Flexible Sensory Representations in Auditory Cortex Driven by Behavioral Relevance

Flexible Sensory Representations in Auditory Cortex Driven by Behavioral Relevance Kato Gillet Isaacson

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Flexible Sensory Representations in Auditory Cortex Driven by Behavioral Relevance 2015 Neuron 10.1016/j.neuron.2015.10.024 Kato2015-flexible-sensory-representations-in-auditory Kato2015-flexible-sensory-representations-in-auditory.txt, the project's staged copy 6cb7869f521d840ead59106d64f1a4ca6df23c44 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 how the cortical representation of a sound changes when the sound is repeatedly heard and means nothing, and changes again when the same sound becomes useful Quotation withheld because this record is not cleared for redistribution. what happens to auditory cortical representations over days of passive experience, which has been hard to ask because the same cells could not be followed that long Quotation withheld because this record is not cleared for redistribution. passive daily exposure suppresses pyramidal responses and recruits inhibition through SOM cells, and task engagement reverses it by suppressing those same SOM cells Quotation withheld because this record is not cleared for redistribution. mechanism Quotation withheld because this record is not cleared for redistribution. experience-dependent habituation of auditory cortex sound representations Quotation withheld because this record is not cleared for redistribution. electrophysiology electrophysiology Quotation withheld because this record is not cleared for redistribution. mechanisms Repeated passive exposure raises SOM interneuron activity, and the added SOM-mediated inhibition suppresses layer 2/3 pyramidal responses to the experienced tone. Quotation withheld because this record is not cleared for redistribution. mechanisms Task engagement suppresses SOM cell firing, releasing pyramidal cells from local inhibition and thereby enlarging the representation of the behaviorally relevant sound. Quotation withheld because this record is not cleared for redistribution. habituation is an active process that recruits inhibition rather than a simple withdrawal of excitation Quotation withheld because this record is not cleared for redistribution. SOM cells are the source, since their tone-evoked excitation grows with the same experience that suppresses everything else Quotation withheld because this record is not cleared for redistribution. PV cells cannot be the source because their excitation falls with experience like the pyramidal cells they would have to inhibit Quotation withheld because this record is not cleared for redistribution. the enhancement is specific to the sustained part of the response, matching where the suppression of pyramidal activity is strongest Quotation withheld because this record is not cleared for redistribution. sound-guided behavior had the opposite effect on SOM cells compared to L2/3 pyramidal cells, with fewer SOM cells excited by the tone during the behaving block Quotation withheld because this record is not cleared for redistribution. filtering sustained sounds while preserving onsets is proposed as the function, since onsets carry the more urgent information Quotation withheld because this record is not cleared for redistribution. Spike inference from calcium signals is sensitivity-limited, so conclusions were re-checked on raw dF/F. what this limits: the spike-inference analyses, with the check stated: ALTHOUGH DATA ANALYSES USING SPIKE INFERENCE CAN BE AFFECTED BY SIGNAL-TO-NOISE RATIO, OUR CONCLUSIONS WERE IDENTICAL WHEN DF/F VALUES WERE USED Quotation withheld because this record is not cleared for redistribution. Residual neuropil contamination of cellular signals cannot be excluded. what this limits: the single-cell specificity of the signals: WE CANNOT EXCLUDE SOME RESIDUAL CONTAMINATION OF CELLULAR SIGNALS BY NEUROPIL RESPONSES, though the conclusions do not depend on the degree of background subtraction Quotation withheld because this record is not cleared for redistribution. The use of prolonged tones and a sound-offset detection task may itself explain the divergence from earlier task-engagement studies. what this limits: the comparison with the earlier task literature: the use of prolonged tones and a sound-offset detection task may itself explain the divergence Quotation withheld because this record is not cleared for redistribution. Other mechanisms contributing to auditory habituation are not excluded. what this limits: the sufficiency of the SOM account: WE DO NOT EXCLUDE THE POSSIBILITY THAT OTHER MECHANISMS COULD CONTRIBUTE TO AUDITORY HABITUATION UNDER DIFFERENT CONDITIONS Quotation withheld because this record is not cleared for redistribution. PV cell excitation after habituation is too sparse for a fine comparison during behaviour what this limits: the PV comparison between behaving and passive conditions: