Distributed coding of choice, action and engagement across the mouse brain — TEI-tagged view

Steinmetz, Zatka-Haas, Carandini, Harris · 2019 · neuroscience · doi:10.1038/s41586-019-1787-x · full text at the source · TEI xml · tagged edition · index · concepts · authors
Where this record's text came from. faceted 2026-08 by reading the locally held text (xml); every evidence string verified as an exact substring of that text before merging. Claim type is existence: the headline contribution is a map of which neurons encoding which variables exist where, not an account of why. The paper also introduces two methodological instruments (reduced-rank kernel regression, combined-conditions choice probability, jPECC), which is captured in claim relations rather than by changing claim_type. Note the small verbatim typo in limitation 2 ("whether visual neurons would be found these “motor” areas") - it is reproduced exactly as printed, not corrected. 2026-08-13 and 2026-08-15 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.
13 quotations on this page contain a reference number where the paper printed a superscript. This journal style prints citations as raised numerals — parameters12,13 — and the extraction flattened them into the line, so the sentence reads parameters 12 , 13 . The digits are citation markers and not part of what the author wrote. The quotes keep them for the same reason they keep every other extraction fault: a quote is reproduced exactly or it proves nothing.
Other fields define these differently the same word, defined another way in another discipline
principal component analysis 5 fields · engagement 4 fields · logistic regression 3 fields · cross-validation 3 fields · bootstrap 2 fields · generalized linear model 2 fields · and 1 more →
Tags on this paper every tag the record carries, whether or not the words occur in the text below — a number is how many papers in the corpus carry that tag; a tag with no number is unique to this paper
choice7 action3 vision engagement10 kernel2 go/nogo2 passive7 Miss midbrain5 forebrain2 neocortex7 VISp ZI timing2 variance54 contralateral4 ipsilateral6 correlate4 electrophysiology110

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content-tags & facets: phenomenon · claim-type · cross-domain-transfer · mechanism · claim-relation · replication-status · limitation — grounded in the full text
phenomenonbrain-wide distribution of neural coding for vision, action, choice and task engagement— “To determine the distribution of neurons encoding vision, choice, action, and behavioral engagement, we recorded neural activity across the brain while mice performed a task that allows distinguishing these processes ( Fig. 1a-c ).”
claim-typeexistence— “By contrast, neurons encoding visual stimuli and upcoming choices occupied restricted regions in neocortex, basal ganglia, and midbrain.”
method — evidence— “We used Neuropixels probes 1 , 2 to record from ~30,000 neurons in 42 brain regions of mice performing a visual discrimination task 3 .”
object of study — evidence— “Nevertheless, neuronal signals that correlate with action do not necessarily correlate with action selection. To carry such choice-related signals, a brain region must contain neurons whose firing selectively predicts the chosen action before the action occurs”
research question — evidence— “Nevertheless, neuronal signals that correlate with action do not necessarily correlate with action selection. To carry such choice-related signals, a brain region must contain neurons whose firing selectively predicts the chosen action before the action occurs”
contribution — evidence— “Vision, choice, action, and behavioral engagement arise from neuronal activity that may be distributed across brain regions. Here we delineate the spatial distribution of neurons underlying these processes. We used Neuropixels probes 1 , 2 to record from ~30,000 neurons in 42 brain regions of mice performing a visual discrimination task 3 .…
transferfrom the joint peristimulus time histogram, from paired-recording analysis: JOINT PERI-EVENT CANONICAL CORRELATION (jPECC) ANALYSIS, A NOVEL EXTENSION OF THE “joint Peristimulus Time Histogram” METHOD MODIFIED TO DETECT CORRELATIONS IN A “communication subspace” BETWEEN TWO POPULATIONS— “Choice signals emerged with similar timing across choice-encoding regions ( Figure 4c-d ). To examine timing, we performed two analyses. First, we trained a decoder to predict the subject’s choice from recorded population activity, after first subtracting the prediction of population activity from the Vision and Action kernels (to yield a…transfers[0]
transferfrom single-neuron choice probability analysis: a version of CHOICE PROBABILITY ANALYSIS FOR TASKS WITH MANY STIMULUS CONDITIONS, CALLED COMBINED-CONDITIONS CHOICE PROBABILITY, which QUANTIFIES THE PROBABILITY THAT A NEURON'S SPIKE COUNT WILL BE GREATER ON TRIALS WITH ONE CHOICE THAN TRIALS OF ANOTHER, FOR MATCHED STIMULUS CONDITIONS— “To further confirm these conclusions, we developed a version of choice probability analysis for tasks with many stimulus conditions, called “combined-conditions choice probability” ( Methods ). This statistic quantifies the probability that a neuron’s spike count will be greater on trials with one choice than trials of another, for matched…transfers[1]
