Cortical Membrane Potential Signature of Optimal States for Sensory Signal Detection — TEI-tagged view

McGinley, David, McCormick · 2015 · neuroscience · doi:10.1016/j.neuron.2015.05.038 · 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. Claim type 'mechanism' rather than 'existence' because the paper offers a synaptic/network account (suppression of slow rhythms then depolarization with hyper-arousal) of an already-classic behavioural relationship; the headline empirical statement is 'Optimal signal detection behavior and sound-evoked responses, at both sub-threshold and spiking levels, occurred at intermediate arousal when pre-decision membrane potentials were stably hyperpolarized.' The Reimer et al. (2014) relation is recorded twice with two types because the paper both aligns itself with that study and reports a specific disagreement with it (layer 2/3 vs deep pyramidal cells). Methods are heavily abbreviated in this source text (Supplemental Methods referenced but not included), so method_evidence comes from the condensed Experimental Procedures section. 2026-08-13, 2026-08-14 and 2026-08-15 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
Other fields define these differently the same word, defined another way in another discipline
reliability 5 fields · behavior 4 fields · standard deviation 3 fields · arousal 2 fields · anesthesia 2 fields · latency 2 fields · and 2 more →
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
arousal10 pupil2 ripple5 eyelid EIS3 derivative membrane potential7 Vm variance54 hyperpolarized depolarization oscillation5 microdilation walking movement5 whisking3 ACtx MG2 layer4 evoked2 spontaneous6 gain10 reliability12 coherence6 detection8 hit false alarm2 latency19 inverted-U U-shaped state anesthesia14 sensory3 behavior19 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
phenomenonarousal-dependent auditory signal detection— “The neural correlates of optimal states for signal detection task performance are largely unknown.”
claim-typemechanism— “Our results suggest a cellular and network basis for this classical result.”
method — evidence— “Whole cell recordings were made from regular or fast spiking neurons in layers 4 and 5 of ACtx using a physiological intracellular solution (see Supplemental Methods : Whole-Cell Recording).”
object of study — evidence— “We recorded membrane potentials of auditory cortical neurons in mice trained on a challenging tone-in-noise detection task while assessing arousal with simultaneous pupillometry and hippocampal recordings. Arousal measures accurately predicted multiple modes of membrane potential activity, including: rhythmic slow oscillations at low arousal…
research question — evidence— “Human and animal studies have reported that performance on signal detection tasks is highly state-dependent, exhibiting an inverted-U dependence on arousal and the activity of neuromodulatory pathways. This relationship, known as the Yerkes-Dodson curve, predicts that optimal performance occurs at intermediate levels of arousal”
contribution — evidence— “Optimal signal detection behavior and sound-evoked responses, at both sub-threshold and spiking levels, occurred at intermediate arousal when pre-decision membrane potentials were stably hyperpolarized. These results reveal a cortical physiological signature of the classically-observed inverted-U relationship between task performance and…
transferfrom psychophysiology: pupillometry as a continuous index of autonomic state, imported to index cortical membrane potential rather than emotion— “We primarily focus on pupil diameter as a measure of state, owing to its ease of measurement, non-invasive nature, and ability to continuously and sensitively track state changes, whether or not they accompany overt walking, whisking, or other movements”transfers[0]
transferfrom hippocampal electrophysiology: sharp-wave ripple rate as an independent state variable, used to validate the pupil rather than as the object of study— “The pupil diameter exhibited a strong inverse relationship to the rate of occurrence of fast ripples ( Figs. 1c–f , S1 ), with a peak coherence of 0.78 ± 0.07 (n=6 cells from N=6 animals) at low frequencies (<0.05 Hz) and a significant coherence at all frequencies up to 1 Hz ( Figs. 1e ). Theta activity was largely associated with walking…transfers[1]
transferfrom signal detection theory: response latency on hit trials plotted against pupil diameter, showing short latency licks and a large decrease in variability of lick latency at intermediate diameters— “Plotting response latency on hit trials versus pupil diameter, for an example session, reveals that intermediate pupil diameters are associated with short latency licks and a large decrease in variability of lick latency. e, Group data demonstrating that lick latency on hit trials is shortest for intermediate pupil diameters (n=30 sessions…transfers[2]
