Task difficulty and performance induce diverse adaptive patterns in gain and shape of primary auditory cortical receptive fields

Atiani, Elhilali, David, Fritz, Shamma · Neuron · 2009 · neuroscience · 10.1016/j.neuron.2008.12.027 · record Atiani2009-task-difficulty-and-performance-induce-diver
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. Claim type is 'mechanism': the contribution is not that plasticity occurs (established in the cited earlier work) but how it is composed - a global gain term plus an orthogonal shape term whose signs differ for near- and far-from-target cells as difficulty changes. The behavioural arm (three ferrets, lick-withdrawal times, probe trials) makes 'behavioural-experiment' a defensible secondary method tag; 'electrophysiology' is primary because the claim rests on 237 single-unit STRFs recorded during behaviour. The 'replicates' relation is recorded on the paper's own wording 'broadly consistent with'; it is convergence with the same laboratory's prior work rather than an independent replication, which is why replication status remains 'untested'. The paper also reports an internal analytic robustness check using David et al. (2008)'s alternative shape measure, and a naive-animal control (93 units, no consistent STRF change without behaviour), neither of which is an external replication. 2026-08-13 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.
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.
Other fields define these differently the same word, defined another way in another discipline
attention 36 fields · plasticity 4 fields · signal-to-noise ratio 4 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 13 concept tags on this paper, 1 are 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.
spectrotemporal receptive fields STRF task difficulty signal-to-noise ratio gain shape contrast matched filter best frequency TORCs lick withdrawal time primary auditory cortex attention probe trials Lowering target SNR raises the attentional effort the animal must invest, and that increased effort drives larger adaptive changes in the receptive field Pulling a tone out of a masker requires simultaneously suppressing the responses driven by the concurrent noise and enhancing sensitivity to the tone Which direction a cell changes is set by what drives it: near cells are enhanced by proximity to the target tone, far cells are suppressed because the behaviourally relevant broadband masker drives them spectrotemporal receptive field TORC ferret plasticity masker conditioned avoidance passive suppression enhancement lick withdrawal time (LWT) performance level STRF gain naïve animals target SNR discrimination rate ANOVA Using ANOVA p-value median mean ANOVA across SNR conditions STRF power normalisation an alternative noise-aware shape-change method 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
conceptconceptsan idea the paper works with, quoted where the paper uses 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.neuron.2008.12.027. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 2 · claim_relations 8 · claim_type 1 · concept_edges 6 · concepts 1 · content_tags 53 · contribution 1 · controls 1 · limitations 10 · mechanisms 8 · 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 6 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
object_of_studyrapid changes in A1 spectrotemporal receptive fields while an animal detects a tone in noise
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object_of_studythe paper's words
research_questionwhether the magnitude of task-driven receptive field plasticity depends on task difficulty, and whether the change is one of gain or of shape
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research_questionthe paper's words
contributionseparates gain from shape change and shows they move in opposite directions, with the sign of the net effect set jointly by task difficulty and by the cell's distance from the target frequency
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contributionthe paper's words
claim_typemechanism
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claim_typethe paper's words
phenomenonrapid task-related plasticity of auditory cortical receptive fields
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
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mechanisms[0]the tagger's words
mechanismsmechanisms
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mechanisms[1]the tagger's words
mechanismsmechanisms
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mechanisms[2]the tagger's words
mechanismsa contrast matched filter: the acoustic features of the foreground are enhanced and those of the background suppressed
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mechanisms[3]1.0the tagger's words
mechanismsthe net effect keeps the target salient by two different routes - enhancing near cells when the tone is loud, suppressing far cells when it is buried
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mechanisms[4]1.0the tagger's words
mechanismsthe active STRF is modelled explicitly as a scaled passive STRF plus an orthogonal shape change, S_d = g.S_b + delta
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mechanisms[5]1.0the tagger's words
mechanismsSTRFs are measured from the reference TORCs alone, never during target presentation, so the measurement does not include the target
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mechanisms[6]0.85the tagger's words
mechanismsattentional load is validated behaviourally by inserting identical intermediate-difficulty probes into easy and hard blocks
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mechanisms[7]1.0the tagger's words
limitationsSTRF frequency resolution is too coarse to exclude that some A1 neurons build narrow matched filters at the exact target frequency.
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limitations[0]the tagger's words
limitationsIt is unknown how far the gain-reduction/shape-enhancement pattern is specific to the tone-in-noise task, its avoidance paradigm, or the fixed noise level.