mechanismA recurrent loop between forebrain and midbrain choice areas keeps their choice signals simultaneous despite their different encoding schemes— “Despite this distinct encoding, we could not distinguish the timing of choice-related signals between these regions, an observation parsimoniously explained by a recurrent loop across them ( Supplementary Fig. 4a ).”mechanisms[0]
mechanismEngagement raises subcortical baseline firing toward the threshold at which movements are triggered, so the same stimulus is more likely to release an action— “Enhanced activity in subcortical areas, by contrast, brings activity in these regions closer to the level at which actions are initiated, providing a potential mechanism for increased probability of action in engaged states ( Supplementary Fig. 4b-d ).”mechanisms[1]
mechanismArousal-linked hyperpolarization and reduced spiking in cortex sharpens the signal-to-noise ratio of sensory representations, explaining why engagement suppresses neocortical activity— “However, reduced spiking activity and hyperpolarization has been associated with increased arousal in multiple cortical areas, an effect that may improve signal-to-noise ratios of sensory representations”mechanisms[2]
mechanismthe criterion the paper sets itself before measuring anything: TO CARRY SUCH CHOICE-RELATED SIGNALS, A BRAIN REGION MUST CONTAIN NEURONS WHOSE FIRING SELECTIVELY PREDICTS THE CHOSEN ACTION BEFORE THE ACTION OCCURS— “Nevertheless, neuronal signals that correlate with action do not necessarily correlate with action selection. To carry such choice-related signals, a brain region must contain neurons whose firing selectively predicts the chosen action before the action occurs”mechanisms[3]
mechanismthe design that makes the criterion testable: THE SAME VISUAL STIMULUS COULD THEREFORE LEAD TO EITHER DIRECTION OF TURN, OR TO NO ACTION, ALLOWING US TO DISSOCIATE THE NEURAL CORRELATES OF THE VISUAL STIMULI, OF ACTION INITIATION AND OF ACTION SELECTION— “This task combined the advantages of two-alternative forced choice and Go/NoGo designs 3 , 31 . On each trial, visual stimuli of varying contrast could appear on the left or right sides, or both, or neither. Mice earned a water reward by turning a wheel with their forepaws to indicate which side had highest contrast ( Fig. 1a-c ). If neither…mechanisms[4]
mechanismthe two encodings that differ without differing in time: IN MIDBRAIN NEARLY ALL CHOICE-SELECTIVE NEURONS (53/54, 98%) PREFERRED CONTRALATERAL CHOICES, while forebrain neurons COULD PREFER EITHER CHOICE, giving A DISTINCTIVE 'BILATERAL' ENCODING OF BOTH CHOICES IN THE FOREBRAIN, VERSUS A 'UNILATERAL' ENCODING OF CONTRALATERAL CHOICES IN THE MIDBRAIN— “Although the encoding of choice emerged with indistinguishable timing in midbrain and forebrain, these regions encoded choice differently ( Fig. 4e-h ). In midbrain (MRN, SCm, SNr, and ZI) nearly all choice-selective neurons (53/54, 98%) preferred contralateral choices ( Fig. 4e , top, and Fig. 4f ). By contrast, choice-selective neurons in…mechanisms[5]
mechanismthe proposed reason engagement helps: ENHANCED ACTIVITY IN SUBCORTICAL AREAS BRINGS ACTIVITY IN THESE REGIONS CLOSER TO THE LEVEL AT WHICH ACTIONS ARE INITIATED, PROVIDING A POTENTIAL MECHANISM FOR INCREASED PROBABILITY OF ACTION IN ENGAGED STATES— “Brain-wide correlates of engagement are characterized by enhanced subcortical activity and suppressed neocortical activity prior to visual stimulus onset. Engagement-related cortical suppression might seem surprising, given that visual cortex is required for performance of this task 3 , 41 . However, reduced spiking activity and…mechanisms[6]
mechanismand what the global pre-movement activity cannot be: THESE SIGNALS MAY COMPRISE FORMS OF COROLLARY DISCHARGE, BUT CANNOT REFLECT SENSORY RE-AFFERENCE AS THEY WERE OBSERVED PRIOR TO MOVEMENT ONSET— “These signals may comprise forms of corollary discharge 39 , but cannot reflect sensory re-afference as they were observed prior to movement onset.”mechanisms[7]
builds-on Guo et al (ref. 45), headplate and recording-chamber implantation method— “The implantation method largely followed the method of Guo et al 45 with some modifications and was described previously 44 .”claim_relations[0]
builds-on Treves and Rolls (ref. 54), sparseness measure, adapted here as the Focality Index— “This measure is an adaptation of the sparseness measure of Treves and Rolls 54 , and would take the value 1 if all neurons were located in a single region, and the value 1/ N if neurons were equally probable in N regions.”claim_relations[1]
extends The "joint Peristimulus Time Histogram" method (refs 35, 36), extended here to jPECC— “Second, we validated this conclusion using joint Peri-Event Canonical Correlation (jPECC) analysis, a novel extension of the “joint Peristimulus Time Histogram” method 35 , 36 modified to detect correlations in a “communication subspace” 37 between two populations.”claim_relations[2]
builds-on The previously described two-alternative unforced choice task (ref. 3)— “The two-alternative unforced choice task design was described previously 3 .”claim_relations[3]
critiques go/nogo designs as evidence about choice-selective neurons— “This ubiquitous presence of non-selective action correlates underscores the importance of multi-alternative tasks for studying the neural correlates of behavioral choice: go/nogo tasks cannot distinguish neurons that nonspecifically fire for any action, from neurons selective for specific choices.”claim_relations[4]
