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

Naumov, Heyd, de Arnal, Koch · Journal of Neurophysiology · 2019 · neuroscience · 10.1152/jn.00594.2018 · record Naumov2019-analysis-of-excitatory-and-inhibitory-neuron
tag layer · index · concepts · authors · עברית
Where this record's text came from. faceted 2026-08 by reading the locally held text (pdftext); every evidence string verified as an exact substring of that text before merging. This is a pdftext extraction, so several sentences contain literal '(cid:N)' artifacts standing in for symbols such as + and -; the Zhao (2011) evidence quote retains '(cid:2)' exactly as extracted, since any tidying would break the substring check. The Koch et al. (2004) relation is tagged extends because the paper deliberately tests the rat HCN1 tonotopic gradient in mouse; the test came out negative (no dorso-ventral gradient in mouse CIC), which is closer to a species-limited disconfirmation than the paper says in so many words, so it is not tagged contradicts. Claim type is existence rather than mechanism because the headline result is a distribution/correlation across cell types, not a causal account; the functional consequences of the Ih differences are discussed only as inference from other literature. 2026-08-14, 2026-08-15 and 2026-08-16 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
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Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 3 concept tags on this paper, all 3 are grounded in a quoted sentence.
inferior colliculus CIC activation time constant 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 GABAergic neurons carry a slow Ih because their current is conducted by channels assembled from HCN2 and HCN4 subunits 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 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 A large, fast-activating Ih depolarizes the membrane after release from hyperpolarization and thereby enables rebound spiking in specific excitatory subclasses onset firing neuron adapting firing neuron current density excitatory neurons inhibitory neurons hyperpolarizing voltage steps adaptation rate holding potential Ih densities half-maximal activation voltage Shapiro–Wilk normality test Shapiro- Wilk normality Levene test for equality of variance Levene variance test Kruskal–Wallis test with Dunn's post hoc nonparametric Kruskal-Wallis test or Friedman test followed by Dunn’s multiple comparisons test one-way ANOVA with Tukey post hoc one-way ANOVA followed by a Tukey’s multiple comparisons test standard error of the mean means ± SE curve fitting (single exponential) applying a single exponential fit to the currents curve fitting (Boltzmann function) fitted with a Boltzmann function curve fitting (double exponential) a double exponential function ... was fit to the successive spike frequency plot electrophysiology

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relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
conceptconceptsan idea the paper works with, quoted where the paper uses it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
controlcontrolsan experiment run to eliminate a rival explanation
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
phenomenonphenomenonthe effect or regularity under study
sensesensesa distinct use of a term, quoted where the paper fixes it

The paper

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1152/jn.00594.2018. The reading view exists for the papers whose licence permits it.

Every tag on this page

claim_relations 4 · claim_type 1 · concept_edges 2 · concepts 3 · content_tags 42 · controls 1 · limitations 8 · mechanisms 5 · method 1 · phenomenon 1 · senses 3

