phenomenon · claim-type · cross-domain-transfer · mechanism · claim-relation · replication-status · limitation — grounded in the full text| phenomenon | excitatory-inhibitory interplay in primary auditory cortex— “In particular, the interplay of excitation and inhibition explained functional properties that were not explicitly modeled, including nonlinear intensity tuning, and responses to sound onset and offset.” | |
| claim-type | mechanism— “Intensity tuning occurs because the relative strength of excitation compared with inhibition differs at different intensities.” | |
| method — evidence | — “Auditory responses were recorded extracellularly from single neurons in A1 of six awake, passively listening ferrets.” | |
| object of study — evidence | — “Intracellular studies have revealed the importance of cotuned excitatory and inhibitory inputs to neurons in auditory cortex, but typical spectrotemporal receptive field models of neuronal processing cannot account for this overlapping tuning. Here, we apply a new nonlinear modeling framework to extracellular data recorded from primary… | |
| research question — evidence | — “Here, we employ a novel nonlinear modeling approach designed to infer separate tuning and response properties of excitation and inhibition from extracellular recordings of neurons in A1 during the presentation of speech stimuli ( David et al. 2009 ). We find that the inferred tuning of excitation and inhibition can predict a variety of… | |
| contribution — evidence | — “We present a description of processing in A1 in a natural stimulus context and describe how many previously observed nonlinearities may arise through the interplay of excitatory and inhibitory synaptic inputs. In particular, the interplay of excitation and inhibition explained functional properties that were not explicitly modeled, including… | |
| transfer | from intracellular physiology: cotuned excitation and inhibition, and the timing and gain differences between them, imported as the model's structure— “Intracellular studies have revealed the importance of cotuned excitatory and inhibitory inputs to neurons in auditory cortex, but typical spectrotemporal receptive field models of neuronal processing cannot account for this overlapping tuning. Here, we apply a new nonlinear modeling framework to extracellular data recorded from primary… | transfers[0] |
| transfer | from speech technology: a sentence database, used as the natural stimulus set— “We recorded the responses of isolated A1 neurons to segments of continuous speech. Speech samples were taken from the Texas Instruments/Massachusetts Institute of Technology (TIMIT) database ( Garofolo 1988 ). Stimuli were sentences (3–4 s) sampled each from a different speaker and balanced across sex. The digital signals were transformed to… | transfers[1] |
| transfer | from systems identification: the linear receptive field itself, and the maximum likelihood machinery used to fit it— “as the standard first-order description of the stimulus selectivity of an auditory neuron, the linear spectrotemporal receptive field (STRF) offers an intuition for how the neuron responds to complex stimuli. Sounds that most closely match the STRF (in both frequency and relative timing) will drive the strongest responses, whereas sounds that… | transfers[2] |
| transfer | from information-theoretic feature search: spike-triggered covariance and maximally informative dimension analysis, which identify additional stimulus features that contribute to the computation underlying the neuronal response— “Two main approaches have been taken to model nonlinear aspects of A1 neuron responses. 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; Sen et al. 2000 ) or maximally… | transfers[3] |
| mechanism | 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.— “Intensity tuning occurs because the relative strength of excitation compared with inhibition differs at different intensities.” | mechanisms[0] |
| mechanism | Onset responses are generated by excitation reaching the cell before inhibition, with the lagging inhibition cutting the response short.— “At sound onset, the excitatory filter output rises earlier than the inhibitory output, producing sharp onset responses that are promptly extinguished by the delayed inhibition.” | mechanisms[1] |
| mechanism | 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.— “However, because the excitatory nonlinearity is amplified compared with that of inhibition, inhibition decreases earlier than excitation, resulting in a peak in the firing rate at sound offset.” | mechanisms[2] |
| mechanism | Sustained firing is generated when, at large filter outputs, the internal nonlinearity amplifies excitation more than inhibition.— “Hence, the nonlinearity amplifies the excitatory response to sounds that match its best frequency compared with the inhibitory response, resulting in sustained activity that is larger than the activity at sound onset.” | mechanisms[3] |
| mechanism | transient onset responses arise from inhibition arriving later than excitation, and also from a difference in their nonlinear gains even when the latencies match— “The GNM can predict the responses to sound onset and offsets, again through the interplay of excitation and inhibition. Onset neurons can produce transient responses through a relative delay of inhibition compared with excitation, which has been suggested through numerous intracellular studies, both in A1 ( Tan and Wehr 2009 ; Wehr and Zador… | mechanisms[4] |
| mechanism | offset responses are described as rebound from inhibition, in the 15% of neurons that show them— “Only 60% of A1 neurons recorded are clear onset cells, and for neurons exhibiting different dynamics, the GNM revealed a different interplay of excitation and inhibition. In particular, 15% of neurons have a clear offset response, which the GNM described by rebound from inhibition. Such a mechanism has been suggested from intracellular… | mechanisms[5] |
| mechanism | and intensity tuning falls out of the separate nonlinearities, which shift the balance between excitation and inhibition as intensity and stimulus dynamics change— “Although the structure of the GNM has no explicit mechanism to capture intensity tuning, it was able to predict A1 intensity tuning in this natural context better than the linear model and as well as or better than a model with explicit intensity tuning ( Ahrens et al. 2008b ). The GNM produced this intensity tuning through the separate… | mechanisms[6] |
| mechanism | with the whole approach resting on approximating the mechanisms rather than the response properties— “The GNM takes a third approach, which, instead of modeling specific nonlinear response properties, introduces nonlinearities to approximate the underlying nonlinear mechanisms known to be present (and likely contribute to the response). It focuses on what is expected to be a dominant nonlinearity in A1 ( Dorrn et al. 2010 ; Sun et al. 2010… | mechanisms[7] |
| builds-on | the generalized nonlinear model (GNM) framework of Butts et al. (2011)— “The GNM framework is composed of nonlinear elements designed to simulate the effects of excitation and inhibition and also can be fit to data using maximum likelihood estimation ( Butts et al. 2011 ).” | claim_relations[0] |
