A cortical circuit for gain control by behavioral state — TEI-tagged view

Fu, Tucciarone, Espinosa, Sheng, Darcy, Nicoll, Huang, Stryker · 2014 · neuroscience · doi:10.1016/j.cell.2014.01.050 · full text at the source · TEI xml · tagged edition · index · concepts · authors
Where this record's text came from. faceted 2026-08 by reading the locally held text (xml); every evidence string verified as an exact substring of that text before merging. method is tagged 'electrophysiology' for the extracellular silicon-probe recordings and slice recordings, but the paper's principal measurement is in vivo two-photon calcium imaging (OGB-1 and GCaMP6s), supplemented by optogenetics, targeted photolytic lesioning and monosynaptic rabies tracing; the controlled method vocabulary has no term for imaging or for circuit-perturbation work, so the label understates the design. Replication status is 'untested' for this paper's own VIP-circuit claim; note separately that the paper does report reproducing Niell and Stryker's (2010) locomotion effect with a different technique, which is recorded as a claim_relation. 2026-08-12, 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.
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
VIP14 SST7 PV22 locomotion8 gain control3 orientation selectivity index disinhibition6 nicotinic basal forebrain2 nucleus of the diagonal band of Broca lateral geniculate nucleus primary visual cortex channelrhodopsin3 rabies virus cross-correlation9 electrophysiology110

Our tag layer for this paper

Cross-disciplinary analysis
content-tags & facets: phenomenon · claim-type · cross-domain-transfer · mechanism · claim-relation · replication-status · limitation — grounded in the full text
phenomenonlocomotion-driven gain modulation of visual cortex— “Notably, the visual response of mouse primary visual cortex (V1) is enhanced by locomotion, a tractable and accessible example of a time-locked change in cortical state.”
claim-typemechanism— “These findings establish a cortical circuit for the enhancement of visual response by locomotion and provide a potential common circuit for the modulation of sensory processing by behavioral state.”
method — evidence— “To determine the impact of activating VIP neurons on the visual response of V1 neurons, we performed extracellular recordings as described in Supplemental Information and alternated trials in which VIP neurons were optogenetically-activated (using a blue light, referred to as LED condition) with control trials (No LED condition)”
object of study — evidence— “The neural circuits that transmit behavioral state to sensory cortex to produce this modulation are unknown.”
research question — evidence— “The neural circuits that transmit behavioral state to sensory cortex to produce this modulation are unknown.”
contribution — evidence— “These findings establish a cortical circuit for the enhancement of visual response by locomotion and provide a potential common circuit for the modulation of sensory processing by behavioral state.”
transferfrom barrel cortex sensory-motor physiology: the SST-hyperpolarisation-during-whisking result, used as evidence that the same circuit may run in another modality— “Considering the similar distribution pattern of VIP neurons through the cortical areas, and the broad projection of NDB to other cortical areas, we examined whether VIP neurons in other primary sensory cortices are also modulated by locomotion.”transfers[0]
transferfrom virology: rabies virus monosynaptic retrograde tracing, imported to find the afferents of a genetically defined cell type— “Interestingly, such retrograde tracing identified the nucleus of the diagonal band of Broca (NDB), a basal forebrain nucleus enriched in cholinergic neurons, as a prominent source of input to upper layer VIP neurons in V1”transfers[1]
mechanismLocomotion drives VIP interneurons, which inhibit SST interneurons, thereby disinhibiting neighbouring excitatory cells and raising their visual gain— “The present study reveals that a specific type of GABAergic cortical neuron, the VIP neuron, transmits the signal of locomotion to the cortex and operates through disinhibition of another GABAergic cortical neuron, the SST cell, to enhance the responses of neighboring cortical excitatory neurons”mechanisms[0]
mechanismAcetylcholine released from basal forebrain depolarises VIP neurons directly via nicotinic receptors, carrying the behavioural-state signal into cortex— “These findings in vivo suggest that acetylcholine (ACh) activates VIP neurons directly through nAChR.”mechanisms[1]
mechanismPV neurons show mixed locomotion responses because their change is a second-order consequence of SST disinhibition rather than direct locomotor drive— “Because VIP neurons principally inhibit SST neurons, which then inhibit PV neurons, which also receive local excitation, the activity of PV neurons during locomotion would be expected to be heterogeneous, as a secondary effect of relieving the inhibition from SST neurons rather than a direct response to locomotary input.”mechanisms[2]
