A cortical circuit for gain control by behavioral state

Fu, Tucciarone, Espinosa, Sheng, Darcy, Nicoll, Huang, Stryker · Cell · 2014 · neuroscience · 10.1016/j.cell.2014.01.050 · record Fu2014-a-cortical-circuit-for-gain-control-by-behav
tag layer · index · concepts · authors · עברית
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.
How to read this page. The article itself is not reproduced here — this paper is not cleared for redistribution, so what follows is the tag layer alone. The evidence sentences are withheld with the article. The table at the foot lists every tag the record carries; each row names the facet it belongs to, and the whose words column says whether the tag records the paper's own wording or the tagger's sentence about the paper. Every tag links to each other paper in the corpus carrying it. The machine-readable edition of the same thing is the TEI file beside this one, where the tags live in standOff, and the article is at its publisher via the link below. 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 WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below.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.

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 15 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.
VIP SST PV locomotion gain control orientation selectivity index disinhibition nicotinic basal forebrain nucleus of the diagonal band of Broca lateral geniculate nucleus primary visual cortex channelrhodopsin rabies virus cross-correlation Locomotion drives VIP interneurons, which inhibit SST interneurons, thereby disinhibiting neighbouring excitatory cells and raising their visual gain Acetylcholine released from basal forebrain depolarises VIP neurons directly via nicotinic receptors, carrying the behavioural-state signal into cortex PV neurons show mixed locomotion responses because their change is a second-order consequence of SST disinhibition rather than direct locomotor drive somatostatin parvalbumin acetylcholine diagonal band optogenetic GCaMP6 OGB-1 orientation selectivity behavioral state locomotion modulation index running state central rather than peripheral orientation selectivity index (OSI) running speed correlation cross-correlation between the calcium signal and running speed Mann-Whitney U test Mann-Whitney U-test paired t-test ANOVA one-way ANOVA Wilcoxon rank-sum test rank-sum test Wilcoxon signed-rank test paired Wilcoxon signed rank test standard error of the mean zero-time cross-correlation 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

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1016/j.cell.2014.01.050. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 9 · claim_type 1 · concept_edges 5 · content_tags 53 · contribution 1 · controls 1 · limitations 10 · mechanisms 9 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 4 · 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 5 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studythe circuit by which locomotion increases visual responses in mouse V1
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object_of_studythe paper's words
research_questionwhat pathway carries behavioural state into sensory cortex to produce the gain change
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research_questionthe paper's words
contributionidentifies VIP interneurons as the necessary and sufficient element, driven by nicotinic input from basal forebrain, and proposes the circuit as a general mechanism for state-dependent cortical gain control
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contributionthe paper's words
claim_typemechanism
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claim_typethe paper's words
phenomenonlocomotion-driven gain modulation of visual cortex
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[0]the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[1]the tagger's words
mechanismsmechanisms
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
mechanisms[2]the tagger's words
mechanismsVIP neurons inhibit SST neurons, releasing neighbouring excitatory cells - a disinhibitory circuit
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mechanisms[3]1.0the tagger's words
mechanismsacetylcholine from the nucleus of the diagonal band depolarises VIP neurons directly through nicotinic receptors
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mechanisms[4]1.0the tagger's words
mechanismsPV heterogeneity is explained as a secondary effect: VIP inhibits SST, SST inhibits PV, and PV also receives local excitation
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mechanisms[5]1.0the tagger's words
mechanismsthe state signal is carried by a pathway distinct from the sensory one, which is why VIP responses persist in darkness
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mechanisms[6]0.95the tagger's words
mechanismsthe kinetics are slow and lack a fast component, so the receptor is likely non-alpha7 nicotinic
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mechanisms[7]0.9the tagger's words
mechanismsphotolytic damage of individual VIP cells by restricting the excitation beam to one cell diameter at a time, sparing nearby cells
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mechanisms[8]0.95the tagger's words
limitationsNicotinic blockade did not fully abolish the locomotion response, so other inputs must contribute.
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what this limits: 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
limitations[0]the tagger's words
limitationsRabies tracing is inefficient, so additional neuromodulatory inputs may have gone undetected.
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what this limits: the completeness of the input map only: inputs MIGHT NOT BE REVEALED DUE TO THE LOW EFFICIENCY OF rabies tracing
limitations[1]the tagger's words
limitationsIn the barrel-cortex experiment whiskers were not immobilised, so sensory input was not fully removed.
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what this limits: 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
limitations[2]the tagger's words
limitationsThe auditory-cortex recordings were made without adequate isolation of auditory input.
