Tonotopic Organization of the Superior Olivary Nucleus in the Chicken Auditory Brainstem

Tonotopic Organization of the Superior Olivary Nucleus in the Chicken Auditory Brainstem Tabor Coleman Rubel Burger

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Tonotopic Organization of the Superior Olivary Nucleus in the Chicken Auditory Brainstem 2012 The Journal of comparative neurology 10.1002/cne.22807 Tabor2012-tonotopic-organization-of-the-superior-oliva Tabor2012-tonotopic-organization-of-the-superior-oliva.txt, the project's staged copy ddbd4e03c31f274648786fb87e990eac0898355a 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 the topographic organisation of frequency and of response types in the chicken superior olivary nucleus Quotation withheld because this record is not cleared for redistribution. whether the tonotopic organisation of the auditory brainstem is preserved in an inhibitory long-range projection nucleus Quotation withheld because this record is not cleared for redistribution. quantifies tonotopy in an inhibitory nucleus, shows it is less precise than in excitatory nuclei of the same species, and gives a predictive model from position to best frequency Quotation withheld because this record is not cleared for redistribution. existence Quotation withheld because this record is not cleared for redistribution. tonotopic organization of an inhibitory auditory brainstem nucleus Quotation withheld because this record is not cleared for redistribution. electrophysiology electrophysiology Quotation withheld because this record is not cleared for redistribution. mechanisms SON response patterns are inherited from its nucleus angularis input, which is why sustained and onset types and diverse tuning shapes appear there Quotation withheld because this record is not cleared for redistribution. mechanisms the narrow hearing range of the chicken makes the frequency-to-place function linear rather than logarithmic Quotation withheld because this record is not cleared for redistribution. mechanisms summation of individually subthreshold spike waveforms raises the apparent threshold of multiple-unit recordings Quotation withheld because this record is not cleared for redistribution. mechanisms the broad, non-restricted SON to nucleus laminaris projection supplies the anatomical substrate by which inhibition laterally sharpens frequency tuning Quotation withheld because this record is not cleared for redistribution. inhibitory neurons in this system sit in a discrete projection nucleus rather than among principal cells, which is what makes their organisation separable at all Quotation withheld because this record is not cleared for redistribution. recording sites are located by electrolytic lesions and electrode tracks, then normalised into a common 3D nucleus so cases can be combined Quotation withheld because this record is not cleared for redistribution. PCA is run first because it was unknown whether the map was linear or nonlinear, single or multiple gradients Quotation withheld because this record is not cleared for redistribution. SON inhibition widens dynamic range and sharpens temporal coding for sound localisation Quotation withheld because this record is not cleared for redistribution. Threshold comparisons between sustained and onset responders may not be valid because the spike-rate criterion may not be equally sensitive across response categories. what this limits: threshold comparisons between response types only: CAUTION SHOULD BE USED WHEN COMPARING THRESHOLDS BETWEEN SUSTAINED AND ONSET RESPONDERS BECAUSE THE SPIKE RATE CRITERION USED TO DETERMINE RESPONSE THRESHOLD differs in effect between them Quotation withheld because this record is not cleared for redistribution. Binaural response properties of SON neurons were not assessed, so a possible contribution of the nucleus laminaris projection remains unexplored. what this limits: any claim about the origin of SON response properties: WE DID NOT ASSESS THE BINAURAL RESPONSE PROPERTIES OF SON NEURONS, so NL's projection remains an unexplored contribution Quotation withheld because this record is not cleared for redistribution. In most animals only a limited region of the SON was sampled, which is why individual maps had to be combined into a standardized SON. what this limits: the tonotopic map as a whole-nucleus claim: IN MOST SUBJECTS WE SAMPLED A LIMITED REGION OF THE SON, which is why individual maps had to be combined Quotation withheld because this record is not cleared for redistribution. the map is explicitly less precise than in excitatory nuclei of the same species, by both error magnitude and r-squared what this limits: any use of this map at the precision expected of excitatory nuclei: the paper states its map is the less precise of the two by both error magnitude and its own comparison Quotation withheld because this record is not cleared for redistribution. prediction error varies widely along the tonotopic axis rather than being uniform what this limits: the tonotopic prediction