Tonotopic organization of human auditory cortex

Tonotopic organization of human auditory cortex Humphries Liebenthal Binder

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Tonotopic organization of human auditory cortex 2010 NeuroImage 10.1016/j.neuroimage.2010.01.046 Humphries2010-tonotopic-organization-of-human-auditory-cor Humphries2010-tonotopic-organization-of-human-auditory-cor.txt, the project's staged copy 2f6d257a4e7b8dd32b96147bda27a8441ab7fa63 not-redistributable neuroscience

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0 spans located by searching the stored text for the quote's exact words, allowing any whitespace between them the spatial arrangement of frequency-selective responses in human auditory cortex Quotation withheld because this record is not cleared for redistribution. how many tonotopic gradients human auditory cortex contains, how they are oriented relative to HG, and how far they vary across subjects Quotation withheld because this record is not cleared for redistribution. two mirror-symmetric gradients running perpendicular to HG rather than along it, a third smaller gradient on lateral posterior STG, and a methodological diagnosis of why the previous literature disagreed Quotation withheld because this record is not cleared for redistribution. existence Quotation withheld because this record is not cleared for redistribution. tonotopic organization of human auditory cortex Quotation withheld because this record is not cleared for redistribution. fmri fmri Quotation withheld because this record is not cleared for redistribution. mechanisms cortical folding pushes the anterior and posterior high-frequency zones close together in 3-D, so volume-based smoothing and inter-subject variation blur them into one, hiding the perpendicular gradients from earlier studies Quotation withheld because this record is not cleared for redistribution. mechanisms flattening the superior temporal plane onto a surface model pulls the buried sulcal regions apart, which is what makes the perpendicular topology visible duplicate of: mechanisms[3] Quotation withheld because this record is not cleared for redistribution. cortical folding puts the anterior and posterior high-frequency zones physically close together, so volume-based analysis and smoothing blur them into one duplicate of: mechanisms[0] Quotation withheld because this record is not cleared for redistribution. flattening the superior temporal plane separates regions that are adjacent in 3-D, making gradients perpendicular to the gyrus visible Quotation withheld because this record is not cleared for redistribution. sparse sampling: the scanner runs only in a 1.6 s silent gap after the 8 s tone sequence, so scanner noise never overlaps the stimulus Quotation withheld because this record is not cleared for redistribution. stochastic alternation between two- and three-tone bursts holds the cortical response up, since auditory neurons adapt after onset Quotation withheld because this record is not cleared for redistribution. tone ratios chosen so the composite produces no harmonic relationship and no missing-fundamental pitch Quotation withheld because this record is not cleared for redistribution. best frequency is interpolated between the six tested bands by weighting adjacent conditions, giving finer resolution than the design has Quotation withheld because this record is not cleared for redistribution. landmark-based surface alignment rather than volumetric alignment, for spatial precision in the temporal lobe Quotation withheld because this record is not cleared for redistribution. non-parametric permutation testing because eight subjects will not support a parametric group test Quotation withheld because this record is not cleared for redistribution. several sessions per subject, to buy single-subject statistical power and a test-retest estimate at the same time Quotation withheld because this record is not cleared for redistribution. The core/belt boundary drawn on the homology map cannot be established from frequency responses alone and is an estimate imported from macaque anatomy and human cytoarchitecture. what this limits: the core/belt boundary drawn on the map: THE EXACT BOUNDARY BETWEEN CORE AND BELT CANNOT BE DETERMINED BASED SOLELY ON FREQUENCY RESPONSE, THUS THIS LINE IS AN ESTIMATE GUIDED BY THE KNOWN TOPOGRAPHICAL ORGANIZATION IN THE MACAQUE Quotation withheld because this record is not cleared for redistribution. The absence of a gradient corresponding to macaque area RT may be a sensitivity failure of the stimuli rather than evidence that the area is absent in humans. what this limits: the claimed absence of an RT-like gradient: THE LACK OF FREQUENCY-SELECTIVE ACTIVATION IN THIS REGION COULD BE DUE TO LOW SENSITIVITY TO THE STIMULI USED IN THE CURRENT STUDY Quotation withheld because this record is not cleared for redistribution. The sample was small enough to force non-parametric permutation testing for the group analysis. what this limits: the group significance test, and the paper says what it did about it: Quotation withheld Quotation withheld because this record is not cleared for redistribution. the tested range truncates the tuning estimate, so points preferring 200 or 6400 Hz were excluded from that analysis