extends reports of widespread action correlates in multiple species— “In contrast, neurons encoding action (requiring an Action kernel) were spread throughout all recorded regions ( Fig. 3e, g ). The distribution of these neurons encoding action was significantly broader that of neurons encoding visual stimuli ( Extended Data Fig. 4f ). The great majority of neurons encoding action did not require an additional…claim_relations[5]
qualifies the classical set of choice-related areas (frontal cortex, striatum, SNr, deep superior colliculus)— “Neurons encoding specific choices were found in a small subset of brain regions ( Fig. 4a,b ). We identified choice-selective neurons as neurons for which the Choice kernel was required to explain their activity, in the nested test described above. These neurons were rare, and were found in frontal cortex (MOs, PL, and MOp), basal ganglia…claim_relations[6]
senseA CRITERION FOR WHAT COUNTS AS EVIDENCE, STATED BEFORE ANYTHING IS MEASURED: NEURONAL SIGNALS THAT CORRELATE WITH ACTION DO NOT NECESSARILY CORRELATE WITH ACTION SELECTION - TO CARRY SUCH CHOICE-RELATED SIGNALS, A BRAIN REGION MUST CONTAIN NEURONS WHOSE FIRING SELECTIVELY PREDICTS THE CHOSEN ACTION BEFORE THE ACTION OCCURS— “Nevertheless, neuronal signals that correlate with action do not necessarily correlate with action selection. To carry such choice-related signals, a brain region must contain neurons whose firing selectively predicts the chosen action before the action occurs”senses[0]
senseAND A WHOLE CLASS OF DESIGNS CANNOT MEET IT: GO/NOGO TASKS CANNOT DISTINGUISH NEURONS THAT NONSPECIFICALLY FIRE FOR ANY ACTION, FROM NEURONS SELECTIVE FOR SPECIFIC CHOICES. Not a result being corrected - a design being retired, and every finding that used it inherits the ambiguity— “This ubiquitous presence of non-selective action correlates underscores the importance of multi-alternative tasks for studying the neural correlates of behavioral choice: go/nogo tasks cannot distinguish neurons that nonspecifically fire for any action, from neurons selective for specific choices.”senses[1]
senseTHE SAME STIMULUS AND THE SAME behavior, DIFFERENT ACTIVITY: BOTH VISUAL STIMULI AND BEHAVIORAL REPORTS WERE IDENTICAL, and EVEN THOUGH THE TWO CONDITIONS WERE MATCHED FOR VISUAL STIMULATION AND (LACK OF) ACTION, THEY WERE ACCOMPANIED IN MANY AREAS BY DIFFERENT ACTIVITY, BOTH BEFORE AND AFTER STIMULUS PRESENTATION. The response is not a property of the stimulus— “We began by comparing two conditions where both visual stimuli and behavioral reports were identical: passive visual responses measured outside the task context, and responses on Miss trials in task context ( Fig. 5a-c ). Even though the two conditions were matched for visual stimulation and (lack of) action, they were accompanied in many…senses[2]
senseand the context effect has OPPOSITE SIGNS in different structures: ACTIVITY IN VISp WAS LOWER IN THE TASK THAN IN THE PASSIVE TRIALS, WHEREAS ACTIVITY IN CP SHOWED THE OPPOSITE MODULATION - WHILE NEOCORTEX AND SENSORY THALAMUS SHOWED A NET DECREASE, OTHER REGIONS SHOWED A CONSISTENT INCREASE. A brain-wide average of this is zero— “Consistent differences were also seen in pre-stimulus firing rates: for instance, activity in VISp was lower in the task (Miss trials) than in the passive trials, whereas activity in CP showed the opposite modulation ( Fig. 5a ). While neocortex and sensory thalamus showed a net decrease in pre-stimulus activity during task context, other…senses[3]
senseTHE GLOBAL SIGNAL IS ABOUT MOVING, NOT ABOUT THE TASK: THE WIDELY DISTRIBUTED ACTIVITY SEEN FOLLOWING TRIAL ONSET WAS THEREFORE PRESENT ONLY WHEN ANIMALS MOVED, AND REGARDLESS OF THE PARTICULAR STIMULUS AND PARTICULAR ACTION— “The widely distributed activity seen following trial onset was therefore present only when animals moved, and regardless of the particular stimulus and particular action.”senses[4]
senseand the ordering of areas reverses with the side of the stimulus: WHEN THE MOUSE SUCCESSFULLY SELECTED AN IPSILATERAL STIMULUS, MOST AREAS WERE AGAIN ACTIVATED, BUT VIS AND SCs WERE NOW AMONGST THE LAST AREAS TO RESPOND, RATHER THAN THE FIRST— “When a mouse successfully selected a visual stimulus contralateral to the recorded hemisphere, activity emerged first in classical visual regions such as visual cortex (VIS) and superficial superior colliculus (SCs), and soon spread to most of the remaining recorded regions ( Fig. 2d ). When the mouse successfully selected an ipsilateral…senses[5]
senseWHAT IS GLOBAL AND WHAT IS NOT, MEASURED RATHER THAN ASSERTED: NEURONS ENCODING ACTION WERE SPREAD THROUGHOUT ALL RECORDED REGIONS, and THE GREAT MAJORITY OF NEURONS ENCODING ACTION DID NOT REQUIRE AN ADDITIONAL CHOICE KERNEL - THEY RESPONDED EQUALLY FOR MOVEMENTS IN EITHER DIRECTION— “In contrast, neurons encoding action (requiring an Action kernel) were spread throughout all recorded regions ( Fig. 3e, g ). The distribution of these neurons encoding action was significantly broader that of neurons encoding visual stimuli ( Extended Data Fig. 4f ). The great majority of neurons encoding action did not require an additional…senses[6]