transferfrom gain modelling: responses sorted by state against responses averaged over all states, where no effect would be a diagonal of slope one— “If the state changes reflected by pupil diameter had no effect, this relationship would be a diagonal line with slope of 1. Variations in pupil diameter exhibited a consistent change in the slope of this relationship, with the highest slope occurring at intermediate pupil diameters ( Fig. 5d–h ) for both ACtx and MG responses (see however, (…transfers[3]
mechanismrising arousal suppresses low-frequency cortical rhythms, letting cortico-cortical and cortico-subcortical circuits interact more reliably and so producing larger, more precise sensory responses— “We hypothesize that this suppression of low frequency rhythms allows cortico-cortical and cortico-subcortical circuits to operate more reliably, enhancing sensory-evoked neuronal interactions, and thereby contributing to precise, reliable, and large sensory responses ( Haider et al., 2010 ; Lee and Dan, 2012 ; Pinto et al., 2013 ).”mechanisms[0]
mechanismat hyper-arousal the depolarization is carried by task-unrelated synaptic activity that raises membrane variance and thereby disrupts sensory-evoked neuronal interactions— “Although depolarization may increase the responsiveness of single cortical neurons, it may have a detrimental effect on sensory evoked neuronal interactions if the depolarization is mediated by synaptic activity that increases membrane variance or is unrelated to the task.”mechanisms[1]
mechanismthe opposing arousal-driven changes in thalamic gain and cortical depolarization combine to generate the U and inverted-U curves relating arousal to neural and behavioural performance— “We propose that together these opposing arousal-associated changes in thalamo-cortical activity produce the U, or inverted U, shaped relationships between neuronal and behavioral responses and arousal observed here ( Fig. 8 ).”mechanisms[2]
mechanismthe lag between cortical depolarization and pupil dilation arises because pupillary and eyelid muscles respond slowly to sympathetic and parasympathetic drive— “Much or all of this lag may represent the slow time course of pupillary/eyelid movement to alterations in sympathetic and parasympathetic activity ( Loewenfeld, 1999 ).”mechanisms[3]
mechanismmembrane potential and its variance trace a U-shaped relationship against arousal level as indexed by normalized pupil diameter— “Thus, plotting the membrane potential of cortical neurons as a group (n=9) against arousal level revealed a U-shaped relationship between average membrane potential/variance with normalized pupil diameter”mechanisms[4]
mechanismso the signal-to-noise ratio of an evoked response is high at that state, from both suppressed background and larger responses— “At the best state for behavioral performance on our sensory detection task, the signal-to-noise ratio of evoked cortical responses is high, owing both to the suppression of spontaneous activities and the enhanced amplitude of sensory evoked responses”mechanisms[5]
mechanismat high arousal the cortex depolarises, which may raise single-neuron responsiveness while degrading the interactions that carry a sensory response— “Although depolarization may increase the responsiveness of single cortical neurons, it may have a detrimental effect on sensory evoked neuronal interactions if the depolarization is mediated by synaptic activity that increases membrane variance or is unrelated to the task. The subsequent decreased reliability of cortical responses may also…mechanisms[6]
mechanismand thalamic responses fall with arousal and fall further with walking, so part of the cortical decline is inherited— “To examine if the variations in ACtx response amplitude also occurred at subcortical levels, we performed MU (n=9) recordings in the medial geniculate body (MG). As observed with ACtx neurons, variations in pupil diameter in the non-walking animal were associated with variations in the amplitude and reliability of the evoked action potential…mechanisms[7]
extends Reimer et al. (2014)— “A recently published study of the mouse visual cortex, which paralleled our own investigations, suggested that the rate of change of pupil diameter was an accurate measure of changes in cortical membrane potential power ( Reimer et al., 2014 ).”claim_relations[0]
contradicts Reimer et al. (2014)— “Indeed, while our deep lying pyramidal neurons were depolarized during micro-arousals without walking ( Fig. 2 ), whole cell recordings from layer 2/3 pyramidal cells in the visual cortex reveal no significant change in membrane potential with non-movement related arousal ( Reimer et al., 2014 ).”claim_relations[1]
replicates Williamson et al. (2014)— “These results are consistent with the large walking-specific decrease in evoked firing rate in MG neurons ( Fig. 5b ) ( Williamson et al., 2014 ).”claim_relations[2]