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what this limits: 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
limitations[3]the tagger's words
limitationsnicotinic blockade did not completely abolish the locomotion response, so other inputs exist
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duplicate of: limitations[0] · what this limits: the same scope as limitations[0] - nicotinic blockade did not completely abolish the locomotion response, so other inputs contribute
limitations[4]1.0the tagger's words
limitationsrabies tracing is low-efficiency, so other neuromodulatory inputs may simply not have been revealed
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what this limits: the same scope as limitations[1] - rabies tracing is low-efficiency, so inputs may simply not have been revealed
limitations[5]1.0the tagger's words
limitationsin the barrel cortex experiment the whiskers were intact and free to move, so the result may not isolate locomotion
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duplicate of: limitations[2] · what this limits: 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
limitations[6]1.0the tagger's words
limitationsthe auditory setup was far from ideal at isolating auditory input
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what this limits: the same scope as limitations[3] - the auditory setup was far from ideal at isolating auditory input
limitations[7]1.0the tagger's words
limitationsthe basal forebrain responds to many things besides locomotion, including females and water
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what this limits: 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
limitations[8]1.0the tagger's words
limitationsother groups report different effects of locomotion on SST neurons
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what this limits: 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
limitations[9]1.0the tagger's words
transfersthe SST-hyperpolarisation-during-whisking result, used as evidence that the same circuit may run in another modality
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transfers[0]0.85the tagger's words
transfersrabies virus monosynaptic retrograde tracing, imported to find the afferents of a genetically defined cell type
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transfers[1]0.9the tagger's words
controlsidentified inhibitory neuron types recorded during stationary periods and locomotion, with and without visual stimulation, and compared against unlabelled neurons in the same recordings
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eliminates: 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 · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
sensesgain 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]1.0the tagger's words
sensesbehavioural 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]0.9the tagger's words
sensescentral rather than peripheral is established by a negative control in LGN, not by an argument
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senses[2]1.0the tagger's words
sensesVIP, 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]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: two-photon calcium imaging in running head-fixed mice, local pharmacology, slice recording, rabies retrograde tracing, optogenetic activation and photolytic ablation
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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: behavioural-state gain control in sensory cortex
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a second reading DIFFERING from the recorded value. Recorded: locomotion-driven gain modulation of visual 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_relationsreplicates → Niell and Stryker, 2010
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claim_relations[0]the tagger's words
claim_relationsbuilds-on → Pfeffer et al., 2013
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claim_relations[1]the tagger's words
claim_relationsextends → Lee et al., 2010
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claim_relations[2]the tagger's words
claim_relationssynthesises-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]1.0the tagger's words
claim_relationsextends → Niell and Stryker's finding that locomotion increases V1 gain without changing selectivity
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claim_relations[4]1.0the tagger's words
claim_relationssupports → Pfeffer et al. on VIP neurons inhibiting SST neurons
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claim_relations[5]1.0the tagger's words
claim_relationscontradicts → the expectation of substantial direct motor cortex input to V1 VIP neurons
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claim_relations[6]1.0the tagger's words
claim_relationsdistinguishes-from → the run-onset neurons of Keller et al., which respond only transiently
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differs in: THE AXIS IS TRANSIENT VERSUS SUSTAINED RESPONSE. 'Quotation withheld, 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.
claim_relations[7]1.0the tagger's words
claim_relationsqualifies → its own cholinergic account, by noting noradrenergic input as a likely additional source
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claim_relations[8]1.0the tagger's words
concept_edgesVIP —causes→ SST
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strength of the relation: 0.95 · note: inhibits. and the same quote records that other groups report different effects on SST - the disagreement travels with the edge
also in: VIP · SST
concept_edges[0]1.0the tagger's words
concept_edgeslocomotion —causes→ VIP
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strength of the relation: 0.95
also in: locomotion · VIP
concept_edges[1]1.0the tagger's words
concept_edgesbasal forebrain —causes→ VIP
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strength of the relation: 0.9 · note: largely through nicotinic inputs; blockade cut the response by more than two thirds but did not abolish it
also in: basal forebrain · VIP
concept_edges[2]1.0the tagger's words
concept_edgesVIP —causes→ gain control
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strength of the relation: 1 · note: the rare corpus case where BOTH necessity and sufficiency are demonstrated rather than inferred 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.
also in: VIP · gain control
concept_edges[3]1.0the tagger's words
concept_edgesSST —causes→ PV
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strength of the relation: 0.8 · note: inhibits; invoked to explain why PV responses are heterogeneous rather than uniform
also in: SST · PV
concept_edges[5]1.0the tagger's words
content_tagscorrelation
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also in: correlation
content_tags.statistics[0]the tagger's words
content_tagsMann-Whitney U test
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content_tags.statistics[1]the tagger's words
content_tagspaired t-test
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also in: paired t-test
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_tagsWilcoxon rank-sum test
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content_tags.statistics[4]the tagger's words
content_tagsWilcoxon signed-rank test
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content_tags.statistics[5]the tagger's words
content_tagsstandard error of the mean
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content_tags.statistics[6]the tagger's words
content_tagszero-time cross-correlation
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content_tags.statistics[7]1.0the tagger's words
content_tagsMann-Whitney U-test
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.statistics[8]1.0the tagger's words
content_tagspaired Wilcoxon signed rank test
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content_tags.statistics[9]0.9the tagger's words
content_tagsone-way ANOVA
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also in: one-way ANOVA
content_tags.statistics[10]1.0the tagger's words
content_tagsVIP 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]the tagger's words
content_tagsVisual stimulation did not increase but significantly reduced the cross-correlation between VIP activity and locomotion.