senseand the negative that makes the positive mean something: CHOICE-SELECTIVE NEURONS WERE NOT FOUND IN VISp, although THIS SET OF REGIONS ENCODING CHOICE OVERLAPPED PARTIALLY WITH THE VISUAL PATHWAY— “Neurons encoding specific choices were found in a small subset of brain regions ( Fig. 4a,b ). We identified choice-selective neurons as neurons for which the Choice kernel was required to explain their activity, in the nested test described above. These neurons were rare, and were found in frontal cortex (MOs, PL, and MOp), basal ganglia…senses[7]
senseTHE SAME VARIABLE ENCODED IN TWO INCOMPATIBLE WAYS IN TWO PLACES: IN MIDBRAIN NEARLY ALL CHOICE-SELECTIVE NEURONS (53/54, 98%) PREFERRED CONTRALATERAL CHOICES, whereas in forebrain A SIZEABLE PROPORTION PREFERRED IPSILATERAL CHOICES (19/48, 40%). A `midbrain encodes choice` tag and a `forebrain encodes choice` tag are the same words for opposite arrangements— “Although the encoding of choice emerged with indistinguishable timing in midbrain and forebrain, these regions encoded choice differently ( Fig. 4e-h ). In midbrain (MRN, SCm, SNr, and ZI) nearly all choice-selective neurons (53/54, 98%) preferred contralateral choices ( Fig. 4e , top, and Fig. 4f ). By contrast, choice-selective neurons in…senses[8]
senseA NULL DOING THE ARGUMENTATIVE WORK: WE COULD NOT DISTINGUISH THE TIMING OF CHOICE-RELATED SIGNALS BETWEEN THESE REGIONS, AN OBSERVATION PARSIMONIOUSLY EXPLAINED BY A RECURRENT LOOP ACROSS THEM - the loop is the simplest account of an absence, and is labeled as such— “Despite this distinct encoding, we could not distinguish the timing of choice-related signals between these regions, an observation parsimoniously explained by a recurrent loop across them”senses[9]
senseand the null was sought two ways: the decoding time course COULD NOT BE STATISTICALLY DISTINGUISHED BETWEEN CHOICE-SELECTIVE REGIONS, and jPECC REVEALED NO LAG FOR ACTIVITY CORRELATIONS BETWEEN FRONTAL AND MIDBRAIN AREAS while it DID reveal A CONSISTENT TIME-LAG between visual and frontal cortex— “Choice signals emerged with similar timing across choice-encoding regions ( Figure 4c-d ). To examine timing, we performed two analyses. First, we trained a decoder to predict the subject’s choice from recorded population activity, after first subtracting the prediction of population activity from the Vision and Action kernels (to yield a…senses[10]
senseTHE MODEL EXPLAINS LITTLE OF WHAT IT PREDICTS WELL: THE FRACTION OF CROSS-VALIDATED VARIANCE THE KERNELS EXPLAINED WAS FREQUENTLY SMALL, EVEN FOR NEURONS WHOSE MEAN RATES THEY ACCURATELY PREDICTED - 50.2% for one example neuron and 13.6% for another— “The fraction of cross-validated variance the kernels explained was frequently small ( Fig. 3c ; Extended Data Fig. 6a ), even for neurons whose mean rates they accurately predicted (50.2% for the neuron in Fig. 3a and 13.6% for the neuron in Fig. 3b ), as expected from trial-to-trial variability and encoding of task-independent…senses[11]
senseTHE ERROR RATE OF THE CLASSIFICATION IS PRINTED: WE USED PARAMETERS WHICH GAVE FALSE-POSITIVE ERROR RATES OF 0.33% ON SHUFFLED DATA - so every count of encoding neurons in this paper has a stated denominator of error— “In determining whether a neuron passed this test, we used parameters which gave false-positive error rates of 0.33% on shuffled data ( Extended Data Fig. 5h ). As our question concerns activity predictive of upcoming movements, we applied this analysis only to pre-movement activity.”senses[12]
senseAN INDEX DEFINED BY THE DIFFERENCE BETWEEN TWO CONTEXTS, AND CHECKED AGAINST FOUR CONFOUNDS: the engagement index COULD NOT BE FULLY EXPLAINED BY VARIABILITY IN PUPIL DIAMETER, IN OVERT MOVEMENTS DETECTABLE BY VIDEO RECORDINGS, IN THE PRESENCE OF A REWARD ON THE PREVIOUS TRIAL, OR IN THE INTER-TRIAL INTERVAL— “Indeed, it was possible to predict whether an animal would respond to the stimulus on a given trial within the task context, by projecting pre-stimulus population activity onto a weight vector given by the difference of pre-stimulus activity in passive and task contexts (“engagement index”, Fig. 5e ). This engagement index differed between Go…senses[13]
senseENGAGEMENT MOVES CORTEX AND SUBCORTEX IN OPPOSITE DIRECTIONS: CORRELATES OF ENGAGEMENT ARE CHARACTERIZED BY ENHANCED SUBCORTICAL ACTIVITY AND SUPPRESSED NEOCORTICAL ACTIVITY PRIOR TO VISUAL STIMULUS ONSET - and the authors flag it, ENGAGEMENT-RELATED CORTICAL SUPPRESSION MIGHT SEEM SURPRISING, GIVEN THAT VISUAL CORTEX IS REQUIRED FOR PERFORMANCE OF THIS TASK— “Engagement-related cortical suppression might seem surprising, given that visual cortex is required for performance of this task 3 , 41 . However, reduced spiking activity and hyperpolarization has been associated with increased arousal in multiple cortical areas, an effect that may improve signal-to-noise ratios of sensory…senses[14]
senseA QUARTER OF THE DATA HAS NO ADDRESS: WE RECORDED FROM 29,134 NEURONS, OF WHICH 22,458 WERE LOCALIZABLE TO ONE OF 42 BRAIN REGIONS - every per-region number in the paper has that denominator behind itsenses[15]
senseand a majority are modulated, in both directions: A SIZEABLE FRACTION OF ALL NEURONS (60.0%, 13,466 NEURONS) HAD SIGNIFICANT MODULATION OF FIRING RATE DURING THE TASK - MOST OF THESE NEURONS (74.3%) CONSISTENTLY INCREASED THEIR ACTIVITY, BUT A SIZEABLE MINORITY (20.2%) CONSISTENTLY DECREASEDsenses[16]