explains-away the rate of change of pupil diameter as the better index of cortical state, which is shown to be a lagged proxy for the diameter itself— “Importantly, we found that the derivative of the pupil was correlated to the pupil diameter 1.3 ± 0.7 seconds later (CC = 0.32 ± 0.02; p<0.001; n=9; Fig. S6k,l ), but not to the pupil diameter at the same time (correlation coefficient CC = 0.0035 ± 0.0043; p=0.44). This lag approximately matched the observed lag between physiological…claim_relations[3]
reconciles the divergent reports of cortical membrane potential in waking animals, by supplying the continuously varying state variable none of them measured— “One possible explanation of these apparently divergent results is that state (arousal) fluctuates continuously and rapidly, resulting in a high…claim_relations[4]
critiques studies of the effects of walking and whisking on sensory responses, which most likely involved large uncontrolled changes in arousalclaim_relations[5]
supports the classical Yerkes-Dodson inverted-U between arousal and performance, by giving it a cellular and network basis
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claim_relations[6]
qualifies the opposing visual and auditory locomotion results, which are left explicitly unresolved
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claim_relations[7]
senseA LITERATURE'S CONTRADICTIONS ATTRIBUTED TO A VARIABLE NOBODY MEASURED: state fluctuates continuously and in seconds, which produces variability between studies and between moments inside one study, and has not been accounted for
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senses[0]
senseand one earlier report is named as having recorded across un-anaesthetised states without quantifying the arousal of the animals at all
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senses[1]
senseA BINARY CATEGORY IS DISSOLVED INTO A CONTINUUM: the same pupil measure had been used to sort trials into aroused and unaroused, and is used here as a graded quantity
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senses[2]
senseAND A WHOLE LITERATURE'S INDEPENDENT VARIABLE IS RE-IDENTIFIED: the often reported effects of movement may not result from movement per se but from arousal, with one exception named and kept
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senses[3]
senseTHE SHAPE OF THE EFFECT BREAKS THE SUMMARY STATISTIC: because the relation is U-shaped the correlation is positive over one half of the range and negative over the other, which forced a sliding window rather than one coefficient
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senses[4]
senseA RIVAL MEASURE IS TESTED AND THEN EXPLAINED AWAY: the derivative of the pupil predicts less than the diameter, and correlates with the diameter a second later, so its explanatory power may be entirely borrowed
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senses[5]
senseTHE MEASURE IS VALIDATED THROUGH A CHANNEL WHERE THE CONFOUND CANNOT OPERATE: pupil dilation could be a response to light, so eyelid opening was measured in darkness, where dilation is saturated, and moved the same way
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senses[6]
senseand the recording artefact is ruled out by timing rather than by argument - the depolarisation begins before the behavioural event and returns after it
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senses[7]
senseTHIRTEEN MEASURES ARE ASKED WHETHER THEY PEAK IN THE SAME PLACE: twelve are statistically indistinguishable and the thirteenth is not, with an interpretation offered for the one
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senses[8]
senseand the state measure is normalised within session because its maximum drifts between days and animals, so every number here is a fraction of that animal's own range
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senses[9]
controleyelid opening measured IN DARKNESS, alongside the pupil recording — held — rules out illumination change as the cause of the pupil dilations - an eye movement changes how much light reaches the pupil, so a dilation time-locked to behaviour could be an optical consequence of looking elsewhere
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controls[0]
differs instudies of the effects of walking and whisking on sensory responses, which most likely involved large uncontrolled changes in arousal — THE AXIS IS AROUSAL AS AN UNCONTROLLED VARIABLE. 'Owing to the association of walking with large pupil diameter that we observed, previous studies on the effects of walking or other movements on sensory responses have most likely involved significant changes in arousal.' A whole literature's independent variable is shown to have carried a second one with it - established by THIS paper's own pupil measurement, which is why nobody could have controlled for it earlier.
