A synaptic and circuit basis for corollary discharge in the auditory cortex

Schneider, Nelson, Mooney · Nature · 2014 · neuroscience · 10.1038/nature13724 · record Schneider2014-a-synaptic-and-circuit-basis-for-corollary-d
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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. The XML source renders citations as bare superscript numbers with surrounding spaces (e.g. "...visual cortical responses 38 ,"), so relation targets are described by reference number rather than author name — the article text itself contains no reference list above the quoted material. method='electrophysiology' covers the core evidence (in vivo intracellular current clamp plus multi-electrode arrays); the study also uses two-photon calcium imaging of GCaMP6s-labelled M2-to-auditory-cortex neurons and optogenetic/pharmacological manipulation, which the single-value method facet cannot express. 2026-08-13, 2026-08-15 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.
8 quotations on this page contain a reference number where the paper printed a superscript. This journal style prints citations as raised numerals — parameters12,13 — and the extraction flattened them into the line, so the sentence reads parameters 12 , 13 . The digits are citation markers and not part of what the author wrote. The quotes keep them for the same reason they keep every other extraction fault: a quote is reproduced exactly or it proves nothing.
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Other fields define these differently the same word, defined another way in another discipline
standard deviation 3 fields · masking 2 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 18 concept tags on this paper, none is grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.
corollary discharge auditory cortex M2 PV+ excitatory neurons reafference membrane potential thalamocortical ChR2 TTX antidromic presynaptic depression onset slope visual cortex locomotion GCaMP6s input resistance masking Motor-related signals from secondary motor cortex excite PV+ interneurons in auditory cortex, which postsynaptically inhibit excitatory cells and thereby suppress both spontaneous and sound-evoked activity Transient motor-driven dampening of sensitivity to self-predictable quiet sounds preserves headroom for responding to unexpected loud stimuli corollary discharge signals M2 ACtx cells movement laser stimulation membrane potential variability peak tone-evoked response body temperature t-test Student’s t-test, paired t-test and Two-sampled t-test Kolmogorov-Smirnov test tests for normality (Kolmogorov-Smirnov) significance level (alpha) ANOVA two-factor ANOVA with repeated measures interaction effect standard deviation population mean +/− s.d. values percentile threshold linear regression / slope linearly related electrophysiology

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second readingalternativesan independent re-reading of a facet, recorded whether or not it agrees
relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
contributioncontributionwhat the paper claims to add
controlcontrolsan experiment run to eliminate a rival explanation
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
object of studyobject_of_studywhat the paper is about — the thing it investigates
phenomenonphenomenonthe effect or regularity under study
research questionresearch_questionthe question the paper sets itself
sensesensesa distinct use of a term, quoted where the paper fixes it
transfertransfersa method or concept carried in from another field

The paper

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alternatives 3 · claim_relations 8 · claim_type 1 · concept_edges 3 · content_tags 69 · contribution 1 · controls 3 · limitations 7 · mechanisms 7 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 5 · transfers 2

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 3 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studyhow a copy of the motor command reaches auditory cortex and what it does to hearing during movement
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object_of_studythe paper's words
research_questionwhich synapses and which circuit carry the motor-related suppression of auditory cortical responses
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research_questionthe paper's words
contributionidentifies the suppression as postsynaptic inhibition from local PV interneurons, driven by a specific set of secondary motor cortex neurons that project to auditory cortex and fire before movement
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contributionthe paper's words
claim_typemechanism
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claim_typethe paper's words
phenomenonmotor-related corollary discharge in auditory cortex
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
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mechanisms[0]the tagger's words
mechanismsmechanisms
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status: the paper's own words: may reflect
mechanisms[1]the tagger's words
mechanismsmotor-related signals excite PV interneurons, which postsynaptically inhibit excitatory cells and suppress both spontaneous and evoked activity
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duplicate of: mechanisms[0]
mechanisms[2]1.0the tagger's words
mechanismsthe inhibitory cells fire well before movement onset and before the excitatory cells fall below baseline, in the order M2 then PV then excitatory
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mechanisms[3]1.0the tagger's words
mechanismsthe movement-related voltage change reverses near the chloride equilibrium potential, which is what identifies the inhibition as postsynaptic
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mechanisms[4]1.0the tagger's words
mechanismsthe suppression may dampen sensitivity to predictable quiet sounds while preserving responsiveness to unexpected loud ones
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mechanisms[5]0.8the tagger's words
mechanismsthe mechanism differs from the disinhibition proposed for locomotion-driven gain increases in visual cortex
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mechanisms[6]1.0the tagger's words
limitationsThe evidence establishes postsynaptic inhibition but does not exclude a presynaptic contribution.
