OSCR

Noise-invariant representations of sound emerge along the canonical cortical hierarchy.

Code ↔ Paper

9 matches between paragraphs of the paper and lines of its authors' code, computed by the harvester (lexical-v1). Click a colored paragraph or line to see its counterpart.

The 9 matches · 2 of them tie a paragraph to a whole file, not to given lines: weak matches, whose lines are not tinted
  1. [1] § Materials and methods › Data analysis › Pairwise correlations. ↔ Single-Cell-Summaries/extras/+noiseCorr/getNoiseCorr.m, lines 1–56 · score 0.67 · Pearson correlation, simultaneously recorded neurons, Noise correlations, coefficient, subtracting, spike
  2. [2] § Results › L5 IT neurons maintain neural manifold geometry ↔ Manifold-Analysis/plotGeometryOutput.m, lines 21–152 · score 0.66 · Wilcoxon signed rank, manifold capacity, manifold dimensionality, radius, geometry, BN
  3. [3] § Results › BN reduces shared neural variability across spatial scales in IT but not ET neural responses ↔ Single-Cell-Summaries/+plotting/corrDistDivide.m, the whole file · a weak match · score 0.64 · Wilcoxon rank sum, short distances, long distances, correlations
  4. [4] § Results › BN attenuates single-neuron responses in L2/3 ↔ Single-Cell-Summaries/supplementalMotion.m, lines 279–332 · score 0.62 · BF centered curves, way ANOVA, intensity tuning, interaction, dimension, stimulus
  5. [5] § Results › BN attenuates single-neuron responses in L2/3 ↔ Single-Cell-Summaries/extras/matchedComparison_old.m, lines 119–157 · score 0.62 · BF centered curves, deconvolved spikes, intensity tuning curves, stimulus
  6. [6] § Materials and methods › Data analysis › Image processing. ↔ Masked-Noise-Decoding/+utils/is_responsive.m, lines 170–244 · score 0.57 · deconvolved spike, evoked responses, baseline, traces, subtraction, window
  7. [7] § Materials and methods › Data analysis › Mutual information. ↔ Single-Cell-Summaries/+plotting/respHist.m, the whole file · a weak match · score 0.53 · mutual information, bin width, histogramming, probability, Stimulus
  8. [8] § Materials and methods › Data analysis › Tuning curves. ↔ Masked-Noise-Decoding/+utils/is_responsive.m, lines 1–73 · score 0.53 · sound onset, response window, FRA, activity, tuning, matrices
  9. [9] § Results › L5 IT neurons maintain neural manifold geometry ↔ Manifold-Analysis/plotManifoldMetrics.m, lines 128–174 · score 0.50 · Wilcoxon signed rank, manifold metrics, BN

