Functional organization of mouse auditory cortex in response to stimulus complexity and brain state — TEI-tagged view

Afrashteh, Jafari, Sun, Kyweriga, Mohajerani · 2022 · neuroscience · doi:10.1101/2022.08.11.503675 · 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 (pdftext); every evidence string verified as an exact substring of that text before merging. Replication status is 'untested' in the external sense: no independent replication is reported. The paper does demonstrate internal reproducibility of its maps within a session, across three weeks, and between animals ('These maps are highly replicable across weeks and between animals.'), which is a within-study consistency check rather than a replication of a prior finding. method uses a label outside the suggested list because wide-field mesoscale calcium imaging is neither electrophysiology nor fMRI. This is a bioRxiv preprint, not peer reviewed, as the header stamped on every page states. State effects reported include better high-frequency sensitivity awake and shrunken fast-FM regions under anaesthesia ('In the awake versus anesthetized state, the amplitude of tonotopic maps was greater in response to high frequencies (i.e., better hearing sensitivity).'). 2026-08-14 and 2026-08-15 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
AI 8 fields · standard deviation 3 fields · bootstrap 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 UF AI8 auditory cortex19 calcium-imaging10
Method, as described: wide-field-calcium-imaging

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 and FM-sweep-rate maps in mouse auditory cortex— “The present study aimed to assess the mouse auditory cortex (AC) responses under varying conditions of stimulus complexity and brain state.”
claim-typeexistence— “Three new regions responsive to high-frequency tones and four new gradients responsive to frequency modulations (FMs) were identified.”
method — evidence— “In this study, we used wide-field Ca2+ imaging through a large cranial window in a transgenic mouse that expresses the genetically encoded calcium indicator GCaMP6 (Nakai et al., 2001; Iadecola, 2004; Tian et al., 2009; Chen et al., 2013; Cramer et al., 2019).”
mechanismIsoflurane reduces cochlear blood flow, blocks acetylcholine receptors and enhances GABA-gated currents, which raises medial olivocochlear efferent inhibition and so depresses auditory sensitivity in the anesthetized state— “The possible mechanisms underlying these impacts are changes in cochlear blood flow (CBF) (Nakashima et al., 2003; Xiao et al., 2014), the blockage of acetylcholine (ACh) receptors (Dilger et al., 1992), and enhanced gamma-aminobutyric acid (GABA)-gated channel currents (Ming et al., 2001), which lead to increased inhibition mediated by…mechanisms[0]
mechanismBehaviourally relevant high frequencies are over-represented in the mouse auditory cortex because the mouse hearing range extends much further upward, which is why the newly found regions are all high-frequency ones— “This finding suggests that the AC tonotopic maps are oriented more toward high frequencies than low frequencies, which is consistent with the wider spectrum of high frequencies in the mouse hearing range (Heffner, 1980).”mechanisms[1]
contradicts Issa et al. (2017), which reported no difference between upward and downward FM sweep maps— “Despite the report of no difference between upward and downward FM sweep maps in a previous study (Issa et al., 2017), we found that the center of the slow sweep rate responsive area in AI shifts 350 µm rostrally between upward and downward FM sweep maps.”claim_relations[0]
extends Issa et al. (2017), whose imaging window was too small to resolve map borders— “Among a few studies of mouse AC organization in encoding FM sweep rates, the Issa et al. (Issa et al., 2017) study suffers from a small imaging window, which makes it difficult to delineate the structure of the resulting maps near the imaging borders.”claim_relations[1]
extends Romero et al. (2020), whose VPAF is placed differently— “The VPAF region identified by Romero et al. (Romero et al., 2020) is in a different location and is primarily responsive to low frequencies.”claim_relations[2]
builds-on Issa et al. (2014) landmark-registration procedure— “The registration deformation field was constructed using landmarks from each experimental session and target landmarks as previously described in Issa et al. (Issa et al., 2014).”claim_relations[3]
senseAI (primary auditory cortex) - AS ONE AREA in the classical map, versus as an area containing TWO high-frequency regions, which is what optical calcium imaging reported— “Using multiscale optical Ca2+ imaging, it was shown that the AI area includes two rostral regions that process highfrequency tones”senses[0]
senseAI (primary auditory cortex) - AND AS SPLIT DIFFERENTLY AGAIN by a third technique: flavoprotein autofluorescence with anatomy makes the dorsal high-frequency part a discrete region— “A recent publication on AC tonotopy, using a combination of flavoprotein autofluorescence imaging and anatomical techniques, demonstrated that AI includes two areas that respond to high-frequency tones and that the dorsal part of the AI high-frequency area is a discrete region from the rest of the AI”senses[1]
senseAI (primary auditory cortex) - AND AS PART OF A WHOLLY NEW LAYOUT: widefield calcium imaging in awake mice is reported as introducing a new layout for auditory cortex— “Using widefield calcium imaging in awake mice, Romero et al. (Romero et al., 2020) also introduced a new layout for AC”senses[2]
