Tonotopic Organization of the Superior Olivary Nucleus in the Chicken Auditory Brainstem — TEI-tagged view

Tabor, Coleman, Rubel, Burger · 2012 · neuroscience · doi:10.1002/cne.22807 · 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 (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.
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, 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
tonotopic9 topographic organization superior olivary nucleus characteristic frequency4 Q10 response threshold3 onset6 sustained4 principal component analysis37 multiple-unit nucleus laminaris nucleus angularis isofrequency4 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 of an inhibitory auditory brainstem nucleus— “We analyzed the topographic organization of response features of neurons in the SON of chickens.”
claim-typeexistence— “These analyses led to three main conclusions: 1) sound frequency is linearly arranged from dorsal (low frequencies) to ventral (high frequencies) in SON; 2) this tonotopic organization is less precise than the organization of the excitatory nuclei in the chicken auditory brainstem; and 3) neurons with different response patterns to pure tone…
method — evidence— “By using in vivo single- and multiple-unit recordings we classified the characteristic frequency (CF), frequency selectivity, threshold, and response patterns of SON neurons.”
object of study — evidence— “The goal of the current study is to analyze the topographic organization of 1) sound frequency and 2) physiologically identified cell types. Specifically, is the tonotopic organization of the brainstem auditory pathway preserved in the SON?”
research question — evidence— “The goal of the current study is to analyze the topographic organization of 1) sound frequency and 2) physiologically identified cell types. Specifically, is the tonotopic organization of the brainstem auditory pathway preserved in the SON?”
contribution — evidence— “These analyses led to three main conclusions: 1) sound frequency is linearly arranged from dorsal (low frequencies) to ventral (high frequencies) in SON; 2) this tonotopic organization is less precise than the organization of the excitatory nuclei in the chicken auditory brainstem; and 3) neurons with different response patterns to pure tone…
transferfrom visual neuroscience: retinotopy as the template for what a sensory map is and why it matters— “Moreover, quantitative maps provide predictive tools to determine the optimal stimulus for a particular neuron based on its location in the tissue. Using this information investigators can 1) relate anatomical, biophysical, and pharmacological characteristics of a neuron to sensory processing based on stimulus selectivity ( Smith and Rubel…transfers[0]
mechanismSON response patterns are inherited from its nucleus angularis input, which is why sustained and onset types and diverse tuning shapes appear there— “Firing patterns of SON neurons to ipsilateral tones appear to be shaped primarily by the inputs from the NA, one of two main excitatory inputs to the SON.”mechanisms[0]
mechanismthe narrow hearing range of the chicken makes the frequency-to-place function linear rather than logarithmic— “This linear relationship, as opposed to a log relationship often observed in auditory centers, is probably a consequence of the limited frequency range of hearing in chickens”mechanisms[1]
mechanismsummation of individually subthreshold spike waveforms raises the apparent threshold of multiple-unit recordings— “This increase reflects multiple-unit recordings with neuronal waveforms that individually were not detected by using our threshold criterion, but summed to form detectable waveforms at higher sound intensities.”mechanisms[2]
mechanismthe broad, non-restricted SON to nucleus laminaris projection supplies the anatomical substrate by which inhibition laterally sharpens frequency tuning— “This projection pattern suggests an anatomical substrate for lateral sharpening of frequency tuning in NL neurons.”mechanisms[3]
mechanisminhibitory neurons in this system sit in a discrete projection nucleus rather than among principal cells, which is what makes their organisation separable at all— “Inhibitory components of many neuronal circuits comprise interneurons residing alongside principal neurons ( Huang et al., 2007 ). Thus, independent evaluation of the organizing features of these circuit components is difficult. However, in the auditory brainstem, inhibitory neurons often reside in nuclei that project to one or more targets…mechanisms[4]
mechanismrecording sites are located by electrolytic lesions and electrode tracks, then normalised into a common 3D nucleus so cases can be combined— “To combine data from all 19 reconstructed SONs, locations of the recording sites were converted into normalized distances along three orthogonal axes in the SON (see Materials and Methods). These normalized coordinates allowed all recording sites to be mapped in a standardized SON.”mechanisms[5]
mechanismPCA is run first because it was unknown whether the map was linear or nonlinear, single or multiple gradients— “Because the presence and arrangement (linear or nonlinear, single or multiple gradients) of the tonotopic organization in SON were unknown, we performed principal component analysis (PCA) on the combined dataset to determine the appropriate dimension(s) to observe the possible tonotopic organization. The first principal component accounted…mechanisms[6]
mechanismSON inhibition widens dynamic range and sharpens temporal coding for sound localisation— “Inhibition from the SON expands the dynamic range and sharpens temporal coding of auditory neurons to enable sound localization”mechanisms[7]
replicates Coleman et al., 2011 (previous characterization of the chicken SON)— “These activity patterns, and the relative proportion of each response type are consistent with our previous characterization of the SON in chickens ( Coleman et al., 2011 ).”claim_relations[0]
builds-on Moiseff and Konishi, 1983; Lachica et al., 1994; Tabor et al., 2011claim_relations[1]
extends Rubel and Parks, 1975 (tonotopy of other chicken brainstem auditory nuclei)claim_relations[2]
extends the tonotopic organisation established for excitatory nuclei of the chicken auditory brainstem
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.