THE VERY SPARSE EXCITATION OF PV CELLS FOLLOWING HABITUATION MADE IT DIFFICULT TO PERFORM A FINE COMPARISON Quotation withheld because this record is not cleared for redistribution. the use of prolonged tones and an offset detection task may itself account for differences from earlier task-engagement studies that used transient sounds what this limits: the disagreement with the task-engagement literature: WE CANNOT RULE OUT THE POSSIBILITY THAT THE USE OF PROLONGED TONES AND A SOUND-OFFSET DETECTION TASK MAY ACCOUNT FOR DIFFERENCES BETWEEN OUR FINDINGS AND PREVIOUS STUDIES Quotation withheld because this record is not cleared for redistribution. the effect of silencing shrank when the same experiment was repeated over days, which is read as compensation downstream what this limits: any silencing result, including this one, as a measure of necessity: WHEN THE CONTRALATERAL CORTICAL SILENCING EXPERIMENTS WERE REPEATED OVER SEVERAL DAYS THE SHIFT OF THE PSYCHOMETRIC CURVE BECAME PROGRESSIVELY SMALLER, SUGGESTING COMPENSATORY CHANGES IN SUBCORTICAL CIRCUITS Quotation withheld because this record is not cleared for redistribution. no increase in passive representations appeared after training, contrary to earlier learning studies what this limits: the comparison with the learning literature: WE DID NOT OBSERVE AN INCREASE IN SENSORY REPRESENTATIONS IN THE PASSIVE CONDITION AFTER TRAINING, contrary to four cited studies Quotation withheld because this record is not cleared for redistribution. the whole GABAergic population showed only variable effects, so the result depends on splitting it by subtype what this limits: the subtype-specific result, which the population-level data did not show: DAILY TONE EXPERIENCE PRODUCED VARIABLE EFFECTS ACROSS THIS CELL POPULATION ... MODEST EFFECTS AT THE POPULATION LEVEL COULD ARISE IF EXPERIENCE HAD DIFFERENT AND POTENTIALLY OPPOSING ACTIONS ON PARTICULAR SUBTYPES Quotation withheld because this record is not cleared for redistribution. Cre-dependent expression of a calcium indicator, which is what makes the cell-type split possible at all Quotation withheld because this record is not cleared for redistribution. a macroscopic tonotopic map used to place the cellular imaging field inside the right area Quotation withheld because this record is not cleared for redistribution. deconvolution of fluorescence into an inferred rate, with the conclusions then rechecked without it Quotation withheld because this record is not cleared for redistribution. a psychometric threshold for tone detection, used as the readout of a cortical silencing experiment Quotation withheld because this record is not cleared for redistribution. sounds of three different durations delivered in RANDOMLY INTERLEAVED trials eliminates: temporal expectation - a fixed duration lets the animal learn when the sound ends, so a response at offset would be an anticipated event rather than a sensory one · outcome (stated by the tagger, no quotation): held Quotation withheld because this record is not cleared for redistribution. habituation habituation Quotation withheld because this record is not cleared for redistribution. inhibition inhibition Quotation withheld because this record is not cleared for redistribution. A WHOLE RESPONSE CLASS EXISTS ONLY IN THE AWAKE ANIMAL: inhibitory responses were present in about a quarter of cells awake and in NONE under anaesthesia, and prolonged tones evoked only transient onset excitation Quotation withheld because this record is not cleared for redistribution. so a literature built on anaesthetised recordings cannot contain the observation this paper is about, whatever it looked for Quotation withheld because this record is not cleared for redistribution. THE DISAGREEMENT WITH PRIOR WORK IS EXPLAINED BY WHAT THE OLDER INSTRUMENT COULD SEE: unit recordings are prone to pick up cells with high firing rates, so earlier experiments may have been sampling the very populations that do not habituate Quotation withheld because this record is not cleared for redistribution. and a second explanation is offered beside it, that interneuron activity is attenuated under anaesthesia, with neither chosen Quotation withheld because this record is not cleared for redistribution. AVERAGING OVER A POPULATION HID THE EFFECT: the whole GABAergic population gave variable results, and the finding only appeared once the two interneuron subtypes were imaged separately Quotation withheld because this record is not cleared for redistribution. THE DESIGN REMOVES THE CONFOUND RATHER THAN CORRECTING FOR IT: detecting the OFFSET of a tone puts the measurement before the licking and before the reward, so neither can contaminate the sound-evoked response Quotation withheld because this record is not cleared for redistribution. AND THE CAUSAL EFFECT SHRINKS WITH REPETITION: the psychometric shift from silencing got progressively smaller over days, so the measured size of a perturbation depends on how many