Analysis of excitatory and inhibitory neuron types in the inferior colliculus based on I h properties

Analysis of excitatory and inhibitory neuron types in the inferior colliculus based on I h properties Naumov Heyd de Arnal Koch

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Analysis of excitatory and inhibitory neuron types in the inferior colliculus based on I h properties 2019 Journal of Neurophysiology 10.1152/jn.00594.2018 Naumov2019-analysis-of-excitatory-and-inhibitory-neuron Naumov2019-analysis-of-excitatory-and-inhibitory-neuron.txt, the project's staged copy 4fc4367f906484ea499fb1ff4b26af13ed1d919b 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 existence Quotation withheld because this record is not cleared for redistribution. Ih and HCN channel diversity across inferior colliculus neuron types Quotation withheld because this record is not cleared for redistribution. electrophysiology electrophysiology Quotation withheld because this record is not cleared for redistribution. mechanisms Differential distribution of HCN channel subtypes across neuron classes (or, alternatively, second-messenger modulation of Ih) produces the observed differences in Ih kinetics and voltage dependence Quotation withheld because this record is not cleared for redistribution. mechanisms GABAergic neurons carry a slow Ih because their current is conducted by channels assembled from HCN2 and HCN4 subunits Quotation withheld because this record is not cleared for redistribution. mechanisms Fast Ih in excitatory neurons is carried by HCN1/HCN2 channels while slow Ih is carried by HCN2 and sometimes HCN4, so subunit composition sets the kinetic class Quotation withheld because this record is not cleared for redistribution. mechanisms Because HCN4 is restricted to GABAergic cells and is cAMP-sensitive, neuromodulators that raise second-messenger levels would selectively retune the integrative properties of the inhibitory population Quotation withheld because this record is not cleared for redistribution. mechanisms A large, fast-activating Ih depolarizes the membrane after release from hyperpolarization and thereby enables rebound spiking in specific excitatory subclasses Quotation withheld because this record is not cleared for redistribution. Slice preparation and recording configuration may have biased sampling toward particular GABAergic cell types, so further subclasses may exist. what this limits: the completeness of the subclass list: BRAIN SLICE PREPARATION AND RECORDING CONFIGURATIONS MAY HAVE BIASED OUR SAMPLE TOWARD ONE NEURON TYPE Quotation withheld because this record is not cleared for redistribution. Low-prevalence GABAergic types could have been missed given the sample size. what this limits: any claim that a type is absent: OTHER GABAERGIC NEURON TYPES WITH LOW PREVALENCE (10% OR LOWER) MAY HAVE ESCAPED OUR SAMPLE DUE TO OUR RELATIVELY LOW SAMPLE SIZE Quotation withheld because this record is not cleared for redistribution. The current-clamp sag measures used to group cells are only partial proxies for Ih amplitude and kinetics. what this limits: the grouping variable itself: THESE TWO PARAMETERS ONLY PARTIALLY CORRELATE WITH Ih AMPLITUDE AND Ih ACTIVATION TIME CONSTANT BUT ALSO DEPEND ON MEMBRANE CAPACITANCE AND THE ACTIVATION OF OTHER VOLTAGE-GATED ION CHANNELS Quotation withheld because this record is not cleared for redistribution. The physiological classification alone may not exhaust the GABAergic populations; molecular profiling could split them further. what this limits: the physiological classification as a final answer: FURTHER PHYSIOLOGICAL MEASUREMENTS OR MOLECULAR PROFILING MAY REVEAL DIFFERENT POPULATIONS OF GABAERGIC IC NEURONS Quotation withheld because this record is not cleared for redistribution. The dividing line between the two groups is a chosen value, and the neurons that do not fit it are named in the same sentence. what this limits: the two Ih groups as natural kinds: the boundary is an activation time constant chosen by the authors at 235 ms, with two intermediate neurons stated not to follow it Quotation withheld because this record is not cleared for redistribution. The inhibitory population is reported as undivided BY THESE MEASUREMENTS, with the authors stating the preparation may have biased the sample toward one type. what this limits: the finding that GABAergic IC neurons form a single physiological class: the authors state this holds for these measurements and that the preparation may have biased the sample toward one type Quotation withheld because this record is not cleared for redistribution. A detection floor is stated: cell types at or below 10% prevalence may simply not be in the sample. what this limits: the absence of a GABAergic subtype in these data as evidence it does not exist: the authors state types at 10% prevalence or lower may have escaped the sample given its size Quotation withheld because this record is not cleared for redistribution. A measurement is declared biased with its direction named, and then argued not to affect the comparison that rests on it. what this limits: the reported time constants for slow-Ih neurons: the authors state Ih was not fully activated at the measurement point, so the taus are underestimates, and argue the group differences survive Quotation withheld because this record is not cleared for redistribution. the dorso-ventral extent of each inferior colliculus normalised from zero to one before being divided into five regions eliminates: differences in absolute size between animals - without normalising, the same anatomical position falls in different regions in different brains · outcome (stated by the tagger, no quotation): held Quotation withheld because this record is not cleared for redistribution. inferior colliculus inferior colliculus Quotation withheld because this record is not cleared for redistribution. CIC CIC Quotation withheld because this record is not cleared for redistribution. activation time constant activation time constant Quotation