claim_relations[3]
confirms earlier suggestions from owl and chicken that frequency is systematically represented in the SON
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claim_relations[4]
supports the group's own earlier characterisation of SON response types and their proportions
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claim_relations[5]
contradicts the expectation that physiologically distinct SON cell types occupy distinct territories
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claim_relations[6]
sensetonotopy 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]
sensethe 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]
sensecharacteristic 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]
senseonset 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]
controleach stimulus presented twenty times in a pseudo-randomised sequence — held — rules out adaptation and order - a fixed sequence would confound the tonotopic response with position in the run
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controls[0]
notecharacteristic frequency —operationalised-as→ response threshold — CF is DEFINED through threshold, and threshold is itself defined as two SDs above spontaneous rate - a two-step operational chain
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concept_edges[1]
noteonset —contrasted-with→ sustained — and they are spatially interspersed rather than segregated, which is the paper's third conclusion
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concept_edges[2]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
for which uselimitation — 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
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limitations[0]
for which uselimitation — 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
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limitations[1]
for which uselimitation — 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
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limitations[2]
for which uselimitation — 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
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limitations[3]
for which uselimitation — the tonotopic prediction at any single site: ERROR MAGNITUDE (|CF_predicted - CF_measured|) VARIED WIDELY AT EACH POSITION ALONG THE CALCULATED TONOTOPIC AXIS
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limitations[4]
duplicate oflimitation — limitations[0]
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limitations[5]
for which uselimitation — 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
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limitations[5]
for which uselimitation — 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
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limitations[6]
for which uselimitation — the spatial claims for suppressed responders: SUPPRESSED RESPONDERS WERE ENCOUNTERED TOO INFREQUENTLY TO ASSESS THEIR SPATIAL DISTRIBUTION
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limitations[7]
for which uselimitation — 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
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limitations[8]
for which uselimitation — 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
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limitations[9]
replicationuntested
limitationThreshold 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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limitations[0]
limitationBinaural response properties of SON neurons were not assessed, so a possible contribution of the nucleus laminaris projection remains unexplored.
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limitations[1]
limitationIn 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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limitations[2]
limitationthe map is explicitly less precise than in excitatory nuclei of the same species, by both error magnitude and r-squared
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limitations[3]
limitationprediction error varies widely along the tonotopic axis rather than being uniform
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limitations[4]
limitationthe threshold criterion may not be equally sensitive across response categories, so between-category threshold comparisons are cautioned against
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limitations[5]
limitationmulti-unit thresholds run higher than single-unit thresholds because sub-threshold waveforms sum
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limitations[6]
limitationsuppressed responders were too rare to analyse spatially
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limitations[7]
limitationin most animals only a limited region of the nucleus was sampled, which is why standardisation was needed
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limitations[8]
limitationwithin an isofrequency slice, position carried no information about CF at all
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limitations[9]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingIn 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]
findingA 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]
findingThe first principal component of the combined dataset accounted for 72% of the variance.
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content_tags.findings[2]
findingLarge 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]
findingWithin 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]
findingNeurons 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]
findingEach 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]
findingSustained responders were more sharply tuned than onset responders.