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limitations[1]the tagger's words
limitationsMost cells could not be held across multiple task conditions, so difficulty comparisons are largely between rather than within cells.
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limitations[2]the tagger's words
limitationsDisentangling stimulus structure from behavioural paradigm requires further experiments; the interpretations offered are conjectures.
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limitations[3]the tagger's words
limitationsThe 0.6-octave boundary dividing near from far cells was an arbitrary choice.
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limitations[4]the tagger's words
limitationsthe near/far dividing line is arbitrary, stated as such, and shown not to change the results
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limitations[5]1.0the tagger's words
limitationsthe standard STRF power normalisation used in the group's previous papers does not separate gain from shape
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limitations[6]1.0the tagger's words
limitationsthe dependence of gain on task difficulty was NOT confirmed when all performance levels were pooled
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limitations[7]1.0the tagger's words
limitationschanges often persisted after the task with no systematic relation to the type of plasticity
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limitations[8]1.0the tagger's words
limitationsmost cells were recorded in only one task condition, so within-cell comparison across difficulties is limited
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limitations[9]0.8the tagger's words
transfersthe finding that harder tasks recruit more attention, and the probe-trial design used to demonstrate it
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transfers[0]1.0the tagger's words
transfersthe matched filter, imported as the model for how a receptive field should adapt to separate foreground from background
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transfers[1]1.0the tagger's words
controlsthe target tone frequency and signal-to-noise ratio held constant throughout a given task
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eliminates: a within-task change in the stimulus itself explaining the receptive-field change attributed to task difficulty · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
conceptsshape
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also in: shape
concepts[5]0.85the paper's words
sensesplasticity here is rapid and reversible - measured as the difference between active and passive STRFs within a session, not as learning
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senses[0]0.95the tagger's words
sensestask difficulty is operationalised as target SNR, and validated separately by lick withdrawal time and by probe-trial performance
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senses[1]1.0the tagger's words
sensesgain and shape are given distinct definitions by an explicit model: gain is the fraction of the STRF that keeps its original shape, shape change is the orthogonal remainder
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senses[2]1.0the tagger's words
sensesnear and far are defined by a 0.6-octave cut from the target frequency, which the paper calls arbitrary and tests
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senses[3]1.0the tagger's words
alternativesmethod: STRFs measured from responses to modulated broadband TORC references during and before a conditioned-avoidance tone-in-noise detection task, at target SNRs from +15 to -10 dB
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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: attention-driven receptive field plasticity in primary auditory cortex
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a second reading DIFFERING from the recorded value. Recorded: rapid task-related plasticity of auditory cortical receptive fields — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsextends → Fritz et al. (2003; 2005; 2007a,b), the group's earlier demonstrations of rapid task-related STRF plasticity with pure-tone targets
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claim_relations[0]the tagger's words
claim_relationsbuilds-on → the "contrast matched filter" hypothesis of Fritz et al. (2007a)
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claim_relations[1]the tagger's words
claim_relationsextends → visual studies of task difficulty and attention (Boudreau et al., 2006; Spitzer et al., 1988; Spitzer and Richmond, 1991), which did not separate gain from shape
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differs in: THE AXIS IS GAIN AGAINST SHAPE. 'Quotation withheld' Two ways a receptive field can change that a single measure cannot separate - so the earlier result is a sum of two effects that this paper's [7] then shows move differently with difficulty.
claim_relations[2]the tagger's words
claim_relationsreplicates → earlier task-related plasticity results from the same group in tone-versus-TORC and tone-versus-tone paradigms (Fritz et al., 2003; 2005; 2007b)
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claim_relations[3]the tagger's words
claim_relationsextends → Fritz et al. on rapid STRF plasticity with pure-tone targets
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claim_relations[4]1.0the tagger's words
claim_relationsqualifies → the contrast matched filter model, by showing the sign of the change depends on difficulty and on distance from target
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differs in: THE AXIS IS DISTANCE FROM THE TARGET FREQUENCY, AND THE SIGN FLIPS ACROSS IT. 'when we grouped the plasticity changes by near and far cells, it became apparent that near cells on average were enhanced (red) whereas far cells were suppressed (blue).' The matched-filter model is qualified rather than refuted: it holds, with a sign that depends on where the cell sits.