at any single site: ERROR MAGNITUDE (|CF_predicted - CF_measured|) VARIED WIDELY AT EACH POSITION ALONG THE CALCULATED TONOTOPIC AXIS Quotation withheld because this record is not cleared for redistribution. the threshold criterion may not be equally sensitive across response categories, so between-category threshold comparisons are cautioned against duplicate of: limitations[0] · what this limits: the same scope as limitations[0] - the threshold criterion may not be equally sensitive across response categories, so between-category threshold comparisons are the affected ones Quotation withheld because this record is not cleared for redistribution. multi-unit thresholds run higher than single-unit thresholds because sub-threshold waveforms sum what this limits: any comparison of thresholds between recording types: THRESHOLDS OF MULTIPLE-UNIT RESPONSES TENDED TO BE SLIGHTLY HIGHER THAN THE THRESHOLDS OF ISOLATED NEURONAL RESPONSES ... THIS INCREASE REFLECTS ... WAVEFORMS THAT INDIVIDUALLY WERE NOT DETECTED Quotation withheld because this record is not cleared for redistribution. suppressed responders were too rare to analyse spatially what this limits: the spatial claims for suppressed responders: SUPPRESSED RESPONDERS WERE ENCOUNTERED TOO INFREQUENTLY TO ASSESS THEIR SPATIAL DISTRIBUTION Quotation withheld because this record is not cleared for redistribution. in most animals only a limited region of the nucleus was sampled, which is why standardisation was needed what this limits: every position reported in the standardised frame: in most animals only a limited region was sampled, so LOCATIONS OF THE RECORDING SITES WERE CONVERTED INTO NORMALIZED DISTANCES ALONG THREE ORTHOGONAL AXES Quotation withheld because this record is not cleared for redistribution. within an isofrequency slice, position carried no information about CF at all what this limits: the tonotopy claim at the scale a user would apply it: within an isofrequency slice, position carries no information about CF - THE STANDARDIZED SON WAS SLICED INTO FIVE SECTIONS to test exactly this Quotation withheld because this record is not cleared for redistribution. retinotopy as the template for what a sensory map is and why it matters Quotation withheld because this record is not cleared for redistribution. each stimulus presented twenty times in a pseudo-randomised sequence eliminates: adaptation and order - a fixed sequence would confound the tonotopic response with position in the run · outcome (stated by the tagger, no quotation): held Quotation withheld because this record is not cleared for redistribution. characteristic frequency characteristic frequency Quotation withheld because this record is not cleared for redistribution. Q10 Q10 Quotation withheld because this record is not cleared for redistribution. response threshold response threshold Quotation withheld because this record is not cleared for redistribution. tonotopy is treated here as a MATTER OF DEGREE with a measured precision, not as present or absent - the paper's second conclusion is that this map is less precise than others in the same species Quotation withheld because this record is not cleared for redistribution. the map is operationalised as a multiple regression of characteristic frequency on normalised 3D position, reported as r-squared - the same operationalisation Muniak2014 used for mouse DCN Quotation withheld because this record is not cleared for redistribution. characteristic frequency is defined as the frequency with the lowest response threshold, and threshold itself as two SDs above spontaneous rate Quotation withheld because this record is not cleared for redistribution. onset versus sustained are response patterns classified from post-stimulus histograms at 20 dB above threshold, and the paper finds they are spatially interspersed rather than segregated Quotation withheld because this record is not cleared for redistribution. method: in vivo single- and multi-unit recording with electrolytic lesion marking, 3D Nissl reconstruction, normalisation to a standardised nucleus, then PCA and multiple regression of CF on position 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: tonotopy in an inhibitory long-range projection nucleus a second reading DIFFERING from the recorded value. Recorded: tonotopic organization of an inhibitory auditory brainstem nucleus — 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. replicates → Coleman et al., 2011 (previous characterization of the chicken SON) Quotation withheld because this record is not cleared for redistribution. builds-on → Moiseff and Konishi, 1983; Lachica et al., 1994; Tabor et al., 2011 Quotation withheld because this record is not cleared for redistribution. extends → Rubel and Parks, 1975 (tonotopy of other chicken brainstem auditory nuclei) Quotation withheld because this record is not cleared for redistribution. extends → the tonotopic organisation established for excitatory nuclei of the chicken auditory brainstem Quotation withheld because this record is