Paper

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The authors' code

MATLAB · 308 lines · 14 KB · MIT · 2 matches

  1. function [resp, resp_tuning] = is_responsive(d, which_method, resp_window, spont_window)
  2. % Rebecca Krall 1/25/22
  3. %
  4. % Takes an experimental struct generated by extract_experimental_data
  5. % and determines if each cell is responsive using different methods to
  6. % easily compare heuristic- and statistics-based methods. This assumes
  7. % that d contains information about the onset of stimulus and the frame
  8. % rate
  9. if nargin < 4
  10. spont_window = 1:14;
  11. end
  12. if nargin < 3
  13. resp_window = 15:29;
  14. end
  15. switch which_method
  16. case 1
  17. % Method 1: Use determine_responsive This normalizes the
  18. % average trace by zscoring to a spontaneous window then
  19. % looking for three consecutive points where the zscore is
  20. % above a threshold
  21. threshold = 3;
  22. for i=1:size(d.spike_traces,1)
  23. resp(i) = determine_responsive(nanmean(d.spike_traces(i,:,:),3),resp_window,spont_window,threshold);
  24. end
  25. resp_tuning = nan(size(d.spike_zscores,1), length(unique(d.inner_index)));
  26. case 2
  27. % Method 2: Use determine responsive based on stimuli This is
  28. % the same as above, but instead of looking at the overall
  29. % response for the neuron it separates it by inner_index
  30. % (frequency for FRA parameter files, intensity for RLF). This
  31. % also returns 'tuning' of the neuron which is a n x m boolean
  32. % matrix where n = number of cells and m = number of stimuli
  33. % and A(n,m) = 1 indicates that cell was responsive to that
  34. % stimuli.
  35. threshold = 3;
  36. for i = 1:size(d.spike_traces, 1)
  37. boo = false;
  38. for j = unique(d.inner_index)
  39. resp_tuning(i,j) = utils.determine_responsive(nanmean(d.spike_traces(i,:,d.inner_index == j),3), resp_window, spont_window, threshold);
  40. boo = boo | resp_tuning(i,j);
  41. end
  42. resp(i) = boo;
  43. end
  44. case 3
  45. % Method 3: Caroline Runyan - Biorxiv "Sound responsiveness was
  46. % calculated based on the mean z-scored deconvolved activity of
  47. % each neuron aligned on sound onset. For each neuron, we
  48. % calculated the difference between the mean activity during
  49. % the sound presentation at a certain location (either 1 or 2
  50. % seconds) and the mean activity in the 240 milliseconds prior
  51. % to sound onset. We calculated sound responsiveness separately
  52. % for each sound location as the mean difference in activity
  53. % between these two windows for each neuron. We then compared
  54. % the observed sound responsiveness of each neuron for each
  55. % sound location to a shuffled distribution. Each cell’s
  56. % activity was shifted randomly by at least 5 seconds in time
  57. % relative to sound location time series, and for 1000
  58. % time-shifted iterations, sound responsiveness to each sound
  59. % location was recalculated. Each sound responsive neuron had a
  60. % positive sound responsiveness value for at least one location
  61. % that was greater than the 95th percentile of that cell’s
  62. % shuffled distribution for that location. All other neurons
  63. % were not considered sound responsive."
  64. % The following code uses Caroline's paper as inspiration but
  65. % not direct methods. Similar to Caroline, a change in z-score
  66. % is calculated between a response window and a window prior to
  67. % sound onset. Then this value is compared to 1000 samples of
  68. % randomly shifted responses (5 - 20 second shifts in the
  69. % trace) to misalign the stimulus and response windows. A cell
  70. % is responsive if the change in zscore for at least one
  71. % frequency is greater than the 98th percentile of the randomly
  72. % shifted dataset
  73. %resp_window = d.stim_onset_frame: d.stim_onset_frame + d.fr/2;
