Differential Receptive Field Properties of Parvalbumin and Somatostatin Inhibitory Neurons in Mouse Auditory Cortex

Differential Receptive Field Properties of Parvalbumin and Somatostatin Inhibitory Neurons in Mouse Auditory Cortex Li Xiong Ibrahim Yuan Tao Zhang

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Differential Receptive Field Properties of Parvalbumin and Somatostatin Inhibitory Neurons in Mouse Auditory Cortex 2015 Cerebral Cortex (New York, N.Y.: 1991) 10.1093/cercor/bht417 Li2015-differential-receptive-field-properties-of-p Li2015-differential-receptive-field-properties-of-p.txt, the project's staged copy 925df939c2af61c87bf927a3a1b54f5dd74cf7cb not-redistributable neuroscience

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0 spans located by searching the stored text for the quote's exact words, allowing any whitespace between them how the two main cortical interneuron types differ in what sounds they respond to, once they are identified genetically rather than by spike shape Quotation withheld because this record is not cleared for redistribution. whether parvalbumin and somatostatin neurons in auditory cortex have different frequency tuning, thresholds and timing from each other and from excitatory cells Quotation withheld because this record is not cleared for redistribution. shows PV cells are broadly tuned with low thresholds and fast responses while SOM cells are as selective as excitatory neurons and markedly delayed, and traces the difference to input range and input-output efficiency Quotation withheld because this record is not cleared for redistribution. existence Quotation withheld because this record is not cleared for redistribution. tonal receptive fields of cortical inhibitory interneuron subtypes Quotation withheld because this record is not cleared for redistribution. electrophysiology electrophysiology Quotation withheld because this record is not cleared for redistribution. mechanisms PV neurons pool a wider range of excitatory inputs and receive stronger subthreshold drive, so more of that input crosses spike threshold, producing broad, weakly selective spike tuning Quotation withheld because this record is not cleared for redistribution. mechanisms SOM spiking is late mainly because the excitatory synaptic input itself arrives late, with longer synaptic integration adding to the delay Quotation withheld because this record is not cleared for redistribution. mechanisms Non-selective local connectivity from excitatory cells onto inhibitory (especially PV) cells pools inputs with many different preferred frequencies onto one interneuron Quotation withheld because this record is not cleared for redistribution. mechanisms SOM neurons bypass the direct ascending (feedforward) pathway and are instead driven through slower horizontal circuits, which is why their input is delayed Quotation withheld because this record is not cleared for redistribution. PV cells pool excitatory input from neurons with a wide range of preferred frequencies because excitatory-to-PV connectivity is unselective Quotation withheld because this record is not cleared for redistribution. PV cells also convert input to output more efficiently, so a broad input becomes an even broader spike tuning Quotation withheld because this record is not cleared for redistribution. PV cells are driven at the same time as excitatory cells, consistent with shared feedforward input from layer 4 Quotation withheld because this record is not cleared for redistribution. SOM cells receive delayed input, suggesting horizontal rather than ascending drive Quotation withheld because this record is not cleared for redistribution. the two types target different parts of the pyramidal cell - perisomatic for PV, distal dendrite for SOM - which is why they can act on different input streams Quotation withheld because this record is not cleared for redistribution. Cells taken as excitatory were unlabelled rather than positively identified, so a minority may be other inhibitory types. what this limits: the excitatory arm of every comparison: THE UNLABELED NEURONS WERE MOST LIKELY EXCITATORY, ALTHOUGH AT A LOW CHANCE (<10%), THEY COULD ALSO BE ONE OF A VARIETY OF UNLABELED INHIBITORY CELL TYPES Quotation withheld because this record is not cleared for redistribution. Intensity thresholds are relative, not absolute, because background noise could not be fully removed. what this limits: every threshold figure: THE INTENSITY THRESHOLDS BEING COMPARED ARE RELATIVE RATHER THAN ABSOLUTE THRESHOLDS, SINCE ... WE COULD NOT COMPLETELY ELIMINATE BACKGROUND NOISE Quotation withheld because this record is not cleared for redistribution. All recordings were made under urethane anaesthesia, and SOM responses in particular may differ in the awake state. what this limits: the SOM results in the awake animal: FUTURE EXPERIMENTS ARE REQUIRED TO INVESTIGATE WHETHER SOM NEURONS ... EXHIBIT HIGHER EVOKED FIRING RATES IN AWAKE STATES THAN IN ANESTHESIA Quotation