not cleared for redistribution. confirms → earlier suggestions from owl and chicken that frequency is systematically represented in the SON Quotation withheld because this record is not cleared for redistribution. supports → the group's own earlier characterisation of SON response types and their proportions Quotation withheld because this record is not cleared for redistribution. contradicts → the expectation that physiologically distinct SON cell types occupy distinct territories Quotation withheld because this record is not cleared for redistribution. characteristic frequency —operationalised-as→ response threshold characteristic frequency response threshold strength of the relation: 1 · note: CF is DEFINED through threshold, and threshold is itself defined as two SDs above spontaneous rate - a two-step operational chain Quotation withheld because this record is not cleared for redistribution. onset —contrasted-with→ sustained onset sustained strength of the relation: 1 · note: and they are spatially interspersed rather than segregated, which is the paper's third conclusion Quotation withheld because this record is not cleared for redistribution. regression regression Quotation withheld because this record is not cleared for redistribution. principal component analysis principal component analysis Quotation withheld because this record is not cleared for redistribution. eigen-decomposition of the covariance matrix eigen-decomposition of the covariance matrix Quotation withheld because this record is not cleared for redistribution. t-test t-test Quotation withheld because this record is not cleared for redistribution. coefficient of determination coefficient of determination Quotation withheld because this record is not cleared for redistribution. standard deviation standard deviation Quotation withheld because this record is not cleared for redistribution. multiple regression analysis of CF on 3D position multiple regression analysis of CF on 3D position Quotation withheld because this record is not cleared for redistribution. principal component analysis principal component analysis Quotation withheld because this record is not cleared for redistribution. coefficient of determination coefficient of determination Quotation withheld because this record is not cleared for redistribution. error magnitude between predicted and measured CF error magnitude between predicted and measured CF Quotation withheld because this record is not cleared for redistribution. In most reconstructed nuclei characteristic frequency ran along the dorsal-ventral axis, low frequencies dorsally and high frequencies ventrally. Quotation withheld because this record is not cleared for redistribution. A linear rather than logarithmic function best described the frequency map in the large majority of individual cases. Quotation withheld because this record is not cleared for redistribution. The first principal component of the combined dataset accounted for 72% of the variance. Quotation withheld because this record is not cleared for redistribution. Large prediction errors and a low r2 suggest the SON frequency map is less precise than the tonotopy of other auditory brainstem nuclei in the same species. Quotation withheld because this record is not cleared for redistribution. Within isofrequency slices the residual scatter in CF was not related to medial-lateral or anterior-posterior position, so the imprecision is not confined to a particular subregion. Quotation withheld because this record is not cleared for redistribution. Neurons with onset and with sustained response patterns are intermingled throughout the nucleus rather than occupying separate territories. Quotation withheld because this record is not cleared for redistribution. Each of the two response types forms a complete tonotopic map, and the two maps are arranged similarly. Quotation withheld because this record is not cleared for redistribution. Sustained responders were more sharply tuned than onset responders. Quotation withheld because this record is not cleared for redistribution. characteristic frequency runs linearly from low dorsally to high ventrally in the SON Quotation withheld because this record is not cleared for redistribution. a linear relation fitted 15 of 19 individual nuclei, a log relation two, and two showed none Quotation withheld because this record is not cleared for redistribution. in the best individual case, 85% of CF variation was accounted for by position Quotation withheld because this record is not cleared for redistribution. across the standardised nucleus the fit was significant but not strong, r-squared 0.62 Quotation withheld because this record is not cleared for redistribution. the first principal component accounted for 72% of the variance Quotation withheld because this record is not cleared for redistribution. the map is less precise than the tonotopy of other auditory brainstem nuclei in the same species Quotation withheld because this record is not cleared for redistribution. within isofrequency slices, position carried no information about CF Quotation withheld because this record is not cleared for redistribution. onset and sustained responders both show a dorsal-low to ventral-high tonotopic organisation Quotation withheld because this record is not cleared for redistribution. 