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claim_relations[5]
differs inthe opposing visual and auditory locomotion results, which are left explicitly unresolved — THE AXIS IS THE CORTICAL AREA, AND THE SIGN REVERSES ACROSS IT. 'Are the effects of arousal similar across cortical areas? In studies of the visual cortex, locomotion is associated with enhanced sensory-evoked visual responses (over non-locomotion), while in auditory cortex locomot[ion suppresses them].' The paper poses the question, gives the opposite findings, and LEAVES IT UNRESOLVED - so 'locomotion modulates sensory responses' is true with two opposite signs and the area is not optional.
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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.
claim_relations[7]
auditstate —operationalised-as→ arousal — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span writes 'state (arousal)'; operationalised-as is loose.
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concept_edges[0]
notestate —operationalised-as→ arousal — A LITERATURE'S CONTRADICTIONS ATTRIBUTED TO A VARIABLE NOBODY MEASURED: it fluctuates continuously and rapidly, producing variability between studies and between moments within one, and has not been accounted for
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.
concept_edges[0]
notepupil —operationalised-as→ state — chosen for being non-invasive and continuous, and for tracking state changes whether or not they come with overt movement
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concept_edges[3]
auditEIS —operationalised-as→ eyelid — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but reflected infrared light is a proxy for lid opening; operationalised-as is loose.
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concept_edges[6]
noteEIS —operationalised-as→ eyelid — and a cheaper proxy is then derived from the light reflected off the eye, so the measure can travel
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concept_edges[6]
auditpupil —contrasted-with→ derivative — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span compares pupil diameter with its derivative as indicators; contrasted-with is loose.
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concept_edges[7]
differs inpupil —contrasted-with→ derivative — THE AXIS IS WHICH PREDICTOR ADDS ANYTHING, AND THE ANSWER IS THAT ONE DOES NOT. 'Factoring in both the pupil diameter and its derivative explained 40+/-6% of the variance of power in the 2-10 Hz band, which was statistically larger than the amount explained by pupil derivative (p<0.0005), but not pupil diameter alone (p=0.36).' Adding the derivative to diameter does not improve on diameter alone - a null reported in the same sentence as the significant comparison, and the pair is only distinguishable because both tests are given.
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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.
concept_edges[7]
notepupil —contrasted-with→ derivative — A RIVAL MEASURE TESTED HEAD TO HEAD: the diameter explains 34 percent of the variance in low-frequency power, the derivative about half that, and both together add nothing over the diameter alone
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concept_edges[7]
auditmicrodilation —causes→ membrane potential — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says microdilations were ASSOCIATED with depolarization; causes overstates it.
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concept_edges[11]
evidential statusmicrodilation —causes→ membrane potential — ASSOCIATIONAL AND TIME-LOCKED, WHICH IS NOT THE SAME AS CAUSAL: microdilations 'were reliably associated with 5-20 mV time-locked depolarization'. Tight temporal coupling in 16 neurons; nothing was manipulated to establish which leads.
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concept_edges[11]
notemicrodilation —causes→ membrane potential — 5 to 20 mV depolarisations time-locked to them, and leading the pupil by about a second
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concept_edges[11]
noteMG —contrasted-with→ ACtx — thalamic responses fall steadily with arousal and fall further during walking, so part of the cortical decline is inherited and part is not
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concept_edges[21]
notewalking —is-a→ movement — since walking only ever happened at large pupil diameters, previous studies of movement effects most likely involved large changes in arousal
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concept_edges[23]
auditwhisking —distinct-from→ arousal — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span argues effects may come from arousal rather than movement; distinct-from is loose.
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concept_edges[25]
weightwhisking —distinct-from→ arousal — 0.9
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concept_edges[25]
notehit —contrasted-with→ false alarm — with hit and miss trials resampled to equal target level distributions, so loudness cannot be what separates them
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concept_edges[29]
auditarousal —causes→ detection — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says intermediate arousal is optimal for detection; causes is loose and unsigned.
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concept_edges[32]
notearousal —causes→ detection — the optimum is 40 to 60 percent of full dilation, for the neurons and for the behaviour alike
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concept_edges[32]
auditevoked —distinct-from→ spontaneous — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span attributes high SNR to suppression of spontaneous activity; distinct-from is loose.