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what this limits: the mechanism of the suppression: the features INDICATE ... POSTSYNAPTIC INHIBITION, BUT DO NOT EXCLUDE THE POSSIBILITY THAT PRESYNAPTIC INHIBITION IS ALSO INVOLVED
limitations[0]the tagger's words
limitationsAnimals were not randomised to groups and investigators were not blinded to group allocation.
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what this limits: every group comparison in the paper: RANDOMIZATION WAS NOT USED TO DETERMINE HOW ANIMALS WERE ALLOCATED TO EXPERIMENTAL GROUPS, NOR WERE INVESTIGATORS BLINDED TO GROUP ALLOCATION
limitations[1]the tagger's words
limitationsOptogenetic terminal stimulation could in principle recruit indirect pathways via antidromic activation (argued against by two control observations).
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what this limits: the terminal-stimulation results: OPTOGENETIC ACTIVATION OF M2 TERMINALS ... TRIGGERS ANTIDROMIC PROPAGATION ... AND THUS EXCITES OTHER TARGETS OF M2 ACtx CELLS
limitations[2]the tagger's words
limitationspostsynaptic inhibition is established but presynaptic inhibition is not excluded outright
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duplicate of: limitations[0] · what this limits: the same scope as limitations[0] - postsynaptic inhibition is established, presynaptic is not excluded
limitations[3]1.0the tagger's words
limitationsM2 could also act indirectly through neuromodulatory cell groups, which the design does not rule out
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what this limits: the direct-pathway interpretation: M2 COULD ALSO ACT INDIRECTLY THROUGH OR IN CONCERT WITH NEUROMODULATORY CELL GROUPS
limitations[4]1.0the tagger's words
limitationsvocalizations were always accompanied by other movements, so no vocalization-specific signal could be isolated
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what this limits: any vocalisation-specific reading: VOCALIZATIONS WERE ALWAYS ACCOMPANIED BY OTHER HEAD AND BODY MOVEMENTS, and the membrane potential dynamics WERE INDISTINGUISHABLE FROM THOSE OBSERVED DURING HEAD MOVEMENTS
limitations[5]1.0the tagger's words
limitationsexcitatory cells were identified by intrinsic and firing properties, with staining in only a subset
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what this limits: results reported by cell class: cells were CLASSIFIED AS PUTATIVE EXCITATORY CELLS BASED ON THEIR INTRINSIC PROPERTIES ... AND, IN A SUBSET OF NEURONS, INTRACELLULAR STAINING
limitations[6]1.0the tagger's words
transferscorollary discharge as a framework, imported from systems where the motor copy is a single identified neuron
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transfers[0]0.8the tagger's words
transfersterminal stimulation used to bypass a pathway rather than to activate a region, isolating where an effect arises
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transfers[1]1.0the tagger's words
controlsrandomisation of animals to experimental groups - NOT USED, and the paper says so
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eliminates: that a difference between groups is a difference in which animals were put in them · outcome (stated by the tagger, no quotation): absent-declared-unreported
controls[0]0.85the tagger's words
controlsblinding of investigators to group allocation - NOT USED, and the paper says so
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eliminates: experimenter expectancy in allocating and handling the animals · outcome (stated by the tagger, no quotation): absent-declared-unreported
controls[1]0.85the tagger's words
controlsblinding to movement/rest and laser on/off obtained from AUTOMATION rather than procedure - membrane potential mean and variance, tone-evoked responses and movement onset/offset all calculated automatically
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eliminates: the experimenter's judgement in scoring the conditions that carry the paper's main claim - the blinding the previous sentence says was not done by allocation is recovered here by taking the human out of the measurement · outcome (stated by the tagger, no quotation): held
controls[2]0.85the tagger's words
sensescorollary discharge here means a motor-related signal that PRECEDES movement, which is what distinguishes it from sensory reafference caused by the movement
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senses[0]1.0the tagger's words
sensesthe local share of the suppression is quantified rather than asserted: about 60 percent arises inside auditory cortex, the rest elsewhere in the pathway
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senses[1]1.0the tagger's words
sensesM2 ACtx cells are a subset defined by their projection target, identified by an intersectional strategy, and behave like the general M2 population
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senses[2]1.0the tagger's words
sensesmovement here spans locomotion, head and body movements, grooming and vocalization, all producing indistinguishable auditory cortical dynamics
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senses[3]1.0the tagger's words
sensesthe head-fixed treadmill preparation was validated against the freely behaving one before being used for the optogenetics
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senses[4]1.0the tagger's words
alternativesclaim_type: mechanism
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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.