mechanismVIP neurons inhibit SST neurons, releasing neighbouring excitatory cells - a disinhibitory circuit— “Our findings that VIP neurons are activated while SST neurons are inhibited during locomotion are in good agreement with the current consensus that VIP neurons mainly innervate SST neurons ( Lee et al., 2013 ), although different effects on SST neurons have also been reported”mechanisms[3]
mechanismacetylcholine from the nucleus of the diagonal band depolarises VIP neurons directly through nicotinic receptors— “In the presence of tetrodotoxin (TTX), the depolarization induced by ACh was abolished by the nAChR blockers MEC & MLA (12.41±0.98mV vs. 2.61±0.26mV, p <0.01, Supplemental Figure 5B and C ), confirming the existence of direct nicotinic cholinergic responses in VIP neurons.”mechanisms[4]
mechanismPV heterogeneity is explained as a secondary effect: VIP inhibits SST, SST inhibits PV, and PV also receives local excitation— “Because VIP neurons principally inhibit SST neurons, which then inhibit PV neurons, which also receive local excitation, the activity of PV neurons during locomotion would be expected to be heterogeneous, as a secondary effect of relieving the inhibition from SST neurons rather than a direct response to locomotary input.”mechanisms[5]
mechanismthe state signal is carried by a pathway distinct from the sensory one, which is why VIP responses persist in darkness— “A signal conveying behavioral state to sensory cortex would ideally not be confounded by the sensory response that it modulates.”mechanisms[6]
mechanismthe kinetics are slow and lack a fast component, so the receptor is likely non-alpha7 nicotinic— “No fast nAChR component like that shown in layer I inhibitory neurons ( Letzkus et al., 2011 ) was evident; our findings showed instead a response with slow kinetics resembling what has been found in ChAT positive bipolar neurons ( Arroyo et al., 2012 ). Therefore, it is likely that VIP neurons are activated by ACh through non-α7”mechanisms[7]
mechanismphotolytic damage of individual VIP cells by restricting the excitation beam to one cell diameter at a time, sparing nearby cellsmechanisms[8]
replicates Niell and Stryker, 2010claim_relations[0]
builds-on Pfeffer et al., 2013
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claim_relations[1]
extends Lee et al., 2010
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claim_relations[2]
synthesises-with Pi et al. 2013 on VIP disinhibition in prefrontal and auditory cortex - a paper independently tagged in this corpus
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claim_relations[3]
extends Niell and Stryker's finding that locomotion increases V1 gain without changing selectivity
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claim_relations[4]
supports Pfeffer et al. on VIP neurons inhibiting SST neurons
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claim_relations[5]
contradicts the expectation of substantial direct motor cortex input to V1 VIP neurons
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claim_relations[6]
distinguishes-from the run-onset neurons of Keller et al., which respond only transiently
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claim_relations[7]
qualifies its own cholinergic account, by noting noradrenergic input as a likely additional source
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claim_relations[8]
sensegain control here means a multiplicative change in visual response with no change in orientation selectivity - the OSI is checked and unchanged in both the locomotion and the optogenetic case
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senses[0]
sensebehavioural state is operationalised as running speed on a floating ball, with stationary and running separated at 1 cm/s
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senses[1]
sensecentral rather than peripheral is established by a negative control in LGN, not by an argument
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senses[2]
senseVIP, SST and PV are genetic labels here, not physiological inferences - the paper says electrophysiology alone cannot distinguish these cell types
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senses[3]
controlidentified inhibitory neuron types recorded during stationary periods and locomotion, with and without visual stimulation, and compared against unlabelled neurons in the same recordings — held — rules out a change that follows from locomotion or from the visual stimulus alone rather than from the cell type - the unlabelled neurons see the same two manipulations
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controls[0]
differs inthe run-onset neurons of Keller et al., which respond only transiently — THE AXIS IS TRANSIENT VERSUS SUSTAINED RESPONSE. 'The neurons identified in that study are unlikely to be VIP neurons because they responded only briefly to some of the onsets and offsets of running, while we find that VIP neurons resp[ond throughout].' The identification of a cell class in an earlier paper is disputed on a response-time-course criterion - so the two literatures may be about different populations, and only the time course says so.