  74. %spont_window = 7 : d.stim_onset_frame-1;
  75. for i = 1:size(d.spike_zscores,1)
  76. trace = squeeze(d.spike_zscores(i,:,:));
  77. delta_spikes = squeeze(nanmean(trace(resp_window,:)) - nanmean(trace(spont_window, :)));
  78. flat_trace = trace(:);
  79. for j = 1:1000
  80. shift = randi([30 600]);
  81. if mod(j,2)
  82. new_trace = [nan(shift,1);flat_trace(1:end-shift)];
  83. else
  84. new_trace = [flat_trace(shift+1:end); nan(shift,1)];
  85. end
  86. new_trace = reshape(new_trace, size(trace));
  87. delta_shift(:,j) = squeeze(nanmean(new_trace(resp_window,:)) - nanmean(new_trace(spont_window, :)));
  88. end
  89. boo = false;
  90. for k = unique(d.inner_index)
  91. zspike = nanmean(delta_spikes(d.inner_index == k));
  92. shifts = nanmean(delta_shift(d.inner_index == k, :));
  93. compare = prctile(shifts, 98);
  94. boo = boo | (compare < zspike);
  95. if compare < zspike
  96. resp_tuning(i, k) = 1;
  97. else
  98. resp_tuning(i,k) = 0;
  99. end
  100. end
  101. resp(i) = boo;
  102. end
  103. case 4
  104. % Method 4 - Walker King 'Complexity of frequency receptive
  105. % fields predicts tonotopic variability across species' A
  106. % two-way ANOVA, with tone frequency and sound level as
  107. % predictors, was used to determine if the evoked activity was
  108. % significantly modulated by sound frequency or intensity (α =
  109. % 0.05). Neurons showing a significant main effect of frequency
  110. % or frequency/level interaction were defined as ‘frequency
  111. % sensitive’, and only these neurons were included in further
  112. % analyses.
  113. % rows = frequencies
  114. % columns = sound intensities
  115. count = 1;
  116. for i = 1:size(d.spike_zscores,1)
  117. for j = unique(d.inner_index)
  118. for k = unique(d.outer_index)
  119. choose = d.outer_index == k & d.inner_index == j;
  120. spont = squeeze(nanmean(d.spike_zscores(i, spont_window, choose),2));
  121. response = squeeze(nanmean(d.spike_zscores(i, resp_window, choose),2));
  122. delta(:,j) = response - spont;
  123. end
  124. end
  125. an = anova2(delta, 5, 'off');
  126. if an(1) < 0.05 | an(3) < 0.05
  127. resp(i) = 1;
  128. else
  129. resp(i) = 0;
  130. end
  131. end
  132. resp_tuning = nan(size(d.spike_zscores,1), length(unique(d.inner_index)));
  133. case 5
  134. % Simple t-test between the mean of the response in the
  135. % spontaneous window and the response window
  136. for i = 1:size(d.spike_zscores,1)
  137. smoothed = smooth(mean(d.spike_traces(i,:,:),3));
  138. [~,loc_temp] = max(smoothed(resp_window,:));
  139. peak_latency = resp_window(1)+loc_temp-1;
  140. for j = unique(d.inner_index)
  141. choose = d.inner_index == j;
  142. current_trace = squeeze(d.spike_zscores(i,:,choose));
  143. tuning_window = [peak_latency-1:peak_latency+1];
  144. peak = mean(current_trace(tuning_window, :));
  145. mean_response = mean(current_trace(resp_window, :));
  146. baseline = mean(current_trace(spont_window, :));
  147. %thresh = mode(peak > 1);
  148. [h,~] = ttest(baseline, mean_response, 'Alpha', .01, 'Tail', 'left');
  149. resp_tuning(i,j) = h; %& thresh;
  150. end
  151. end
  152. resp = (sum(resp_tuning,2) > 0)';
  153. case 6
  154. for i = 1:size(d.spike_zscores,1)
  155. z = squeeze(d.spike_zscores(i,:,:));
  156. tr = squeeze(d.spike_traces(i,:,:));
  157. [delta_spikes, lat] = determine_baseline_subtracted_peak(tr, z);
  158. flat_tr = tr(:);
  159. flat_z = z(:);
  160. for j = 1:1000
  161. shift = randi([30 600]);
  162. if mod(j,2)
  163. new_tr = [nan(shift,1);flat_tr(1:end-shift)];
  164. new_z = [nan(shift,1);flat_z(1:end-shift)];
  165. else
  166. new_tr = [flat_tr(shift+1:end); nan(shift,1)];
  167. new_z = [flat_z(shift+1:end); nan(shift,1)];
  168. end
  169. shift_tr = reshape(new_tr, size(tr));
  170. shift_z = reshape(new_z, size(z));
  171. [delta_shift(:,j), ~] = determine_baseline_subtracted_peak(shift_tr, shift_z, lat);