alternatives.claim_typethe tagger's words
alternativesmethod: sharp-electrode intracellular recording in freely behaving and head-fixed mice, with optogenetic activation and silencing, multielectrode arrays and two-photon imaging
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a second reading DIFFERING from the recorded value. Recorded: electrophysiology — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.methodthe tagger's words
alternativesphenomenon: suppression of auditory cortical responses before and during movement
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a second reading DIFFERING from the recorded value. Recorded: motor-related corollary discharge in auditory cortex — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationscontradicts → the disinhibitory mechanism proposed for locomotion-dependent increases in visual cortical responses (ref. 38), the negative-rescaling account of state-dependent auditory cortical changes (ref. 23) and presynaptic depression from state-dependent thalamic activity (ref. 30)
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claim_relations[0]the tagger's words
claim_relationsbuilds-on → the prior anatomical finding that the M2-to-auditory-cortex projection is almost completely ipsilateral (ref. 20)
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claim_relations[1]the tagger's words
claim_relationsextends → observations of movement-related modulation of auditory cortex in monkeys and humans (refs. 1, 10, 11, 18, 37), for which a direct motor-cortical role had not been tested
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claim_relations[2]the tagger's words
claim_relationscontradicts → sensory reafference and peripheral masking as explanations for movement-related auditory suppression
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claim_relations[3]1.0the tagger's words
claim_relationsdistinguishes-from → the disinhibitory account of locomotion-driven gain increases in visual cortex
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differs in: THE AXIS IS THE CIRCUIT MECHANISM, WITH BOTH NAMED. 'This mechanism contrasts with A DISINHIBITORY MECHANISM implicated in locomotion-dependent increases in visual cortical responses, WITH A parallel negative rescaling of excitatory and inhibitory synaptic drive that has been advanced to account for STATE-DEPENDENT CHANGES.' Two accounts of behaviour-driven cortical change that predict opposite things about inhibition - so a result reported as 'state-dependent modulation' is compatible with either until the inhibitory drive is measured.
claim_relations[4]1.0the tagger's words
claim_relationsextends → anesthetized-mouse work showing motor cortical synapses suppress tone-evoked auditory responses
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claim_relations[5]0.9the tagger's words
claim_relationssupports → corollary discharge accounts described across species from insects to humans
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claim_relations[6]0.9the tagger's words
claim_relationsbuilds-on → anatomical tracing showing the M2-to-auditory projection is almost entirely ipsilateral
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claim_relations[7]1.0the tagger's words
concept_edgesPV+ —causes→ excitatory neurons
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strength of the relation: 1 · note: SIGN: suppressive, and the firing order M2 then PV then excitatory matches the proposed direction Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span reports PV+ rates rising while excitatory rates fell; causes is loose here.
concept_edges[3]1.0the tagger's words
concept_edgespresynaptic depression —operationalised-as→ onset slope
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strength of the relation: 1 · note: CRITERION, and the assay was VALIDATED FIRST: 20 Hz stimulation was shown to change the slope before the absence of a change was claimed as a null
concept_edges[6]1.0the tagger's words
concept_edgesvisual cortex —contrasted-with→ auditory cortex
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strength of the relation: 1 · note: SIGN REVERSAL: the same manipulation raises gain in one sensory cortex and lowers it in the other
concept_edges[9]1.0the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[0]the tagger's words
content_tagsKolmogorov-Smirnov test
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content_tags.statistics[1]the tagger's words
content_tagssignificance level (alpha)
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content_tags.statistics[2]the tagger's words
content_tagsANOVA
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also in: ANOVA
content_tags.statistics[3]the tagger's words
content_tagsinteraction effect
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content_tags.statistics[4]the tagger's words
content_tagsstandard deviation
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content_tags.statistics[5]the tagger's words
content_tagspercentile threshold
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content_tags.statistics[6]the tagger's words
content_tagslinear regression / slope
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content_tags.statistics[7]the tagger's words
content_tagslinearly related
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content_tags.statistics[8]0.7the tagger's words
content_tagsJust before and during locomotion, head movements, grooming and other movements, auditory cortical excitatory neurons showed less variable membrane potentials and a slight depolarization.
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content_tags.findings[0]the tagger's words
content_tagsThese membrane potential changes began on average more than 200 ms before locomotion started and persisted about as long after it ended.
