Inferring the role of inhibition in auditory processing of complex natural stimuli

Schinkel-Bielefeld, David, Shamma, Butts · Journal of Neurophysiology · 2012 · neuroscience · 10.1152/jn.01173.2011 · record Schinkel-Bielefeld2012-inferring-the-role-of-inhibition-in-auditory
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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. method is tagged electrophysiology because the evidence base is extracellular single-unit recording in awake ferrets, but the paper's actual novelty is computational: a nonlinear modelling framework (GNM) fitted to those spike trains and compared against a linear-nonlinear STRF model; no separate 'modelling' value exists in the method vocabulary. Claim type could also be read as 'method' (a new modelling framework); mechanism was chosen because the paper's headline result is causal — nonlinear A1 properties arise from the interplay of excitation and inhibition. A fourth possible relation (tension with Scholl et al. 2010, who found offset responses arising from a distinct set of inputs) was left out because the paper reports it as a competing account of prior intracellular data rather than as a contradiction of its own claim. 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.

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Other fields define these differently the same word, defined another way in another discipline
prediction 24 fields · context 10 fields · performance 5 fields · model 4 fields · cross-validation 3 fields · spike 2 fields · and 4 more →

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 38 concept tags on this paper, none is grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.

STRF GNM model nonlinear averaging excitation inhibition cotuned intracellular extracellular A1 neuron spike intensity tuning onset offset speech stimulus context mechanism prediction performance generality efficient proof guarantee STC MID multilinear gain latency input measure framework tonotopic isolation resolution natural Intensity tuning arises because the excitatory/inhibitory balance shifts with sound level: at high intensities the filtered inputs land on the saturating part of the internal nonlinearities, so the excitatory lead no longer produces a large net drive. Onset responses are generated by excitation reaching the cell before inhibition, with the lagging inhibition cutting the response short. Offset responses are generated when the excitatory nonlinearity is amplified relative to the inhibitory one, so inhibition falls away before excitation does and firing rebounds. Sustained firing is generated when, at large filter outputs, the internal nonlinearity amplifies excitation more than inhibition. generalized nonlinear model (GNM) onset-triggered average silence rate and sound rate effective time kernel level-rate correlation coefficient cross-validated LL firing rate sound pressure level maximum likelihood estimation cross-validation cross-validated LL (LL x ) t-test t -test paired t-test paired t -test correlation correlation coefficient principal components analysis principal components analysis and k -means clustering median median difference entropy entropy as a measure of their overall width explained variance (r-squared) k -means clustering electrophysiology

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second reading	`alternatives`	an independent re-reading of a facet, recorded whether or not it agrees
relation to other work	`claim_relations`	how this paper stands to another: agrees, contradicts, extends, qualifies
claim type	`claim_type`	the shape of the claim: existence, mechanism, method, critique, prediction, null
concept edge	`concept_edges`	an asserted relation between two concepts, with the sentence asserting it
content tag	`content_tags`	a finding, definition, quantity, statistic or variable named in the paper
contribution	`contribution`	what the paper claims to add
control	`controls`	an experiment run to eliminate a rival explanation
limitation	`limitations`	what the paper says its evidence cannot support
mechanism	`mechanisms`	how the reported effect is said to come about
method	`method`	how the paper produced its evidence
object of study	`object_of_study`	what the paper is about — the thing it investigates
phenomenon	`phenomenon`	the effect or regularity under study
research question	`research_question`	the question the paper sets itself
sense	`senses`	a distinct use of a term, quoted where the paper fixes it
transfer	`transfers`	a 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/10.1152/jn.01173.2011. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 9 · claim_type 1 · concept_edges 5 · content_tags 67 · contribution 1 · controls 1 · limitations 9 · mechanisms 8 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 15 · transfers 4

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 5 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facet	tag	field	conf.	whose words