senserevision of the map - attributed by the paper to TECHNIQUE rather than to anatomy: 'technological advancements in neuroimaging and recording have led to significant updates in the tonotopic organization of the belt area'— “However, during the past decades, technological advancements in neuroimaging and recording have led to significant updates in the tonotopic organization of the belt area”senses[3]
controla deliberately LARGE craniotomy, 3 to 4 mm across, over the auditory cortical regions — held — rules out that the map is shaped by the size of the recording window rather than by the cortex - a small window would truncate the functional organisation being measured— “In this study, a large craniotomy (3-4 mm in diameter) was created over auditory cortical regions to rule out all uncertainties associated with the size of the recording window.”controls[0]
controlresponsive pixels were identified first and used to exclude the unresponsive ones from analysis — held — rules out that unresponsive pixels dilute the signal and flatten the functional map— “Responsive pixels were identified and used to exclude unresponsive pixels.”controls[1]
auditUF —part-of→ AI — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but measurements 'suggest' UF should be considered subparts of AAF and AI; hedged.— “For instance, electrophysiological measurements suggest the “UF” area should be considered as subparts of the AAF and AI”concept_edges[0]
differs inUF —part-of→ AI — WHICH TECHNIQUE IS ASKED. 'ELECTROPHYSIOLOGICAL MEASUREMENTS suggest the UF area should be considered as SUBPARTS of the AAF and AI' - so on that instrument UF is not an area at all but pieces of two others. The record already carries THREE readings of AI as senses, one per technique: optical calcium imaging splits it into two high-frequency regions, flavoprotein autofluorescence makes the dorsal part discrete, widefield imaging introduces a new layout entirely— “For instance, electrophysiological measurements suggest the “UF” area should be considered as subparts of the AAF and AI”concept_edges[0]
noteUF —part-of→ AI — AN AREA THAT DISSOLVES INTO PARTS OF TWO OTHERS DEPENDING ON THE METHOD. The boundary is not a fact the techniques disagree about - each technique constitutes a different partition, and all of them are current.— “For instance, electrophysiological measurements suggest the “UF” area should be considered as subparts of the AAF and AI”concept_edges[0]
weightUF —part-of→ AI — 0.7— “For instance, electrophysiological measurements suggest the “UF” area should be considered as subparts of the AAF and AI”concept_edges[0]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
outcome (stated by the tagger, no quotation)control — heldcontrols[1]
for which uselimitation — any lateralisation claim: A CRANIOTOMY WAS MADE ONLY OVER THE RIGHT AC— “In this study, a craniotomy was made only over the right AC.”limitations[0]
for which uselimitation — the sleep interpretation of the state effects: REPLICATION OF OUR FINDINGS DURING NATURAL SLEEP WILL CLARIFY WHETHER CHANGES IN HEARING SENSITIVITY RESULT FROM USING ISOFLURANE OR ARE ASSOCIATED WITH SLEEP PROCESSES— “Replication of our findings during natural sleep will clarify whether changes in hearing sensitivity result from using isoflurane or are associated with sleep processes.”limitations[1]
for which uselimitation — the same scope as limitations[1]: THE IMPACT OF SLEEP COMPARED TO ANESTHETIZED AND AWAKE STATES SHOULD BE STUDIED IN THE FUTURE— “The impact of sleep compared to anesthetized and awake states should be studied in the future.”limitations[2]
for which uselimitation — any area label in the mouse auditory cortex literature: the paper states the tonotopic organisation of the belt area has been significantly updated as neuroimaging and recording advanced— “However, during the past decades, technological advancements in neuroimaging and recording have led to significant updates in the tonotopic organization of the belt area.”limitations[3]
for which uselimitation — the UF area as a unit in the mouse auditory map: electrophysiological measurements are reported as suggesting it should be considered subparts of AAF and AI— “For instance, electrophysiological measurements suggest the “UF” area should be considered as subparts of the AAF and AI”limitations[4]
for which uselimitation — the tonotopic maps in this paper: for each frequency a threshold level was identified and used to determine the map— “For each frequency, the threshold level was identified (red squares) and used to determine the tonotopic map.”limitations[5]
replicationuntested
limitationImaging was confined to the right auditory cortex, so lateralization of vocal processing could not be assessed.— “In this study, a craniotomy was made only over the right AC.”limitations[0]
limitationIsoflurane was used as a model of the sleeping brain, and whether the state effects are anaesthetic or sleep-related is unresolved.— “Replication of our findings during natural sleep will clarify whether changes in hearing sensitivity result from using isoflurane or are associated with sleep processes.”limitations[1]
limitationThe comparison of natural sleep against awake and anaesthetized states is left to future work.— “The impact of sleep compared to anesthetized and awake states should be studied in the future.”limitations[2]