times it has been done Quotation withheld because this record is not cleared for redistribution. the frequency-specificity check is what separates a change in the representation from a change in the animal - the effect appears only at the experienced frequency Quotation withheld because this record is not cleared for redistribution. and the conclusion is held against three analysis choices - inferred spikes against raw fluorescence, the amount of neuropil subtracted, and imaging against electrophysiology Quotation withheld because this record is not cleared for redistribution. claim_type: mechanism a second reading reaching the same value. Recorded: mechanism — note: a second reading reaching the same value from a different sentence. Not applied - a person decides. Quotation withheld because this record is not cleared for redistribution. method: chronic two-photon imaging of GCaMP6s in identified pyramidal, layer 4, PV and SOM populations across five days of tone exposure and during alternating behaving and passive blocks, plus optogenetic cortical silencing during a sound-offset detection task 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. Quotation withheld because this record is not cleared for redistribution. phenomenon: habituation of a cortical sensory representation to a repeated, meaningless stimulus, and its reversal when the stimulus becomes behaviourally relevant a second reading DIFFERING from the recorded value. Recorded: experience-dependent habituation of auditory cortex sound representations — 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. contradicts → Bao et al. (2001), Kilgard and Merzenich (1998) and Zhang et al. (2001) on adult A1 being refractory to passive experience differs in: the state of the animal and the recording: ACUTE RECORDINGS FROM ANESTHETIZED ANIMALS report A1 representations LARGELY REFRACTORY TO EXPERIENCE-DEPENDENT CHANGES IN ADULTHOOD, against chronic imaging in the awake mouse here Quotation withheld because this record is not cleared for redistribution. contradicts → Noreña et al. (2006) on extensive sound exposure in cat auditory cortex differs in: the LEVEL of the pathway at which the change appears: UNLIKE THE RESULTS WE SHOW, THIS EXTENSIVE SOUND EXPOSURE GENERATED A LOSS OF RESPONSIVENESS AT THE LEVEL OF THALAMOCORTICAL INPUT Quotation withheld because this record is not cleared for redistribution. builds-on → Pfeffer et al. (2013) on SOM-to-PV inhibitory connectivity Quotation withheld because this record is not cleared for redistribution. contradicts → the view that long-term changes in adult auditory cortex require associative conditioning differs in: whether reinforcement is required: long-term changes ARE BELIEVED TO REQUIRE ASSOCIATIVE CONDITIONING - PAIRING TONES OF A PARTICULAR FREQUENCY WITH FOOT SHOCKS OR STIMULATION OF SUBCORTICAL NEUROMODULATORY SYSTEMS - where this paper uses passive exposure alone Quotation withheld because this record is not cleared for redistribution. reconciles → earlier electrophysiological reports of no adult experience effect, by attributing them to which cells unit recording samples and to anaesthesia differs in: which cells were recorded: ONE POSSIBILITY REFLECTS DIFFERENCES IN THE CELL POPULATIONS STUDIED - WE FOUND THAT EXPERIENCE HAD UNIQUE EFFECTS ON SOUND REPRESENTATIONS OF L2/3 PYRAMIDAL CELLS Quotation withheld because this record is not cleared for redistribution. qualifies → cat studies reporting loss of thalamocortical responsiveness after months of exposure, which differ in where the change sits differs in: the exposure duration and the level affected: TONE BURSTS MAINTAINED WITHOUT INTERRUPTION FOR 1.5-5 MONTHS, and the effect was A LOSS OF RESPONSIVENESS AT THE LEVEL OF THALAMOCORTICAL INPUT rather than a cortical change Quotation withheld because this record is not cleared for redistribution. qualifies → the inconsistent literature on whether auditory cortex is needed for simple sound detection, by measuring a threshold shift rather than an all-or-none loss Quotation withheld because this record is not cleared for redistribution. supports → the parallel result in visual cortex, where silencing raises contrast detection threshold by a similar factor Quotation withheld because this record is not cleared for redistribution. extends → studies of task engagement in auditory cortex that reached opposite conclusions and did not separate cell types Quotation withheld because this record is not cleared for redistribution. habituation —causes→ inhibition habituation inhibition direction on the measure: increases · strength of the relation: 1 · note: twenty minutes a day for five days, and the whole effect is carried by a rise in inhibition Audit 2026-09-28: DOUBTFUL - read against its span and kept, but this span describes reduced representations after exposure; the increase in inhibition is stated in a neighbouring sentence. Quotation withheld because this record is not cleared for redistribution. L4 —distinct-from→ L2/3 L4 L2/3 strength of the relation: 1 · note: the thalamorecipient layer barely moves, which is the argument that the change is made in cortex rather than inherited Quotation withheld because this record is not cleared for redistribution. SOM —contrasted-with→ PV SOM PV strength of the relation: 1 · note: SOM cells are the most excitable population before any experience, at a ratio of 12.5 against 0.34 for PV cells Quotation withheld because this record is not cleared for redistribution. behavior —causes→ SOM behavior SOM strength of the relation: 1 · note: AND THE CIRCUIT RUNS BACKWARDS: engagement suppresses SOM responses in 62.3 percent of cells against 1.4 percent the other way Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says behaviour had the opposite effect on SOM cells; 'causes SOM' drops the effect. Quotation withheld because this record is not cleared for redistribution. behaving —contrasted-with→ passive behaving passive differs in: THE AXIS IS THE PROPORTION OF CELLS FAVOURING EACH STATE, AND NEITHER SIDE IS EMPTY. 'Quotation withheld.' A statement that behaviour enhances responses is true of half the cells and false of a fifth - so the population claim and the cell claim are different claims, and only the percentages distinguish them. · strength of the relation: 1 · note: excited cells rise during the task, and 51.6 percent of them are stronger while behaving against 19.4 percent while passive Quotation withheld because this record is not cleared for redistribution. silencing —causes→ psychometric silencing psychometric strength of the relation: 1 · note: AND THE EFFECT SHRINKS WITH REPETITION: the shift got progressively smaller over days, read as compensation downstream - so the size of a perturbation depends on how often it has been applied Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says contralateral silencing shifted the psychometric curve; 'causes psychometric' is terse. Quotation withheld because this record is not cleared for redistribution. onset —contrasted-with→ sustained onset sustained strength of the relation: 1 · note: the proposed function: filter out the sustained, keep the onset, because onsets carry the urgent information Quotation withheld because this record is not cleared for redistribution. t-test t-test Quotation withheld because this record is not cleared for redistribution. Wilcoxon signed-rank test Wilcoxon signed-rank test Quotation withheld because this record is not cleared for redistribution. ANOVA ANOVA Quotation withheld because this record is not cleared for redistribution. ROC analysis ROC analysis Quotation withheld because this record is not cleared for redistribution. correlation correlation Quotation withheld because this record is not cleared for redistribution. one-way ANOVA one-way ANOVA Quotation withheld because this record is not cleared for redistribution. SEM SEM Quotation withheld because this record is not cleared for redistribution. Five days of brief daily passive exposure to the same tone sharply reduced the fraction of layer 2/3 pyramidal cells excited by that tone and roughly doubled the fraction inhibited by it. Quotation withheld because this record is not cleared for redistribution. Cortical habituation is an active process that recruits inhibition rather than a passive loss of excitatory drive. Quotation withheld because this record is not cleared for redistribution. Layer 4 thalamorecipient neurons showed only weak, balanced changes with the same exposure, so the habituation in layer 2/3 is generated in cortex rather than inherited from subcortical inputs. Quotation withheld because this record is not cleared for redistribution. The experience-dependent increase in inhibition of L2/3 pyramidal cells appears to reflect selectively enhanced sound-evoked activity in SOM interneurons. Quotation withheld because this record is not cleared for redistribution. When mice performed a sound-offset detection task with the habituated tone, pyramidal cell excitatory representations grew again, partially undoing habituation. Quotation withheld because this record is not cleared for redistribution. Photoinactivating auditory cortex raised the sound-detection threshold from about 41 to 52 dB. Quotation withheld because this record is not cleared for redistribution. Under anaesthesia inhibitory responses were entirely absent and tones evoked only transient onset or offset excitation. Quotation withheld because this record is not cleared for redistribution. awake responses to prolonged tones were varied - sustained, ramping, transient, and inhibitory Quotation withheld because this record is not cleared for redistribution. and the same patterns appeared in electrophysiology, so they are not an artefact of imaging Quotation withheld because this record is not cleared for redistribution. the transient shift was quantified with an onset response index built from the first and last second of the tone Quotation withheld because