object_of_study	how the interplay of excitation and inhibition shapes A1 responses to natural sounds, and whether it can be recovered from extracellular spikes alone Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`object_of_study`		the paper's words
research_question	whether a model that separates excitatory from inhibitory contributions predicts A1 spike trains better than the linear receptive field, and recovers properties matching intracellular measurements Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`research_question`		the paper's words
contribution	a nonlinear model whose inferred excitation and inhibition predict spikes better than the linear model and reproduce intensity tuning, onset and offset responses that were never explicitly built in Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`contribution`		the paper's words
claim_type	mechanism Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_type`		the paper's words
phenomenon	excitatory-inhibitory interplay in primary auditory cortex Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`phenomenon`		the paper's words
method	electrophysiology Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: electrophysiology	`method`		the paper's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: Intensity tuning arises because the excitatory/inhibitory balance shifts with sound level: at high intensities the filtered inputs land on the saturating part of the internal nonlinearities, so the excitatory lead no longer produces a large net drive.	`mechanisms[0]`		the tagger's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: Onset responses are generated by excitation reaching the cell before inhibition, with the lagging inhibition cutting the response short.	`mechanisms[1]`		the tagger's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: Offset responses are generated when the excitatory nonlinearity is amplified relative to the inhibitory one, so inhibition falls away before excitation does and firing rebounds.	`mechanisms[2]`		the tagger's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: Sustained firing is generated when, at large filter outputs, the internal nonlinearity amplifies excitation more than inhibition.	`mechanisms[3]`		the tagger's words
mechanisms	transient onset responses arise from inhibition arriving later than excitation, and also from a difference in their nonlinear gains even when the latencies match Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[4]`	1.0	the tagger's words
mechanisms	offset responses are described as rebound from inhibition, in the 15% of neurons that show them Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[5]`	1.0	the tagger's words
mechanisms	and intensity tuning falls out of the separate nonlinearities, which shift the balance between excitation and inhibition as intensity and stimulus dynamics change Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[6]`	1.0	the tagger's words
mechanisms	with the whole approach resting on approximating the mechanisms rather than the response properties Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[7]`	1.0	the tagger's words
limitations	The model's excitatory and inhibitory terms cannot be claimed to correspond to actual synaptic currents or conductances. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: every interpretation of the model's terms as physiology: THE LABELS FOR EACH TERM ARE MEANT TO DESCRIBE THE INTUITION BEHIND THE STRUCTURE OF THE MODEL, BUT WE CANNOT CLAIM THAT THESE TERMS NECESSARILY REPRESENT ACTUAL excitatory and inhibitory currents	`limitations[0]`		the tagger's words
limitations	The model's constrained structure sacrifices generality and may perform poorly for nonlinearities it does not embody. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the model's performance outside the nonlinearities it encodes: THIS LACK OF GENERALITY COULD HURT THE GNM PERFORMANCE WHEN CONSIDERING NONLINEAR PROCESSING OF FORMS NOT CAPTURED BY ITS STRUCTURE	`limitations[1]`		the tagger's words
limitations	The current model cannot represent separate offset-tuned inputs as an alternative source of offset responses. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the account of offset responses: the model cannot represent separate offset-tuned inputs as the alternative source, so that explanation is untested rather than excluded	`limitations[2]`		the tagger's words
limitations	The fit does not prove that the proposed nonlinear mechanisms actually produce the observed A1 nonlinearities. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the mechanistic reading of the model fit: ALTHOUGH THIS IS BY NO MEANS PROOF THAT SUCH NONLINEAR MECHANISMS ARE RESPONSIBLE FOR OBSERVED A1 NONLINEARITIES, IT DOES PROVIDE A UNIFIED FRAMEWORK	`limitations[3]`		the tagger's words
