Sleep Differentially Affects Early and Late Neuronal Responses to Sounds in Auditory and Perirhinal Cortices — TEI-tagged view

Sela, Krom, Bergman, Regev, Nir · 2020 · sleep-neuroscience · doi:10.1523/jneurosci.1186-19.2020 · 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 (html); every evidence string verified as an exact substring of that text before merging. Replication status refers to the paper's headline claim (robust PRC attenuation), which it describes as "the first direct neuronal evidence for attenuation of auditory responses in natural sleep"; the secondary auditory-cortex result is however explicitly a replication — "Neuronal responses in AC showed modest (∼10%) differences in response gain across vigilance states, replicating previous studies." The fourth claim_relation names the thalamic gating model as target although the quoted sentence says only "a discrete “gate” at a specific brain region"; the model is named earlier in the Introduction ("Early studies ( Gücer, 1979 ; Livingstone and Hubel, 1981 ) contributed to the “thalamic gating” model ( Steriade and McCarley, 1990 )"). The corpus should note that the sample is male rats only — the paper states this in Materials and Methods but never frames it as a limitation, so it is not recorded as one. 2026-08-12 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.
A publisher correction exists for this article. PUBLISHER CORRECTION NOT FETCHED. A publisher correction notice exists for this article and is NOT present in the staged text: searching the staged body for 'correction', 'erratum', 'retraction' and 'publisher' returns nothing in Sela2020 and, in Pho2018, only a statistical use of the word ('no correction for multiple comparisons'). Every tag on this record is therefore anchored to the UNCORRECTED text. The correction has not been fetched and the text has NOT been re-staged, by a project decision of 2026-08-13. This is record-level provenance and carries no quote, because there is nothing in the staged text to quote.
Other fields define these differently the same word, defined another way in another discipline
standard deviation 3 fields · confidence interval 2 fields · response latency 2 fields
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
sleep3 sensory disconnection response latency6 perirhinal cortex5 auditory cortex19 thalamic gating targeted memory reactivation 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
phenomenonsleep attenuation of auditory responses in high-order cortex— “The robust attenuation of late responses prevalent in PRC represents a novel neural correlate of sensory disconnection during sleep, opening new avenues for investigating the mediating mechanisms.”
claim-typeexistence— “We find that PRC exhibits robust attenuation of response magnitude in both NREM sleep and REM sleep, whereas AC neurons are only modestly affected by sleep.”
method — evidence— “Here we recorded spiking activity in 435 neuronal clusters evoked by acoustic stimuli in the perirhinal cortex (PRC) and in AC of freely behaving male rats across wakefulness and sleep. Keywords:”
object of study — evidence— “For the above reasons, we set out to directly compare auditory responses in PRC neurons across wakefulness and natural sleep.”
research question — evidence— “However, investigation of sensory responses downstream from sensory cortical regions at the level of neuronal spiking activity is still missing.”
contribution — evidence— “In addition, we find that the response latency of each neuronal cluster (typically <20 ms in AC and >40 ms in PRC) is correlated with the degree of NREM sleep attenuation, and better predicts attenuation than does cortical region.”