senseWHAT THE GLOBAL SIGNAL CANNOT BE, ON A TIMING ARGUMENT: THESE SIGNALS MAY COMPRISE FORMS OF COROLLARY DISCHARGE, BUT CANNOT REFLECT SENSORY RE-AFFERENCE AS THEY WERE OBSERVED PRIOR TO MOVEMENT ONSET
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senses[17]
senseand the finding may belong to the trained animal rather than to the structure: WHETHER VISUAL NEURONS WOULD BE FOUND these ‘motor’ AREAS IN UNTRAINED ANIMALS IS NOT CLEAR FROM THESE DATA
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senses[18]
controlthe visual contrast matched between the conditions being compared — held — rules out the activity difference reflecting different visual drive - CONTRALATERAL VISUAL STIMULUS CONTRASTS WERE MATCHED so that DIFFERENCES IN ACTIVITY DO NOT REFLECT DIFFERING VISUAL DRIVE
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controls[0]
controlthe task-context comparison restricted to trials where the animal did nothing, against passive presentation — held — rules out movement explaining the context difference - BOTH VISUAL STIMULI AND BEHAVIORAL REPORTS WERE IDENTICAL, and the conditions were MATCHED FOR VISUAL STIMULATION AND (LACK OF) ACTION
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controls[1]
controlthe engagement index computed from PRE-STIMULUS activity only — held — rules out the index being a movement correlate - WE EXAMINED ONLY PRE-STIMULUS ACTIVITY, WHICH COULD NOT BE CONFLATED WITH THE LARGE NON-SPECIFIC RESPONSES RELATED TO MOVEMENTS
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controls[2]
controlthe engagement effect tested against pupil diameter, video-detectable movement, previous reward and inter-trial interval — held — rules out arousal, movement, reward history and timing - the effect COULD NOT BE FULLY EXPLAINED BY VARIABILITY IN PUPIL DIAMETER, IN OVERT MOVEMENTS DETECTABLE BY VIDEO RECORDINGS, IN THE PRESENCE OF A REWARD ON THE PREVIOUS TRIAL, OR IN THE INTER-TRIAL INTERVAL
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controls[3]
controlthe kernel test's threshold set from shuffled data — held — rules out the encoding counts being a product of the test - WE USED PARAMETERS WHICH GAVE FALSE-POSITIVE ERROR RATES OF 0.33% ON SHUFFLED DATA
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controls[4]
controlthe whole choice result repeated with a second, independent statistic — held — rules out the conclusion depending on the kernel model - a version of CHOICE PROBABILITY ANALYSIS FOR TASKS WITH MANY STIMULUS CONDITIONS was developed and EMPLOYING THIS METHOD, WE OBTAINED SIMILAR RESULTS
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controls[5]
controlthe analysis restricted to activity before the movement — held — rules out activity caused by the movement being read as predicting it - AS OUR QUESTION CONCERNS ACTIVITY PREDICTIVE OF UPCOMING MOVEMENTS, WE APPLIED THIS ANALYSIS ONLY TO PRE-MOVEMENT ACTIVITY
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controls[6]
differs ingo/nogo designs as evidence about choice-selective neurons — what the design can separate - GO/NOGO TASKS CANNOT DISTINGUISH NEURONS THAT NONSPECIFICALLY FIRE FOR ANY ACTION, FROM NEURONS SELECTIVE FOR SPECIFIC CHOICES
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claim_relations[4]
differs inthe classical set of choice-related areas (frontal cortex, striatum, SNr, deep superior colliculus) — two regions nobody expected - BUT ALSO UNEXPECTED REGIONS INCLUDING THE MIDBRAIN RETICULAR NUCLEUS AND ZONA INCERTA
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claim_relations[6]
noteaction —distinct-from→ choice — NEURONAL SIGNALS THAT CORRELATE WITH ACTION DO NOT NECESSARILY CORRELATE WITH ACTION SELECTION
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concept_edges[0]
auditgo/nogo —contrasted-with→ choice — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span argues for multi-alternative over go/no-go tasks; contrasted-with is loose.
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concept_edges[2]
notego/nogo —contrasted-with→ choice — GO/NOGO TASKS CANNOT DISTINGUISH NEURONS THAT NONSPECIFICALLY FIRE FOR ANY ACTION, FROM NEURONS SELECTIVE FOR SPECIFIC CHOICES
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concept_edges[2]
differs inmidbrain —contrasted-with→ forebrain — THE AXIS IS WHAT THEY SHARE AND WHAT THEY DO NOT, BOTH NAMED IN ONE CLAUSE. 'Although the encoding of choice emerged with indistinguishable timing in midbrain and forebrain, these regions encoded choice differently.' Identical on timing, different on code. A bare 'contrasted-with' records the difference and drops the fact that the obvious explanation - one region leading the other - is ruled out in the same sentence.
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Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
concept_edges[9]
notemidbrain —contrasted-with→ forebrain — ALTHOUGH THE ENCODING OF CHOICE EMERGED WITH INDISTINGUISHABLE TIMING IN MIDBRAIN AND FOREBRAIN, THESE REGIONS ENCODED CHOICE DIFFERENTLY - a 'BILATERAL' ENCODING OF BOTH CHOICES IN THE FOREBRAIN, VERSUS a 'UNILATERAL' ENCODING OF CONTRALATERAL CHOICES IN THE MIDBRAIN 2026-08-20 the project re-read worst chars/edge queue): SOUND - the passage is built as this contrast and says so twice - 'By contrast' and 'in the forebrain…'. Both endpoints named, type correct. KEEP.