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claim_relations[7]
noteVIP —causes→ SST — inhibits. and the same quote records that other groups report different effects on SST - the disagreement travels with the edge
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concept_edges[0]
weightVIP —causes→ SST — 0.95
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concept_edges[0]
weightlocomotion —causes→ VIP — 0.95
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concept_edges[1]
notebasal forebrain —causes→ VIP — largely through nicotinic inputs; blockade cut the response by more than two thirds but did not abolish it
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concept_edges[2]
weightbasal forebrain —causes→ VIP — 0.9
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concept_edges[2]
auditVIP —causes→ gain control — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but VIP activation is necessary and sufficient for the locomotion-induced INCREASE in visual response; 'gain control' is not in the span.
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concept_edges[3]
noteVIP —causes→ gain control — the rare corpus case where BOTH necessity and sufficiency are demonstrated rather than inferred
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concept_edges[3]
noteSST —causes→ PV — inhibits; invoked to explain why PV responses are heterogeneous rather than uniform
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concept_edges[5]
weightSST —causes→ PV — 0.8
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concept_edges[5]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
for which uselimitation — the sufficiency of the nicotinic account: THE RESPONSE OF VIP NEURONS TO LOCOMOTION ... WAS NOT COMPLETELY ABOLISHED BY NACHRS ANTAGONISTS, INDICATING THAT other inputs contribute
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limitations[0]
for which uselimitation — the completeness of the input map only: inputs MIGHT NOT BE REVEALED DUE TO THE LOW EFFICIENCY OF rabies tracing
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limitations[1]
for which uselimitation — the barrel-cortex control specifically: ALTHOUGH THE MICE WERE RUNNING IN DARKNESS WITH NO VISUAL INPUT, THE WHISKERS WERE INTACT AND THE MICE WERE FREE TO MOVE THEM - the sensory input was not actually removed
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limitations[2]
for which uselimitation — the auditory-cortex recordings only: auditory input was not adequately isolated, so the modality-general reading of those data is not supported by that experiment
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limitations[3]
duplicate oflimitation — limitations[0]
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limitations[4]
for which uselimitation — the same scope as limitations[0] - nicotinic blockade did not completely abolish the locomotion response, so other inputs contribute
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limitations[4]
for which uselimitation — the same scope as limitations[1] - rabies tracing is low-efficiency, so inputs may simply not have been revealed
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limitations[5]
duplicate oflimitation — limitations[2]
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limitations[6]
for which uselimitation — the same scope as limitations[2] - in the barrel-cortex experiment the whiskers were intact and free to move, so sensory input was not fully removed
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limitations[6]
for which uselimitation — the same scope as limitations[3] - the auditory setup was far from ideal at isolating auditory input
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limitations[7]
for which uselimitation — the locomotion-specificity of the basal forebrain account: THE BASAL FOREBRAIN RESPONDS TO MANY DIFFERENT STIMULI AND MAY INFLUENCE CORTICAL RESPONSE BY MODULATING DIFFERENT POPULATIONS OF NEURONS
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limitations[8]
for which uselimitation — the agreement with the wider literature on SST cells: DIFFERENT EFFECTS ON SST NEURONS HAVE ALSO BEEN REPORTED elsewhere, so the consensus this result matches is not unanimous
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limitations[9]
replicationuntested
limitationNicotinic blockade did not fully abolish the locomotion response, so other inputs must contribute.
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limitations[0]
limitationRabies tracing is inefficient, so additional neuromodulatory inputs may have gone undetected.
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limitations[1]
limitationIn the barrel-cortex experiment whiskers were not immobilised, so sensory input was not fully removed.
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limitations[2]
limitationThe auditory-cortex recordings were made without adequate isolation of auditory input.