what this limits: tuning estimates at the edges of the tested range: POINTS WERE ALSO EXCLUDED IF THEIR LARGEST RESPONSE WAS TO THE LOWEST (200 Hz) OR HIGHEST (6400 Hz) FREQUENCY Quotation withheld because this record is not cleared for redistribution. an absence the authors decline to resolve: no anterior RT-like gradient, which may be the stimuli or may be the species what this limits: the same scope as limitations[1], left open: ALTERNATIVELY, IT IS POSSIBLE THAT AREA RT DOES NOT EXIST IN HUMANS - a sensitivity failure and a real absence are not distinguished Quotation withheld because this record is not cleared for redistribution. the sharper-tuning reading is undercut by the authors themselves, because the effect tracks overall response strength what this limits: the sharper-tuning reading: THESE PATTERNS GENERALLY APPEAR TO BE CORRELATED WITH THE OVERALL STRENGTH OF THE RESPONSE Quotation withheld because this record is not cleared for redistribution. the core/belt boundary cannot be fixed from frequency response alone, so the line drawn is an estimate imported from other evidence what this limits: the same scope as limitations[0] - the core/belt line, imported from macaque and cytoarchitecture rather than measured here Quotation withheld because this record is not cleared for redistribution. apparent parietal operculum activation is an artifact of volume-to-surface mapping, not a finding what this limits: the parietal operculum activation, which is not a finding: THIS ACTIVATION IS PRIMARILY AN ARTIFACT OF THE MAPPING BETWEEN VOLUME AND SURFACE SPACES DUE TO PARTIAL VOLUME EFFECTS Quotation withheld because this record is not cleared for redistribution. high variability across subjects and hemispheres in where the low-frequency border between gradients falls what this limits: any claim about where the border between gradients falls: THE LOCATION OF THE LOW-FREQUENCY BORDER BETWEEN THE TWO GRADIENTS VARIED IN RELATION TO HG, with A HIGH DEGREE OF VARIABILITY ACROSS SUBJECTS AND ACROSS HEMISPHERES Quotation withheld because this record is not cleared for redistribution. inter-session variability attributed partly to head movement and partly to unmeasured attention and alertness what this limits: voxel counts and preferred frequency across sessions: SOME INTER-SESSION VARIABILITY WAS SEEN IN THE NUMBER OF SIGNIFICANTLY ACTIVATED VOXELS AND THEIR PREFERRED FREQUENCY Quotation withheld because this record is not cleared for redistribution. eight subjects, seven of them male what this limits: the population every map describes: FMRI DATA WERE COLLECTED FROM 8 RIGHT-HANDED SUBJECTS (7 MALE, 1 FEMALE; AGES 20-37 YEARS) Quotation withheld because this record is not cleared for redistribution. no task was performed, so attentional state during scanning is uncontrolled by design what this limits: the attentional state the tonotopic maps were measured in: SUBJECTS WERE INSTRUCTED TO LIE STILL AND LISTEN TO THE TONES. NO SPECIFIC TASK WAS PERFORMED DURING THE EXPERIMENT Quotation withheld because this record is not cleared for redistribution. the core/belt field layout, used as the template against which a human fMRI map is read and labelled Quotation withheld because this record is not cleared for redistribution. retinotopy used as the contrast case for how a receptive-field gradient behaves at an areal boundary Quotation withheld because this record is not cleared for redistribution. the intensity of every tone sequence normalised to A-weighted 76.3 dB eliminates: a loudness difference between sequences - otherwise a tonotopic response could be a response to how loud each frequency happened to be · outcome (stated by the tagger, no quotation): held Quotation withheld because this record is not cleared for redistribution. cortical folding cortical folding Quotation withheld because this record is not cleared for redistribution. physical proximity physical proximity Quotation withheld because this record is not cleared for redistribution. tonotopy - a gradient of preferred frequency mirroring the cochlea; explicitly a map of one stimulus dimension, not of any other response property Quotation withheld because this record is not cleared for redistribution. gradient - as a field bounded by high and low bands, not as a line between two poles; the paper says this reading is itself a source of the literature's disagreement Quotation withheld because this record is not cleared for redistribution. frequency-selective - defined operationally as a significant F test across the six tested frequencies, carrying no claim about tuning shape Quotation withheld because this record is not cleared for redistribution. tuning bandwidth - here a ratio of preferred to non-preferred BOLD amplitude, which the paper scare-quotes and then declines to read as tuning width Quotation withheld because this record is not cleared for redistribution. core and belt - cytoarchitectonic and connectional divisions, explicitly not divisions defined by the frequency map being measured Quotation withheld because this record is not cleared for redistribution. corresponds to / homologous to - used hedged throughout for human-macaque field matches, never as identity Quotation withheld because this record is not cleared for redistribution. method: fMRI with sparse