withheld because this record is not cleared for redistribution. urethane anaesthesia may suppress SOM cells specifically; in visual cortex their evoked rates fall about tenfold relative to the awake running state while PV rates do not what this limits: the SOM-versus-PV comparison specifically, because the anaesthetic is not neutral between them: SENSORY-EVOKED Quotation withheld ... WHILE THE FIRING RATES OF PV NEURONS ARE NOT DIFFERENT Quotation withheld because this record is not cleared for redistribution. intensity thresholds are relative, not absolute, because background noise could not be fully eliminated duplicate of: limitations[1] · what this limits: the same scope as limitations[1] - thresholds are relative to the noise floor of this rig Quotation withheld because this record is not cleared for redistribution. the unlabelled comparison cells are only most likely excitatory, with under a 10% chance of being an unlabelled interneuron type duplicate of: limitations[0] · what this limits: the same scope as limitations[0] - the comparison group is defined by absence of a label Quotation withheld because this record is not cleared for redistribution. the Gaussian bandwidth measure fits many PV cells poorly because their tuning is plateau-shaped, and 67% fall below R squared 0.3 what this limits: the bandwidth statistic for PV cells: THE PLATEAU SHAPE OF FREQUENCY TUNING OF MANY PV CELLS MAKES IT FIT POORLY INTO A GAUSSIAN CURVE (67% OF PV CELLS DISPLAYING R2 < 0.3), though INCLUDING ALL CELLS DID NOT AFFECT OUR CONCLUSION Quotation withheld because this record is not cleared for redistribution. whether SOM receptive fields look different in awake animals is left for future work duplicate of: limitations[2] · what this limits: the same scope as limitations[2] - the receptive-field claims for SOM cells are anaesthetised ones Quotation withheld because this record is not cleared for redistribution. Cre driver lines crossed to a reporter, so that a cell's type is known before it is recorded rather than inferred after Quotation withheld because this record is not cleared for redistribution. imaging-guided targeted recording of identified interneuron types, and the expectation that PV cells are less selective Quotation withheld because this record is not cleared for redistribution. cells whose Gaussian fit gave R-squared below 0.2 excluded from the bandwidth measure, AND the whole analysis re-run including every cell eliminates: that the bandwidth difference between cell types is an artefact of fitting a Gaussian to the plateau-shaped tuning of PV cells - and the re-run shows the exclusion is not doing the work · outcome (stated by the tagger, no quotation): held Quotation withheld because this record is not cleared for redistribution. SOM SOM Quotation withheld because this record is not cleared for redistribution. urethane urethane Quotation withheld because this record is not cleared for redistribution. PV and SOM here are GENETIC identities - Cre driver lines crossed to a fluorescent reporter and confirmed by dye co-localisation - not waveform classes Quotation withheld because this record is not cleared for redistribution. fast-spiking is NOT interchangeable with PV: 13% of SOM cells here had trough-to-peak intervals under 0.5 ms, and the cited work reports the PV and pyramidal waveform distributions overlap much more than assumed Quotation withheld because this record is not cleared for redistribution. spike-width classification is valid HERE and the paper says for what: 'for cell-attached recording in layer 2/3' - a scope, not a general licence Quotation withheld because this record is not cleared for redistribution. bandwidth is measured three ways in this paper - BW10, BW50% of an envelope fit, and the sigma of a Gaussian fit - and the third is adopted only to match the study being disputed Quotation withheld because this record is not cleared for redistribution. intensity threshold here is relative to an unremovable noise floor, so the numbers are comparable within this study and not against absolute-threshold work Quotation withheld because this record is not cleared for redistribution. claim_type: existence a second reading reaching the same value. Recorded: existence — note: a second reading reaching the same value from a different sentence. Not applied - a person decides. Quotation withheld because this record is not cleared for redistribution. method: in vivo two-photon imaging-guided cell-attached and whole-cell recording from fluorescently labelled PV and SOM neurons in layer 2/3 of premapped A1 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. Quotation withheld because this record is not cleared for redistribution. phenomenon: the tonal receptive field: which frequency and intensity combinations drive a cortical neuron a second reading DIFFERING from the recorded value. Recorded: tonal receptive fields of cortical inhibitory interneuron subtypes — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides. Quotation