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content_tags.findings[1]the tagger's words
content_tagsBecause the cortical changes precede movement onset by hundreds of milliseconds, they cannot be caused by sensory reafference from the movement, and could reflect a motor-related signal.
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content_tags.findings[2]the tagger's words
content_tagsUnlike visual cortex, where locomotion boosts responses, tone-evoked synaptic responses of auditory cortical excitatory neurons were significantly reduced during movement.
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content_tags.findings[3]the tagger's words
content_tagsRoughly 60 percent of the movement-related suppression of tone-evoked responses originates within the auditory cortex itself rather than upstream, since optogenetically evoked thalamocortical responses were suppressed in proportion to tone responses.
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content_tags.findings[4]the tagger's words
content_tagsThe rising slope of thalamocortical EPSPs was unchanged between movement and rest, ruling out presynaptic depression of thalamic terminals as the cause of suppression.
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content_tags.findings[5]the tagger's words
content_tagsThe movement-related membrane potential change reversed near -72 mV, close to the chloride equilibrium potential reported for mouse auditory cortical pyramidal neurons.
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content_tags.findings[6]the tagger's words
content_tagsAround movement, PV+ and other fast-spiking interneurons in auditory cortex increased their firing while putative excitatory neurons decreased theirs.
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content_tags.findings[7]the tagger's words
content_tagsThe inhibitory neurons' firing rose several hundred milliseconds before movement onset and before excitatory cells dropped below baseline, consistent with inhibition leading the suppression.
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content_tags.findings[8]the tagger's words
content_tagsExcitatory neurons in secondary motor cortex increased their firing about 870 ms before movement onset, earlier than any of the auditory cortical changes.
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content_tags.findings[9]the tagger's words
content_tagsOptogenetically activating M2 axon terminals in the auditory cortex of a resting mouse reproduced the movement signature: reduced membrane potential variability, reduced tone-evoked responses and slight depolarization.
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content_tags.findings[10]the tagger's words
content_tagsThe terminal-activation effect was not an indirect antidromic one: it survived pharmacological silencing of M2 cell bodies with TTX.
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content_tags.findings[11]the tagger's words
content_tagsThe auditory cortical change began within 7 ms of terminal stimulation, faster than the roughly 12 ms antidromic conduction time back to M2, again excluding an indirect route.
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content_tags.findings[12]the tagger's words
content_tagsSilencing ipsilateral M2 restored rest-like membrane potential dynamics in auditory cortex within about 70 ms of laser onset.
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content_tags.findings[13]the tagger's words
content_tagsTones presented after M2 silencing, while the mouse was still moving, showed a strong (~40 percent) recovery of tone-evoked responses.
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content_tags.findings[14]the tagger's words
content_tagsauditory cortical excitatory neurons are suppressed before and during movement, due in part to increased activity of local PV+ interneurons
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content_tags.findings[15]1.0the tagger's words
content_tagsthe change led locomotion by more than 200 ms and outlasted it by about as much
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content_tags.findings[16]1.0the tagger's words
content_tagstone-evoked synaptic responses were suppressed during movement, opposite to the visual cortex
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content_tags.findings[17]1.0the tagger's words
content_tagsPV and other fast-spiking cells increased firing before movement while excitatory cells decreased
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content_tags.findings[18]1.0the tagger's words
content_tagsthe firing order was PV at -805 ms, other inhibitory cells at -605 ms and excitatory cells at -490 ms relative to movement onset
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content_tags.findings[19]1.0the tagger's words
content_tagsM2 cells defined by their projection to auditory cortex behaved like the general M2 population
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content_tags.findings[20]1.0the tagger's words
content_tagsactivating M2 terminals in resting mice reproduced the movement-like dynamics
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content_tags.findings[21]1.0the tagger's words
content_tagssilencing ipsilateral M2 restored rest-like dynamics within about 70 ms, while contralateral silencing did nothing until movement stopped
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content_tags.findings[22]1.0the tagger's words
content_tagsAPPORTIONED, NOT EXCLUDED: about 60 percent of the suppression arises inside auditory cortex, measured from the slope relating optogenetic to tone suppression
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content_tags.findings[23]1.0the tagger's words
content_tagsASSAY VALIDATED BEFORE THE NULL: high-frequency stimulation was shown to reduce EPSP onset slope before the absence of such a change during movement was claimed
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content_tags.findings[24]1.0the tagger's words
content_tagsCONTROL NULL: EPSP onset slope did not differ between movement and rest, so thalamic terminals are not depressed during movement
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content_tags.findings[25]1.0the tagger's words
content_tagsCONFOUND EXCLUDED TWO WAYS: the terminal-activation effect survived TTX silencing of M2 somata and was faster than antidromic propagation