  172. end
  173. boo = false;
  174. for k = unique(d.inner_index)
  175. zspike = nanmean(delta_spikes(d.inner_index == k));
  176. shifts = nanmean(delta_shift(d.inner_index == k,:));
  177. compare = prctile(shifts, 98);
  178. boo = boo | (compare < zspike);
  179. if compare < zspike
  180. resp_tuning(i, k) = 1;
  181. else
  182. resp_tuning(i,k) = 0;
  183. end
  184. end
  185. resp(i) = boo;
  186. end
  187. case 7
  188. % Method 7 - Kato et al, 2017 (Network-Level Control of
  189. % Frequency Tuning in Auditory Cortex) This method is adapted
  190. % from Kato's paper, which uses dF/F, to be applicable to
  191. % deconvolved spikes. Sound-evoked responses are based on two
  192. % criteria:
  193. % (1) At least 50% of trials at a cell's best frequency must
  194. % have 3 consecutive frames exceed 0.5 SD from baseline
  195. % (2) Average tone responses (across all stimuli) must have 3
  196. % consecutive frames exceed 3 SD from baseline
  197. respThres = 0.5; % min no. of SDs above baseline during resp window
  198. respThres_avr = 3; % response threshold for Criteria #1
  199. frAboveThres = 3; % min no. of consecutive frames above respThres
  200. propOfTrials = 0.4; % proportion of trials where response is above threshold
  201. nTrials = size(d.spike_zscores,3); % no. of trials
  202. if isempty(d.inner_sequence) && sum(d.inner_index)==numel(d.inner_index)
  203. inner_sequence = 1;
  204. else
  205. inner_sequence = d.inner_sequence;
  206. end
  207. % Extract BF based on avr z-scored response across resp_window
  208. respWindTuning = zeros(size(d.spike_zscores,1),numel(d.inner_sequence));
  209. for stim = 1:numel(d.inner_sequence)
  210. tStim = find(d.inner_index==stim);
  211. dums = d.spike_zscores(:,resp_window,tStim);
  212. avrTrialResp = mean(dums,3,'omitnan'); % [cells * frames in resp_wind]
  213. respWindTuning(:,stim) = mean(avrTrialResp,2,'omitnan');
  214. end
  215. if inner_sequence==1 & sum(d.inner_index)==numel(d.inner_index)
  216. BF = ones(1,size(d.spike_zscores,1)); % essentially making all cells have same BF (work around)
  217. else
  218. maxResp = max(respWindTuning,[],2,'omitnan');
  219. for neuron = 1:size(respWindTuning,1)
  220. BF(neuron) = find(respWindTuning(neuron,:)==maxResp(neuron));
  221. end
  222. end
  223. % Determine if cell is responsive based on response to BF
  224. % NOTE: only 1D stimuli are applicable for this method
  225. for neuron = 1:size(d.spike_zscores,1)
  226. spikes = squeeze(d.spike_traces(neuron,:,:));
  227. psth_avr = mean(d.spike_traces(neuron,:,:),3,'omitnan');
  228. psth_zscored_avr = smooth((psth_avr-nanmean(psth_avr(spont_window)))/nanstd(psth_avr(spont_window)));
  229. BF_trials = find(d.inner_index==BF(neuron));
  230. % Grab traces
  231. dums = spikes(spont_window,BF_trials);
  232. psth_BF = squeeze(d.spike_traces(neuron,:,BF_trials)); % [frames * trials]
  233. psth_zscored_BF = (psth_BF-nanmean(psth_avr(spont_window))) ./ nanstd(dums(:));
  234. for trial = 1:numel(BF_trials)
  235. psth_smooth(:,trial) = smooth(psth_zscored_BF(:,trial));
  236. end
  237. psth_zscored_smoothBF = psth_smooth;
  238. % Criteria #1
  239. idx_criteriaOne = psth_zscored_smoothBF(resp_window,:) >= respThres;
  240. [~,c] = find(movsum(idx_criteriaOne,frAboveThres) >= frAboveThres);
  241. trials_idx = unique(c);
  242. % Criteria #2
  243. idx_criteriaTwo = psth_zscored_avr(resp_window) >= respThres_avr;
  244. frForCriteriaTwo = find(movsum(idx_criteriaTwo,frAboveThres) >= frAboveThres);
  245. criteriaOne = (numel(trials_idx) / (numel(BF_trials))) >= propOfTrials;
  246. criteriaTwo = ~isempty(frForCriteriaTwo);
  247. if criteriaOne & criteriaTwo
  248. resp(neuron) = 1;
  249. else
  250. resp(neuron) = 0;
  251. end
  252. resp_tuning=[]; % will change this later
  253. end
  254. %%
  255. end % switch/case
  256. end % function

