Corticostriatal neurons in auditory cortex drive decisions during auditory discrimination — TEI-tagged view

Znamenskiy, Zador · 2013 · neuroscience · doi:10.1038/nature12077 · 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. Method is recorded as electrophysiology because tetrode/optical-fibre array recordings plus optogenetic manipulation in behaving rats carry the evidence; the study is equally a behavioural experiment (a two-alternative-choice cloud-of-tones discrimination task) and the vocabulary cannot hold both. Prior work is cited only by superscript number in this text, so claim relations targets are described rather than named. 2026-08-12 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.
9 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.
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logistic regression 3 fields · cross-validation 3 fields · bootstrap 2 fields · confidence interval 2 fields · Confidence intervals 2 fields
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corticostriatal auditory cortex19 striatum2 perceptual decision-making2 cloud-of-tones two alternative choice3 tonotopic9 Channelrhodopsin-23 archaerhodopsin-3 psychometric curve2 chronometric choice biases anti-bias antidromic2 compensatory plasticity2 logistic regression30 ideal observer2 space constant superior colliculus reinforcement learning10 electrophysiology110

Our tag layer for this paper

Cross-disciplinary analysis
content-tags & facets: phenomenon · claim-type · cross-domain-transfer · mechanism · claim-relation · replication-status · limitation — grounded in the full text
phenomenonauditory perceptual decision-making— “We hypothesized that the projection from the auditory cortex to the striatum carries acoustic information that drives behavioural choices during auditory discrimination.”
claim-typemechanism— “We directly demonstrate the role of corticostriatal projection neurones in auditory decisions by specifically manipulating the activity of these neurones in rats performing an auditory frequency discrimination task.”
method — evidence— “We used tetrode recordings to characterize the activity of individual neurones in the auditory cortex and the auditory striatum while rats performed the task.”
object of study — evidence— “We hypothesized that the projection from the auditory cortex to the striatum carries acoustic information that drives behavioural choices during auditory discrimination.”
research question — evidence— “The neural pathways by which information about the acoustic world reaches the auditory cortex are well characterised, but how auditory representations are transformed into motor commands is not known.”
contribution — evidence— “Targeted Channelrhodopsin-2 1 , 2 (ChR2)-mediated stimulation of corticostriatal neurones during the task biased decisions in the direction predicted by the frequency tuning of the stimulated neurones, whereas archaerhodopsin-3 3 (Arch)-mediated inactivation biased decisions in the opposite direction.”
transferfrom primate visual neurophysiology: the random-dot-motion logic of a graded two-alternative discrimination, remade for frequency using clouds of tones— “We trained rats to discriminate low and high frequency “cloud-of-tones” stimuli in a two alternative choice task”transfers[0]
transferfrom psychophysics and signal detection: the ideal observer, applied to single neurons so that cortical and striatal selectivity can be compared on one scale— “These observations indicate that neurones in the striatum preserve the information available to perform the ”cloud-of-tones” task.”transfers[1]
transferfrom virology: a retrogradely transported herpes virus carrying Cre, which turns an anatomical projection into a genetically addressable cell class— “In the first strategy, we targeted ChR2 specifically to corticostriatal neurones using a Herpes Simplex Virus-1 (HSV1) engineered to express Cre recombinase 19 , 20 , 21 ( Fig. 2a ). This virus, when injected into the auditory striatum, was transported retrogradely along the axons and drove Cre expression in corticostriatal neurones. We then…transfers[2]
mechanismThe auditory cortex-to-striatum projection carries frequency-specific acoustic information that is read out to select the motor response, so changing spiking in that projection changes the choice.— “We hypothesized that the projection from the auditory cortex to the striatum carries acoustic information that drives behavioural choices during auditory discrimination.”mechanisms[0]
mechanismBecause exciting corticostriatal cells suppresses surrounding cortex rather than exciting it, the behavioural bias must be delivered by their long-range axons to the striatum, not by recruiting other cortical output pathways.— “Excitation of corticostriatal neurones resulted in general suppression of local cortical activity ( Fig. S6 ), favouring the hypothesis that behavioural effects of photostimulation are mediated by long-range rather than local outputs of these neurones.”mechanisms[1]