limitations	the fit is explicitly not proof that these mechanisms are responsible for the observed nonlinearities Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the same scope as limitations[3], from the other direction: THERE IS NO GUARANTEE THAT FOCUSING ON INTERNAL NONLINEAR MECHANISMS WOULD NECESSARILY DESCRIBE THE OBSERVED RESPONSE NONLINEARITIES, WE FOUND THAT THEY DID	`limitations[4]`	1.0	the tagger's words
limitations	the model cannot distinguish rebound from a separate set of offset-tuned inputs, which more recent work reports Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the rebound account of offset responses: THE GNM IN ITS CURRENT FORM DOES NOT ADDRESS THE POSSIBILITY OF SEPARATE OFFSET-TUNED INPUTS	`limitations[5]`	1.0	the tagger's words
limitations	some intensity tuning may be inherited from the inputs to A1 rather than generated there Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the claim that the tuning is generated in A1: IT IS POSSIBLE THAT SOME ASPECTS OF INTENSITY TUNING ARE INHERITED FROM THE INPUTS TO A1	`limitations[6]`	1.0	the tagger's words
limitations	and the structure that makes it efficient also costs it the generality of methods that search for stimulus dimensions without assuming a form Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the generality of the model class: it FOCUSES ON WHAT IS EXPECTED TO BE A DOMINANT mechanism rather than searching the space of stimulus dimensions, which is what makes it efficient and what bounds it	`limitations[7]`	1.0	the tagger's words
limitations	with the intensity tuning measured here agreeing only qualitatively with the same property measured using pure tones Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: any comparison of these intensity-tuning values with pure-tone measurements: Quotation withheld, DUE TO THE EFFECT OF STIMULUS CONTEXT ON A1 RESPONSES	`limitations[8]`	1.0	the tagger's words
transfers	cotuned excitation and inhibition, and the timing and gain differences between them, imported as the model's structure Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[0]`	1.0	the tagger's words
transfers	a sentence database, used as the natural stimulus set Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[1]`	1.0	the tagger's words
transfers	the linear receptive field itself, and the maximum likelihood machinery used to fit it Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[2]`	1.0	the tagger's words
transfers	spike-triggered covariance and maximally informative dimension analysis, which identify additional stimulus features that contribute to the computation underlying the neuronal response Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[3]`	1.0	the tagger's words
controls	the equaliser calibrated before each experiment to achieve flat gain for the acoustic properties of that earphone insertion Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. eliminates: a spectral tilt introduced by how the earphone happened to sit - re-calibrating per experiment means the stimulus spectrum is the one intended rather than the one the fitting produced · outcome (stated by the tagger, no quotation): held	`controls[0]`	0.85	the tagger's words
senses	THE STANDARD MODEL STRUCTURALLY CANNOT SEE THE PHENOMENON: the linear receptive field assumes responses are an average over the neuron's many inputs, and SUCH AVERAGING NECESSARILY HIDES the effects of excitation and inhibition where their tuning overlaps Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[0]`	1.0	the tagger's words
senses	so a property that intracellular work established is invisible to the measure the field uses extracellularly - not undetected, but hidden by construction Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[1]`	1.0	the tagger's words
senses	AND THE FIT IS DECLARED NOT TO BE PROOF: there is no guarantee that focusing on internal mechanisms would describe the observed nonlinearities, and this is BY NO MEANS PROOF that they are responsible Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[2]`	1.0	the tagger's words
senses	AN OUT-OF-DESIGN PREDICTION USED AS EVIDENCE: the model has NO EXPLICIT MECHANISM for intensity tuning and predicts it better than the linear model and as well as a model built for it Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[3]`	1.0	the tagger's words
senses	THREE modeling APPROACHES COMPARED, WITH THE TRADE-OFF NAMED: searching for extra stimulus features is general but leaves it unclear what mechanism is being captured, and this model's structure costs generality while buying data efficiency Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[4]`	1.0	the tagger's words
senses	with the weakness of the first approach stated in its own terms - no constraints are introduced on the type of computation, so the higher-order description is hard to tie to known processing Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[5]`	1.0	the tagger's words