withheld because this record is not cleared for redistribution. contradicts → Moore and Wehr (2013) on PV frequency tuning breadth Quotation withheld because this record is not cleared for redistribution. replicates → Moore and Wehr (2013) on PV intensity tuning Quotation withheld because this record is not cleared for redistribution. builds-on → Wu et al. (2008) intracellular study of rat auditory cortex layer 4 Quotation withheld because this record is not cleared for redistribution. contradicts → Moore and Wehr 2013, which reported PV neurons as sharply tuned as excitatory cells in the same species and area Quotation withheld because this record is not cleared for redistribution. critiques → spike-waveform identification of inhibitory neurons in extracellular recording Quotation withheld because this record is not cleared for redistribution. confirms → fast-spiking-cell reports of broad frequency tuning, some with morphological confirmation Quotation withheld because this record is not cleared for redistribution. extends → visual cortex findings that PV neurons are less stimulus-selective than excitatory cells Quotation withheld because this record is not cleared for redistribution. builds-on → their own layer 4 intracellular work, where input range was similar but spike range still broader in fast-spiking cells Quotation withheld because this record is not cleared for redistribution. fast-spiking —distinct-from→ PV fast-spiking PV strength of the relation: 0.9 · note: CRITERION: the two are commonly equated, and the cited work shows PV and pyramidal waveform distributions overlap much more than assumed. elsewhere in the paper, 13% of SOM cells here also spike narrowly Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span reports a study questioning PV-FS correspondence; distinct-from overstates it. Quotation withheld because this record is not cleared for redistribution. PV —contrasted-with→ SOM PV SOM strength of the relation: 1 · note: CRITERION: broader tuning, lower threshold, stronger and earlier responses - four separable measures, all differing Quotation withheld because this record is not cleared for redistribution. layer 4 —causes→ PV layer 4 PV strength of the relation: 0.8 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but PV neurons receive feedforward inputs 'LIKELY from layer 4'; causes is hedged. Quotation withheld because this record is not cleared for redistribution. urethane —causes→ SOM urethane SOM status: BORROWED: reported of the VISUAL cortex and cited, not measured in this paper's auditory recordings · direction on the measure: reduces · strength of the relation: 0.8 · note: SIGN: suppressive, and selectively so - about tenfold in visual cortex while PV rates are unchanged. The main scope limit on this record Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says SOM firing is reduced under anesthesia; the endpoint 'SOM' drops 'firing'. Quotation withheld because this record is not cleared for redistribution. Gaussian —operationalised-as→ tonal receptive field Gaussian tonal receptive field strength of the relation: 0.5 — the record marks this weak · note: a measure adopted for comparability that fits the object badly - 67% of PV cells below R squared 0.3 because their tuning is plateau-shaped Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but frequency tuning was FITTED with a Gaussian, which fits PV cells poorly; operationalised-as is loose. Quotation withheld because this record is not cleared for redistribution. PV —operationalised-as→ feedforward inhibition PV feedforward inhibition strength of the relation: 0.9 · note: RE-GROUNDS another row of this record: the old span named ONE endpoint. Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but PV neurons 'CAN PROVIDE' feedforward inhibition; operationalised-as is loose. Quotation withheld because this record is not cleared for redistribution. ANOVA ANOVA Quotation withheld because this record is not cleared for redistribution. post hoc test post hoc test Quotation withheld because this record is not cleared for redistribution. Kolmogorov-Smirnov test Kolmogorov-Smirnov test Quotation withheld because this record is not cleared for redistribution. bootstrap bootstrap Quotation withheld because this record is not cleared for redistribution. standard error of the mean standard error of the mean Quotation withheld because this record is not cleared for redistribution. standard deviation standard deviation Quotation withheld because this record is not cleared for redistribution. curve fitting (Gaussian fit, coefficient of determination) curve fitting (Gaussian fit, coefficient of determination) Quotation withheld because this record is not cleared for redistribution. p-value p-value Quotation withheld because this record is not cleared for redistribution. one-way analysis of variance one-way analysis of variance Quotation withheld because this record is not cleared for redistribution. K–S test K–S test Quotation withheld because