is_responsive.m at commit 520d1f2, under MIT · at the source

Overview

Authors: Tomas Suarez Omedas1,2,3, Ross S. Williamson2,3,4,5,6
  1. Neuroscience Institute, Carnegie Mellon University, Pittsburgh, Pennsylvania, United States of America
  2. Pittsburgh Hearing Research Center, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America
  3. Center for the Neural Basis of Cognition, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America
  4. Department of Otolaryngology, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America
  5. Department of Neurobiology, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America
  6. Department of Bioengineering, University of Pittsburgh, Pittsburgh, Pennsylvania, United States of America
Institutions: University of Pittsburgh (United States); Center for the Neural Basis of Cognition (United States); Carnegie Mellon University (United States)
Journal: PLoS biology, volume 24, issue 7, article e3003915
Dates: received 19 December 2025; accepted 8 July 2026; published online 20 July 2026
Type: Research article · Language: English
License: CC BY
Identifiers: DOI 10.1371/journal.pbio.3003915 · PMID 42475408 · PMCID PMC13399537 · OpenAlex W7169786066
Open access: gold, a free copy (OpenAlex)
Status: code verified
Categories: mouse (organism)
Methods: Connectivity, Statistics, Smoothing, state filtering, decompositions, Machine learning, Preprocessing, fMRI & imaging, Single-unit activity, calcium imaging
MeSH: Auditory Cortex*, Auditory Perception*, Acoustic Stimulation, Animals, Auditory Pathways, Male, Mice, Neurons, Noise, Sound (* major topic)
Journal subjects: Biology and Life Sciences, Neuroscience, Neuronal Tuning, Computational Biology, Computational Neuroscience, Single Neuron Function, Cell Biology, Cellular Types, Animal Cells, Neurons, Cellular Neuroscience, Physical sciences, Mathematics, Geometry, Non-Euclidean geometry, Cognitive Science, Cognitive Psychology, Perception, Sensory Perception, Psychology, Social Sciences, Cognition, Engineering and Technology, Signal Processing, White Noise, Behavior
Topic: Neural dynamics and brain function (Cognitive Neuroscience, Neuroscience), according to OpenAlex
Funding: National Institute on Deafness and Other Communication Disorders (R21DC018327, R01DC020459); Esther A. and Joseph Klingenstein Fund (EAJK Fund) (Klingenstein-Simons Fellowship Award in Neuroscience)
Citations: not cited yet (Europe PMC); 115 references in the paper

Abstract

Neurons in the auditory system must represent behaviorally relevant sounds in the presence of background noise (BN) to support noise-invariant perception and behavior. Although the primary auditory cortex (ACtx) has been implicated in constructing noise-invariant representations, it remains unclear which excitatory subpopulations within ACtx carry out this transformation from noise-dependent to noise-invariant coding. To address this, we presented pure tones with and without continuous BN to head-fixed mice and used two-photon calcium imaging to record sound-evoked activity from three major excitatory subpopulations in ACtx: layer (L)2/3 intratelencephalic (IT) neurons, L5 IT neurons, and L5 extratelencephalic (ET) neurons. L2/3 IT neurons exhibited strong noise dependence at the level of single-neuron responses, pairwise interactions, and population representations. In contrast, deep-layer pathways showed greater noise invariance, with L5 IT neurons preserving stable representations most consistently and L5 ET neurons exhibiting more limited invariance at the population level. These findings reveal a functional division of labor in ACtx, in which superficial neurons remain noise-dependent and deep-layer broadcast pathways, particularly L5 IT, preferentially carry noise-invariant representations, suggesting that excitatory subpopulations contribute differentially to the construction and propagation of noise-invariant codes.

Reproduced under the paper's license (CC BY), from the paper cited above.

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Its files are read in the Code ↔ Paper reader above, with 9 matches between paragraphs and lines of code.

WilliamsonLab-Pitt/BackgroundNoise-Characterization

License: MIT
State: the link answers, verified on 27 September 2026
Evidence: files inventoried
Commit: 520d1f2cb42eeda3c726caa693fe0d57b209129d, 27 June 2026
Languages: MATLAB (290)
Size: 370 files, 290 scripts
Software Heritage: not archived
Found in: “Data Availability”
Holds: README, license file, tests
Not found: CITATION.cff, environment file, continuous integration, documentation
Availability: 1 check, the latest on 27 September 2026: the link answers
  • 27 September 2026: the link answers
292 files

RichieHakim/ROICaT

License: GPL-3.0
State: the link answers, verified on 27 September 2026
Evidence: files inventoried
Commit: 487bb6f3c0b9021f10d8fd7581921949d1674c5b, 26 September 2026
Languages: Python (37), Jupyter (9), Shell (2)
Size: 91 files, 48 scripts
Software Heritage: not archived
Found in: the text, “Image processing.”
Holds: README, license file, environment (pyproject.toml, docs/requirements.txt), tests, continuous integration, documentation, 9 notebooks
Not found: CITATION.cff
Tools: NumPy (33 files), PyTorch (22 files), Matplotlib (19 files), SciPy (17 files), Pillow (10 files), scikit-learn (10 files), pandas (9 files), OpenCV (4 files), Numba (2 files), UMAP (2 files), h5py (1 file), scikit-image (1 file), seaborn (1 file)
Availability: 1 check, the latest on 27 September 2026: the link answers
  • 27 September 2026: the link answers
50 files

The paper's code and data availability statement is in the Data section.