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content_tags.findings[26]1.0the tagger's words
content_tagsDECOUPLING EXPERIMENT: with ipsilateral M2 silenced the cortex returned to a rest-like state while the mouse was still moving, and tones then recovered about 40 percent of their response
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content_tags.findings[27]1.0the tagger's words
content_tagsthe decoupling separates the neural state from the movement itself and shows the state controls the gain
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content_tags.findings[28]1.0the tagger's words
content_tagssilencing M2 in a resting mouse slightly enhanced tone responses, so even baseline M2 activity is mildly suppressive
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content_tags.findings[29]1.0the tagger's words
content_tagscorollary discharge signals
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content_tags.definitions[0]the tagger's words
content_tagsM2 ACtx cells
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also in: M2 ACtx cells
content_tags.definitions[1]the tagger's words
content_tagscorollary discharge
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content_tags.definitions[2]0.8the tagger's words
content_tagsonset slope
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also in: onset slope
content_tags.definitions[3]1.0the tagger's words
content_tagsantidromic
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also in: antidromic
content_tags.definitions[4]1.0the tagger's words
content_tagsmovement
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also in: movement
content_tags.variables[0]the tagger's words
content_tagslaser stimulation
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content_tags.variables[1]the tagger's words
content_tagsmembrane potential variability
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content_tags.variables[2]the tagger's words
content_tagspeak tone-evoked response
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content_tags.variables[3]the tagger's words
content_tagsbody temperature
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content_tags.variables[4]the tagger's words
content_tagsmembrane potential
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content_tags.variables[6]1.0the tagger's words
content_tagsonset slope
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also in: onset slope
content_tags.variables[7]1.0the tagger's words
content_tagslocomotion
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also in: locomotion
content_tags.variables[8]1.0the tagger's words
content_tagsslope = 0.6 — Linear relation between suppression of optogenetically evoked and tone-evoked responses, implying ~60% of suppression is cortex-local
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content_tags.quantities[0]the tagger's words
content_tags>200 ms — Average lead time by which auditory cortical membrane potential changes precede locomotion onset
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content_tags.quantities[1]the tagger's words
content_tags−805 ms for PV+ cells, −605 ms for VGAT+ cells, and −490 ms for excitatory cells — Times relative to movement onset at which each cell population's firing rate changed
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content_tags.quantities[2]the tagger's words
content_tags−870 ms — Time relative to movement onset at which M2 excitatory neuron firing increased
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content_tags.quantities[3]the tagger's words
content_tags~ −72 mV — Reversal potential of the movement-related membrane potential change, near the chloride equilibrium potential
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content_tags.quantities[4]the tagger's words
content_tags<7 ms — Latency of auditory cortical change after M2 terminal stimulation, versus ~12 ms antidromic propagation time
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content_tags.quantities[5]the tagger's words
content_tags~500 ms — Delay after light onset at which silencing M2 arrested the animal's movement
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content_tags.quantities[6]the tagger's words
content_tags~40 percent — Recovery of tone-evoked response when ipsilateral M2 was silenced during ongoing movement
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content_tags.quantities[7]the tagger's words
content_tags14.25 min — Mean duration of the intracellular recordings in freely behaving mice
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content_tags.quantities[8]the tagger's words
content_tagsn = 37 cells from 11 mice — Sample for the treadmill comparison of membrane potential variance at rest versus movement
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content_tags.quantities[9]the tagger's words
content_tagsp = 8e-9 — Significance of the drop in tone-evoked voltage area from rest to movement
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content_tags.quantities[10]the tagger's words
content_tagsmore than 200 ms — how far the auditory cortical change precedes locomotion onset
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content_tags.quantities[11]1.0the tagger's words
content_tagsabout 60 percent — share of movement-related suppression arising locally in auditory cortex
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content_tags.quantities[12]1.0the tagger's words
content_tags-72 mV — reversal potential of the movement-related change, near the chloride equilibrium
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content_tags.quantities[13]1.0the tagger's words
content_tagsunder 7 ms against about 12 ms — onset of the terminal-activation effect against antidromic propagation time
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content_tags.quantities[14]1.0the tagger's words
content_tagsabout 40 percent — recovery of tone-evoked responses when M2 is silenced during ongoing movement
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content_tags.quantities[15]1.0the tagger's words
content_tagsabout 70 ms — delay before ipsilateral M2 silencing restored rest-like cortical dynamics
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content_tags.quantities[16]1.0the tagger's words

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