senses	THE SAME PROPERTY MEASURED WITH TWO STIMULI GIVES DIFFERENT NUMBERS, AND THE PAPER SAYS WHY IT SHOULD: intensity tuning agrees qualitatively but NOT quantitatively with the pure-tone measurement, because context and stimulus dynamics change the excitatory-inhibitory balance Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[6]`	1.0	the tagger's words
senses	and a new measure is introduced specifically so that models can be compared on it, rather than only on spike prediction Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[7]`	1.0	the tagger's words
senses	A RIVAL MECHANISM FOR THE SAME DATA, NAMED AND LEFT OPEN: offset responses are described here as rebound from inhibition, while more recent work finds a distinct set of offset-tuned inputs Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[8]`	1.0	the tagger's words
senses	and the population is heterogeneous - only 60% are clear onset cells and 15% have a clear offset response, so one mechanism cannot stand for the class Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[9]`	1.0	the tagger's words
senses	THE STIMULUS CHOICE MAKES CERTAIN MODEL PROPERTIES RELEVANT: speech is made of discrete syllables separated by gaps, which is what makes onset, offset and sustained responses matter to overall performance Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[10]`	1.0	the tagger's words
senses	and the experiment that would settle it is specified - tones ramped in intensity at different speeds, compared against the model or measured intracellularly Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[11]`	1.0	the tagger's words
senses	with an alternative source acknowledged: some intensity tuning may be INHERITED from the inputs to A1 rather than computed there Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[12]`	1.0	the tagger's words
senses	AND THE VALIDATION IS AGREEMENT WITH A DIFFERENT METHOD: properties inferred from extracellular spikes are consistent with previous intracellular observations Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[13]`	1.0	the tagger's words
senses	the class membership is assigned by a property bundle - tonotopic organization, latency and simple frequency tuning verify a neuron as belonging to the area Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[14]`	1.0	the tagger's words
alternatives	claim_type: method Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. a second reading DIFFERING from the recorded value. Recorded: mechanism — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.	`alternatives.claim_type`		the tagger's words
alternatives	method: a generalised nonlinear model with separate filters and nonlinearities for putative excitatory and inhibitory elements, fitted to spikes evoked by continuous speech Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. 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.method`		the tagger's words
alternatives	phenomenon: cotuned excitation and inhibition, whose effects cancel in any measure that averages over a neuron's inputs Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. a second reading DIFFERING from the recorded value. Recorded: excitatory-inhibitory interplay in primary auditory cortex — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.	`alternatives.phenomenon`		the tagger's words
claim_relations	builds-on → the generalized nonlinear model (GNM) framework of Butts et al. (2011) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[0]`		the tagger's words
claim_relations	builds-on → the A1 recording dataset and procedures of David et al. (2009) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[1]`		the tagger's words
claim_relations	extends → the multilinear / input-nonlinearity model of A1 intensity tuning (Ahrens et al. 2008b) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[2]`		the tagger's words
claim_relations	critiques → the linear spectrotemporal receptive field, which averages over inputs and therefore cannot represent cotuned excitation and inhibition Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: THE AXIS IS WHETHER INHIBITION IS COTUNED WITH EXCITATION. 'Quotation withheld.' A model that averages over inputs cannot represent two opposing drives with the same tuning - so the objection is about what the FORMALISM can hold, not about whether its fits are good.	`claim_relations[3]`	1.0	the tagger's words
claim_relations	extends → intracellular findings of cotuned excitation and inhibition, to extracellular recordings and natural stimuli Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[4]`	1.0	the tagger's words
