Tonotopic Organization of the Superior Olivary Nucleus in the Chicken Auditory Brainstem

Tabor, Coleman, Rubel, Burger · The Journal of comparative neurology · 2012 · neuroscience · 10.1002/cne.22807 · record Tabor2012-tonotopic-organization-of-the-superior-oliva
tag layer · index · concepts · authors · עברית
Where this record's text came from. faceted 2026-08 by reading the locally held text (xml); every evidence string verified as an exact substring of that text before merging. Claim type is 'existence' because the three headline conclusions assert that a particular spatial arrangement exists in SON; the paper also has a strong methodological component (3D quantitative mapping with PCA and multilinear regression offered as a broadly applicable tool), which the single claim type facet cannot represent. Two of the nineteen cases showed no tonotopic organization at all, which the paper attributes to sampling; the facets do not capture this partial negative result. 2026-08-13 and 2026-08-25 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 WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below.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 An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason.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
principal component analysis 5 fields · standard deviation 3 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 13 concept tags on this paper, 3 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.
tonotopic topographic organization superior olivary nucleus characteristic frequency Q10 response threshold onset sustained principal component analysis multiple-unit nucleus laminaris nucleus angularis isofrequency SON response patterns are inherited from its nucleus angularis input, which is why sustained and onset types and diverse tuning shapes appear there the narrow hearing range of the chicken makes the frequency-to-place function linear rather than logarithmic summation of individually subthreshold spike waveforms raises the apparent threshold of multiple-unit recordings the broad, non-restricted SON to nucleus laminaris projection supplies the anatomical substrate by which inhibition laterally sharpens frequency tuning inhibitory GABA chicken brainstem MRA coefficient of determination 3D reconstruction sound localization characteristic frequency (CF) center of mass of recording sites tonotopic map body temperature normalized distances response patterns normalized medial-lateral coordinate normalized dorsal-ventral coordinate axis normalized anterior-posterior coordinate regression Multilinear regression analysis (MRA) Principle component analysis (PCA) eigen-decomposition of the covariance matrix eigenvectors and eigenvalues of the covariance matrix t-test standard deviation mean ± SD multiple regression analysis of CF on 3D position error magnitude between predicted and measured CF 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

The paper

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Every tag on this page

alternatives 2 · claim_relations 7 · claim_type 1 · concept_edges 2 · concepts 3 · content_tags 59 · contribution 1 · controls 1 · limitations 10 · mechanisms 8 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 4 · transfers 1

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 2 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studythe topographic organisation of frequency and of response types in the chicken superior olivary nucleus
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object_of_studythe paper's words
research_questionwhether the tonotopic organisation of the auditory brainstem is preserved in an inhibitory long-range projection nucleus
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research_questionthe paper's words
contributionquantifies tonotopy in an inhibitory nucleus, shows it is less precise than in excitatory nuclei of the same species, and gives a predictive model from position to best frequency
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contributionthe paper's words
claim_typeexistence
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claim_typethe paper's words
phenomenontonotopic organization of an inhibitory auditory brainstem nucleus
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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
mechanismsmechanisms
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mechanisms[2]the tagger's words
mechanismsmechanisms
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mechanisms[3]the tagger's words
mechanismsinhibitory neurons in this system sit in a discrete projection nucleus rather than among principal cells, which is what makes their organisation separable at all
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mechanisms[4]1.0the tagger's words
mechanismsrecording sites are located by electrolytic lesions and electrode tracks, then normalised into a common 3D nucleus so cases can be combined
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mechanisms[5]1.0the tagger's words
mechanismsPCA is run first because it was unknown whether the map was linear or nonlinear, single or multiple gradients
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mechanisms[6]1.0the tagger's words
mechanismsSON inhibition widens dynamic range and sharpens temporal coding for sound localisation
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mechanisms[7]1.0the tagger's words
limitationsThreshold comparisons between sustained and onset responders may not be valid because the spike-rate criterion may not be equally sensitive across response categories.
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what this limits: threshold comparisons between response types only: CAUTION SHOULD BE USED WHEN COMPARING THRESHOLDS BETWEEN SUSTAINED AND ONSET RESPONDERS BECAUSE THE SPIKE RATE CRITERION USED TO DETERMINE RESPONSE THRESHOLD differs in effect between them
limitations[0]the tagger's words
limitationsBinaural response properties of SON neurons were not assessed, so a possible contribution of the nucleus laminaris projection remains unexplored.