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concept_edges[9]
notepassive —contrasted-with→ Miss — BOTH VISUAL STIMULI AND BEHAVIORAL REPORTS WERE IDENTICAL, and yet THEY WERE ACCOMPANIED IN MANY AREAS BY DIFFERENT ACTIVITY, BOTH BEFORE AND AFTER STIMULUS PRESENTATION
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concept_edges[11]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
outcome (stated by the tagger, no quotation)control — heldcontrols[1]
outcome (stated by the tagger, no quotation)control — heldcontrols[2]
outcome (stated by the tagger, no quotation)control — heldcontrols[3]
outcome (stated by the tagger, no quotation)control — heldcontrols[4]
outcome (stated by the tagger, no quotation)control — heldcontrols[5]
outcome (stated by the tagger, no quotation)control — heldcontrols[6]
for which uselimitation — every effect size and null in the survey: THE SAMPLE SIZES (n = 10 MICE; n = 39 RECORDING SESSIONS; n = 29,134 NEURONS) WERE NOT DETERMINED WITH A POWER ANALYSIS
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limitations[0]
for which uselimitation — the same scope as limitations[7] - whether visual responses in the newly implicated motor choice areas would exist in untrained animals cannot be determined here
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limitations[1]
for which uselimitation — the mechanistic and anatomical completeness of the survey: FUTURE WORK WILL BE REQUIRED TO DETERMINE THE CIRCUIT MECHANISMS THAT ENFORCE THESE PRINCIPLES
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limitations[2]
for which uselimitation — the contrast-condition comparisons: THE EXPERIMENTER WAS NOT BLINDED TO CONTRAST CONDITION EITHER DURING DATA ACQUISITION OR DURING ANALYSIS
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limitations[3]
for which uselimitation — the choice-probability control analysis: SIX SESSIONS HAD TO BE EXCLUDED FOR HAVING FEWER THAN 10 TRIALS INCLUDE-ABLE IN THE CP ANALYSIS
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limitations[4]
for which uselimitation — reading 'this neuron encodes action' as a statement about that neuron's spiking rather than about its trial-averaged rate
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[5]
for which uselimitation — any per-region count or map: WE RECORDED FROM 29,134 NEURONS ... OF WHICH 22,458 WERE LOCALIZABLE TO ONE OF 42 BRAIN REGIONS
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[6]
for which uselimitation — reading the visual responses in midbrain and striatum as a property of those structures rather than of trained animals
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limitations[7]
for which uselimitation — the brain-wide reading of the survey: future work must determine HOW THEY EXTEND TO AREAS SUCH AS CEREBELLUM AND BRAINSTEM OMITTED FROM THE CURRENT SURVEY
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limitations[8]
for which uselimitation — reading the recurrent loop as observed rather than as the simplest account of an absence
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limitations[9]
replicationuntested
limitationSample sizes were not set by a power analysis.
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limitations[0]
limitationWhether visual responses in the newly implicated 'motor' choice areas would exist in untrained animals cannot be determined from these data.
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limitations[1]
limitationCircuit mechanisms, omitted regions (cerebellum, brainstem) and generality to other choice tasks all remain for future work.
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limitations[2]
limitationThe experimenter was not blinded to the stimulus contrast condition during acquisition or analysis.
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limitations[3]
limitationSix of 39 sessions had to be dropped from the choice-probability control analysis for too few usable trials.
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limitations[4]
limitationTHE FRACTION OF CROSS-VALIDATED VARIANCE THE KERNELS EXPLAINED WAS FREQUENTLY SMALL, EVEN FOR NEURONS WHOSE MEAN RATES THEY ACCURATELY PREDICTED (50.2% for one and 13.6% for another)
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limitations[5]
limitationa quarter of the neurons could not be placed - WE RECORDED FROM 29,134 NEURONS, OF WHICH 22,458 WERE LOCALIZABLE TO ONE OF 42 BRAIN REGIONS
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limitations[6]
limitationthe training may have made the finding - WHETHER VISUAL NEURONS WOULD BE FOUND these ‘motor’ AREAS IN UNTRAINED ANIMALS IS NOT CLEAR FROM THESE DATA
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limitations[7]
limitationthe survey is not the brain - future work must determine HOW THEY EXTEND TO AREAS SUCH AS CEREBELLUM AND BRAINSTEM OMITTED FROM THE CURRENT SURVEY, and THE EXTENT TO WHICH SIMILAR PRINCIPLES GOVERN THE NEURAL CORRELATES OF DIFFERENT CHOICE TASKS
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limitations[8]
limitationthe timing result is a null - WE COULD NOT DISTINGUISH THE TIMING OF CHOICE-RELATED SIGNALS BETWEEN THESE REGIONS, and the recurrent loop is offered as the PARSIMONIOUS explanation of that null rather than as a measurement
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limitations[9]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingNeurons whose firing rose non-specifically at the moment of action initiation were present in nearly every one of the 42 recorded brain regions.
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content_tags.findings[0]
findingNeurons coding the visual stimulus and the upcoming choice were not brain-wide but restricted to particular neocortical, basal ganglia and midbrain regions.
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content_tags.findings[1]
findingMidbrain choice neurons were almost all activated before contralateral turns and suppressed before ipsilateral ones, whereas forebrain choice neurons could prefer either direction.
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content_tags.findings[2]
findingPre-stimulus activity across the brain predicted whether the mouse would respond on a given trial, with subcortical activity raised and neocortical activity lowered during engagement.
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content_tags.findings[3]
findingTHE TIME COURSE OF CHOICE DECODING COULD NOT BE STATISTICALLY DISTINGUISHED BETWEEN CHOICE-SELECTIVE REGIONS IN FRONTAL CORTEX, STRIATUM, AND MIDBRAIN
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content_tags.findings[4]
findingChoice-selective neurons were found in frontal cortex, striatum, substantia nigra pars reticulata, higher-order thalamus and deep superior colliculus, and unexpectedly also in the midbrain reticular nucleus and zona incerta.