this record is not cleared for redistribution. five days of twenty minutes' exposure cut excited cells from 19.9 to 6.1 percent, doubled inhibited cells, and reduced their ratio tenfold Quotation withheld because this record is not cleared for redistribution. which makes habituation an active recruitment of inhibition rather than a withdrawal of excitation Quotation withheld because this record is not cleared for redistribution. THE FREQUENCY-SPECIFICITY CHECK separates a change in the representation from a change in the animal - the effect appears only at the experienced frequency Quotation withheld because this record is not cleared for redistribution. and the conclusions held with raw fluorescence instead of inferred spikes and across different amounts of neuropil subtraction Quotation withheld because this record is not cleared for redistribution. layer 4 thalamorecipient cells barely changed, and their excited-to-inhibited ratio not at all Quotation withheld because this record is not cleared for redistribution. which is the argument that the change is made in cortex rather than inherited from below Quotation withheld because this record is not cleared for redistribution. AVERAGING OVER THE WHOLE INHIBITORY POPULATION HID THE EFFECT, and opposing actions on subtypes was the suspected reason Quotation withheld because this record is not cleared for redistribution. PV cells moved with the pyramidal cells, which rules them out as the source of the new inhibition Quotation withheld because this record is not cleared for redistribution. SOM cells moved the opposite way, with 58 percent showing larger responses on day five against 9.8 percent smaller Quotation withheld because this record is not cleared for redistribution. and the enhancement was strongest in the sustained component, matching where pyramidal suppression was strongest Quotation withheld because this record is not cleared for redistribution. SOM cells were already the most excitable population before any experience, at a ratio of 12.5 against 0.34 for PV cells Quotation withheld because this record is not cleared for redistribution. THE TASK IS BUILT SO THE CONFOUND CANNOT ARISE: detecting the offset of a tone puts the measurement before the licking and before the reward Quotation withheld because this record is not cleared for redistribution. silencing the contralateral cortex raised the detection threshold by about 10 dB, from 40.7 to 51.9 Quotation withheld because this record is not cleared for redistribution. and illuminating the ipsilateral cortex in the same mice did nothing, ruling out the light itself Quotation withheld because this record is not cleared for redistribution. AND THE EFFECT SHRANK WITH REPETITION over days, read as compensation in subcortical circuits Quotation withheld because this record is not cleared for redistribution. so subcortical structures alone suffice for detection while cortex improves sensitivity Quotation withheld because this record is not cleared for redistribution. and the size of the shift matches what silencing visual cortex does to contrast detection Quotation withheld because this record is not cleared for redistribution. no increase in passive representations followed training, contrary to earlier learning studies Quotation withheld because this record is not cleared for redistribution. but during task engagement excited cells rose and 51.6 percent of them were stronger while behaving against 19.4 percent while passive Quotation withheld because this record is not cleared for redistribution. with block order and baseline fluorescence both checked Quotation withheld because this record is not cleared for redistribution. layer 4 did not move during behaviour either, localising the engagement effect to cortex as well Quotation withheld because this record is not cleared for redistribution. and SOM cells moved the opposite way again, suppressed in 62 percent of cells against 1.4 percent enhanced Quotation withheld because this record is not cleared for redistribution. PV cells could not be compared finely because their excitation was too sparse after habituation Quotation withheld because this record is not cleared for redistribution. THE DISAGREEMENT WITH EARLIER WORK IS ATTRIBUTED TO WHAT UNIT RECORDING SAMPLES - cells with high firing rates, which may be the very populations that do not habituate Quotation withheld because this record is not cleared for redistribution. and another possibility is the difference in cortical activity between anesthetized and awake animals, since the activity of GABAergic interneurons can be strongly attenuated in the anesthetized state Quotation withheld because this record is not cleared for redistribution. cat studies after months of exposure changed the thalamocortical input instead, which is a different location for the effect Quotation withheld because this record is not cleared for redistribution. the proposed function is filtering sustained sounds while preserving