mechanismNREM sleep imposes a graded, cumulative attenuation that builds up along the processing hierarchy rather than a single discrete gate at one anatomical station— “The robust correlation of NREM sleep attenuation with response latency suggests that NREM sleep may exert a gradual attenuation of cortical responses, rather than imposing a discrete “gate” at a specific brain region.”mechanisms[0]
mechanismBreakdown of cortical effective connectivity in NREM sleep means signals lose more strength at each additional synapse, so responses arriving later are attenuated more— “The fact that late responses exhibit stronger attenuation in NREM sleep is in line with the notion of breakdown of cortical effective connectivity during NREM sleep ( Massimini et al., 2005 ) that may lead to greater attenuation with increasing synaptic connections.”mechanisms[1]
mechanismthe disconnection is not at the sensory front end: auditory cortex responds nearly as strongly asleep as awake, so whatever gates perception acts further downstream— “In addition, several recent studies in the auditory system showed that responses in the primary and secondary auditory cortex (AC) are comparable across wakefulness and natural sleep”mechanisms[2]
mechanismperirhinal cortex is the candidate site because it is multimodal, downstream of several primary cortices, and carries intrinsic inhibition controlling flow to the hippocampus— “Moreover, it has been shown to contain intrinsic inhibitory mechanisms that affect information flow to the hippocampus ( de Curtis and Paré, 2004 ; Pelletier et al., 2004 ).”mechanisms[3]
mechanismthe operative variable is TIME not PLACE: whatever suppresses the response acts on late activity wherever it occurs, not on a particular region— “Beyond the separation according to cortical region, response latency in each neuronal cluster was correlated with the degree of NREM sleep attenuation, such that late (>40 ms) responses in all monitored regions diminished during NREM sleep.”mechanisms[4]
mechanismthe older explanation - thalamic gating - was already in trouble, because olfactory responses that bypass the thalamus are attenuated too— “However, this view was challenged by sleep attenuation of responses in the olfactory cortex that are not relayed through thalamus ( Murakami et al., 2005 ).”mechanisms[5]
replicates reports of preserved auditory cortical responses across vigilance states (Issa and Wang, 2008; Nir et al., 2015)— “In line with previous studies ( Issa and Wang, 2008 ; Nir et al., 2015 ), spiking responses in AC neurons exhibited similar profiles across vigilance states, whereas PRC neurons exhibited weaker responses in both NREM and REM sleep ( Fig. 4 ).”claim_relations[0]
extends Krom et al. (2018), the same early/late distinction observed in humans under light anesthesia— “In addition, a recent study comparing auditory responses at the neuronal level in humans across wakefulness and light anesthesia found a similar distinction between relatively preserved early responses in AC versus robust attenuation of late responses in association cortex ( Krom et al., 2018 ).”claim_relations[1]
contradicts Pereira et al. (2007), increased hippocampal responses to somatosensory microstimulation during sleep— “This discrepancy with the robust attenuation we find here in rat PRC may reflect differences in sensory modalities, or differences between natural stimuli used here versus ultra-short electrical stimulation.”claim_relations[2]
contradicts the "thalamic gating" model (Steriade and McCarley, 1990), which posits a discrete gate rather than graded attenuation— “The robust correlation of NREM sleep attenuation with response latency suggests that NREM sleep may exert a gradual attenuation of cortical responses, rather than imposing a discrete “gate” at a specific brain region.”claim_relations[3]
replicates prior findings that auditory cortex responses are comparable across wakefulness and sleep— “Neuronal responses in AC showed modest (∼10%) differences in response gain across vigilance states, replicating previous studies.”claim_relations[4]
contradicts the thalamic gating model of sleep-related sensory disconnection— “However, this view was challenged by sleep attenuation of responses in the olfactory cortex that are not relayed through thalamus ( Murakami et al., 2005 ).”claim_relations[5]
extends human imaging work showing attenuation downstream of sensory cortex during sleep— “However, investigation of sensory responses downstream from sensory cortical regions at the level of neuronal spiking activity is still missing.”claim_relations[6]
sensesleep - behavioural: the state in which an animal stops responding to what happens around it— “Reduced behavioral responses to external sensory stimuli are an essential definition of sleep ( Carskadon and Dement, 2011 ) and are ubiquitous across the animal kingdom ( Cirelli and Tononi, 2008 ).”senses[0]
sensesleep - neural: a set of vigilance states, NREM and REM, that attenuate responses by different amounts— “By contrast, PRC neuronal responses were robustly attenuated by 47% and 36% during NREM sleep and REM sleep, respectively.”senses[1]