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what this limits: any claim about the origin of SON response properties: WE DID NOT ASSESS THE BINAURAL RESPONSE PROPERTIES OF SON NEURONS, so NL's projection remains an unexplored contribution
limitations[1]the tagger's words
limitationsIn most animals only a limited region of the SON was sampled, which is why individual maps had to be combined into a standardized SON.
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what this limits: the tonotopic map as a whole-nucleus claim: IN MOST SUBJECTS WE SAMPLED A LIMITED REGION OF THE SON, which is why individual maps had to be combined
limitations[2]the tagger's words
limitationsthe map is explicitly less precise than in excitatory nuclei of the same species, by both error magnitude and r-squared
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what this limits: any use of this map at the precision expected of excitatory nuclei: the paper states its map is the less precise of the two by both error magnitude and its own comparison
limitations[3]1.0the tagger's words
limitationsprediction error varies widely along the tonotopic axis rather than being uniform
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what this limits: the tonotopic prediction at any single site: ERROR MAGNITUDE (|CF_predicted - CF_measured|) VARIED WIDELY AT EACH POSITION ALONG THE CALCULATED TONOTOPIC AXIS
limitations[4]1.0the tagger's words
limitationsthe threshold criterion may not be equally sensitive across response categories, so between-category threshold comparisons are cautioned against
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duplicate of: limitations[0] · what this limits: the same scope as limitations[0] - the threshold criterion may not be equally sensitive across response categories, so between-category threshold comparisons are the affected ones
limitations[5]1.0the tagger's words
limitationsmulti-unit thresholds run higher than single-unit thresholds because sub-threshold waveforms sum
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what this limits: any comparison of thresholds between recording types: THRESHOLDS OF MULTIPLE-UNIT RESPONSES TENDED TO BE SLIGHTLY HIGHER THAN THE THRESHOLDS OF ISOLATED NEURONAL RESPONSES ... THIS INCREASE REFLECTS ... WAVEFORMS THAT INDIVIDUALLY WERE NOT DETECTED
limitations[6]1.0the tagger's words
limitationssuppressed responders were too rare to analyse spatially
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what this limits: the spatial claims for suppressed responders: SUPPRESSED RESPONDERS WERE ENCOUNTERED TOO INFREQUENTLY TO ASSESS THEIR SPATIAL DISTRIBUTION
limitations[7]1.0the tagger's words
limitationsin most animals only a limited region of the nucleus was sampled, which is why standardisation was needed
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what this limits: every position reported in the standardised frame: in most animals only a limited region was sampled, so LOCATIONS OF THE RECORDING SITES WERE CONVERTED INTO NORMALIZED DISTANCES ALONG THREE ORTHOGONAL AXES
limitations[8]0.9the tagger's words
limitationswithin an isofrequency slice, position carried no information about CF at all
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what this limits: the tonotopy claim at the scale a user would apply it: within an isofrequency slice, position carries no information about CF - THE STANDARDIZED SON WAS SLICED INTO FIVE SECTIONS to test exactly this
limitations[9]1.0the tagger's words
transfersretinotopy as the template for what a sensory map is and why it matters
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transfers[0]0.85the tagger's words
controlseach stimulus presented twenty times in a pseudo-randomised sequence
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eliminates: adaptation and order - a fixed sequence would confound the tonotopic response with position in the run · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
conceptscharacteristic frequency
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concepts[3]0.85the paper's words
conceptsQ10
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also in: Q10
concepts[4]0.85the paper's words
conceptsresponse threshold
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concepts[5]0.85the paper's words
sensestonotopy is treated here as a MATTER OF DEGREE with a measured precision, not as present or absent - the paper's second conclusion is that this map is less precise than others in the same species
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senses[0]1.0the tagger's words
sensesthe map is operationalised as a multiple regression of characteristic frequency on normalised 3D position, reported as r-squared - the same operationalisation Muniak2014 used for mouse DCN
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senses[1]1.0the tagger's words
sensescharacteristic frequency is defined as the frequency with the lowest response threshold, and threshold itself as two SDs above spontaneous rate
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senses[2]1.0the tagger's words
sensesonset versus sustained are response patterns classified from post-stimulus histograms at 20 dB above threshold, and the paper finds they are spatially interspersed rather than segregated
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senses[3]1.0the tagger's words
alternativesmethod: in vivo single- and multi-unit recording with electrolytic lesion marking, 3D Nissl reconstruction, normalisation to a standardised nucleus, then PCA and multiple regression of CF on position