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content_tags.findings[5]
findingOn trials where the mouse missed a contralateral stimulus and did not act, activity appeared in a visual pathway of visual areas, basal ganglia and midbrain but failed to propagate globally.
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content_tags.findings[6]
findingSixty per cent of all recorded neurons (13,466) showed significant task-related firing-rate modulation.
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content_tags.findings[7]
findingCanonical-correlation analysis of simultaneous populations showed visual cortex leading frontal cortex and midbrain, but no lead-lag relationship at all between frontal cortex and midbrain.
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content_tags.findings[8]
findingThe engagement index separated Go from Miss trials consistently across recordings, an effect not fully explained by pupil diameter, overt movement, previous reward or inter-trial interval.
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content_tags.findings[9]
findingMore than half of midbrain choice-selective neurons increased firing before one choice and fell below baseline before the other.
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content_tags.findings[10]
findingNEURONS ENCODING ACTION WERE SPREAD THROUGHOUT ALL RECORDED REGIONS, and THE GREAT MAJORITY DID NOT REQUIRE AN ADDITIONAL CHOICE KERNEL - THEY RESPONDED EQUALLY FOR MOVEMENTS IN EITHER DIRECTION
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content_tags.findings[11]
findingCHOICE-SELECTIVE NEURONS WERE NOT FOUND IN VISp, and were found in TWO SUBCORTICAL NUCLEI WHICH UNEXPECTEDLY CONTAINED NEURONS SELECTIVE FOR CHOICE (MRN AND ZI)
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content_tags.findings[12]
findingIN MIDBRAIN NEARLY ALL CHOICE-SELECTIVE NEURONS (53/54, 98%) PREFERRED CONTRALATERAL CHOICES, while in forebrain A SIZEABLE PROPORTION PREFERRED IPSILATERAL CHOICES (19/48, 40%)
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content_tags.findings[13]
findingTHE WIDELY DISTRIBUTED ACTIVITY SEEN FOLLOWING TRIAL ONSET WAS PRESENT ONLY WHEN ANIMALS MOVED, AND REGARDLESS OF THE PARTICULAR STIMULUS AND PARTICULAR ACTION
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content_tags.findings[14]
findingEVEN THOUGH THE TWO CONDITIONS WERE MATCHED FOR VISUAL STIMULATION AND (LACK OF) ACTION, THEY WERE ACCOMPANIED IN MANY AREAS BY DIFFERENT ACTIVITY, BOTH BEFORE AND AFTER STIMULUS PRESENTATION
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content_tags.findings[15]
findingWHILE NEOCORTEX AND SENSORY THALAMUS SHOWED A NET DECREASE IN PRE-STIMULUS ACTIVITY DURING TASK CONTEXT, OTHER REGIONS SHOWED A CONSISTENT INCREASE
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content_tags.findings[16]
findingthe time course of choice decoding COULD NOT BE STATISTICALLY DISTINGUISHED BETWEEN CHOICE-SELECTIVE REGIONS IN FRONTAL CORTEX, STRIATUM, AND MIDBRAIN
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content_tags.findings[17]
findingWE RECORDED FROM 29,134 NEURONS, OF WHICH 22,458 WERE LOCALIZABLE TO ONE OF 42 BRAIN REGIONS
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content_tags.findings[18]
findingA SIZEABLE FRACTION OF ALL NEURONS (60.0%, 13,466 NEURONS) HAD SIGNIFICANT MODULATION OF FIRING RATE DURING THE TASK - 74.3% CONSISTENTLY INCREASED, 20.2% CONSISTENTLY DECREASED
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content_tags.findings[19]
findingTHE FRACTION OF CROSS-VALIDATED VARIANCE THE KERNELS EXPLAINED WAS FREQUENTLY SMALL, EVEN FOR NEURONS WHOSE MEAN RATES THEY ACCURATELY PREDICTED
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content_tags.findings[20]
quantity29,134 — neurons recorded in total, of which 22,458 could be assigned to one of 42 brain regions
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content_tags.quantities[0]
quantity60.0% — share of recorded neurons with significant task-related firing-rate modulation
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content_tags.quantities[1]
quantity53/54, 98% — proportion of midbrain choice-selective neurons preferring contralateral choices
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content_tags.quantities[2]
quantity19/48, 40% — proportion of forebrain choice-selective neurons preferring ipsilateral choices, significantly more than in midbrain
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content_tags.quantities[3]
quantity0.33% — false-positive rate of the kernel selectivity test estimated on shuffled data
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content_tags.quantities[4]
quantity0.079 / 0.069 / 0.040 — THIS FOCALITY INDEX WAS 0.079 FOR CHOICE, 0.069 FOR VISUAL KERNELS AND 0.040 FOR ACTION KERNELS - the differences between Choice and Move, and between Contralateral Vision and Action, WERE STATISTICALLY SIGNIFICANT
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content_tags.quantities[5]
quantity1.7 ± 2.5% — incorrect-choice rate on high-contrast single-stimulus trials, with 10.1% misses
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content_tags.quantities[6]
quantity94.1% ± 0.6% — proportion of passive-replay trials with high-contrast stimuli in which the mouse made no wheel movement
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content_tags.quantities[7]
quantity88.7 μm — median absolute error of the longitudinal probe-to-atlas alignment, close to the 70 micrometer probe shank width
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content_tags.quantities[8]
quantity29/54, 54% — midbrain choice-selective neurons showing directionally opposed activity (enhanced for one choice, suppressed below baseline for the other)
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content_tags.quantities[9]
quantity53/54, 98% — midbrain choice-selective neurons preferring contralateral choices
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content_tags.quantities[10]
quantity0.33% — the FALSE-POSITIVE ERROR RATE ON SHUFFLED DATA that the kernel test's parameters were set to give
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content_tags.quantities[11]
quantity29,134 — neurons recorded, OF WHICH 22,458 WERE LOCALIZABLE TO ONE OF 42 BRAIN REGIONS
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content_tags.quantities[12]
quantity60.0%, 13,466 neurons — the fraction with SIGNIFICANT MODULATION OF FIRING RATE DURING THE TASK
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content_tags.quantities[13]
quantity50.2% ... 13.6% — cross-validated variance explained for two example neurons whose mean rates the model predicted accurately
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content_tags.quantities[14]
variablecontrast condition (independent)
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content_tags.variables[0]
variablefiring rate (dependent)
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content_tags.variables[2]
variablecontralateral visual stimulus contrast (control)
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content_tags.variables[4]