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 2 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facettagfieldconf.whose words
claim_typeexistence
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claim_typethe paper's words
phenomenonIh and HCN channel diversity across inferior colliculus neuron types
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phenomenonthe paper's words
methodelectrophysiology
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methodthe paper's words
mechanismsmechanisms
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mechanisms[0]the tagger's words
mechanismsmechanisms
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mechanisms[1]the tagger's words
mechanismsmechanisms
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mechanisms[2]the tagger's words
mechanismsmechanisms
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mechanisms[3]the tagger's words
mechanismsmechanisms
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mechanisms[4]the tagger's words
limitationsSlice preparation and recording configuration may have biased sampling toward particular GABAergic cell types, so further subclasses may exist.
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what this limits: the completeness of the subclass list: BRAIN SLICE PREPARATION AND RECORDING CONFIGURATIONS MAY HAVE BIASED OUR SAMPLE TOWARD ONE NEURON TYPE
limitations[0]the tagger's words
limitationsLow-prevalence GABAergic types could have been missed given the sample size.
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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
limitations[1]the tagger's words
limitationsThe current-clamp sag measures used to group cells are only partial proxies for Ih amplitude and kinetics.
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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
limitations[2]the tagger's words
limitationsThe physiological classification alone may not exhaust the GABAergic populations; molecular profiling could split them further.
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what this limits: the physiological classification as a final answer: FURTHER PHYSIOLOGICAL MEASUREMENTS OR MOLECULAR PROFILING MAY REVEAL DIFFERENT POPULATIONS OF GABAERGIC IC NEURONS
limitations[3]the tagger's words
limitationsThe dividing line between the two groups is a chosen value, and the neurons that do not fit it are named in the same sentence.
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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
limitations[4]1.0the tagger's words
limitationsThe inhibitory population is reported as undivided BY THESE MEASUREMENTS, with the authors stating the preparation may have biased the sample toward one type.
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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
limitations[5]1.0the tagger's words
limitationsA detection floor is stated: cell types at or below 10% prevalence may simply not be in the sample.
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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
limitations[6]1.0the tagger's words
limitationsA measurement is declared biased with its direction named, and then argued not to affect the comparison that rests on it.
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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
limitations[7]1.0the tagger's words
controlsthe dorso-ventral extent of each inferior colliculus normalised from zero to one before being divided into five regions
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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
controls[0]0.85the tagger's words
conceptsinferior colliculus
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concepts[0]the paper's words
conceptsCIC
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also in: CIC
concepts[1]the paper's words
conceptsactivation time constant
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concepts[2]the paper's words
sensesneuron 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
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senses[0]1.0the tagger's words
sensesGABAergic neurons - a single undivided population UNDER THESE MEASUREMENTS, which the paper distinguishes from the claim that no further subclasses exist
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senses[1]1.0the tagger's words
sensesinferior colliculus subdivisions - adopted from an atlas classification based on Nissl staining, not derived from the physiology being reported
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senses[2]1.0the tagger's words
claim_relationsreplicates → Zhao et al. (2011)
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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.
claim_relations[0]the tagger's words
claim_relationscontradicts → Ono et al. (2005)
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claim_relations[1]the tagger's words
claim_relationsextends → Koch et al. (2004)
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claim_relations[2]the tagger's words
claim_relationsreplicates → Ono et al. (2017)
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claim_relations[3]the tagger's words
concept_edgesEVIDENCE CORRECTED (RE-GROUNDED) · CIC —part-of→ inferior colliculus
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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
concept_edges[0]the tagger's words
concept_edgesCIC —part-of→ inferior colliculus
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strength of the relation: 1 · note: RE-GROUNDS another row of this record: the old span named ONE endpoint.
concept_edges[1]0.9the tagger's words
content_tagsShapiro–Wilk normality test
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content_tags.statistics[0]the tagger's words
content_tagsLevene test for equality of variance
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content_tags.statistics[1]the tagger's words
content_tagsKruskal–Wallis test with Dunn's post hoc
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content_tags.statistics[2]the tagger's words
content_tagsone-way ANOVA with Tukey post hoc
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content_tags.statistics[3]the tagger's words
content_tagsstandard error of the mean
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content_tags.statistics[4]the tagger's words
content_tagscurve fitting (single exponential)
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content_tags.statistics[5]the tagger's words
content_tagscurve fitting (Boltzmann function)
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content_tags.statistics[6]the tagger's words
content_tagscurve fitting (double exponential)