sampling, six multi-tone frequency bands from 200 to 6400 Hz, analysed on flattened cortical surfaces with landmark-based alignment a second reading DIFFERING from the recorded value. Recorded: fmri — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides. Quotation withheld because this record is not cleared for redistribution. phenomenon: tonotopy - a cortical gradient of preferred frequency mirroring the cochlea a second reading DIFFERING from the recorded value. Recorded: tonotopic organization of human auditory cortex — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides. Quotation withheld because this record is not cleared for redistribution. replicates → Formisano et al. (2003); Talavage et al. (2004); Woods et al. (2009) Quotation withheld because this record is not cleared for redistribution. contradicts → Formisano et al. (2003), on the orientation of the gradients relative to HG differs in: the ORIENTATION of the gradients relative to HG: THESE GRADIENTS ARE ORIENTED APPROXIMATELY ALONG THE AXIS OF HG, WHEREAS THE GRADIENTS WE OBSERVED WERE PERPENDICULAR TO THE LONG AXIS OF HG Quotation withheld because this record is not cleared for redistribution. contradicts → Woods et al. (2009), on labelling the posterior lateral gradient as parabelt differs in: where the parabelt is and whether it is tonotopic at all: THERE IS LITTLE EVIDENCE FOR TONOTOPIC ORGANIZATION IN THE PARABELT OF MONKEYS, and RECENT HUMAN CYTOARCHITECTONIC DATA SUGGEST THAT THE PARABELT IS LOCATED MORE VENTRALLY IN THE UPPER BANK OF THE SUPERIOR TEMPORAL SULCUS Quotation withheld because this record is not cleared for redistribution. partially-supports → Formisano et al. 2003: two mirror-symmetric gradients, but running along the axis of HG Quotation withheld because this record is not cleared for redistribution. refines → Talavage et al. 2004: three gradients described as narrow lines between high and low poles Quotation withheld because this record is not cleared for redistribution. supports → Woods et al. 2009: one low-frequency area on HG flanked by two high-frequency areas Quotation withheld because this record is not cleared for redistribution. contradicts → Woods et al. 2009: the posterior lateral field is parabelt differs in: which area the same patch of cortex IS - the two papers show A SIMILAR DIAGRAM and label it differently: AREA CL here, POSTERIOR PARABELT there, and the criterion offered is that IN THE MONKEY, THIS GRADIENT DOES NOT EXTEND INTO ADJACENT AREAS; THEREFORE, ITS BOUNDARIES ARE EASILY DETERMINED Quotation withheld because this record is not cleared for redistribution. contradicts → the early two-frequency studies: a single gradient running along the axis of HG differs in: the analysis method, named as decisive across the whole prior literature: ONE THAT WE BELIEVE IS CRITICAL IS THAT Quotation withheld Quotation withheld because this record is not cleared for redistribution. qualifies → their own reading that these central regions are more sharply tuned differs in: which side of the difference moves: the sharper tuning is DRIVEN PRIMARILY BY INCREASES IN RESPONSE TO THE PREFERRED FREQUENCY RATHER THAN DECREASES IN THE RESPONSE TO THE NON-PREFERRED FREQUENCIES, so the pattern IS CORRELATED WITH THE OVERALL STRENGTH OF THE RESPONSE and need not be sharper tuning Quotation withheld because this record is not cleared for redistribution. supports → macaque evidence that area CM prefers higher than average frequencies without a gradient Quotation withheld because this record is not cleared for redistribution. tonotopy —same-as→ cochleotopy tonotopy cochleotopy strength of the relation: 1 · note: given as alternate names in one clause, unhedged - a positive control for the join key Quotation withheld because this record is not cleared for redistribution. frequency-selective response —operationalised-as→ F test frequency-selective response F test strength of the relation: 1 Quotation withheld because this record is not cleared for redistribution. surface-based —contrasted-with→ volume-based analysis surface-based volume-based analysis differs in: THE AXIS IS THE ANALYSIS METHOD, AND IT IS OFFERED AS THE EXPLANATION OF THE DISAGREEMENT. 'Quotation withheld' The authors name the one difference they think decides the result out of many they could have named - which is the analyst decision this project exists to make legible. · strength of the relation: 1 · note: the contrast the paper says explains the whole literature's disagreement Quotation withheld because this record is not cleared for redistribution. cortical folding —causes→ physical proximity cortical folding physical proximity direction on the measure: increases · strength of the relation: 0.95 Quotation withheld because this record is not cleared for redistribution. posterior lateral gradient —same-as→ area CL posterior lateral gradient area CL strength of the relation: 0.65 — the record marks this weak · note: most likely corresponds to Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the gradient 'MOST LIKELY corresponds to' area CL; same-as is hedged. Quotation withheld because this record is not cleared for redistribution. posterior lateral gradient —distinct-from→ posterior parabelt posterior lateral gradient posterior parabelt strength of the relation: 0.85 · note: a labelling dispute with Woods et al., settled by appeal to a different evidence type Quotation withheld because this record is not cleared for redistribution. core —distinct-from→ belt core belt differs in: THE AXIS IS WHAT DEFINES THE BOUNDARY. 