claim_relations	distinguishes-from → feature-search methods such as spike-triggered covariance and maximally informative dimensions, which are more general and less interpretable Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: THE AXIS IS WHETHER THE NONLINEARITY IS DISCOVERED FROM THE DATA. 'One approach has been to identify additional stimulus features (in addition to the linear STRF) that contribute to the computation underlying the neuronal response, for example using SPIKE-TRIGGERED COVARIANCE (STC) or MAXIMALLY INFORMATIVE DIMENSION (MID) ANALYSIS.' Its twin at [6] carries the other family, which builds in nonlinearities chosen in advance. NOTE: the target also calls these methods 'more general and less interpretable' AND THIS SPAN SAYS NEITHER - the axis recorded here is the one the sentence supports, and the target's extra claim remains unanchored.	`claim_relations[5]`	1.0	the tagger's words
claim_relations	distinguishes-from → multilinear models that add components for specific known nonlinearities Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: THE AXIS IS WHETHER THE NONLINEARITY IS DISCOVERED OR SPECIFIED IN ADVANCE. 'A second approach has taken an opposite tack, by incorporating model components to capture specific nonlinear elements of A1 processing, such as intensity tuning and dependence on stimulus context. This approach has been CAST AS A MULTILINEAR MODEL.' One family searches for features the data reveal; the other builds in nonlinearities the modeller already believes in - so the two can fit the same neuron and disagree about what it does.	`claim_relations[6]`	1.0	the tagger's words
claim_relations	supports → the account in which relative timing of excitation and inhibition produces transient onset responses Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[7]`	1.0	the tagger's words
claim_relations	qualifies → its own rebound account of offset responses, against a report of separate offset-tuned inputs Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[8]`	1.0	the tagger's words
concept_edges	excitation —contrasted-with→ inhibition Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span concerns co-tuned excitation and inhibition; contrasted-with is loose. also in: excitation · inhibition	`concept_edges[1]`	1.0	the tagger's words
concept_edges	STRF —operationalised-as→ stimulus Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 · note: sounds matching it drive the strongest responses, which is the intuition the whole field runs on Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the STRF describes stimulus selectivity; operationalised-as is loose. also in: STRF · stimulus	`concept_edges[3]`	1.0	the tagger's words
concept_edges	GNM —causes→ intensity tuning Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 · note: AN OUT-OF-DESIGN PREDICTION USED AS EVIDENCE: NO EXPLICIT MECHANISM for it, and it predicts it better than the linear model and as well as a model built for it Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the model REPRODUCED A1 intensity tuning through its nonlinearities; causes is loose. also in: GNM · intensity tuning	`concept_edges[16]`	1.0	the tagger's words
concept_edges	input —causes→ intensity tuning Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 · note: with the alternative source acknowledged - some of it may be INHERITED from the inputs rather than computed here Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says intensity tuning may be inherited from inputs; causes states a possibility. also in: input · intensity tuning	`concept_edges[19]`	1.0	the tagger's words
concept_edges	inhibition —causes→ onset Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 · note: inhibition delayed relative to excitation, as intracellular work suggested - AND a gain difference can do it with matched latencies Audit 2026-09-28: DOUBTFUL - read against its span and kept, but onset responses arise from the interplay of excitation and inhibition; causes is loose. also in: inhibition · onset	`concept_edges[23]`	1.0	the tagger's words
content_tags	maximum likelihood estimation Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: maximum likelihood estimation	`content_tags.statistics[0]`		the tagger's words
content_tags	cross-validation Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: cross-validation	`content_tags.statistics[1]`		the tagger's words
content_tags	t-test Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: t-test	`content_tags.statistics[2]`		the tagger's words
content_tags	paired t-test Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: paired t-test	`content_tags.statistics[3]`		the tagger's words
content_tags	correlation Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: correlation	`content_tags.statistics[4]`		the tagger's words
content_tags	principal components analysis Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: principal components analysis	`content_tags.statistics[5]`		the tagger's words
content_tags	median Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: median	`content_tags.statistics[6]`		the tagger's words
content_tags	entropy Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: entropy	`content_tags.statistics[7]`		the tagger's words
content_tags	explained variance (r-squared) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: explained variance (r-squared)	`content_tags.statistics[8]`		the tagger's words