Tracing map

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What the map holds:

  • 2 repositories of the authors' code, each at its verified commit, with its license and how the link was found in the paper;
  • 338 scripts, each with its path and the digest of its content;
  • 9 matches between paragraphs of the paper and lines of the code (method lexical-v1);
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Its JSON (tracing-map.json) is deposited on Zenodo with its DOI once the map is validated.

Data

Datasets cited

Data Availability

The data generated in this study have been deposited in an open-access Zenodo repository and can be accessed here: https://doi.org/10.5281/zenodo.21111739. Custom scripts used to analyze the data have been deposited to the same Zenodo repository, as well as in a GitHub repository, which can be found here: https://github.com/WilliamsonLab-Pitt/BackgroundNoise-Characterization.

Reproduced under the paper's license (CC BY), from the paper cited above.

Versions

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Version 1, 27 September 2026: the first record

Recorded: type, language, journal, volume, issue, pages, dates, 2 authors, 10 MeSH terms, 2 funders, 112 references.

Cite

This paper

Suarez Omedas, T., & Williamson, R. S. (2026). Noise-invariant representations of sound emerge along the canonical cortical hierarchy. PLoS biology, 24(7), e3003915. https://doi.org/10.1371/journal.pbio.3003915

BibTeX

@article{suarezomedas2026noise,
author = {Suarez Omedas, Tomas and Williamson, Ross S.},
title = {{Noise-invariant representations of sound emerge along the canonical cortical hierarchy}},
journal = {PLoS biology},
year = {2026},
month = jul,
volume = {24},
number = {7},
pages = {e3003915},
publisher = {PLOS},
issn = {1544-9173},
doi = {10.1371/journal.pbio.3003915},
url = {https://doi.org/10.1371/journal.pbio.3003915},
pmid = {42475408},
pmcid = {PMC13399537}
}

RIS

TY - JOUR
AU - Suarez Omedas, Tomas
AU - Williamson, Ross S.
TI - Noise-invariant representations of sound emerge along the canonical cortical hierarchy
T2 - PLoS biology
J2 - PLoS Biol
PY - 2026
DA - 2026/07/20
VL - 24
IS - 7
SP - e3003915
SN - 1544-9173
PB - PLOS
DO - 10.1371/journal.pbio.3003915
UR - https://doi.org/10.1371/journal.pbio.3003915
LA - en
ER -

CSL-JSON

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"volume": "24",
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"page": "e3003915",
"DOI": "10.1371/journal.pbio.3003915",
"PMID": "42475408",
"PMCID": "PMC13399537",
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Convergence-divergence circuits for multimodal integration of innate and learned opponent valences.
Journal: Frontiers in systems neuroscience
In common: UMAP, Numba, scikit-image, 9 other tools, 2 references
[7] doi:10.1038/s41593-026-02232-0 [code]
Entorhinal cortex represents task-relevant remote locations independently of CA1.
Journal: Nature neuroscience
In common: Optimization Toolbox, Numba, OpenCV, 11 other tools, mouse
[8] doi:10.1016/j.patter.2026.101590 [code]
Density-based longitudinal neuron tracking in high-density electrophysiological recordings.
Journal: Patterns (New York, N.Y.)
In common: Violinplot-Matlab, Optimization Toolbox, UMAP, 10 other tools
[9] doi:10.1038/s41467-026-76581-6 [code]
Thalamocortical bursts encode reward contingencies and drive associative learning.
Journal: Nature communications
In common: Violinplot-Matlab, Curve Fitting Toolbox, OpenCV, 8 other tools, mouse, 1 reference
[10] doi:10.1016/j.isci.2026.116825 [code]
Social hierarchy shapes behavioral and transcriptional responses to chronic stress and ketamine in male mice.
Journal: iScience
In common: UMAP, Numba, OpenCV, 10 other tools, mouse

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