mechanismA decision bottleneck: the small number of projection neurons that actually drive the choice provides finite bandwidth, which caps the effective pool size and hence psychophysical performance.— “Our results suggest that the finite bandwidth provided by a small number of projection neurones that drive decisions constitutes an additional constraint limiting the effective pool size and therefore psychophysical performance.”mechanisms[2]
mechanismsufficiency and necessity are separated explicitly, and the direction of the necessity test is predicted before it is run— “If corticostriatal spikes are important under normal conditions, then suppressing these spikes during the task should lead to an “anti-bias,” i.e. a bias in the direction opposite that induced by ChR2 activation.”mechanisms[3]
mechanismcell-type specificity is achieved by an intersection: a retrograde virus delivers Cre from the target region, and a Cre-dependent virus supplies the opsin in the source region— “In the first strategy, we targeted ChR2 specifically to corticostriatal neurones using a Herpes Simplex Virus-1 (HSV1) engineered to express Cre recombinase 19 , 20 , 21 ( Fig. 2a ). This virus, when injected into the auditory striatum, was transported retrogradely along the axons and drove Cre expression in corticostriatal neurones. We then…mechanisms[4]
mechanisma failed non-specific manipulation is reported first, with three candidate explanations, and becomes the argument for targeting— “The lack of correspondence between the direction of stimulation-evoked choice biases and frequency tuning was surprising and has several possible…mechanisms[5]
mechanismfull inactivation is deliberately avoided because compensatory plasticity would confound it, so only a patch of the tonotopic map is silenced
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mechanisms[6]
mechanismthe same pathway is driven at two points - somata in cortex and axon terminals in striatum - so the result cannot depend on where the light went
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mechanisms[7]
mechanismrecurrent cortical activity is pharmacologically removed while the axons are stimulated, which isolates the long-range output as the carrier of the effect
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mechanisms[8]
mechanismthe behavioural effect is converted into stimulus units by fitting a logistic model, so a neural manipulation and an acoustic change become comparable
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mechanisms[9]
mechanismstimulation is restricted to a quarter of trials and still rewarded normally, to stop the animal adapting to the manipulation itself
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mechanisms[10]
mechanismtone intensity is randomised across trials specifically to block a loudness-based strategy
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mechanisms[11]
mechanismthe effect size is tied to an anatomical count by combining fluorescence, a measured inactivation space constant, and cell density
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mechanisms[12]
mechanismuninjected animals receiving identical light serve as the control for both opsins separately
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mechanisms[13]
mechanismlocal suppression during corticostriatal excitation is used to argue the behavioural effect travels by the long-range output rather than through local circuitry
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mechanisms[14]
builds-on perceptual decision experiments in primate area MT (ref. 16)
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claim_relations[0]
extends prior characterisations of auditory striatal neurones in anaesthetised and passively listening animals (refs. 14, 15)
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claim_relations[1]
replicates previous results on the laminar location of corticostriatal neurones (ref. 22)
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claim_relations[2]
replicates previously reported response-time increases on difficult trials in rats (ref. 18)
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claim_relations[3]
builds-on the known topography of corticostriatal projections (ref. 23)
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claim_relations[4]
extends work on striatal regions receiving motor and prefrontal input, by asking what the sensory-recipient striatum does
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claim_relations[5]
critiques non-specific optogenetic activation of mixed cortical populations as a way of testing a pathway's role
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claim_relations[6]
critiques complete-inactivation lesion designs, whose interpretation compensatory plasticity can obscure
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claim_relations[7]
builds-on primate area MT microstimulation experiments, whose logic the task design is modelled on
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claim_relations[8]
extends estimates of how few neurons animals can be trained to detect, from artificial detection to normal perception