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content_tags.statistics[7]the tagger's words
content_tagsEvery 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.
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content_tags.findings[0]the tagger's words
content_tagsImmunostaining placed the fast-gating HCN1 channel around excitatory neurons and the slow-gating HCN4 channel perisomatically around most inhibitory neurons.
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content_tags.findings[1]the tagger's words
content_tagsGABAergic somata lacked HCN1 labelling.
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content_tags.findings[2]the tagger's words
content_tagsAbout 15% of glutamatergic neurons fired only at stimulus onset, and this pattern always co-occurred with a large, fast Ih.
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content_tags.findings[3]the tagger's words
content_tagsIn 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.
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content_tags.findings[4]the tagger's words
content_tagsOnly the onset-firing excitatory neurons formed a distinct class; GABAergic neurons were statistically indistinguishable from many excitatory types on membrane properties and Ih.
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content_tags.findings[5]the tagger's words
content_tagsHCN1 immunoreactivity showed no systematic dorso-ventral (tonotopic) gradient across the mouse central nucleus.
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content_tags.findings[6]the tagger's words
content_tags80% of fast-Ih adapting excitatory neurons showed rebound spiking, whereas few GABAergic or slow-Ih excitatory neurons did.
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content_tags.findings[7]the tagger's words
content_tagsFiring rate versus injected-current relationships did not differ between GABAergic and any excitatory group.
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content_tags.findings[8]the tagger's words
content_tagsGABAergic neurons adapted their spike rate rapidly within the first 200 ms and little thereafter.
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content_tags.findings[9]the tagger's words
content_tags96% of EYFP-positive (GABAergic) neurons carried a perisomatic HCN4 ring.
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content_tags.findings[10]the tagger's words
content_tagsThe onset-firing excitatory neurons' membrane properties resemble those of principal neurons of the lateral and medial superior olive.
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content_tags.findings[11]the tagger's words
content_tagsIh current density was indistinguishable between GABAergic and slow-tau excitatory neurons but significantly larger in fast-tau excitatory neurons.
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content_tags.findings[12]the tagger's words
content_tagsonset firing neuron
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content_tags.definitions[0]the tagger's words
content_tagsadapting firing neuron
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content_tags.definitions[1]the tagger's words
content_tagscurrent density
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content_tags.definitions[2]the tagger's words
content_tagsexcitatory neurons
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content_tags.definitions[5]the tagger's words
content_tagsinhibitory neurons
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content_tags.definitions[6]the tagger's words
content_tagshyperpolarizing voltage steps
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content_tags.variables[0]the tagger's words
content_tagsadaptation rate
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content_tags.variables[4]the tagger's words
content_tagsholding potential
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content_tags.variables[7]the tagger's words
content_tagsIh densities
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also in: Ih densities
content_tags.variables[8]the tagger's words
content_tagsactivation time constant
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content_tags.variables[9]the tagger's words
content_tagshalf-maximal activation voltage
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content_tags.variables[10]the tagger's words
content_tags33%, 36%, 25% — share of recorded neurons that were GABAergic with slow Ih, excitatory with fast Ih, and excitatory with slow Ih
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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.
content_tags.quantities[0]the tagger's words
content_tags6% — share of neurons with no detectable Ih, all of them excitatory
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
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.
content_tags.quantities[1]the tagger's words
content_tags96% (74 out of 77) — proportion of GABAergic (EYFP-positive) neurons showing a perisomatic HCN4 ring
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
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.
content_tags.quantities[2]the tagger's words
content_tags88% — share of inferior colliculus neurons with an adapting firing pattern, irrespective of transmitter type
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[3]the tagger's words
content_tags15% — share of glutamatergic neurons showing onset spiking, always paired with large fast Ih
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[4]the tagger's words
content_tagsP = 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)
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
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.
content_tags.quantities[5]the tagger's words
content_tags69 ± 11 ms vs 167 ± 32 ms — weighted adaptation time constants for GABAergic neurons versus slow-Ih excitatory neurons
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
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.
content_tags.quantities[6]the tagger's words
content_tags235 ms — activation time-constant cutoff used to split excitatory neurons into fast-tau and slow-tau Ih groups
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
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.
content_tags.quantities[7]the tagger's words
content_tags22–28 days, ~1:1 — age of the mice used for patch-clamp recording and the male/female ratio of the sample
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[8]the tagger's words
content_tagsn = 5 — number of inferior colliculi in which HCN1 mean gray value was quantified along the dorso-ventral axis, with no gradient found
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
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.
content_tags.quantities[9]the tagger's words

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