| builds-on | the A1 recording dataset and procedures of David et al. (2009)— “As described in detail in David et al. (2009) , animals were implanted with a steel headpost to allow for stable recording.” | claim_relations[1] |
| extends | the multilinear / input-nonlinearity model of A1 intensity tuning (Ahrens et al. 2008b)— “Although the structure of the GNM has no explicit mechanism to capture intensity tuning, it was able to predict A1 intensity tuning in this natural context better than the linear model and as well as or better than a model with explicit intensity tuning ( Ahrens et al. 2008b ).” | claim_relations[2] |
| critiques | the linear spectrotemporal receptive field, which averages over inputs and therefore cannot represent cotuned excitation and inhibition— “However, recent intracellular studies in auditory cortex challenge the applicability of such straightforward stimulus processing to primary auditory cortex (A1) neurons. These studies show that inhibitory inputs onto A1 neurons are often cotuned with excitation ( Dorrn et al. 2010 ; Sun et al. 2010 ; Wehr and Zador 2003 ) and implicate the… | claim_relations[3] |
| extends | intracellular findings of cotuned excitation and inhibition, to extracellular recordings and natural stimuli— “Intracellular studies have revealed the importance of cotuned excitatory and inhibitory inputs to neurons in auditory cortex, but typical spectrotemporal receptive field models of neuronal processing cannot account for this overlapping tuning. Here, we apply a new nonlinear… | claim_relations[4] |
| distinguishes-from | feature-search methods such as spike-triggered covariance and maximally informative dimensions, which are more general and less interpretable | claim_relations[5] |
| distinguishes-from | multilinear models that add components for specific known nonlinearities Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | claim_relations[6] |
| supports | the account in which relative timing of excitation and inhibition produces transient onset responses Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | claim_relations[7] |
| qualifies | its own rebound account of offset responses, against a report of separate offset-tuned inputs Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | claim_relations[8] |
| sense | 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 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. | senses[0] |
| sense | 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 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. | senses[1] |
| sense | 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 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. | senses[2] |
| sense | 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 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. | senses[3] |
| sense | 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 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. | senses[4] |
| sense | 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 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. | senses[5] |
| sense | 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 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. | senses[6] |
| sense | and a new measure is introduced specifically so that models can be compared on it, rather than only on spike prediction 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. | senses[7] |
| sense | 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 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. | senses[8] |
| sense | 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 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. | senses[9] |
| sense | 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 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. | senses[10] |
| sense | 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 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. | senses[11] |
| sense | with an alternative source acknowledged: some intensity tuning may be INHERITED from the inputs to A1 rather than computed there 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. | senses[12] |
| sense | AND THE VALIDATION IS AGREEMENT WITH A DIFFERENT METHOD: properties inferred from extracellular spikes are consistent with previous intracellular observations 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. | senses[13] |
| sense | 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 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. | senses[14] |
| control | the equaliser calibrated before each experiment to achieve flat gain for the acoustic properties of that earphone insertion — held — rules out 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 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. | controls[0] |
| differs in | the linear spectrotemporal receptive field, which averages over inputs and therefore cannot represent cotuned excitation and inhibition — THE AXIS IS WHETHER INHIBITION IS COTUNED WITH EXCITATION. 'recent intracellular studies in auditory cortex challenge the applicability of such straightforward stimulus processing to primary auditory cortex (A1) neurons. These studies show that inhibitory inputs onto A1 neurons are often cotuned with excitation.' 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. 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. Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | claim_relations[3] |
| differs in | feature-search methods such as spike-triggered covariance and maximally informative dimensions, which are more general and less interpretable — 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. 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. Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | claim_relations[5] |
| differs in | multilinear models that add components for specific known nonlinearities — 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. 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. Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | claim_relations[6] |
| audit | excitation —contrasted-with→ inhibition — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span concerns co-tuned excitation and inhibition; contrasted-with is loose. 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[1] |
| audit | STRF —operationalised-as→ stimulus — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the STRF describes stimulus selectivity; operationalised-as is loose. 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[3] |
| note | STRF —operationalised-as→ stimulus — sounds matching it drive the strongest responses, which is the intuition the whole field runs on 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[3] |
| audit | GNM —causes→ intensity tuning — 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. 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[16] |
| note | GNM —causes→ intensity tuning — 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 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[16] |
| audit | input —causes→ intensity tuning — 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. 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[19] |
| note | input —causes→ intensity tuning — with the alternative source acknowledged - some of it may be INHERITED from the inputs rather than computed here 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[19] |