onsets, which carry the more urgent information Quotation withheld because this record is not cleared for redistribution. prior work on task engagement had reached opposite conclusions and had not separated cell types Quotation withheld because this record is not cleared for redistribution. and the prolonged tones and offset task are named as a possible reason for the difference Quotation withheld because this record is not cleared for redistribution. the standing view had been that adult auditory cortex changes only with associative conditioning Quotation withheld because this record is not cleared for redistribution. difficulty in monitoring the dynamics of sensory representations in the same animal over long time scales has hampered investigation of the long-term effects of passive sensory experience Quotation withheld because this record is not cleared for redistribution. areas were told apart by tonotopic pattern before any cell was imaged, and all experiments were confined to the 10 to 20 kHz region Quotation withheld because this record is not cleared for redistribution. about 31 percent of pyramidal cells responded to at least one frequency Quotation withheld because this record is not cleared for redistribution. the exposure protocol was 200 trials a day of 5 to 9 second tones for five consecutive days Quotation withheld because this record is not cleared for redistribution. onset response index onset response index Quotation withheld because this record is not cleared for redistribution. change index (CI) change index (CI) Quotation withheld because this record is not cleared for redistribution. habituation habituation Quotation withheld because this record is not cleared for redistribution. change index change index Quotation withheld because this record is not cleared for redistribution. deconvolution deconvolution Quotation withheld because this record is not cleared for redistribution. TRF TRF Quotation withheld because this record is not cleared for redistribution. engagement in the task engagement in the task Quotation withheld because this record is not cleared for redistribution. interneuron subtypes interneuron subtypes Quotation withheld because this record is not cleared for redistribution. ipsilateral silencing ipsilateral silencing Quotation withheld because this record is not cleared for redistribution. excitation excitation Quotation withheld because this record is not cleared for redistribution. inhibition inhibition Quotation withheld because this record is not cleared for redistribution. change index change index Quotation withheld because this record is not cleared for redistribution. psychometric psychometric Quotation withheld because this record is not cleared for redistribution. TRF TRF Quotation withheld because this record is not cleared for redistribution. fraction of excited cells fraction of excited cells Quotation withheld because this record is not cleared for redistribution. fraction of inhibited cells fraction of inhibited cells Quotation withheld because this record is not cleared for redistribution. ten-fold — reduction in the ratio of excited to inhibited layer 2/3 pyramidal cells after five days of passive tone exposure Quotation withheld because this record is not cleared for redistribution. 10 dB — rise in the sound level detection threshold caused by acute optogenetic silencing of contralateral auditory cortex Quotation withheld because this record is not cleared for redistribution. 58.0% — proportion of SOM cells whose tone-evoked responses were significantly larger on Day 5 than Day 1 (versus 9.8% showing the opposite) Quotation withheld because this record is not cleared for redistribution. 19.9 ± 2.7% / 6.1 ± 1.1% — fraction of layer 2/3 pyramidal cells excited by the exposed tone on Day 1 versus Day 5 Quotation withheld because this record is not cleared for redistribution. 0% — inhibitory responses under anaesthesia, against about a quarter of cells awake - a response class the anaesthetised preparation cannot produce Quotation withheld because this record is not cleared for redistribution. ten-fold — the drop in the ratio of excited to inhibited cells over five days of passive exposure Quotation withheld because this record is not cleared for redistribution. 58.0% — SOM cells with larger responses on day five, against 9.8 percent smaller - the opposite direction to every other population Quotation withheld because this record is not cleared for redistribution. 12.5 ± 0.4 — excited-to-inhibited ratio in SOM cells before any experience, against 0.34 in PV cells Quotation withheld because this record is not cleared for redistribution. 51.9 ± 4.0 dB — detection threshold with cortex silenced, against 40.7 dB without Quotation withheld because this record is not cleared for redistribution. 31.0 ± 4.4% — pyramidal cells responding to at least one frequency, from 1480 cells in 8 mice 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.neuron.2015.10.024.