this record is not cleared for redistribution. Tamhane Tamhane Quotation withheld because this record is not cleared for redistribution. Bootstrap sampling Bootstrap sampling Quotation withheld because this record is not cleared for redistribution. Parvalbumin (PV) inhibitory neurons in layer 2/3 auditory cortex have broader tonal receptive fields, lower intensity thresholds and stronger tone-evoked spiking than somatostatin (SOM) neurons. Quotation withheld because this record is not cleared for redistribution. SOM neurons are as sharply frequency-tuned as excitatory neurons even though they too receive a broad range of excitatory input. Quotation withheld because this record is not cleared for redistribution. Both the synaptic input onset and the spike onset of SOM neurons occur significantly later than those of PV and excitatory neurons. Quotation withheld because this record is not cleared for redistribution. Measured 10 dB above threshold, PV receptive fields were twice as broad as those of SOM and excitatory neurons, which did not differ from each other. Quotation withheld because this record is not cleared for redistribution. SOM neurons had significantly higher intensity thresholds than PV and excitatory neurons. Quotation withheld because this record is not cleared for redistribution. SOM neurons fired with significantly less precise timing, showing larger first-spike jitter than PV and excitatory neurons. Quotation withheld because this record is not cleared for redistribution. PV neurons converted almost their whole synaptic input range into spike output (highest spike-range/input-range ratio), whereas SOM neurons converted the least. Quotation withheld because this record is not cleared for redistribution. Tone-evoked subthreshold depolarizations (EPSPs) were significantly larger in PV neurons than in SOM or excitatory neurons. Quotation withheld because this record is not cleared for redistribution. Every recorded PV cell had a spike trough-to-peak interval below 0.5 ms while every recorded excitatory cell was above 0.5 ms, so waveform width separates the two classes in layer 2/3. Quotation withheld because this record is not cleared for redistribution. PV neurons fired spontaneously at a significantly higher rate than excitatory cells. Quotation withheld because this record is not cleared for redistribution. The three cell groups were recorded at statistically indistinguishable cortical depths, so the broad PV tuning is not a depth-sampling artefact. Quotation withheld because this record is not cleared for redistribution. PV receptive fields were about twice as broad as SOM and excitatory ones, which did not differ from each other Quotation withheld because this record is not cleared for redistribution. the same result held for a second bandwidth measure taken from an envelope fit Quotation withheld because this record is not cleared for redistribution. PV cells fired much faster to tones than SOM or excitatory cells, and had higher spontaneous rates Quotation withheld because this record is not cleared for redistribution. SOM cells had higher intensity thresholds, so they may not inhibit at low sound levels Quotation withheld because this record is not cleared for redistribution. SOM cells had larger first-spike jitter than PV or excitatory cells Quotation withheld because this record is not cleared for redistribution. spikes occur in the order PV, excitatory, SOM, with SOM markedly delayed Quotation withheld because this record is not cleared for redistribution. all PV cells had trough-to-peak intervals under 0.5 ms and all excitatory cells above, but 13% of SOM cells also fell under Quotation withheld because this record is not cleared for redistribution. PV cells were less intensity-tuned than excitatory cells Quotation withheld because this record is not cleared for redistribution. ADOPTED THE RIVAL'S METHOD: fitting bandwidth by Gaussian as the disputing study did, the disagreement survived - PV tuning was still broader Quotation withheld because this record is not cleared for redistribution. three candidate causes remain for the disagreement: indirect optogenetic versus direct visual targeting, contamination in tungsten electrode recording, and general versus premapped A1 Quotation withheld because this record is not cleared for redistribution. CONTROL NULL: the three cell groups did not differ in characteristic frequency distribution Quotation withheld because this record is not cleared for redistribution. CONTROL NULL: the three groups did not differ in cortical depth, so the tuning difference is not a sampling bias Quotation withheld because this record is not cleared for redistribution. CONTROL: the imaging laser and chiller were isolated and no scanning occurred during recording, so scanner noise could not contaminate auditory responses Quotation withheld because this record is not cleared for redistribution. characteristic frequency (CF) characteristic frequency (CF) Quotation withheld because this