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content_tags.findings[1]the tagger's words
content_tagsSST neuron activity was on average negatively correlated with locomotion at time zero.
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content_tags.findings[2]the tagger's words
content_tagsBlocking 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]the tagger's words
content_tagsMonosynaptic 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]the tagger's words
content_tagsOptogenetically 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]the tagger's words
content_tagsThe 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]the tagger's words
content_tagsVIP neuron responses in primary auditory cortex were also associated with locomotor activity.
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content_tags.findings[7]the tagger's words
content_tagsRunning increases visual responses without changing orientation selectivity.
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content_tags.findings[8]the tagger's words
content_tagsVIP neurons track locomotion in complete darkness, when neighbouring neurons show only spontaneous activity
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content_tags.findings[9]1.0the tagger's words
content_tagsVIP neurons are the majority of locomotion-responsive cells in V1 in the absence of visual stimulation
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content_tags.findings[10]1.0the tagger's words
content_tagsvisual stimulation does not increase, and in fact reduces, the coupling between VIP activity and locomotion
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content_tags.findings[11]1.0the tagger's words
content_tagsVIP, PV and SST neurons differ significantly from one another in their locomotion coupling
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content_tags.findings[12]1.0the tagger's words
content_tagsSST neurons are suppressed by locomotion
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content_tags.findings[13]1.0the tagger's words
content_tagsnicotinic antagonists dramatically reduced VIP responses to locomotion without blocking nearby non-VIP visual responses
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content_tags.findings[14]1.0the tagger's words
content_tagsthe nucleus of the diagonal band is a prominent monosynaptic input to upper-layer V1 VIP neurons
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content_tags.findings[15]1.0the tagger's words
content_tagsat most very sparse direct input from motor cortex was found
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content_tags.findings[16]1.0the tagger's words
content_tagsoptogenetic VIP activation in stationary mice raised visual responses by about a third without changing selectivity
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content_tags.findings[17]1.0the tagger's words
content_tagsphotolytic damage to VIP cells cut the locomotion enhancement from 86% to 12%
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content_tags.findings[18]1.0the tagger's words
content_tagsVIP activation in anaesthetised mice does not change visual responses, as the circuit predicts
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content_tags.findings[19]1.0the tagger's words
content_tagslocomotion modulation index
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content_tags.definitions[0]the tagger's words
content_tagsrunning state
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also in: running state
content_tags.definitions[1]the tagger's words
content_tagscentral rather than peripheral
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content_tags.definitions[3]1.0the tagger's words
content_tagsbehavioral state
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content_tags.variables[0]the tagger's words
content_tagsorientation selectivity index (OSI)
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content_tags.variables[5]the tagger's words
content_tagsrunning speed
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also in: running speed
content_tags.variables[7]1.0the tagger's words
content_tagslocomotion modulation index
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content_tags.variables[8]0.9the tagger's words
content_tagsorientation selectivity index
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content_tags.variables[9]1.0the tagger's words
content_tags37±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]the tagger's words
content_tags0.47±0.03, n=28 — Mean zero-lag cross-correlation between VIP-neuron calcium signal and running speed in darkness
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[1]the tagger's words
content_tags20/28 versus 3/77 — VIP neurons versus non-VIP neurons whose zero-lag correlation with locomotion exceeded 0.4
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[2]the tagger's words
content_tags0.58±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
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[3]the tagger's words
content_tags155.0±34.8% — Average increase in the GCaMP6 signal of VIP neurons aligned to the start of running events
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[4]the tagger's words
content_tags32.4±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
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[5]the tagger's words
content_tags86±16% versus 12±6% — Locomotion-induced enhancement of non-VIP neuron activity before versus after photolytic damage of nearby VIP neurons
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[6]the tagger's words
content_tags12.41±0.98mV vs. 2.61±0.26mV — Acetylcholine-induced depolarisation of VIP neurons in slices before versus after nicotinic receptor blockade under TTX
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[7]the tagger's words
content_tags37 plus or minus 7% increase in calcium response with running — reproduction of the electrophysiological effect by imaging
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[8]1.0the tagger's words
content_tagsVIP 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 record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[9]1.0the tagger's words
content_tags20/28 VIP versus 3/77 non-VIP neurons above 0.4 correlation — how specific the locomotion response is to VIP cells
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[10]1.0the tagger's words
content_tags32.4 plus or minus 8.3% — increase in visual response from optogenetic VIP activation
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[11]1.0the tagger's words
content_tags86 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 record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[12]1.0the tagger's words
content_tags0 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 record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[13]1.0the tagger's words

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