sensedisconnection - not an absence of processing: discriminative sensory processing continues while perception does not follow— “During sleep, external stimuli rarely affect perception ( Nir and Tononi, 2010 ), but it is also clear that some discriminative sensory processing persists ( Andrillon et al., 2016 ).”senses[2]
controlneurons with peak latencies in the 20-40 ms band DROPPED from the latency-based split, keeping only <20 ms and >40 ms — held — rules out ambiguous cases deciding the early/late contrast - a neuron near the boundary can fall either side on noise, and those are exactly the cases a threshold chosen after looking would assign favourablycontrols[0]
controlthe first second after each auditory stimulus EXCLUDED before smoothing when detecting OFF periods — held — rules out stimulus-induced firing reductions being counted as spontaneous OFF periods - the sound itself suppresses firing, so an OFF-period detector run over that window would find the stimulus and report sleepcontrols[1]
differs inthe "thalamic gating" model (Steriade and McCarley, 1990), which posits a discrete gate rather than graded attenuation — THE AXIS IS RESPONSE LATENCY. 'The robust correlation of NREM sleep attenuation with response latency suggests that NREM sleep may exert A GRADUAL ATTENUATION of cortical responses, RATHER THAN IMPOSING A DISCRETE GATE at a specific brain region.' A gate predicts a step at one locus; the graded alternative predicts a monotone relationship with latency - so the two accounts differ on a measurable parameter rather than on a location.claim_relations[3]
retractedEVIDENCE CORRECTED (RE-GROUNDED) · perirhinal cortex —contrasted-with→ auditory cortex — RE-GROUNDED. The old span named neither area. This sentence names both and states the contrast with its direction - modest changes in auditory cortex, robust attenuation in perirhinal.
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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[0]
auditRETRACTED (THE 'to' ENDPOINT IS THE TAGGER'S PHRASE) · sleep —causes→ sensory disconnection — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says responses were attenuated during sleep; the endpoint 'sensory disconnection' is interpretive.
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concept_edges[2]
retractedRETRACTED (THE 'to' ENDPOINT IS THE TAGGER'S PHRASE) · sleep —causes→ sensory disconnection — THE 'to' ENDPOINT IS THE TAGGER'S PHRASE. The edge is 'sleep --causes--> sensory disconnection'. The span: 'WERE ROBUSTLY…' 'Sensory disconnection' does not occur - as a word or at all. What the sentence reports is a MEASURED ATTENUATION with two numbers attached; 'sensory disconnection' is the interpretation those numbers are usually put to, supplied from knowledge of the literature. TRUE OF THE FIELD, UNSUPPORTED BY THE SPAN - and note the edge also drops the 47% and 36%, which are the only things the sentence actually establishes. Found by the neighbour query.
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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[2]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
outcome (stated by the tagger, no quotation)control — heldcontrols[1]
for which uselimitation — any claim about behaviourally relevant sounds: WE USED A PASSIVE AUDITORY STIMULATION SCHEME WHERE SOUNDS WERE NOT ASSOCIATED WITH SPECIFIC BEHAVIORS OR REWARDS
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limitations[0]
for which uselimitation — the latency-attenuation relationship outside NREM: IN REM SLEEP, ATTENUATION DID NOT CLEARLY FOLLOW RESPONSE LATENCY AS WAS THE CASE DURING NREM SLEEP (DATA NOT SHOWN)
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limitations[1]
for which uselimitation — the REM explanation only: ALTHOUGH SPECULATIVE, IT IS POSSIBLE THAT, DURING REM SLEEP, ONGOING MNEMONIC ACTIVITY DURING DREAMING INTERFERES WITH BOTTOM-UP SENSORY PROCESSES
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limitations[2]
for which uselimitation — the population the result speaks for: EXPERIMENTS WERE PERFORMED IN 8 LONG-EVANS AND 2 WISTAR ADULT MALE RATS
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limitations[3]
for which uselimitation — the causal reading: the finding REPRESENTS A NOVEL NEURAL CORRELATE OF SENSORY DISCONNECTION DURING SLEEP, with the mediating mechanism left to future work
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limitations[4]
for which uselimitation — any comparison with the anaesthetised perirhinal literature: SEPARATE STUDIES DURING ANESTHESIA DID NOT REVEAL SIGNIFICANT AUDITORY RESPONSES AS ONE MOVES VENTRALLY TOWARD RHINAL cortex, while waking studies find reliable responses
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limitations[5]
for which uselimitation — the evidence base this study joins: SURPRISINGLY, ONLY FEW STUDIES INVESTIGATED SENSORY RESPONSES DURING NATURAL SLEEP AT THE NEURONAL LEVEL
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limitations[6]
replicationuntested
limitationStimulation was passive, so effects of behaviourally relevant or rewarded sounds were not assessed.