| audit | inhibition —causes→ onset — 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. 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[23] |
| note | inhibition —causes→ onset — inhibition delayed relative to excitation, as intracellular work suggested - AND a gain difference can do it with matched latencies 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[23] |
| outcome (stated by the tagger, no quotation) | control — held | controls[0] |
| for which use | limitation — 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 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. | limitations[0] |
| for which use | limitation — 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 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. | limitations[1] |
| for which use | limitation — 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 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. | limitations[2] |
| for which use | limitation — 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 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. | limitations[3] |
| for which use | limitation — 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 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. | limitations[4] |
| for which use | limitation — the rebound account of offset responses: THE GNM IN ITS CURRENT FORM DOES NOT ADDRESS THE POSSIBILITY OF SEPARATE OFFSET-TUNED INPUTS 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. | limitations[5] |
| for which use | limitation — 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 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. | limitations[6] |
| for which use | limitation — 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 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. | limitations[7] |
| for which use | limitation — any comparison of these intensity-tuning values with pure-tone measurements: SUCH MEASURED INTENSITY TUNING QUALITATIVELY, BUT NOT QUANTITATIVELY, AGREES WITH THAT MEASURED WITH PURE TONES (DATA NOT SHOWN), DUE TO THE EFFECT OF STIMULUS CONTEXT ON A1 RESPONSES Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | limitations[8] |
| replication | untested | |
| limitation | The model's excitatory and inhibitory terms cannot be claimed to correspond to actual synaptic currents or conductances. 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. | limitations[0] |
| limitation | The model's constrained structure sacrifices generality and may perform poorly for nonlinearities it does not embody. 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. | limitations[1] |
| limitation | The current model cannot represent separate offset-tuned inputs as an alternative source of offset responses. Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | limitations[2] |
| limitation | The fit does not prove that the proposed nonlinear mechanisms actually produce the observed A1 nonlinearities. 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. | limitations[3] |
| limitation | the fit is explicitly not proof that these mechanisms are responsible for the observed nonlinearities 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. | limitations[4] |
| limitation | the model cannot distinguish rebound from a separate set of offset-tuned inputs, which more recent work reports 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. | limitations[5] |
| limitation | some intensity tuning may be inherited from the inputs to A1 rather than generated there 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. | limitations[6] |
| limitation | 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 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. | limitations[7] |
| limitation | with the intensity tuning measured here agreeing only qualitatively with the same property measured using pure tones 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. | limitations[8] |
finding · quantity · variable · definition · key-term — grounded in the full text| finding | The nonlinear GNM predicts A1 spike trains during speech significantly better than the linear STRF-based model across the recorded population. 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.findings[0] |
| finding | 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 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.findings[1] |
| finding | Inhibition is more broadly tuned in frequency than excitation in the large majority of cells. 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.findings[2] |
| finding | 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 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.findings[3] |
| finding | The GNM also reproduces neuronal responses to sound onsets and offsets better than the linear model. 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.findings[4] |
| finding | Among neurons with sufficient data, 60% were onset cells with little sustained firing and 24% responded with strong sustained activity. 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.findings[5] |
| finding | Because it is a linear measure, the spectrotemporal receptive field cannot detect overlapping excitation and inhibition. 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.findings[6] |
| finding | 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 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.findings[7] |
| finding | the linear receptive field is the standard first-order description, and sounds matching it drive the strongest responses Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | content_tags.findings[8] |
| finding | 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 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.findings[9] |
| finding | THE VALIDATION IS AGREEMENT WITH A DIFFERENT METHOD: properties inferred from extracellular spikes are consistent with previous intracellular observations 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.findings[10] |
| finding | 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 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.findings[11] |
| finding | the stimulus was continuous speech from a sentence database, balanced across speaker sex 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.findings[12] |
| finding | with a consequence of the upsampling disclosed - all stimulus power was below 8 kHz 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.findings[13] |
| finding | and two sets of neurons heard different stimulus sets, which is disclosed rather than smoothed over 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.findings[14] |
| finding | A NEW MEASURE IS INTRODUCED so that models can be compared on intensity tuning itself, not only on spike prediction 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.findings[15] |
| finding | THE SAME PROPERTY MEASURED WITH TWO STIMULI GIVES DIFFERENT NUMBERS, AND THE PAPER SAYS WHY IT SHOULD 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.findings[16] |
| finding | 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 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.findings[17] |
| finding | and the settling experiment is specified - tones ramped in intensity at different speeds, compared against the model or measured intracellularly 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.findings[18] |