limitationThe map of this cortex has been revised repeatedly, and the paper attributes the revisions to advances in technique rather than to new anatomy.— “However, during the past decades, technological advancements in neuroimaging and recording have led to significant updates in the tonotopic organization of the belt area.”limitations[3]
limitationAn area in the classical map is reported as dissolvable into parts of two others, on the evidence of one technique.— “For instance, electrophysiological measurements suggest the “UF” area should be considered as subparts of the AAF and AI”limitations[4]
limitationThe tonotopic map rests on a per-frequency threshold identified by the experimenters.— “For each frequency, the threshold level was identified (red squares) and used to determine the tonotopic map.”limitations[5]
limitationThe paper's own account of why the map keeps changing is an account of instruments, not of findings - decades of technological advancement producing significant updates to the organisation. On that account the current map is the current technique's map, and the paper is adding a fourth.— “However, during the past decades, technological advancements in neuroimaging and recording have led to significant updates in the tonotopic organization of the belt area”limitations[6]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingThe newly identified auditory cortex regions responded mainly to higher frequencies and lay in dorsal and caudal auditory cortex.— “The newly identified regions were generally responsive to higher frequencies and were in dorsal and caudal AC regions.”content_tags.findings[0]
findingApart from a few sparse pixels, tonotopic maps were similar over time and between animals.— “Except for a few sparse pixels, the tonotopic maps were quite similar over time and between animals.”content_tags.findings[1]
findingHigh-frequency responses in AI-H and UF were larger in awake than in anesthetized mice.— “In this study, the response amplitude at high frequencies (the H region of AI and UF) was greater in the tonotopic maps from awake compared to anesthetized mice.”content_tags.findings[2]
findingUnder anesthesia the cortical area encoding slow FM sweep rates expanded and the area encoding fast sweep rates shrank across AAF, DA and UF.— “The second difference was shown in the AAF-DA-UF regions, in which the response to slower sweep rates was expanded and the response to faster sweep rates was reduced in maps from the anesthetized compared to the awake state.”content_tags.findings[3]
findingContrary to an earlier report of no difference, upward and downward FM sweep maps do differ: the slow-rate centre in AI sits about 350 micrometres more rostral for downward sweeps.— “Despite the report of no difference between upward and downward FM sweep maps in a previous study (Issa et al., 2017), we found that the center of the slow sweep rate responsive area in AI shifts 350 µm rostrally between upward and downward FM sweep maps.”content_tags.findings[4]
findingUltrasonic vocalizations drove stronger responses than pure tones specifically in the regions tuned to fast FM sweep rates (AII, DP, PAF, VPAF and a zone between UF and AAF), and not in the high-frequency tonotopic regions.— “In our study, vocal stimuli elicited a stronger response than tonal stimuli in both anesthetized and awake states, and a higher V/T ratio was obtained in the regions (AII, DP, PAF, VPAF, and a region between UF and AAF) linked to the processing of faster FM sweep rates compared to regions that respond to high frequencies.”content_tags.findings[5]
findingBest frequency changes with distance along a tonotopic gradient as a sigmoid: middle frequencies change steeply over a short stretch of cortex while low and high frequencies occupy longer stretches.— “This sigmoid function reflects steeper frequency changes in a shorter distance for middle frequencies versus lower and higher frequencies.”content_tags.findings[6]
findingEVIDENCE CORRECTED (RE-GROUNDED - THE POINTER WAS WRONG, NOT THE CLAIM) · Tonotopic maps were almost identical for averaging windows from 100 ms to 1000 ms after tone onset.— “Maps were almost identical from 100 ms after the onset of tone presentation up to 1000 ms.”content_tags.findings[7]
retractedfinding — RE-GROUNDED - THE POINTER WAS WRONG, NOT THE CLAIM. The old evidence was verbatim, anchored and correctly cut, and it contained NONE of this claim's numbers (100, 1000). The paper states the claim in one sentence elsewhere and the tagger did not use it. the project, 2026-08-23: 'WE HAVE SPENT THE DAY VERIFYING THAT QUOTES ARE REAL AND NONE OF IT ASKS WHETHER THEY ARE RELEVANT.' Confirmed by reading the claim, the old span and the candidate together; the candidate carries every number AND the claim's own content words.content_tags.findings[7]
findingA window larger than 2 mm in each axis is needed to include every region of the tonotopic organisation.— “The distance between the farthest landmarks in the tonotopic map supports the idea that a window larger than 2 mm in each axis is necessary to include all cortical regions involved in the AC tonotopic organization.”content_tags.findings[8]
quantity12 and 6 — Number of mice recorded awake and anesthetized respectively— “Awake and isoflurane-anesthetized recordings were performed for 12 and 6 mice, respectively.”content_tags.quantities[0]