variablechoice
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content_tags.variables[6]
variableengagement
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content_tags.variables[7]
variablepupil area (covariate)
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content_tags.variables[8]
variablemotion energy (covariate)
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content_tags.variables[9]
variableprevious trial reward outcome (covariate)
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content_tags.variables[10]
variableinter-trial interval (covariate)
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content_tags.variables[11]
definitionchoice
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content_tags.definitions[0]
definitionchoice-related signal
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content_tags.definitions[1]
definitioncombined-conditions Choice Probability (ccCP)
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content_tags.definitions[3]
definitionjoint Peri-Event Canonical Correlation (jPECC)
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content_tags.definitions[4]
definitionengagement index
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content_tags.definitions[5]
definitionFocality index
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content_tags.definitions[6]
definitioncombined-conditions choice probability
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content_tags.definitions[7]
definition‘Action’ kernel
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content_tags.definitions[8]
definition‘Choice’ kernel
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content_tags.definitions[9]
key-termschoice, action, vision, engagement, kernel, go/nogo, passive, Miss, midbrain, forebrain, neocortex, VISp, ZI, timing, contralateral, ipsilateral
Statistics — cross-field join keys
field-invariant statistical terms, used in their statistical sense (gated on a number / symbol / test). A p-value or a standard deviation means the SAME thing in physics, biology, psychology and sociology — so these are the strongest cross-field join keys. Grounded verbatim.
ANOVA p as written: multi-way ANOVA
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content_tags.statistics[0]
ANOVA F as written: two-way ANOVA
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content_tags.statistics[1]
nested ANOVA as written: nested multiple ANOVA test
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content_tags.statistics[2]
chi-squared test p as written: Fisher’s exact test
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content_tags.statistics[3]
t-test p as written: paired t-test
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content_tags.statistics[4]
Bonferroni correction as written: Bonferroni-corrected alpha value
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content_tags.statistics[5]
cross-validation as written: held-out data
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content_tags.statistics[6]
regression as written: reduced rank regression
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content_tags.statistics[7]
logistic regression as written: L1-regularized logistic regression
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content_tags.statistics[8]
generalized linear model as written: GLM with binomial link function
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content_tags.statistics[9]
bootstrap
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content_tags.statistics[10]
principal component analysis as written: principal components analysis
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content_tags.statistics[11]
canonical correlation analysis λ
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content_tags.statistics[12]
ROC / Mann-Whitney U statistic as written: area under a receiver operating characteristic (ROC) curve
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content_tags.statistics[13]
permutation test as written: shuffle test
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content_tags.statistics[14]
two-way ANOVA
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content_tags.statistics[15]
Fisher's exact test
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content_tags.statistics[16]
cross-validated
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content_tags.statistics[17]
Second readings
a facet read a second time, independently, from its own sentence — and whether the two readings agree
claim typeexistence — a second reading reaching the same value; recorded: existence
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alternatives.claim_type
methodfit each neuron with kernels for vision, action and choice, then ask by a nested cross-validated test whether the kernel under test is REQUIRED, with a declared false-positive rate on shuffled data — a second reading DIFFERING from the recorded value; recorded: electrophysiology
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alternatives.method
phenomenonactivity that appears everywhere in the brain before a movement, in regions with no established role in choosing it — a second reading DIFFERING from the recorded value; recorded: brain-wide distribution of neural coding for vision, action, choice and task engagement
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.phenomenon

When we have changed our mind. Some entries in the table begin with a word in capitals and the reason for it. EVIDENCE CORRECTED — the finding still stands, but we had quoted the wrong sentence from the paper, and the quotation has been replaced with the right one. RETRACTED — the entry itself was wrong; it stays on the page, marked, with the reason, rather than being deleted. WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below. Each mark carries its own reason, in brackets, because the reasons differ from one entry to the next. An entry we have WITHDRAWN is removed from this page and kept in the record, with its reason. A line reading Audit and a date is a later re-reading of the entry against its whole sentence: DOUBTFUL means it was kept, and the reason says what is loose about it. Quotation marks inside the tagger's sentence. Where a tag is the tagger's own sentence, a phrase in quotation marks inside it is borrowed from the paper — the tagger's wording around it is not. The paper's own sentence is the evidence quote for that row; on this page it is withheld with the article, and the article is at its publisher. Nothing inside quotation marks anywhere on this page is ours: if it is quoted, it is the paper's. Whose words a concept carries. A concept entry records a term the paper itself uses, so the term, the sense given for it, and its evidence are the paper's own words, quoted where the paper fixes them. This is the opposite of a sense entry in the tag table, which is the tagger's sentence about how the paper uses a term. Where a concept entry could not be grounded in the paper's words, the entry says so on its own row.

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