| finding | with an alternative source acknowledged - some intensity tuning may be inherited from the inputs to the area 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.findings[19] |
| finding | 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 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.findings[20] |
| finding | transient onsets arise from delayed inhibition, AND from a gain difference even when the latencies are the same 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.findings[21] |
| finding | only 60% of neurons are clear onset cells and 15% have a clear offset response 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.findings[22] |
| finding | 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 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.findings[23] |
| finding | with the motivation stated as a recognition - linear models MISS SIGNIFICANT ASPECTS of the processing 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.findings[24] |
| finding | 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 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.findings[25] |
| finding | the second approach adds components for specific known nonlinearities and can be estimated efficiently 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.findings[26] |
| finding | 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 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.findings[27] |
| finding | 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 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.findings[28] |
| finding | the properties that were NOT explicitly modeled are exactly the ones offered as evidence 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.findings[29] |
| quantity | 94 — single units recorded extracellularly from primary auditory cortex of awake ferrets 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.quantities[0] |
| quantity | P = 4 × 10 −12 — significance of the GNM's cross-validated log-likelihood advantage over the linear model across neurons 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.quantities[1] |
| quantity | r 2 (GN) = 0.12 and r 2 (LN) = 0.07 — explained variance of the two models for the example neuron 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.quantities[2] |
| quantity | 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 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.quantities[3] |
| quantity | 83% — proportion of the stimulus used as training data, the rest held out for cross-validation 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.quantities[4] |
| quantity | 40% — median increase in frequency-tuning width (in octaves) of inhibition relative to excitation 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.quantities[5] |
| quantity | 60% / 24% — share of analysable neurons classed as onset cells versus sustained-response cells 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.quantities[6] |
| quantity | 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 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.quantities[7] |
| quantity | 90% isolation — the spike-sorting criterion, so the analysed population is already selected on cluster quality 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.quantities[8] |
| quantity | 8 kHz — the upper bound on stimulus power, a consequence of upsampling that the paper points out itself 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.quantities[9] |
| variable | cross-validated LL (dependent) 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.variables[1] |
| variable | firing rate (dependent) 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.variables[2] |
| variable | sound pressure level (control) 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.variables[3] |
| variable | intensity tuning 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.variables[4] |
| variable | prediction 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.variables[5] |
| variable | latency 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.variables[6] |
| variable | gain 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.variables[7] |
| definition | generalized nonlinear model (GNM) 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.definitions[0] |
| definition | onset-triggered average 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.definitions[1] |
| definition | silence rate and sound 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.definitions[2] |
| definition | effective time kernel 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.definitions[3] |
| definition | level-rate correlation coefficient 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.definitions[4] |
| definition | STRF 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.definitions[5] |
| definition | cotuned 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.definitions[6] |
| definition | GNM 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.definitions[7] |
| definition | isolation 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.definitions[8] |
| key-terms | 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 |
| maximum likelihood estimation LL | 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] |
| cross-validation LL x | as written: cross-validated LL (LL x ) 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] |
| t-test P | 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] |
| paired t-test P | 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] |
| correlation | as written: correlation coefficient 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] |
| principal components analysis | as written: principal components analysis and k -means clustering 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] |
| median | as written: median difference 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] |
| entropy H | as written: entropy as a measure of their overall width 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] |
| explained variance (r-squared) r 2 | as written: percent of explained variance (or r 2 ) 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] |
| principal components analysis | 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] |
| k -means clustering | 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] |
| claim type | method — a second reading DIFFERING from the recorded value; recorded: mechanism Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | alternatives.claim_type |
| method | a generalised nonlinear model with separate filters and nonlinearities for putative excitatory and inhibitory elements, fitted to spikes evoked by continuous speech — a second reading DIFFERING from the recorded value; recorded: electrophysiology Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher. | alternatives.method |
| phenomenon | cotuned excitation and inhibition, whose effects cancel in any measure that averages over a neuron's inputs — a second reading DIFFERING from the recorded value; recorded: excitatory-inhibitory interplay in primary auditory 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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