'Quotation withheld' The two regions are not distinguished by the property the paper measures - they are distinguished by a different property entirely, so a tonotopic result cannot confirm or refute the division it is described against. · strength of the relation: 1 · note: the boundary is stipulated from cytoarchitecture and connectivity, not from the phenomenon being mapped Quotation withheld because this record is not cleared for redistribution. retinotopic gradients —contrasted-with→ receptive field gradients in auditory cortex retinotopic gradients receptive field gradients in auditory cortex strength of the relation: 1 · note: a structural contrast between sensory systems sharing almost no vocabulary Quotation withheld because this record is not cleared for redistribution. general linear model general linear model Quotation withheld because this record is not cleared for redistribution. F-test F-test Quotation withheld because this record is not cleared for redistribution. multiple comparison correction multiple comparison correction Quotation withheld because this record is not cleared for redistribution. ANOVA ANOVA Quotation withheld because this record is not cleared for redistribution. permutation test permutation test about this quote: QUOTE RECOVERED FROM THE PAPER'S OWN CHARACTERS. The recorded quote was LETTERS-ONLY IDENTICAL to this stretch of the staged text but not an exact substring of it - spaces, hyphens, ligature glyphs or mathematical operators differed, so it could never anchor. This is the file's own text between the same two letters: not produced by a rule about the damage, copied out of the file. Round-trip checked (the recovered span is letters-only equal to the original) and length-checked. The text as originally recorded is kept in evidence as extracted. Quotation withheld because this record is not cleared for redistribution. correlation correlation Quotation withheld because this record is not cleared for redistribution. F test across six frequency conditions F test across six frequency conditions Quotation withheld because this record is not cleared for redistribution. cluster thresholding cluster thresholding Quotation withheld because this record is not cleared for redistribution. non-parametric permutation test non-parametric permutation test Quotation withheld because this record is not cleared for redistribution. between-session correlation coefficient between-session correlation coefficient Quotation withheld because this record is not cleared for redistribution. Two mirror-symmetric frequency gradients run anterior-posterior from a low-frequency zone on lateral Heschl's gyrus out to high-frequency zones in front of and behind it. Quotation withheld because this record is not cleared for redistribution. These principal gradients lie perpendicular to HG, overturning the prevailing assumption that they run parallel to it. Quotation withheld because this record is not cleared for redistribution. A third, smaller gradient was found on the lateral posterior superior temporal gyrus. Quotation withheld because this record is not cleared for redistribution. The human maps are argued to be closely homologous to nonhuman primate auditory cortex, with the two main gradients corresponding to macaque A1 and R. Quotation withheld because this record is not cleared for redistribution. Individual tonotopic maps were highly reproducible across separate scanning sessions. Quotation withheld because this record is not cleared for redistribution. Inspection of unnormalized data suggests the regional differences are driven mainly by a larger response to the preferred frequency rather than a smaller response to non-preferred ones. about this quote: QUOTE RECOVERED FROM THE PAPER'S OWN CHARACTERS. The recorded quote was LETTERS-ONLY IDENTICAL to this stretch of the staged text but not an exact substring of it - spaces, hyphens, ligature glyphs or mathematical operators differed, so it could never anchor. This is the file's own text between the same two letters: not produced by a rule about the damage, copied out of the file. Round-trip checked (the recovered span is letters-only equal to the original) and length-checked. The text as originally recorded is kept in evidence as extracted. Quotation withheld because this record is not cleared for redistribution. two mirror-symmetric gradients run roughly perpendicular to HG, not parallel as previously assumed Quotation withheld because this record is not cleared for redistribution. a low-frequency zone on lateral HG is flanked anteriorly and posteriorly by high-frequency zones in every subject Quotation withheld because this record is not cleared for redistribution. a third, smaller gradient runs along lateral posterior STG, mainly in the left hemisphere Quotation withheld because this record is not cleared for redistribution. frequency-selective responses were confined to the superior temporal plane in both hemispheres Quotation withheld because this record is not cleared for redistribution. maps