withheld because this record is not cleared for redistribution. neuron type - defined by Ih properties measured in acute slices, with the group boundary set at a chosen activation time constant rather than at an observed discontinuity Quotation withheld because this record is not cleared for redistribution. GABAergic neurons - a single undivided population UNDER THESE MEASUREMENTS, which the paper distinguishes from the claim that no further subclasses exist Quotation withheld because this record is not cleared for redistribution. inferior colliculus subdivisions - adopted from an atlas classification based on Nissl staining, not derived from the physiology being reported Quotation withheld because this record is not cleared for redistribution. replicates → Zhao et al. (2011) about this quote: 2026-08-16: FAILED GLYPH INSIDE THIS QUOTE. The staged text reads "are also GAD67(cid:2) and thus GABAergic". (cid:2) IS A SIGN CHARACTER AND I AM NOT ASSERTING WHICH ONE. The sentence's own logic points to a plus - GABAergic neurons are GAD67-positive, and the clause concludes "and thus GABAergic" - but plus and minus are OPPOSITE readings of the same tag, and supplying the sign from domain knowledge would be putting a true outside fact into a quote. Read it as: the cells are GAD67-[sign unreadable in the staged file], which the authors take to establish they are GABAergic. The quote is an exact substring of the staged file; the file is incomplete here, not the tag. Quotation withheld because this record is not cleared for redistribution. contradicts → Ono et al. (2005) Quotation withheld because this record is not cleared for redistribution. extends → Koch et al. (2004) Quotation withheld because this record is not cleared for redistribution. replicates → Ono et al. (2017) Quotation withheld because this record is not cleared for redistribution. EVIDENCE CORRECTED (RE-GROUNDED) · CIC —part-of→ inferior colliculus CIC inferior colliculus EVIDENCE CORRECTED: RE-GROUNDED. The old span cited a classification scheme. This sentence states the containment in the paper's own phrase - 'neurons in the central nucleus of inferior colliculus (CIC)' - which is the anatomy the edge records, with the abbreviation defined in place. — strength of the relation: 1 Quotation withheld because this record is not cleared for redistribution. CIC —part-of→ inferior colliculus CIC inferior colliculus strength of the relation: 1 · note: RE-GROUNDS another row of this record: the old span named ONE endpoint. Quotation withheld because this record is not cleared for redistribution. Shapiro–Wilk normality test Shapiro–Wilk normality test Quotation withheld because this record is not cleared for redistribution. Levene test for equality of variance Levene test for equality of variance Quotation withheld because this record is not cleared for redistribution. Kruskal–Wallis test with Dunn's post hoc Kruskal–Wallis test with Dunn's post hoc Quotation withheld because this record is not cleared for redistribution. one-way ANOVA with Tukey post hoc one-way ANOVA with Tukey post hoc Quotation withheld because this record is not cleared for redistribution. standard error of the mean standard error of the mean about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:8). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. curve fitting (single exponential) curve fitting (single exponential) Quotation withheld because this record is not cleared for redistribution. curve fitting (Boltzmann function) curve fitting (Boltzmann function) Quotation withheld because this record is not cleared for redistribution. curve fitting (double exponential) curve fitting (double exponential) about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5), (cid:6). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. Every GABAergic inferior colliculus neuron sampled showed a slowly activating hyperpolarization-activated current of moderate amplitude, whereas some excitatory neurons showed a fast-activating, large-amplitude current. Quotation withheld because this record is not cleared for redistribution. Immunostaining placed the fast-gating HCN1 channel around excitatory neurons and the slow-gating HCN4 channel perisomatically around most inhibitory neurons. Quotation withheld because this record is not cleared for redistribution. GABAergic somata lacked HCN1 labelling. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. About 15% of glutamatergic neurons fired only at stimulus onset, and this pattern always co-occurred with a large, fast Ih. Quotation withheld because this record is not cleared for redistribution. In contrast to many other brain regions, membrane properties and firing patterns were similar in GABAergic neurons and about a third of the excitatory neurons here. Quotation withheld because this record is not cleared for redistribution. Only the onset-firing excitatory neurons formed a distinct class; GABAergic neurons were statistically indistinguishable from many excitatory types on membrane properties and Ih. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. HCN1 immunoreactivity showed no systematic dorso-ventral (tonotopic) gradient across the mouse central nucleus. Quotation withheld because this record is not cleared for redistribution. 80% of fast-Ih adapting excitatory neurons showed rebound spiking, whereas few GABAergic or slow-Ih excitatory neurons did. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. Firing rate versus injected-current relationships did not differ between GABAergic and any excitatory group. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. GABAergic neurons adapted their spike rate rapidly within the first 200 ms and little thereafter. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. 96% of EYFP-positive (GABAergic) neurons carried a perisomatic HCN4 ring. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. The onset-firing excitatory neurons' membrane properties resemble those of principal neurons of the lateral and medial superior olive. Quotation withheld because this record is not cleared for redistribution. Ih current density was indistinguishable between GABAergic and slow-tau excitatory neurons but significantly larger in fast-tau excitatory neurons. about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. onset firing neuron onset firing neuron Quotation withheld because this record is not cleared for redistribution. adapting firing neuron adapting firing neuron Quotation withheld because this record is not cleared for redistribution. current density current density Quotation withheld because this record is not cleared for redistribution. excitatory neurons excitatory neurons about this quote: RULED, NOT PREVIOUSLY ADJUDICATED. This quote carries an unmapped font glyph and was not covered by the 2026-08-25 ruling, which reached only rows flagged with glyph damage. The same reasoning applies and the same decision follows: THE QUOTE IS NOT EDITED. The unmapped characters are the ones carrying the claim, and no code-to-character table is possible - in one sentence of this corpus the same code is '=' once and 'tau' twice. Repair needs the source PDF and is a re-staging job, costed and refused (0 of 114 appendable; a per-record re-stage fixes 21 and breaks 223). Three sibling rows were found ALREADY ruled under this same pass and are untouched. Quotation withheld because this record is not cleared for redistribution. inhibitory neurons inhibitory neurons about this quote: RULED, NOT PREVIOUSLY ADJUDICATED. This quote carries an unmapped font glyph and was not covered by the 2026-08-25 ruling, which reached only rows flagged with glyph damage. The same reasoning applies and the same decision follows: THE QUOTE IS NOT EDITED. The unmapped characters are the ones carrying the claim, and no code-to-character table is possible - in one sentence of this corpus the same code is '=' once and 'tau' twice. Repair needs the source PDF and is a re-staging job, costed and refused (0 of 114 appendable; a per-record re-stage fixes 21 and breaks 223). Three sibling rows were found ALREADY ruled under this same pass and are untouched. Quotation withheld because this record is not cleared for redistribution. hyperpolarizing voltage steps hyperpolarizing voltage steps about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. adaptation rate adaptation rate about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. holding potential holding potential about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. Ih densities Ih densities Quotation withheld because this record is not cleared for redistribution. activation time constant activation time constant Quotation withheld because this record is not cleared for redistribution. half-maximal activation voltage half-maximal activation voltage Quotation withheld because this record is not cleared for redistribution. 33%, 36%, 25% — share of recorded neurons that were GABAergic with slow Ih, excitatory with fast Ih, and excitatory with slow Ih about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. 6% — share of neurons with no detectable Ih, all of them excitatory about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:5). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. 96% (74 out of 77) — proportion of GABAergic (EYFP-positive) neurons showing a perisomatic HCN4 ring about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. 88% — share of inferior colliculus neurons with an adapting firing pattern, irrespective of transmitter type Quotation withheld because this record is not cleared for redistribution. 15% — share of glutamatergic neurons showing onset spiking, always paired with large fast Ih Quotation withheld because this record is not cleared for redistribution. P = 0.98 — one-way ANOVA result showing mean firing rates at 200 pA above threshold did not differ across the four neuron groups (115–117 Hz) about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5), (cid:6), (cid:8). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. · value source note: The value is written with the paper's own symbol (=). Our copy of the text prints it as '(cid:6)' ('P (cid:6) 0.98'), because the PDF's text layer maps its maths characters to the wrong codes. The value is read from the PDF, page 10. Quotation withheld because this record is not cleared for redistribution. 69 ± 11 ms vs 167 ± 32 ms — weighted adaptation time constants for GABAergic neurons versus slow-Ih excitatory neurons about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:5), (cid:6), (cid:8). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. 235 ms — activation time-constant cutoff used to split excitatory neurons into fast-tau and slow-tau Ih groups about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:2), (cid:9). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. Quotation withheld because this record is not cleared for redistribution. 22–28 days, ~1:1 — age of the mice used for patch-clamp recording and the male/female ratio of the sample Quotation withheld because this record is not cleared for redistribution. n = 5 — number of inferior colliculi in which HCN1 mean gray value was quantified along the dorso-ventral axis, with no gradient found about this quote: DAMAGED GLYPHS IN THIS QUOTE: (cid:6). These are raw PDF font glyph indices that leaked out in place of the characters they stand for - usually a mathematical operator. The quote is verbatim to the staged text and its anchor is sound; the STAGED TEXT is what is damaged. NOT repaired: the cid-to-character mapping is defined by the font embedded in that particular PDF, so guessing it would produce a clean and unverifiable quote. Repair needs the source PDF and is a re-staging job. · value source note: The value is written with the paper's own symbol (=). Our copy of the text prints it as '(cid:6)' ('n (cid:6) 5'), because the PDF's text layer maps its maths characters to the wrong codes. The value is read from the PDF, page 5. 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/10.1152/jn.00594.2018.