73% of isolated neurons were sustained, 25% onset and 2% suppressed Quotation withheld because this record is not cleared for redistribution. sustained responders are more frequency-selective than onset responders 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. response threshold response threshold Quotation withheld because this record is not cleared for redistribution. center of mass of recording sites center of mass of recording sites Quotation withheld because this record is not cleared for redistribution. tonotopic map tonotopic map Quotation withheld because this record is not cleared for redistribution. characteristic frequency characteristic frequency Quotation withheld because this record is not cleared for redistribution. response threshold response threshold Quotation withheld because this record is not cleared for redistribution. Q10 Q10 Quotation withheld because this record is not cleared for redistribution. Q10 Q10 Quotation withheld because this record is not cleared for redistribution. response threshold response threshold Quotation withheld because this record is not cleared for redistribution. body temperature body temperature Quotation withheld because this record is not cleared for redistribution. characteristic frequency characteristic frequency Quotation withheld because this record is not cleared for redistribution. normalized distances normalized distances Quotation withheld because this record is not cleared for redistribution. response patterns response patterns Quotation withheld because this record is not cleared for redistribution. normalized medial-lateral coordinate normalized medial-lateral coordinate Quotation withheld because this record is not cleared for redistribution. normalized dorsal-ventral coordinate axis normalized dorsal-ventral coordinate axis Quotation withheld because this record is not cleared for redistribution. normalized anterior-posterior coordinate normalized anterior-posterior coordinate Quotation withheld because this record is not cleared for redistribution. 310 multiple-units and 349 isolated neurons; CF 90 to 5,000 Hz — size of the recorded sample and the range of characteristic frequencies encountered Quotation withheld because this record is not cleared for redistribution. Seventy-three percent — proportion of isolated neurons with sustained (primary-like or pauser-like) responses, against 25% onset Quotation withheld because this record is not cleared for redistribution. 72% — variance in the combined location-plus-CF dataset captured by the first principal component Quotation withheld because this record is not cleared for redistribution. r 2 = 0.62, P < 0.01 — strength of the linear relation between CF and 3D location in the standardized nucleus Quotation withheld because this record is not cleared for redistribution. 85% — variation in CF explained by recording location in the best single case Quotation withheld because this record is not cleared for redistribution. 60% — variability in the standardized dataset accounted for by the multilinear regression, taken as the precision of the map Quotation withheld because this record is not cleared for redistribution. CF = 0.01 + 0.32 / + 4.09 v − 0.55 a — the fitted predictive equation giving characteristic frequency in kHz from normalized 3D coordinates Quotation withheld because this record is not cleared for redistribution. r-squared 0.62 for the standardised nucleus — how much CF variation position accounts for - directly comparable with Muniak2014's 0.87 for mouse DCN, since the operationalisation is the same Quotation withheld because this record is not cleared for redistribution. 85% — the best individual case Quotation withheld because this record is not cleared for redistribution. 72% on the first principal component — how one-dimensional the arrangement is - Muniak2014 reported 75% by the same method Quotation withheld because this record is not cleared for redistribution. 310 multiple-units and 349 isolated neurons, CF from 90 to 5,000 Hz — the sample and its frequency span Quotation withheld because this record is not cleared for redistribution. 73% sustained, 25% onset, 2% suppressed — response type proportions Quotation withheld because this record is not cleared for redistribution. 12 to 75 sites per animal — sampling density Quotation withheld because this record is not cleared for redistribution. two SDs above spontaneous rate — the stipulated threshold criterion Quotation withheld because this record is not cleared for redistribution. two-thirds of peak voltage — the stipulated cut for counting multi-unit events Quotation withheld because this record is not cleared for redistribution.

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