are highly reproducible across sessions within a subject Quotation withheld because this record is not cleared for redistribution. the unnormalized data suggest the difference is driven mainly by a larger preferred response rather than a smaller non-preferred one Quotation withheld because this record is not cleared for redistribution. no distinct frequency gradient in area CM, agreeing with macaque evidence for an overall high-frequency preference there Quotation withheld because this record is not cleared for redistribution. prior imaging results and their interpretations varied considerably Quotation withheld because this record is not cleared for redistribution. four previous studies reached four different conclusions, one of them denying the gradients are gradients at all WHOSE RESULT THIS IS: one study reported - THE EVIDENCE FOR THIS ROW ATTRIBUTES THE WORK TO SOMEBODY ELSE. The cited name is the subject of a result verb and the sentence carries no first-person marker, so the finding, quantity or definition is the CITED author's rather than this record's. The claim is true and the quote is verbatim - what is wrong is whose result the corpus says it is. Nothing edited, nothing withdrawn: the row is marked so a page can name the source. Where the record is a commentary or review, its claim relations layer usually already names the paper being discussed - the record knows at one layer and forgets at another. Quotation withheld because this record is not cleared for redistribution. tuning bandwidth tuning bandwidth Quotation withheld because this record is not cleared for redistribution. frequency-selective response frequency-selective response Quotation withheld because this record is not cleared for redistribution. tonotopy tonotopy Quotation withheld because this record is not cleared for redistribution. frequency-selective response frequency-selective response Quotation withheld because this record is not cleared for redistribution. frequency of maximum response frequency of maximum response Quotation withheld because this record is not cleared for redistribution. mean BOLD signal mean BOLD signal Quotation withheld because this record is not cleared for redistribution. 8 — number of subjects scanned, with sex split and age range Quotation withheld because this record is not cleared for redistribution. 392 trials total — trials per scanning session, from 7 runs of 56 trials Quotation withheld because this record is not cleared for redistribution. .834 (SD = 0.083) — mean correlation of a subject's tonotopy map between scanning sessions, i.e. test-retest stability Quotation withheld because this record is not cleared for redistribution. 20 to 30 mm 3 — range of the per-subject cluster-size threshold used for multiple-comparison correction about this quote: QUOTE RECOVERED FROM THE PAPER'S OWN CHARACTERS. The recorded quote was LETTERS-ONLY IDENTICAL to this stretch of the staged text but not an exact substring of it - spaces, hyphens, ligature glyphs or mathematical operators differed, so it could never anchor. This is the file's own text between the same two letters: not produced by a rule about the damage, copied out of the file. Round-trip checked (the recovered span is letters-only equal to the original) and length-checked. The text as originally recorded is kept in evidence as extracted. Quotation withheld because this record is not cleared for redistribution. 76.3 dB — A-weighted intensity to which every tone sequence was normalised Quotation withheld because this record is not cleared for redistribution. 9.6 seconds — trial length: an 8-second sound followed by a 1.6-second silent gap in which the scan was acquired Quotation withheld because this record is not cleared for redistribution. 200, 400, 800, 1600, 3200, 6400 Hz — the six centre frequencies tested Quotation withheld because this record is not cleared for redistribution. 8 right-handed subjects, 7 male and 1 female, ages 20-37 — the sample Quotation withheld because this record is not cleared for redistribution. .834, SD 0.083 — mean between-session correlation of the tonotopy maps Quotation withheld because this record is not cleared for redistribution. better than 20 dB HL from 250 to 8000 Hz — audiometric inclusion criterion Quotation withheld because this record is not cleared for redistribution. 20 to 30 mm3 — cluster threshold, which varied by subject because smoothness was estimated per subject Quotation withheld because this record is not cleared for redistribution. 8 s stimulus then a 1.6 s scanning gap, 9.6 s per trial — the sparse-sampling timing Quotation withheld because this record is not cleared for redistribution. 3000 Hz — Formisano's highest frequency, offered as the reason that study may have missed medial anterior high-frequency cortex Quotation withheld because this record is not cleared for redistribution. 3600 Hz, 4 Hz presentation rate — Woods' ceiling and rate, against 6400 Hz and 10 Hz here Quotation withheld because this record is not cleared for redistribution.

The full text of this article is not redistributable, so it is not included here. The evidence sentences are withheld with it; the tag layer and its grounding status remain. The article itself is at https://doi.org/10.1016/j.neuroimage.2010.01.046.