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limitations[3]
limitationnicotinic blockade did not completely abolish the locomotion response, so other inputs exist
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limitations[4]
limitationrabies tracing is low-efficiency, so other neuromodulatory inputs may simply not have been revealed
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limitations[5]
limitationin the barrel cortex experiment the whiskers were intact and free to move, so the result may not isolate locomotion
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limitations[6]
limitationthe auditory setup was far from ideal at isolating auditory input
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limitations[7]
limitationthe basal forebrain responds to many things besides locomotion, including females and water
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limitations[8]
limitationother groups report different effects of locomotion on SST neurons
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limitations[9]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingVIP neurons in mouse V1 are activated in tight correlation with locomotion even in complete darkness, unlike nearly all neighbouring non-VIP neurons.
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content_tags.findings[0]
findingVisual stimulation did not increase but significantly reduced the cross-correlation between VIP activity and locomotion.
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content_tags.findings[1]
findingSST neuron activity was on average negatively correlated with locomotion at time zero.
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content_tags.findings[2]
findingBlocking nicotinic cholinergic receptors locally in V1 removed most of the VIP locomotion response, while blocking glutamatergic input did not.
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content_tags.findings[3]
findingMonosynaptic rabies tracing identified the cholinergic nucleus of the diagonal band of Broca in basal forebrain as a direct input source to upper-layer V1 VIP neurons.
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content_tags.findings[4]
findingOptogenetically driving VIP neurons in stationary mice reproduced the locomotion-induced gain increase, and selectively killing VIP neurons removed it.
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content_tags.findings[5]
findingThe gain-and-loss experiments together establish VIP activation as both necessary and sufficient for the locomotion-driven enhancement of excitatory visual responses.
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content_tags.findings[6]
findingVIP neuron responses in primary auditory cortex were also associated with locomotor activity.
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content_tags.findings[7]
findingRunning increases visual responses without changing orientation selectivity.
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content_tags.findings[8]
findingVIP neurons track locomotion in complete darkness, when neighbouring neurons show only spontaneous activity
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content_tags.findings[9]
findingVIP neurons are the majority of locomotion-responsive cells in V1 in the absence of visual stimulation
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content_tags.findings[10]
findingvisual stimulation does not increase, and in fact reduces, the coupling between VIP activity and locomotion
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content_tags.findings[11]
findingVIP, PV and SST neurons differ significantly from one another in their locomotion coupling
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content_tags.findings[12]
findingSST neurons are suppressed by locomotion
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content_tags.findings[13]
findingnicotinic antagonists dramatically reduced VIP responses to locomotion without blocking nearby non-VIP visual responses
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content_tags.findings[14]
findingthe nucleus of the diagonal band is a prominent monosynaptic input to upper-layer V1 VIP neurons
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content_tags.findings[15]
findingat most very sparse direct input from motor cortex was found
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content_tags.findings[16]
findingoptogenetic VIP activation in stationary mice raised visual responses by about a third without changing selectivity
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content_tags.findings[17]
findingphotolytic damage to VIP cells cut the locomotion enhancement from 86% to 12%
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content_tags.findings[18]
findingVIP activation in anaesthetised mice does not change visual responses, as the circuit predicts
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content_tags.findings[19]
quantity37±7% — Average increase in the calcium response of imaged V1 neurons during running, replicating the electrophysiological locomotion effect
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content_tags.quantities[0]
quantity0.47±0.03, n=28 — Mean zero-lag cross-correlation between VIP-neuron calcium signal and running speed in darkness
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content_tags.quantities[1]
quantity20/28 versus 3/77 — VIP neurons versus non-VIP neurons whose zero-lag correlation with locomotion exceeded 0.4
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content_tags.quantities[2]
quantity0.58±0.03 (n=21), 0.30±0.06 (n=40), −0.19±0.05 (n=11) — Zero-lag locomotion cross-correlations for VIP, PV and SST neurons respectively
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content_tags.quantities[3]
quantity155.0±34.8% — Average increase in the GCaMP6 signal of VIP neurons aligned to the start of running events
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content_tags.quantities[4]
quantity32.4±8.3%, n=19, p <0.02 — Increase in the visual response of layer 2/3 neurons produced by optogenetic activation of VIP neurons in stationary mice
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content_tags.quantities[5]