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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 in an inhibitory long-range projection nucleus
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a second reading DIFFERING from the recorded value. Recorded: tonotopic organization of an inhibitory auditory brainstem nucleus — 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 → Coleman et al., 2011 (previous characterization of the chicken SON)
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claim_relations[0]the tagger's words
claim_relationsbuilds-on → Moiseff and Konishi, 1983; Lachica et al., 1994; Tabor et al., 2011
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claim_relations[1]the tagger's words
claim_relationsextends → Rubel and Parks, 1975 (tonotopy of other chicken brainstem auditory nuclei)
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claim_relations[2]the tagger's words
claim_relationsextends → the tonotopic organisation established for excitatory nuclei of the chicken auditory brainstem
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claim_relations[3]1.0the tagger's words
claim_relationsconfirms → earlier suggestions from owl and chicken that frequency is systematically represented in the SON
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claim_relations[4]0.9the tagger's words
claim_relationssupports → the group's own earlier characterisation of SON response types and their proportions
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claim_relations[5]1.0the tagger's words
claim_relationscontradicts → the expectation that physiologically distinct SON cell types occupy distinct territories
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claim_relations[6]1.0the tagger's words
concept_edgescharacteristic frequency —operationalised-as→ response threshold
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strength of the relation: 1 · note: CF is DEFINED through threshold, and threshold is itself defined as two SDs above spontaneous rate - a two-step operational chain
concept_edges[1]1.0the tagger's words
concept_edgesonset —contrasted-with→ sustained
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strength of the relation: 1 · note: and they are spatially interspersed rather than segregated, which is the paper's third conclusion
also in: onset · sustained
concept_edges[2]1.0the tagger's words
content_tagsregression
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also in: regression
content_tags.statistics[0]the tagger's words
content_tagsprincipal component analysis
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content_tags.statistics[1]the tagger's words
content_tagseigen-decomposition of the covariance matrix
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content_tags.statistics[2]the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[3]the tagger's words
content_tagscoefficient of determination
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content_tags.statistics[4]the tagger's words
content_tagsstandard deviation
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content_tags.statistics[5]the tagger's words
content_tagsmultiple regression analysis of CF on 3D position
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content_tags.statistics[6]1.0the tagger's words
content_tagsprincipal component analysis
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content_tags.statistics[7]1.0the tagger's words
content_tagscoefficient of determination
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content_tags.statistics[8]1.0the tagger's words
content_tagserror magnitude between predicted and measured CF
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content_tags.statistics[9]1.0the tagger's words
content_tagsIn most reconstructed nuclei characteristic frequency ran along the dorsal-ventral axis, low frequencies dorsally and high frequencies ventrally.
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content_tags.findings[0]the tagger's words
content_tagsA linear rather than logarithmic function best described the frequency map in the large majority of individual cases.
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content_tags.findings[1]the tagger's words
content_tagsThe first principal component of the combined dataset accounted for 72% of the variance.
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content_tags.findings[2]the tagger's words
content_tagsLarge prediction errors and a low r2 suggest the SON frequency map is less precise than the tonotopy of other auditory brainstem nuclei in the same species.
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content_tags.findings[3]the tagger's words
content_tagsWithin isofrequency slices the residual scatter in CF was not related to medial-lateral or anterior-posterior position, so the imprecision is not confined to a particular subregion.
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content_tags.findings[4]the tagger's words
content_tagsNeurons with onset and with sustained response patterns are intermingled throughout the nucleus rather than occupying separate territories.
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content_tags.findings[5]the tagger's words
content_tagsEach of the two response types forms a complete tonotopic map, and the two maps are arranged similarly.
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content_tags.findings[6]the tagger's words
content_tagsSustained responders were more sharply tuned than onset responders.