content_tags	principal components analysis Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: principal components analysis	`content_tags.statistics[9]`	1.0	the tagger's words
content_tags	k -means clustering Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: k -means clustering	`content_tags.statistics[10]`	1.0	the tagger's words
content_tags	The nonlinear GNM predicts A1 spike trains during speech significantly better than the linear STRF-based model across the recorded population. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[0]`		the tagger's words
content_tags	The excitation and inhibition inferred from extracellular data are cotuned: they peak at the same frequency in 44% of neurons and within half an octave in a further 35%. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[1]`		the tagger's words
content_tags	Inhibition is more broadly tuned in frequency than excitation in the large majority of cells. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[2]`		the tagger's words
content_tags	Nonmonotonic intensity tuning emerges from the model even though no intensity-tuning term was built into it, matching a model that fits intensity tuning explicitly. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[3]`		the tagger's words
content_tags	The GNM also reproduces neuronal responses to sound onsets and offsets better than the linear model. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[4]`		the tagger's words
content_tags	Among neurons with sufficient data, 60% were onset cells with little sustained firing and 24% responded with strong sustained activity. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[5]`		the tagger's words
content_tags	Because it is a linear measure, the spectrotemporal receptive field cannot detect overlapping excitation and inhibition. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[6]`		the tagger's words
content_tags	THE STANDARD MODEL STRUCTURALLY CANNOT SEE THE PHENOMENON: averaging over a neuron's inputs NECESSARILY HIDES excitation and inhibition where their tuning overlaps Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. WHOSE RESULT THIS IS: studies show - THE SUBJECT OF THIS RESULT IS AN INDEFINITE STUDY, NOT THIS PAPER. 'Previous studies reported', 'studies in guinea pig showed', 'recent studies have reported' - somebody else's work under a common noun rather than a name, with a citation in the sentence and no first-person marker. The quote is verbatim and the claim is true; only the attribution is wrong. FOUND BY MEASURING MY OWN CUE'S RECALL: the first sweep required a NAMED author as the subject ('Sperling (1960) demonstrated') and found 17. Reading eleven citation-bearing findings on 'Plakke2016' by hand showed four the cue could not see - 'A study examining recognition found', 'studies of typical human subjects have demonstrated', 'A follow up study found' - so the named cue's 17 was a floor and not a count. A CUE'S FALSE POSITIVES TELL YOU NOTHING ABOUT ITS FALSE NEGATIVES, and the only way to find these was to read a record's whole layer.	`content_tags.findings[7]`	1.0	the tagger's words
content_tags	the linear receptive field is the standard first-order description, and sounds matching it drive the strongest responses Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[8]`	1.0	the tagger's words
content_tags	and the gap is named precisely - existing nonlinear models address some aspects and NONE has addressed the interplay of excitation and inhibition Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[9]`	1.0	the tagger's words
content_tags	THE VALIDATION IS AGREEMENT WITH A DIFFERENT METHOD: properties inferred from extracellular spikes are consistent with previous intracellular observations Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[10]`	1.0	the tagger's words
content_tags	only clusters with 90% isolation were used, and neurons were verified as belonging to the area by tonotopic organization, latency and simple frequency tuning Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[11]`	1.0	the tagger's words
content_tags	the stimulus was continuous speech from a sentence database, balanced across speaker sex Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[12]`	1.0	the tagger's words
content_tags	with a consequence of the upsampling disclosed - all stimulus power was below 8 kHz Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[13]`	1.0	the tagger's words
content_tags	and two sets of neurons heard different stimulus sets, which is disclosed rather than smoothed over Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[14]`	1.0	the tagger's words
content_tags	A NEW MEASURE IS INTRODUCED so that models can be compared on intensity tuning itself, not only on spike prediction Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[15]`	1.0	the tagger's words