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concept_edges[35]
noteevoked —distinct-from→ spontaneous — and the optimum is a ratio: background suppressed AND response enhanced, not either alone
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concept_edges[35]
auditdepolarization —causes→ reliability — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span discusses possible detrimental effects of depolarization; causes is loose.
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concept_edges[36]
notedepolarization —causes→ reliability — the proposed cost of the depolarised state - single neurons more responsive, their interactions less so
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concept_edges[36]
weightdepolarization —causes→ reliability — 0.7
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concept_edges[36]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
for which uselimitation — the thalamo-cortical account of the U-shaped relationship: FURTHER EXAMINATION OF THIS HYPOTHESIS, PARTICULARLY THE INFLUENCE OF BEHAVIORAL CONTEXT, REMAINS TO BE DONE
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limitations[0]
for which uselimitation — the visual-versus-auditory locomotion discrepancy: IT IS CURRENTLY UNCLEAR WHETHER THESE APPARENTLY OPPOSING RESULTS REFLECT VARIATIONS BETWEEN SENSORY SYSTEMS IN CORTICAL OR SUBCORTICAL INFLUENCES OF LOCOMOTION AND/OR AROUSAL
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limitations[1]
for which uselimitation — the neuromodulatory account: THE INFLUENCE OF NOREPINEPHRINE, WHICH IS THOUGHT TO CONTROL PUPIL DIAMETER, ON CORTICAL INHIBITORY CIRCUITS REMAINS TO BE DETERMINED
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limitations[2]
for which uselimitation — the generalisation across cortical areas: in visual cortex LOCOMOTION IS ASSOCIATED WITH ENHANCED SENSORY-EVOKED VISUAL RESPONSES, while in auditory cortex the effect runs the other way
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limitations[3]
for which uselimitation — the same scope as limitations[3], with a second candidate source: ANOTHER POSSIBLE SOURCE OF VARIATION BETWEEN STUDIES ... IS THE LAMINAR POSITION AND CELL TYPE OF THE RECORDED NEURONS
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limitations[4]
for which uselimitation — the proposed U-shaped account: FURTHER EXAMINATION OF THIS HYPOTHESIS, PART[icularly under behavioural context], remains to be done
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limitations[5]
for which uselimitation — the claim that the measures share an optimal arousal level: WE DID NOT FIND STATISTICALLY SIGNIFICANT DIFFERENCE BETWEEN 12 OF THE 13 BEHAVIORAL OR PHYSIOLOGICAL measures - so one of thirteen peaks elsewhere, and it is the behavioural one
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limitations[6]
evidential statusmechanism — the paper's own words: We hypothesize that
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mechanisms[0]
evidential statusmechanism — the paper's own words: We propose that
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mechanisms[2]
replicationuntested
no type for thispupil derivative → pupil diameter — THE INDEXING / PROXY RELATION - 'from' STANDS IN FOR 'to'. We have no type for it: 'causes' asserts production, and 'operationalised-as' says the authors CHOSE a measure, whereas a proxy claim says THIS STANDS IN FOR WHAT WE ACTUALLY CARE ABOUT AND WE KNOW IT IS NOT THE SAME THING. Measured at 8% of papers. A PROXY FOR ITSELF, DISPLACED IN TIME: 'the explanatory power of the pupil derivative occurs because it serves as a proxy for the pupil diameter approximately 1 second later'. The two endpoints are the same measurement at different lags, which is a substitution no vocabulary of concepts would catch.
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unhoused_relations[0]
limitationThe proposed thalamo-cortical account of the U-shaped relationship has not been tested across behavioural contexts.
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limitations[0]
limitationThe paper cannot resolve why locomotion enhances visual but suppresses auditory cortical responses.
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limitations[1]
limitationThe neuromodulatory pathway linking pupil-indexed arousal to the cortical inhibitory circuitry was not established.