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content_tags.findings[7]the tagger's words
content_tagscharacteristic frequency runs linearly from low dorsally to high ventrally in the SON
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content_tags.findings[8]1.0the tagger's words
content_tagsa linear relation fitted 15 of 19 individual nuclei, a log relation two, and two showed none
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content_tags.findings[9]1.0the tagger's words
content_tagsin the best individual case, 85% of CF variation was accounted for by position
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content_tags.findings[10]1.0the tagger's words
content_tagsacross the standardised nucleus the fit was significant but not strong, r-squared 0.62
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content_tags.findings[11]1.0the tagger's words
content_tagsthe first principal component accounted for 72% of the variance
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content_tags.findings[12]1.0the tagger's words
content_tagsthe map is less precise than the tonotopy of other auditory brainstem nuclei in the same species
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content_tags.findings[13]1.0the tagger's words
content_tagswithin isofrequency slices, position carried no information about CF
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content_tags.findings[14]1.0the tagger's words
content_tagsonset and sustained responders both show a dorsal-low to ventral-high tonotopic organisation
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content_tags.findings[15]1.0the tagger's words
content_tags73% of isolated neurons were sustained, 25% onset and 2% suppressed
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content_tags.findings[16]1.0the tagger's words
content_tagssustained responders are more frequency-selective than onset responders
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content_tags.findings[17]1.0the tagger's words
content_tagscharacteristic frequency (CF)
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content_tags.definitions[0]the tagger's words
content_tagsresponse threshold
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content_tags.definitions[1]the tagger's words
content_tagscenter of mass of recording sites
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content_tags.definitions[3]the tagger's words
content_tagstonotopic map
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also in: tonotopic map
content_tags.definitions[4]the tagger's words
content_tagscharacteristic frequency
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content_tags.definitions[5]1.0the tagger's words
content_tagsresponse threshold
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content_tags.definitions[6]1.0the tagger's words
content_tagsQ10
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also in: Q10
content_tags.definitions[7]1.0the tagger's words
content_tagsQ10
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also in: Q10
content_tags.variables[3]the tagger's words
content_tagsresponse threshold
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content_tags.variables[4]the tagger's words
content_tagsbody temperature
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content_tags.variables[5]the tagger's words
content_tagscharacteristic frequency
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content_tags.variables[6]1.0the tagger's words
content_tagsnormalized distances
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content_tags.variables[7]1.0the tagger's words
content_tagsresponse patterns
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content_tags.variables[8]1.0the tagger's words
content_tagsnormalized medial-lateral coordinate
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content_tags.variables[9]the tagger's words
content_tagsnormalized dorsal-ventral coordinate axis
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content_tags.variables[10]the tagger's words
content_tagsnormalized anterior-posterior coordinate
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content_tags.variables[11]the tagger's words
content_tags310 multiple-units and 349 isolated neurons; CF 90 to 5,000 Hz — size of the recorded sample and the range of characteristic frequencies encountered
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content_tags.quantities[0]the tagger's words
content_tagsSeventy-three percent — proportion of isolated neurons with sustained (primary-like or pauser-like) responses, against 25% onset
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content_tags.quantities[1]the tagger's words
content_tags72% — variance in the combined location-plus-CF dataset captured by the first principal component
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content_tags.quantities[2]the tagger's words
content_tagsr 2 = 0.62, P < 0.01 — strength of the linear relation between CF and 3D location in the standardized nucleus
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content_tags.quantities[3]the tagger's words
content_tags85% — variation in CF explained by recording location in the best single case
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content_tags.quantities[4]the tagger's words
content_tags60% — variability in the standardized dataset accounted for by the multilinear regression, taken as the precision of the map
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[5]the tagger's words
content_tagsCF = 0.01 + 0.32 / + 4.09 v − 0.55 a — the fitted predictive equation giving characteristic frequency in kHz from normalized 3D coordinates
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[6]the tagger's words
content_tagsr-squared 0.62 for the standardised nucleus — how much CF variation position accounts for - directly comparable with Muniak2014's 0.87 for mouse DCN, since the operationalisation is the same
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[7]1.0the tagger's words
content_tags85% — the best individual case
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[8]1.0the tagger's words
content_tags72% on the first principal component — how one-dimensional the arrangement is - Muniak2014 reported 75% by the same method
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[9]1.0the tagger's words
content_tags310 multiple-units and 349 isolated neurons, CF from 90 to 5,000 Hz — the sample and its frequency span
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[10]1.0the tagger's words
content_tags73% sustained, 25% onset, 2% suppressed — response type proportions
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_tags12 to 75 sites per animal — sampling density
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_tagstwo SDs above spontaneous rate — the stipulated threshold criterion
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_tagstwo-thirds of peak voltage — the stipulated cut for counting multi-unit events
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

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