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limitations[0]
limitationThe latency-attenuation relationship found in NREM sleep did not hold in REM sleep.
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limitations[1]
limitationThe proposed REM-sleep explanation is offered as speculation.
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limitations[2]
limitationTen rats, male only, in two strains.
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limitations[3]
limitationThe result is a correlate, and the paper says the mediating mechanism remains to be investigated.
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limitations[4]
limitationPrior evidence about perirhinal auditory responses is inconsistent, with anaesthetised preparations failing to find them at all.
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limitations[5]
limitationThe literature the study enters is thin: few studies have examined sensory responses in natural sleep at the neuronal level at all.
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limitations[6]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingAuditory cortex responses changed only about 10% across vigilance states, whereas perirhinal cortex responses were cut by 47% in NREM sleep and 36% in REM sleep.
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content_tags.findings[0]
findingMedian gain factors quantified the perirhinal attenuation as -47% in NREM sleep and -36% in REM sleep, with confidence intervals excluding zero.
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content_tags.findings[1]
findingAuditory cortex neurons showed only small median gain changes of -12% in NREM sleep and -6% in REM sleep.
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content_tags.findings[2]
findingAcross all recorded neurons, the later a neuron's peak response latency, the more its response was attenuated in NREM sleep, a strong negative correlation.
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content_tags.findings[3]
findingNeurons whose latency did not match their region behaved according to latency, not anatomy: late-responding auditory cortex neurons were attenuated by 43% while early-responding perirhinal neurons were attenuated by only 3%.
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content_tags.findings[4]
findingEarly-responding neurons showed modest gain change in NREM sleep (-5.4%) against robust attenuation (-41%) for late-responding neurons, an eightfold difference.
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content_tags.findings[5]
finding85% of neurons peaking under 20 ms were in auditory cortex and 89% of those peaking over 40 ms were in perirhinal cortex.
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content_tags.findings[6]
findingBaseline firing rates in auditory cortex were 21-26% higher in wakefulness and REM sleep than in NREM sleep.
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content_tags.findings[7]
findingWithin NREM sleep, late-responding neurons responded more strongly to sounds delivered during local OFF periods than during ON periods.
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content_tags.findings[8]
findingShort-latency neurons tended to have narrower action potential waveforms than long-latency neurons.
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content_tags.findings[9]
findingAuditory cortex responses change only about 10% between waking and sleep - replicating earlier work.
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content_tags.findings[10]
findingPerirhinal responses fall by 47% in NREM and 36% in REM sleep.
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content_tags.findings[11]
findingLate responses are suppressed wherever they occur, so latency rather than anatomy is the better predictor of sleep attenuation.
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content_tags.findings[12]
findingSleep does not abolish sensory processing - some discriminative processing survives even though stimuli rarely reach perception.