record is not cleared for redistribution. intensity threshold intensity threshold Quotation withheld because this record is not cleared for redistribution. Bandwidth of TRF Bandwidth of TRF Quotation withheld because this record is not cleared for redistribution. intensity selectivity index (ISI) intensity selectivity index (ISI) Quotation withheld because this record is not cleared for redistribution. onset latency of spike response onset latency of spike response Quotation withheld because this record is not cleared for redistribution. characteristic frequency characteristic frequency Quotation withheld because this record is not cleared for redistribution. tonal receptive field tonal receptive field Quotation withheld because this record is not cleared for redistribution. intensity selectivity index intensity selectivity index Quotation withheld because this record is not cleared for redistribution. trough-to-peak interval trough-to-peak interval Quotation withheld because this record is not cleared for redistribution. bandwidth of spike TRF bandwidth of spike TRF Quotation withheld because this record is not cleared for redistribution. response onset latency response onset latency Quotation withheld because this record is not cleared for redistribution. body temperature body temperature Quotation withheld because this record is not cleared for redistribution. distribution of characteristic frequencies distribution of characteristic frequencies Quotation withheld because this record is not cleared for redistribution. cortical depth cortical depth Quotation withheld because this record is not cleared for redistribution. BW10 BW10 Quotation withheld because this record is not cleared for redistribution. intensity threshold intensity threshold Quotation withheld because this record is not cleared for redistribution. jitter jitter Quotation withheld because this record is not cleared for redistribution. trough-to-peak interval trough-to-peak interval Quotation withheld because this record is not cleared for redistribution. 27 PV, 16 SOM, 22 excitatory — number of neurons recorded in the cell-attached spike dataset Quotation withheld because this record is not cleared for redistribution. PV, 75 ± 6 Hz; SOM, 28 ± 4 Hz; Exc, 39 ± 6 Hz — average tone-evoked firing rates of the three cell types Quotation withheld because this record is not cleared for redistribution. PV, 1.43 ± 0.12 octave; SOM, 0.67 ± 0.06 octave; Exc, 0.64 ± 0.03 octave — receptive-field bandwidth 10 dB above threshold (BW10) per cell type Quotation withheld because this record is not cleared for redistribution. PV, 36 ± 1.93 dB SPL; SOM, 47.86 ± 2.22 dB SPL; Exc, 40 ± 0.97 dB SPL — average intensity threshold of the tonal receptive field per cell type Quotation withheld because this record is not cleared for redistribution. PV (0.89 ± 0.06), SOM (0.43 ± 0.05), excitatory (0.61 ± 0.04) — ratio of spike response frequency range to synaptic input frequency range at 60 dB SPL Quotation withheld because this record is not cleared for redistribution. PV, 16.59 ± 0.58 ms; SOM, 26.44 ± 0.91 ms; Exc, 18.64 ± 0.88 ms — average spike onset latency to CF tones at 60 dB SPL Quotation withheld because this record is not cleared for redistribution. PV, 1.54 ± 0.12 ms; SOM, 3.21 ± 0.36 ms; Exc, 1.75 ± 0.19 ms — average jitter (SD) of first-spike timing for CF tones at 60 dB SPL Quotation withheld because this record is not cleared for redistribution. 13% — share of SOM neurons whose spike trough-to-peak interval was shorter than 0.5 ms (the PV-like range) Quotation withheld because this record is not cleared for redistribution. 67% of PV cells displaying R 2 < 0.3 — proportion of PV cells whose plateau-shaped tuning fitted a Gaussian curve poorly Quotation withheld because this record is not cleared for redistribution. 0.5 ms — trough-to-peak cut separating PV from excitatory cells here, crossed by 13% of SOM cells Quotation withheld because this record is not cleared for redistribution. 27, 16, 22 — PV, SOM and excitatory neurons recorded Quotation withheld because this record is not cleared for redistribution. 1.17 vs 0.52 vs 0.55 octave — Gaussian-fit tuning bandwidth for PV, SOM and excitatory cells using the disputed study's own method Quotation withheld because this record is not cleared for redistribution. 67% — PV cells whose plateau-shaped tuning fits a Gaussian below R squared 0.3 Quotation withheld because this record is not cleared for redistribution. under 10% — chance that an unlabelled comparison cell was actually an interneuron Quotation withheld because this record is not cleared for redistribution. 287 — frequency-intensity combinations presented per cell Quotation withheld because this record is not cleared for redistribution.

The full text of this article is not redistributable, so it is not included here. The evidence sentences are withheld with it; the tag layer and its grounding status remain. The article itself is at https://doi.org/10.1093/cercor/bht417.