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content_tags.findings[7]
findingcharacteristic frequency runs linearly from low dorsally to high ventrally in the SON
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content_tags.findings[8]
findinga linear relation fitted 15 of 19 individual nuclei, a log relation two, and two showed none
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content_tags.findings[9]
findingin the best individual case, 85% of CF variation was accounted for by position
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content_tags.findings[10]
findingacross the standardised nucleus the fit was significant but not strong, r-squared 0.62
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content_tags.findings[11]
findingthe first principal component accounted for 72% of the variance
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content_tags.findings[12]
findingthe map is less precise than the tonotopy of other auditory brainstem nuclei in the same species
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content_tags.findings[13]
findingwithin isofrequency slices, position carried no information about CF
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content_tags.findings[14]
findingonset and sustained responders both show a dorsal-low to ventral-high tonotopic organisation
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content_tags.findings[15]
finding73% of isolated neurons were sustained, 25% onset and 2% suppressed
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content_tags.findings[16]
findingsustained responders are more frequency-selective than onset responders
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content_tags.findings[17]
quantity310 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]
quantitySeventy-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]
quantity72% — variance in the combined location-plus-CF dataset captured by the first principal component
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content_tags.quantities[2]
quantityr 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]
quantity85% — variation in CF explained by recording location in the best single case
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content_tags.quantities[4]
quantity60% — variability in the standardized dataset accounted for by the multilinear regression, taken as the precision of the map
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content_tags.quantities[5]
quantityCF = 0.01 + 0.32 / + 4.09 v − 0.55 a — the fitted predictive equation giving characteristic frequency in kHz from normalized 3D coordinates
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content_tags.quantities[6]
quantityr-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
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content_tags.quantities[7]
quantity85% — the best individual case
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content_tags.quantities[8]
quantity72% on the first principal component — how one-dimensional the arrangement is - Muniak2014 reported 75% by the same method
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content_tags.quantities[9]
quantity310 multiple-units and 349 isolated neurons, CF from 90 to 5,000 Hz — the sample and its frequency span
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content_tags.quantities[10]
quantity73% sustained, 25% onset, 2% suppressed — response type proportions
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content_tags.quantities[11]
quantity12 to 75 sites per animal — sampling density
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content_tags.quantities[12]
quantitytwo SDs above spontaneous rate — the stipulated threshold criterion
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content_tags.quantities[13]
quantitytwo-thirds of peak voltage — the stipulated cut for counting multi-unit events
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content_tags.quantities[14]
variableQ10 (dependent)
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content_tags.variables[3]
variableresponse threshold (dependent)
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content_tags.variables[4]
variablebody temperature (control)
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content_tags.variables[5]
variablecharacteristic frequency
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content_tags.variables[6]
variablenormalized distances
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content_tags.variables[7]
variableresponse patterns
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content_tags.variables[8]
variablenormalized medial-lateral coordinate (independent)
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content_tags.variables[9]
variablenormalized dorsal-ventral coordinate axis (independent)
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content_tags.variables[10]
variablenormalized anterior-posterior coordinate (independent)
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content_tags.variables[11]
definitioncharacteristic frequency (CF)
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content_tags.definitions[0]
definitionresponse threshold
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content_tags.definitions[1]
definitioncenter of mass of recording sites
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content_tags.definitions[3]
definitiontonotopic map
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content_tags.definitions[4]
definitioncharacteristic 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.definitions[5]
definitionresponse threshold
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]
definitionQ10
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[7]
key-termstonotopic, topographic organization, superior olivary nucleus, characteristic frequency, Q10, response threshold, onset, sustained, isofrequency, inhibitory, GABA, nucleus laminaris, nucleus angularis, chicken, brainstem, principal component analysis, MRA, coefficient of determination, 3D reconstruction, sound localization
Concepts
terms this paper carries, each with the sentence it was cut from — the layer that joins one paper to another. Terms with no quote yet are on CONCEPTS.html.
conceptcharacteristic frequency — the frequency with the lowest response threshold
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.
concepts[3]
conceptQ10 — not defined in this paper's own words; CF divided by bandwidth 10 dB above threshold, the selectivity measure
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.
concepts[4]
conceptresponse threshold — not defined in this paper's own words; intensity at which rate exceeds spontaneous by two SDs
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.
concepts[5]
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.
regression as written: Multilinear regression analysis (MRA)
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]
principal component analysis PCA as written: Principle component analysis (PCA)
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content_tags.statistics[1]
eigen-decomposition of the covariance matrix as written: eigenvectors and eigenvalues of the covariance matrix
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]
t-test P as written: Student's t-test or r 2 correlation
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]
coefficient of determination r 2 as written: the coefficient of determination (r 2 )
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]
standard deviation SD 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[5]
multiple regression analysis of CF on 3D position
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]
principal component 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[7]
coefficient of determination
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]
error magnitude between predicted and measured CF
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[9]
Second readings
a facet read a second time, independently, from its own sentence — and whether the two readings agree
methodin 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 — 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 in an inhibitory long-range projection nucleus — a second reading DIFFERING from the recorded value; recorded: tonotopic organization of an inhibitory auditory brainstem nucleus
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alternatives.phenomenon

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