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limitations[2]
limitationwhether the opposing visual and auditory results reflect sensory systems, subcortical influences or the arousal level during stillness is stated to be unclear
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limitations[3]
limitationlaminar position and cell type are named as a second possible source of between-study variation, with their layer 5 cells behaving unlike layer 2/3 visual cells
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limitations[4]
limitationthe influence of behavioural context on the proposed account remains to be done
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limitations[5]
limitationthe peak of one measure out of thirteen, behavioural discriminability, sits at a significantly smaller pupil diameter than the rest
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limitations[6]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingPupil diameter showed a strong inverse relationship to the rate of fast ripples, with a peak coherence of about 0.78 at low frequencies.
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content_tags.findings[0]
findingAuditory cortical membrane potential and its variance follow a U-shaped function of arousal, being most hyperpolarized and least variable at intermediate pupil diameter.
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content_tags.findings[1]
findingBrief spontaneous pupil microdilations in still animals are locked to 5-20 mV depolarizations of cortical neurons.
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content_tags.findings[2]
findingThe cortical depolarization leads the pupil dilation by roughly one second, so the membrane potential change comes first.
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content_tags.findings[3]
findingSound-evoked synaptic responses in auditory cortex are largest at intermediate pupil diameter and shrink with either constriction or further dilation.
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content_tags.findings[4]
findingBehavioural hit rate on the tone-in-noise detection task follows an inverted-U in pupil diameter at every tone intensity.
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content_tags.findings[5]
findingJust before correct detections the membrane potential was more hyperpolarized and less variable than before false alarms.
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content_tags.findings[6]
findingPupil diameter predicts trial-to-trial cortical response variability at least as well as walking status and absorbs walking's predictive power.
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content_tags.findings[7]
findingThe optimal arousal band, for both neural coding and behaviour, corresponds to pupils dilated to 40-60% of maximum.
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content_tags.findings[8]
findingThalamic (medial geniculate) sound responses decline steadily as pupil diameter increases and are suppressed further during walking.
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content_tags.findings[9]
findingRaw pupil diameter explains about twice as much of the low-frequency membrane-potential power as the rate of change of pupil diameter does.
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content_tags.findings[10]
findingMice only walked when their pupils were large, so locomotion in this preparation is always accompanied by high arousal.
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content_tags.findings[11]
findingAPPARENTLY DIVERGENT RESULTS ARE ATTRIBUTED TO A VARIABLE NOBODY ACCOUNTED FOR: state fluctuates continuously and rapidly, giving variability between studies and between moments within individual studies
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content_tags.findings[12]
findingwith the divergent reports themselves laid out first, across species and sensory systems
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content_tags.findings[13]
findingand one earlier study named as having recorded across un-anaesthetised states without quantifying arousal
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content_tags.findings[14]
findingwalking increases visual cortical responses and decreases auditory ones, which is the contradiction in view
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content_tags.findings[15]
findingA BINARY CATEGORY IS DISSOLVED INTO A CONTINUUM: the same pupil measure had been used to sort trials into aroused and unaroused
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content_tags.findings[16]
findingpupil diameter tracks hippocampal ripple rate inversely, supporting the view that it may be a useful indicator of brain state
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content_tags.findings[17]
findingTHE LIGHT CONFOUND IS REMOVED BY CHANGING CHANNEL: eyelid opening measured in darkness, where dilation is saturated, moves the same way
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content_tags.findings[18]
findingand a proxy is derived from infrared light reflected from the eye, termed eye-indexed state
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content_tags.findings[19]
findingthe pupil is normalised within session because its maximum drifts between days and animals
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content_tags.findings[20]
findingbrief dilations are time-locked to 5 to 20 mV depolarisations that lead them by about a second
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content_tags.findings[21]
findingmembrane potential and its variance both trace a U against arousal, most hyperpolarised and least variable in the middle
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content_tags.findings[22]
findingTHE SHAPE OF THE EFFECT BREAKS THE SUMMARY STATISTIC: the correlation is positive over one half of the range and negative over the other, forcing a sliding window
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content_tags.findings[23]
findingand layer 5 multi-unit firing shows the same U, lowest at intermediate pupil
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content_tags.findings[24]
findingTHE RECORDING ARTEFACT IS RULED OUT BY TIMING: the depolarisation starts before the behavioural event and returns after it
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content_tags.findings[25]
findingand spike threshold barely tracks the pupil, varying instead with the rate of rise into threshold