content_tags	THE SAME PROPERTY MEASURED WITH TWO STIMULI GIVES DIFFERENT NUMBERS, AND THE PAPER SAYS WHY IT SHOULD Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[16]`	1.0	the tagger's words
content_tags	AN OUT-OF-DESIGN PREDICTION USED AS EVIDENCE: the model has no explicit mechanism for intensity tuning and predicts it better than the linear model Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[17]`	1.0	the tagger's words
content_tags	and the settling experiment is specified - tones ramped in intensity at different speeds, compared against the model or measured intracellularly Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[18]`	1.0	the tagger's words
content_tags	with an alternative source acknowledged - some intensity tuning may be inherited from the inputs to the area Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[19]`	1.0	the tagger's words
content_tags	THE STIMULUS CHOICE MAKES CERTAIN MODEL PROPERTIES RELEVANT: speech is made of syllables separated by gaps, so onset, offset and sustained responses matter to performance Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[20]`	1.0	the tagger's words
content_tags	transient onsets arise from delayed inhibition, AND from a gain difference even when the latencies are the same Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[21]`	1.0	the tagger's words
content_tags	only 60% of neurons are clear onset cells and 15% have a clear offset response Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. WHOSE RESULT THIS IS: Qin et al. 2007 ), although more recent work found - THE EVIDENCE FOR THIS ROW ATTRIBUTES THE WORK TO SOMEBODY ELSE. The cited name is the subject of a result verb and the sentence carries no first-person marker, so the finding, quantity or definition is the CITED author's rather than this record's. The claim is true and the quote is verbatim - what is wrong is whose result the corpus says it is. Nothing edited, nothing withdrawn: the row is marked so a page can name the source. Where the record is a commentary or review, its claim relations layer usually already names the paper being discussed - the record knows at one layer and forgets at another.	`content_tags.findings[22]`	1.0	the tagger's words
content_tags	A RIVAL MECHANISM NAMED AND LEFT OPEN: the model cannot address separate offset-tuned inputs, and could be extended to find them Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[23]`	1.0	the tagger's words
content_tags	with the motivation stated as a recognition - linear models MISS SIGNIFICANT ASPECTS of the processing Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[24]`	1.0	the tagger's words
content_tags	and the weakness of the feature-search approach stated in its own terms - no constraints on the type of computation, so the mechanism captured is unclear Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[25]`	1.0	the tagger's words
content_tags	the second approach adds components for specific known nonlinearities and can be estimated efficiently Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[26]`	1.0	the tagger's words
content_tags	THE TRADE-OFF NAMED IN BOTH DIRECTIONS: the structure could hurt performance for nonlinearities it does not have, and will be much more efficient with data when they match Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[27]`	1.0	the tagger's words
content_tags	AND THE FIT IS DECLARED NOT TO BE PROOF: no guarantee it would work, and BY NO MEANS PROOF that these mechanisms are responsible Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[28]`	1.0	the tagger's words
content_tags	the properties that were NOT explicitly modeled are exactly the ones offered as evidence Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[29]`	1.0	the tagger's words
content_tags	generalized nonlinear model (GNM) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: generalized nonlinear model (GNM)	`content_tags.definitions[0]`		the tagger's words
content_tags	onset-triggered average Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: onset-triggered average	`content_tags.definitions[1]`		the tagger's words
content_tags	silence rate and sound rate Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: silence rate and sound rate	`content_tags.definitions[2]`		the tagger's words
content_tags	effective time kernel Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: effective time kernel	`content_tags.definitions[3]`		the tagger's words
content_tags	level-rate correlation coefficient Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: level-rate correlation coefficient	`content_tags.definitions[4]`		the tagger's words
content_tags	STRF Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: STRF	`content_tags.definitions[5]`	1.0	the tagger's words