quantityR2 > 0.93 and 0.90 — Adjusted R-squared of the linear fit to the inverse-sigmoid frequency-distance relation, awake and anesthetized— “The associations were highly linear (R2 > 0.93 and 0.90 for awake and anesthetized states, respectively) and confirmed the sigmoid relationship between the best frequency and distance from the low-frequency landmark in a tonotopic gradient.”content_tags.quantities[1]
quantityR2 > 0.95 and 0.87 — Fit quality of the inverse-sigmoid function for the FM sweep-rate/distance relation, awake and anesthetized— “A sigmoid function was obtained in the plots (Figures 6I and 6J for awake and anesthetized, respectively) after fitting an inverse-sigmoid function to each graph (R2 > 0.95 and 0.87 for awake and anesthetized states, respectively).”content_tags.quantities[2]
quantity1.36 and 1.51 — Median vocalization-to-tone response ratio in the USV-favouring pixels, awake and anesthetized— “The median values of the V/T ratio for the regions favoring USVs (i.e., pixels with V/T > 1) were 1.36 and 1.51 for awake and anesthetized states, respectively.”content_tags.quantities[3]
quantity2 mm and 1.9 mm — Extent of the auditory cortex tonotopic map along the rostral-caudal and dorsal-ventral axes— “The distance between the centers of the two farthest regions was 2 mm on the rostral-caudal axis and 1.9 mm on the dorsal-ventral axis.”content_tags.quantities[4]
quantityAI-L p = 0.0054, AI-H p = 0.0171, AAF-L p = 0.0181, UF p = 0.0216, DP p = 0.0165 — Median p-values for the five regions whose best frequency differed between awake and anesthetized maps— “and DP (median p-values shown in S1C table: AI-L p = 0.0054, AI-H p = 0.0171, AAF-L p = 0.0181, UF p = 0.0216, DP p = 0.0165).”content_tags.quantities[5]
evidence status notequantity — THE QUOTE CONTINUES A SENTENCE THAT THIS COPY CUTS IN TWO - a bioRxiv watermark stands between it and the words it follows, which read 'the high-frequency regions of AI, UF,'.content_tags.quantities[5]
quantity350 µm — Rostral shift of the AI slow-sweep-rate landmark in the downward compared with the upward FM map— “In downward compared to upward sweep rate maps, the center of the slow sweep rate region in AI is shifted 350 µm rostrally.”content_tags.quantities[6]
quantity25 times — Repetitions of each frequency-SPL pair per imaging session— “Each frequency-SPL pair was played 25 times per imaging session.”content_tags.quantities[7]
quantity33.3ms (30 Hz) — Temporal resolution / frame rate of the wide-field calcium imaging— “For the mesoscale calcium imaging data collection, 12-bit images were captured with 33.3ms temporal resolution (30 Hz frame rate) with a charge-coupled device (CCD) camera”content_tags.quantities[8]
variabletone frequency and sound pressure level (independent)— “Pure tones were generated at low (4 kHz), medium (11.3 kHz), and high (32 kHz) frequencies; and at low (40 dB), medium (60 dB), and high (80 dB) sound pressure levels (SPL).”content_tags.variables[1]
variablebest frequency (dependent)content_tags.variables[3]
variablepixel responsiveness (dependent)content_tags.variables[4]
variablebody temperature (control)content_tags.variables[5]
variablesound pressure level (control)content_tags.variables[6]
variablefrequency coverage (control)content_tags.variables[7]
variablesweep direction (independent)content_tags.variables[9]
variablesweep duration (independent)content_tags.variables[10]
definitioncore auditory areacontent_tags.definitions[0]
definitionbelt auditory region
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definitiontonotopic organization
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definitionfrequency sweep rate
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definitionV/T ratio
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definitionresponsive pixel
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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.
concepttonotopic organization
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concepts[0]
conceptUF
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concepts[1]
conceptAI
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concepts[2]
conceptauditory cortex
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concepts[3]
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.
Kruskal-Wallis test as written: Kruskal-Wallis test at each pixel
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chi-squared goodness-of-fit test as written: Chi-square goodness-of-fit test
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bootstrap as written: the bootstrap method
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Wilcoxon rank-sum test as written: one-sided Wilcoxon rank-sum test
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p-value p as written: Statistical significance was determined at a p-value < 0.05
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standard error of the mean SEM as written: standard error of the mean (SEM)
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regression as written: linear ridge regression
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coefficient of determination R2 as written: adjusted R2
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explained variance R2 as written: the explained variance (R2) map
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singular value decomposition SVD as written: singular value decomposition (SVD)
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standard deviation as written: mean + 2×standard deviation
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