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content_tags.findings[26]
findingA RIVAL MEASURE IS TESTED HEAD TO HEAD: raw pupil diameter and the rate of change of pupil diameter are both compared to membrane power at 2-10 Hz, and factoring in both is not statistically better than the diameter alone
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content_tags.findings[27]
findingAND THEN EXPLAINED AWAY: the derivative correlates with the diameter 1.3 s later and not with it at the same moment
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content_tags.findings[28]
findingevoked spikes and synaptic responses are largest at moderate pupil diameters and smaller in both directions
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content_tags.findings[29]
findingwith evoked rate, reliability and gain all following the same inverse U
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content_tags.findings[30]
findingand the gain model is built so that no effect is a diagonal of slope one
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content_tags.findings[31]
findingthalamic responses fall steadily as the pupil dilates and are suppressed further during walking
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content_tags.findings[32]
findingTHE PUPIL SUBSUMES THE OLDER VARIABLE: walking status adds 20 percent explained variance, pupil adds 28, and both together (29) are not significantly more than pupil alone
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content_tags.findings[33]
findinglick latency is shortest and least variable at intermediate diameters, for hits and false alarms alike
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content_tags.findings[34]
findingmembrane potentials are most hyperpolarised and least variable before hits, with the largest evoked responses
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content_tags.findings[35]
findingand hit and miss trials are resampled to equivalent target tone level distributions
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content_tags.findings[36]
findingTHIRTEEN MEASURES ARE ASKED WHETHER THEY PEAK IN THE SAME PLACE: twelve are indistinguishable and only behavioural discriminability sits lower
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content_tags.findings[37]
findingthe optimum is 40 to 60 percent of full dilation, for the neurons and the behaviour alike
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content_tags.findings[38]
findingwhich suggests a cellular and network basis for the classical inverted-U
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content_tags.findings[39]
findingand at the best state the signal-to-noise ratio of evoked cortical responses is high, owing both to the suppression of spontaneous activities and the enhanced amplitude of sensory evoked responses
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content_tags.findings[40]
findingA LITERATURE'S INDEPENDENT VARIABLE IS RE-IDENTIFIED: reported effects of movement may not result from movement per se but from arousal
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content_tags.findings[41]
findingwith previous walking studies described as having most likely involved large uncontrolled changes in arousal
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content_tags.findings[42]
findingand one arousal-independent walking effect, on thalamic responses and decision bias, named and kept
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content_tags.findings[43]
findinglaminar position and cell type are named as a second source of between-study variation
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content_tags.findings[44]
findingand whether the opposing visual and auditory results reflect systems, subcortical influence or arousal during stillness is left open
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content_tags.findings[45]
findingthe proposed cost is that depolarisation may raise single-neuron responsiveness while harming sensory-evoked neuronal interactions
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content_tags.findings[46]
findingand the influence of behavioural context on the account remains to be done
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content_tags.findings[47]
findingthe classical Yerkes-Dodson curve predicts optimal performance at intermediate arousal
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content_tags.findings[48]
quantity0.78 ± 0.07 — peak coherence between pupil diameter and hippocampal fast-ripple rate at low frequencies
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content_tags.quantities[0]
quantityR=−0.94 +/− 0.03 — correlation between pupil diameter and hippocampal ripple rate
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content_tags.quantities[1]
quantity5–20 mV — amplitude of the membrane-potential depolarizations time-locked to pupil microdilations
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content_tags.quantities[2]
quantity1.1 ± 0.1 s — lead time by which cortical depolarization precedes pupil dilation or eyelid opening
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content_tags.quantities[3]
quantity34±7% versus 18±4% — variance in 2-10 Hz membrane-potential power explained by pupil diameter versus by its temporal derivative
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content_tags.quantities[4]
quantity−74 ± 1.2 mV — average membrane potential corresponding to zero change on the arousal heatmap
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content_tags.quantities[5]
quantity37 ± 17%, 160 ± 120%, and 94 ± 55% — gain in explained variance for d-prime, bias and lick latency when pupil diameter is known in addition to tone intensity
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content_tags.quantities[6]
quantity~0.15 mm to ~1.8 mm — range over which pupil diameter varied in waking mice under constant low light
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content_tags.quantities[7]
quantity40–60% — fraction of full dilation that marks the optimal arousal band for neural coding and behaviour
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content_tags.quantities[8]
quantity40–60% — of full dilation - the optimum, and the same range for the neurons and for the behaviour
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content_tags.quantities[9]