claim_relations[5]1.0the tagger's words
claim_relationsextends → visual studies showing task difficulty enhances attentional modulation, which did not separate gain from shape
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differs in: THE AXIS IS GAIN AGAINST SHAPE, on a second target. Same sentence as [2]: 'THESE MEASUREMENTS DID NOT DISTINGUISH RECEPTIVE FIELD GAIN FROM SHAPE CHANGES.' One limitation of one earlier method, disqualifying it for two different claims this paper makes - which is why the sentence supports two rows rather than one.
claim_relations[6]1.0the tagger's words
claim_relationsqualifies → their own gain-versus-difficulty result, which did not survive pooling across performance levels
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differs in: THE AXIS IS WHETHER THE CELLS ARE POOLED ACROSS PERFORMANCE LEVELS. 'Quotation withheld.' The paper's own effect survives one analysis choice and not another, and it says which - an analyst decision reported as a result rather than buried in methods.
claim_relations[7]1.0the tagger's words
concept_edgesEVIDENCE CORRECTED (RE-GROUNDED) · task difficulty —operationalised-as→ signal-to-noise ratio
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EVIDENCE CORRECTED: RE-GROUNDED. The old span named neither endpoint; this clause states the operationalisation outright - difficulty IS the signal-to-noise ratio in this design. — strength of the relation: 1
concept_edges[0]0.85the tagger's words
concept_edgesgain —contrasted-with→ shape
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strength of the relation: 1 · note: the paper's central result: they move in OPPOSITE directions in the same cells - gain decreased while shape changes were excitatory
also in: gain · shape
concept_edges[1]1.0the tagger's words
concept_edgestask difficulty —causes→ attention
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strength of the relation: 0.9 · note: and it is tested rather than assumed, via probe trials of identical difficulty in easy and hard blocks Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span states a hypothesis about attentional resources; causes states it as a finding.
concept_edges[2]1.0the tagger's words
concept_edgesattention —operationalised-as→ probe trials
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strength of the relation: 0.9 Audit 2026-09-28: DOUBTFUL - read against its span and kept, but probe trials were inserted to test attention; operationalised-as is loose.
concept_edges[3]1.0the tagger's words
concept_edgestask difficulty —operationalised-as→ signal-to-noise ratio
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strength of the relation: 1 · note: RE-GROUNDS another row of this record: the relation was always stated, the endpoint names were not in the span.
concept_edges[5]1.0the tagger's words
concept_edgesSTRF —operationalised-as→ TORCs
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strength of the relation: 0.95 · note: RE-GROUNDS another row of this record: the old span named ONE endpoint.
also in: STRF · TORCs
concept_edges[6]0.9the tagger's words
content_tagsANOVA
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also in: ANOVA
content_tags.statistics[0]the tagger's words
content_tagsp-value
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also in: p-value
content_tags.statistics[1]the tagger's words
content_tagsmedian
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also in: median
content_tags.statistics[2]the tagger's words
content_tagsmean
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also in: mean
content_tags.statistics[3]the tagger's words
content_tagsANOVA across SNR conditions
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content_tags.statistics[4]0.9the tagger's words
content_tagsSTRF power normalisation
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content_tags.statistics[5]1.0the tagger's words
content_tagsan alternative noise-aware shape-change method
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content_tags.statistics[6]1.0the tagger's words
content_tagsEngaging in the detection task reduced overall STRF gain in the majority of recorded cells relative to the passive state.
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content_tags.findings[0]the tagger's words
content_tagsShape changes, unlike gain changes, were excitatory and frequency-specific, concentrated in cells tuned near the target tone.
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content_tags.findings[1]the tagger's words
content_tagsThe combined gain and shape adaptations sharpened the target-versus-noise representation differently at high and low SNR.
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content_tags.findings[2]the tagger's words
content_tagsThe magnitude of receptive-field adaptation tracked behavioural performance, being greatest in the best-performing sessions.
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content_tags.findings[3]the tagger's words
content_tagsSuppression of receptive fields grew as the task became harder and reached significance only in the low-SNR condition.
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content_tags.findings[4]the tagger's words
content_tagsBehaviourally, lick withdrawal time lengthened as target SNR fell, in all three ferrets.
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content_tags.findings[5]the tagger's words
content_tagsIdentical intermediate probe trials were discriminated better within hard blocks than within easy blocks.
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content_tags.findings[6]the tagger's words
content_tagsNaive animals exposed to the identical stimuli showed some minor but no consistent receptive-field changes.