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content_tags.findings[13]
quantity435 — Number of neuronal clusters whose sound-evoked spiking was recorded
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content_tags.quantities[0]
quantity47% and 36% — Attenuation of perirhinal cortex responses during NREM sleep and REM sleep respectively
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content_tags.quantities[1]
quantityr = −0.55, p = 10 −15 — Correlation between a neuron's peak response latency and its NREM sleep gain attenuation
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content_tags.quantities[2]
quantity85% / 89% — Proportion of sub-20 ms neurons located in auditory cortex, and of over-40 ms neurons located in perirhinal cortex
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content_tags.quantities[3]
quantity−43% ... −3% — NREM gain factor for latency-atypical neurons: late-responding AC neurons versus early-responding PRC neurons
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content_tags.quantities[4]
quantity8-fold — Difference in NREM attenuation between early- and late-responding neurons recorded on the same microwire
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content_tags.quantities[5]
quantity21%–26% — Excess baseline firing rate of auditory cortex neurons in wakefulness or REM sleep relative to NREM sleep
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content_tags.quantities[6]
quantityn = 10 — Number of adult male rats implanted with microwire arrays
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content_tags.quantities[7]
quantityn = 178 ... n = 132 — Numbers of responsive neuronal clusters assigned to auditory cortex and to perirhinal cortex by histology
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content_tags.quantities[8]
quantity15 ± 6.3%, 41 ± 6.2%, and 8 ± 2.4% — Proportion of experiment time spent in quiet wake, NREM sleep and REM sleep
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content_tags.quantities[9]
quantity435 neuronal clusters — size of the recorded population across both regions
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content_tags.quantities[10]
quantity47% (NREM) and 36% (REM) — attenuation of perirhinal auditory responses relative to wakefulness
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content_tags.quantities[11]
quantity~10% — the much smaller change in auditory cortex response gain across vigilance states
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content_tags.quantities[12]
quantity<20 ms in AC, >40 ms in PRC — typical response latencies, and the axis along which attenuation is organised
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content_tags.quantities[13]
quantity10 rats — 8 Long-Evans and 2 Wistar adult males
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content_tags.quantities[14]
variablevigilance state (independent)
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content_tags.variables[0]
variablecortical region (independent)
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content_tags.variables[1]
variableresponse latency (moderator)
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content_tags.variables[4]
definitiongain factor
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content_tags.definitions[0]
definitionearly- and late-responding neurons
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content_tags.definitions[1]
definitionspontaneous firing rate
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content_tags.definitions[2]
definitionthalamic gating
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content_tags.definitions[3]
definitionneuronal ON/OFF periods
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content_tags.definitions[4]
definitionsleep
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content_tags.definitions[5]
definitionperirhinal cortex
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content_tags.definitions[7]
key-termssleep, NREM, REM, auditory cortex, perirhinal cortex, response latency, attenuation, thalamic gating, sensory disconnection, spiking activity, vigilance states, targeted memory reactivation
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.
conceptsleep — not defined in this paper's own words; a state defined by reduced behavioural response to external stimuli, found across the animal kingdom
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concepts[0]
conceptsensory disconnection — not defined in this paper's own words; the failure of sensory input to reach perception during sleep - the phenomenon the attenuation is offered as a correlate of
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concepts[1]
conceptresponse latency — not defined in this paper's own words; how long after the stimulus a neuron fires - here the variable that predicts sleep attenuation better than anatomy does
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concepts[2]
conceptperirhinal cortex — not defined in this paper's own words; a multimodal region downstream of primary sensory cortices, supporting perception and recognition memory and gating flow to the hippocampus
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concepts[3]
conceptauditory cortex — not defined in this paper's own words; the primary and secondary sensory region whose responses barely differ between waking and sleep
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concepts[4]
conceptthalamic gating — not defined in this paper's own words; the older account in which thalamic burst firing and spindles block signals from reaching cortex during sleep
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.
concepts[5]
concepttargeted memory reactivation — not defined in this paper's own words; re-presenting training cues during sleep, which does reach the hippocampus - the evidence that sound can still move association 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.
concepts[6]
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.
Mann-Whitney U test as written: two-tailed 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[0]
Mann-Whitney test (one-tailed) as written: a one-tailed Mann–Whitney 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[1]
false discovery rate correction for multiple comparisons as written: corrected via false discovery rate
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]
significance level (alpha) α as written: with base α of 0.001
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 signed-rank test as written: via signed-rank tests
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]
correlation r as written: Pearson correlation coefficients
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]
confidence interval CI as written: [lower and upper] 95% CIs around the median
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]
z score z as written: expressed as z scores relative to baseline
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]
standard deviation SD as written: mean ± SD
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]
correlation r
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]
proportion %
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 typeexistence — a second reading reaching the same value; recorded: existence
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
phenomenonsensory disconnection during sleep, and where along the cortical hierarchy it takes effect — a second reading DIFFERING from the recorded value; recorded: sleep attenuation of auditory responses in high-order 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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