content_tags	cotuned Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. WHOSE RESULT THIS IS: studies show - THE SUBJECT OF THIS RESULT IS AN INDEFINITE STUDY, NOT THIS PAPER. 'Previous studies reported', 'studies in guinea pig showed', 'recent studies have reported' - somebody else's work under a common noun rather than a name, with a citation in the sentence and no first-person marker. The quote is verbatim and the claim is true; only the attribution is wrong. FOUND BY MEASURING MY OWN CUE'S RECALL: the first sweep required a NAMED author as the subject ('Sperling (1960) demonstrated') and found 17. Reading eleven citation-bearing findings on 'Plakke2016' by hand showed four the cue could not see - 'A study examining recognition found', 'studies of typical human subjects have demonstrated', 'A follow up study found' - so the named cue's 17 was a floor and not a count. A CUE'S FALSE POSITIVES TELL YOU NOTHING ABOUT ITS FALSE NEGATIVES, and the only way to find these was to read a record's whole layer. also in: cotuned	`content_tags.definitions[6]`	1.0	the tagger's words
content_tags	GNM Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: GNM	`content_tags.definitions[7]`	1.0	the tagger's words
content_tags	isolation Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: isolation	`content_tags.definitions[8]`	1.0	the tagger's words
content_tags	cross-validated LL Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: cross-validated LL	`content_tags.variables[1]`		the tagger's words
content_tags	firing rate Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: firing rate	`content_tags.variables[2]`		the tagger's words
content_tags	sound pressure level Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: sound pressure level	`content_tags.variables[3]`		the tagger's words
content_tags	intensity tuning Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: intensity tuning	`content_tags.variables[4]`	1.0	the tagger's words
content_tags	prediction Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: prediction	`content_tags.variables[5]`	1.0	the tagger's words
content_tags	latency Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: latency	`content_tags.variables[6]`	1.0	the tagger's words
content_tags	gain Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: gain	`content_tags.variables[7]`	1.0	the tagger's words
content_tags	94 — single units recorded extracellularly from primary auditory cortex of awake ferrets Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[0]`		the tagger's words
content_tags	P = 4 × 10 −12 — significance of the GNM's cross-validated log-likelihood advantage over the linear model across neurons Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[1]`		the tagger's words
content_tags	r 2 (GN) = 0.12 and r 2 (LN) = 0.07 — explained variance of the two models for the example neuron Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[2]`		the tagger's words
content_tags	0.40 bits per spike / 0.30 bits per spike — cross-validated log-likelihood of the GNM versus the linear model for the example neuron Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[3]`		the tagger's words
content_tags	83% — proportion of the stimulus used as training data, the rest held out for cross-validation 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_tags	40% — median increase in frequency-tuning width (in octaves) of inhibition relative to excitation 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_tags	60% / 24% — share of analysable neurons classed as onset cells versus sustained-response 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_tags	60% — the share of recorded neurons that are clear onset cells, against 15% with a clear offset response - the heterogeneity a single mechanism would have to cover Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. WHOSE RESULT THIS IS: Qin et al. 2007 ), although more recent work found - THE EVIDENCE FOR THIS ROW ATTRIBUTES THE WORK TO SOMEBODY ELSE. The cited name is the subject of a result verb and the sentence carries no first-person marker, so the finding, quantity or definition is the CITED author's rather than this record's. The claim is true and the quote is verbatim - what is wrong is whose result the corpus says it is. Nothing edited, nothing withdrawn: the row is marked so a page can name the source. Where the record is a commentary or review, its claim relations layer usually already names the paper being discussed - the record knows at one layer and forgets at another.	`content_tags.quantities[7]`	1.0	the tagger's words
content_tags	90% isolation — the spike-sorting criterion, so the analysed population is already selected on cluster quality Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[8]`	1.0	the tagger's words
content_tags	8 kHz — the upper bound on stimulus power, a consequence of upsampling that the paper points out itself Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[9]`	1.0	the tagger's words

Interpretive choices

Tags that could not be placed in the text:

none — every tag was located in the text

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.