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content_tags.findings[7]the tagger's words
content_tagsThe gain reduction was a broad, difficulty-independent effect present in near and far cells alike, in contrast to the difficulty-dependent shape changes.
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content_tags.findings[8]the tagger's words
content_tagsGoing from high to low SNR targets suppressed firing to the reference noise by roughly a third.
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content_tags.findings[9]the tagger's words
content_tagsgain decreased in most cells during task performance while shape changes were excitatory and specific
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content_tags.findings[10]1.0the tagger's words
content_tagsgrouped by distance from the target, near cells were on average enhanced and far cells suppressed
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content_tags.findings[11]1.0the tagger's words
content_tagssuppression became significant only in the low SNR tasks, at about -13%
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content_tags.findings[12]1.0the tagger's words
content_tagsthe same tone placed far from a cell's BF produced suppression where near placement produced enhancement
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content_tags.findings[13]1.0the tagger's words
content_tagsthe trend of increasing suppression at low SNR repeated in this subset, with the STRF changes amplified
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content_tags.findings[14]1.0the tagger's words
content_tagsnaive animals given identical stimuli showed no consistent STRF changes
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content_tags.findings[15]1.0the tagger's words
content_tagsraising difficulty suppressed firing to the reference noise by about a third overall
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content_tags.findings[16]1.0the tagger's words
content_tagsprobe trials of identical difficulty were detected about 10% better inside hard blocks than inside easy ones
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content_tags.findings[17]1.0the tagger's words
content_tagsgain was at or below 1 in most cells, with medians near 0.9 for near-high-SNR and 0.7 for far-low-SNR
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content_tags.findings[18]1.0the tagger's words
content_tagscontrast matched filter
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content_tags.definitions[0]the tagger's words
content_tagslick withdrawal time (LWT)
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content_tags.definitions[1]the tagger's words
content_tagsTORC
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also in: TORC
content_tags.definitions[5]the tagger's words
content_tagscontrast matched filter
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content_tags.definitions[6]1.0the tagger's words
content_tagslick withdrawal time
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content_tags.definitions[8]1.0the tagger's words
content_tagsperformance level
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content_tags.variables[2]the tagger's words
content_tagsSTRF gain
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also in: STRF gain
content_tags.variables[4]the tagger's words
content_tagslick withdrawal time
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content_tags.variables[5]the tagger's words
content_tagsnaïve animals
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content_tags.variables[6]the tagger's words
content_tagstarget SNR
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also in: target SNR
content_tags.variables[8]1.0the tagger's words
content_tagsdiscrimination rate
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content_tags.variables[10]1.0the tagger's words
content_tags237 — single cells in the population analysis of receptive-field plasticity, from two behaving ferrets
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content_tags.quantities[0]the tagger's words
content_tags93, 57, and 87 — single units in the high, mid and low SNR difficulty groups respectively
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content_tags.quantities[1]the tagger's words
content_tagsmean = −13.1 % — average amplitude change at best frequency in the low-SNR (hardest) condition
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content_tags.quantities[2]the tagger's words
content_tags10–30% — reduction in STRF gain caused by task engagement
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content_tags.quantities[3]the tagger's words
content_tags0.9 and 0.7 — median gain values for near cells in high-SNR tasks and far cells in low-SNR tasks (values below 1 mean suppression)
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_tags350 ms — lick withdrawal time after target onset in the easiest (pure tone) detection task
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_tags0.6 octaves — best-frequency distance criterion separating near cells from far cells
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_tags85 — cells from the top-third best-performing sessions analysed separately
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_tags93 units from 3 naïve animals — control sample recorded with identical stimuli but no behavioural task
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[8]the tagger's words
content_tags0.6 octaves — the near/far dividing line, called arbitrary and shown not to matter
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_tags112 near versus 125 far cells — the balance the cut produced
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_tags93, 57 and 87 single units in high, mid and low SNR — the difficulty groups
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_tags-13.1%, p < 0.05 — mean suppression, significant only at low SNR
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_tags58.6% versus 69.2% — probe detection in easy versus hard blocks at identical SNR
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
content_tags93 units from 3 naive animals — the no-behaviour control
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[14]1.0the tagger's words
content_tags350 ms — lick withdrawal time for a pure-tone target
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[15]0.85the tagger's words

Interpretive choices

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What was deliberately not tagged, and why, is recorded per paper by the tagger. Where a paper has no such record, this page says so rather than implying that nothing was declined.

© 2026 Shir Sivroni · ORCID 0009-0000-2597-2824