — a second reading DIFFERING from the recorded value; recorded: electrophysiology
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.method
phenomenontonotopy as a discontinuous, laminar organisation — a second reading DIFFERING from the recorded value; recorded: tonotopic organization in the inferior colliculus
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.phenomenon

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.

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