quantity34±7% — variance in low-frequency membrane power explained by pupil diameter, against about half that for its derivative
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content_tags.quantities[10]
quantity1.3 ± 0.7 seconds — the lag by which the pupil derivative leads the diameter - which is why it looked like a better predictor
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content_tags.quantities[11]
quantity1.1 ± 0.1 s — by how much the cortical depolarisation precedes the pupil dilation it is locked to
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content_tags.quantities[12]
quantity20 ± 8% — explained variance added by knowing walking status, against 28 percent for pupil diameter and nothing for both together
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content_tags.quantities[13]
quantity12 of the 13 — measures whose optimum could not be distinguished from one another
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content_tags.quantities[14]
variabletarget tone level (independent)
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content_tags.variables[0]
variableintracellular injection of current (independent)
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content_tags.variables[1]
variablespontaneous activity (dependent)
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content_tags.variables[2]
variablebehavioral performance (dependent)
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content_tags.variables[3]
variablepupil diameter (covariate)
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content_tags.variables[4]
variablepupil
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content_tags.variables[7]
variablemembrane potential
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content_tags.variables[8]
variablegain
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content_tags.variables[9]
variablelatency
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content_tags.variables[10]
variablereliability
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content_tags.variables[11]
definitionmicrodilations
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content_tags.definitions[0]
definitioneye-indexed state
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content_tags.definitions[1]
definitionYerkes-Dodson curve
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content_tags.definitions[2]
definitioncanonical wakeful state
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content_tags.definitions[3]
definitionEIS
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content_tags.definitions[4]
definitionmicrodilation
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content_tags.definitions[5]
definitiongain
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content_tags.definitions[6]
definitionarousal
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content_tags.definitions[7]
key-termsarousal, pupil, ripple, eyelid, EIS, derivative, membrane potential, Vm, variance, oscillation, microdilation, walking, movement, whisking, ACtx, MG, layer, evoked, spontaneous, gain, reliability, coherence, detection, hit, false alarm, latency, inverted-U, U-shaped, state, sensory
Concepts
terms this paper carries, each with the sentence it was cut from — the layer that joins one paper to another. Terms with no quote yet are on CONCEPTS.html.
conceptpupil — not defined in this paper's own words; how arousal is read, non-invasively and moment to moment
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concepts[1]
concepthyperpolarized — not defined in this paper's own words; the state of the optimum - stable, quiet, and below rest
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concepts[9]
conceptevoked — not defined in this paper's own words; the sound response, largest and most reliable at the optimum
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concepts[19]
concepthit — not defined in this paper's own words; the trial type before which the cortex is most hyperpolarised and least variable
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concepts[25]
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.
coherence
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content_tags.statistics[0]
correlation CC as written: correlation coefficient CC
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content_tags.statistics[1]
ANOVA as written: ANOVA models with posthoc pairwise comparisons
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content_tags.statistics[2]
standard deviation S.D.
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content_tags.statistics[3]
standard error of the mean SEM
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content_tags.statistics[4]
bootstrap confidence interval CI as written: 68% bootstrap confidence interval
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content_tags.statistics[5]
resampling with replacement as written: resampled (1000 times with replacement)
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content_tags.statistics[6]
signal detection theory measures (d-prime and criterion) d′ as written: standard signal detection theory measures
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content_tags.statistics[7]
F-test F as written: F 1,7 =28.3
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content_tags.statistics[8]
explained variance as written: increase in explained variance
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]
cross-correlation as written: average of the correlation coefficient between all pairs of responses
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]
coherence
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[11]
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[12]
explained variance
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[13]
resampled
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[14]
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
methodpupil diameter as a continuous state measure, validated against hippocampal ripple rate and against eyelid opening in darkness, related to membrane potential, evoked responses and trial outcome — 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
phenomenonthe inverted-U between arousal and performance, and its cortical signature — a second reading DIFFERING from the recorded value; recorded: arousal-dependent auditory signal detection
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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