quantity86±16% versus 12±6% — Locomotion-induced enhancement of non-VIP neuron activity before versus after photolytic damage of nearby VIP neurons
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content_tags.quantities[6]
quantity12.41±0.98mV vs. 2.61±0.26mV — Acetylcholine-induced depolarisation of VIP neurons in slices before versus after nicotinic receptor blockade under TTX
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content_tags.quantities[7]
quantity37 plus or minus 7% increase in calcium response with running — reproduction of the electrophysiological effect by imaging
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.quantities[8]
quantityVIP 0.58 plus or minus 0.03, PV 0.30 plus or minus 0.06, SST -0.19 plus or minus 0.05 — zero-time cross-correlation with locomotion by cell type
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.quantities[9]
quantity20/28 VIP versus 3/77 non-VIP neurons above 0.4 correlation — how specific the locomotion response is to VIP cells
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.quantities[10]
quantity32.4 plus or minus 8.3% — increase in visual response from optogenetic VIP activation
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.quantities[11]
quantity86 plus or minus 16% before versus 12 plus or minus 6% after photo damage — locomotion enhancement lost when VIP cells are destroyed
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.quantities[12]
quantity0 to 1 neurons per mouse in motor cortex against more than 50 labelled VIP neurons — the near-absence of direct motor input
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.quantities[13]
variablebehavioral state (independent)
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[0]
variableorientation selectivity index (OSI) (dependent)
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[5]
variablerunning speed
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[7]
variablelocomotion modulation index
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[8]
variableorientation selectivity index
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.variables[9]
definitionlocomotion modulation index
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[0]
definitionrunning state
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[1]
definitioncentral rather than peripheral
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.definitions[3]
key-termsVIP, somatostatin, parvalbumin, locomotion, gain control, disinhibition, primary visual cortex, basal forebrain, nicotinic, acetylcholine, diagonal band, rabies virus, channelrhodopsin, optogenetic, GCaMP6, OGB-1, cross-correlation, orientation selectivity, lateral geniculate nucleus, behavioral state
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.
correlation as written: cross-correlation between the calcium signal and running speed
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[0]
Mann-Whitney U test p
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[1]
paired t-test p
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[2]
ANOVA p as written: one-way ANOVA
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[3]
Wilcoxon rank-sum test p as written: rank-sum test
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[4]
Wilcoxon signed-rank test p as written: paired Wilcoxon signed rank test
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[5]
standard error of the mean SEM as written: mean±SEM
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[6]
zero-time cross-correlation
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[7]
Mann-Whitney U-test
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[8]
paired Wilcoxon signed rank test
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[9]
one-way ANOVA
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
content_tags.statistics[10]
Second readings
a facet read a second time, independently, from its own sentence — and whether the two readings agree
claim typemechanism — a second reading reaching the same value; recorded: mechanism
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.claim_type
methodtwo-photon calcium imaging in running head-fixed mice, local pharmacology, slice recording, rabies retrograde tracing, optogenetic activation and photolytic ablation — a second reading DIFFERING from the recorded value; recorded: electrophysiology
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.method
phenomenonbehavioural-state gain control in sensory cortex — a second reading DIFFERING from the recorded value; recorded: locomotion-driven gain modulation of visual cortex
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.phenomenon

When we have changed our mind. Some entries in the table begin with a word in capitals and the reason for it. EVIDENCE CORRECTED — the finding still stands, but we had quoted the wrong sentence from the paper, and the quotation has been replaced with the right one. RETRACTED — the entry itself was wrong; it stays on the page, marked, with the reason, rather than being deleted. WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below. Each mark carries its own reason, in brackets, because the reasons differ from one entry to the next. An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason. A line reading Audit and a date is a later re-reading of the entry against its whole sentence: DOUBTFUL means it was kept, and the reason says what is loose about it. Quotation marks inside the tagger's sentence. Where a tag is the tagger's own sentence, a phrase in quotation marks inside it is borrowed from the paper — the tagger's wording around it is not. The paper's own sentence is the evidence quote for that row; on this page it is withheld with the article, and the article is at its publisher. Nothing inside quotation marks anywhere on this page is ours: if it is quoted, it is the paper's. Whose words a concept carries. A concept entry records a term the paper itself uses, so the term, the sense given for it, and its evidence are the paper's own words, quoted where the paper fixes them. This is the opposite of a sense entry in the tag table, which is the tagger's sentence about how the paper uses a term. Where a concept entry could not be grounded in the paper's words, the entry says so on its own row.

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