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claim_relations[9]
supports the account in which correlated firing limits the effective size of the neuronal pool underlying psychophysical performance
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claim_relations[10]
senseoptogenetic activation - non-specific across a cortical population versus targeted to a projection-defined class; the first gives biases unrelated to tuning and degrades performance, the second gives biases predicted by tuning and spares it
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senses[0]
sensesufficiency versus necessity - activation showing the pathway CAN drive choices, inactivation showing it normally DOES; the paper separates the two claims and tests them in that order
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senses[1]
sensestimulating the pathway at the soma versus at the axon terminal; the same cells reached from either end, which is what rules out a location artefact
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senses[2]
sensea behavioural effect measured as a choice bias versus as a shift in response time; the chronometric curve moves as if acoustic evidence had been added
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senses[3]
sensethe neuronal pool limiting psychophysical performance - as constrained by correlated firing versus by the bandwidth of the few projection neurons that drive decisions
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senses[4]
sensestriatal regions receiving motor and prefrontal input, whose functions are well studied, versus those receiving direct sensory cortical input, whose function was not established
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senses[5]
controlthe fit re-run 100 times from RANDOM starting parameters, keeping the solution with the smallest mean square error — held — rules out a local minimum as the reported fit - a single optimisation run reports wherever it happens to stop, and its parameters would be a property of the starting point
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controls[0]
differs incomplete-inactivation lesion designs, whose interpretation compensatory plasticity can obscure — THE AXIS IS THE EXTENT OF THE INACTIVATION, AND THE REASON IS NAMED. 'Rather than attempting to inactivate all corticostriatal cells - a technically difficult feat whose interpretation might be obscured by the compensatory plasticity common to lesion studies - we sought instead to silence corticostriatal neurones WITHIN A RESTRICTED REGION of the tonotopic map.' The critique of whole-population lesions is not that they fail but that the preparation answers back.
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claim_relations[7]
auditEVIDENCE CORRECTED (RE-GROUNDED) · corticostriatal —causes→ choice biases — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'biased subjects' choices', not 'choice biases'.
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concept_edges[0]
retractedEVIDENCE CORRECTED (RE-GROUNDED) · corticostriatal —causes→ choice biases — RE-GROUNDED. The old span named neither endpoint. This clause names both and states the dependence - the bias scales with how many corticostriatal neurones were silenced.
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concept_edges[0]
weightEVIDENCE CORRECTED (RE-GROUNDED) · corticostriatal —causes→ choice biases — 0.95
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concept_edges[0]
differs inChannelrhodopsin-2 —contrasted-with→ archaerhodopsin-3 — THE AXIS IS THE SIGN OF THE MANIPULATION, AND THE BIAS FOLLOWS IT. 'TARGETED CHANNELRHODOPSIN-2 (ChR2)-MEDIATED STIMULATION of corticostriatal neurones during the task biased decisions in the direction predicted by the frequency tuning of the stimulated neurones, WHEREAS ARCHAERHODOPSIN-3 (ARCH)-mediated inactivation biased decisions in T[HE OPPOSITE DIRECTION].' Two opsins are not being compared as tools - they are the two signs of one experiment, and the double dissociation is the evidence.
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concept_edges[1]
retractedEVIDENCE CORRECTED (RE-GROUNDED) · corticostriatal —part-of→ auditory cortex — RE-GROUNDED. The old span named neither endpoint. This sentence names both and places one inside the other - corticostriatal neurones IN the auditory cortex.
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concept_edges[2]
weightEVIDENCE CORRECTED (RE-GROUNDED) · corticostriatal —part-of→ auditory cortex — 0.9
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concept_edges[2]
auditchoice biases —operationalised-as→ logistic regression — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'subjects' choices' / 'behavioural effect', not 'choice biases'.
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concept_edges[4]
weightchoice biases —operationalised-as→ logistic regression — 0.95
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concept_edges[4]
weightstriatum —causes→ superior colliculus — 0.75
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concept_edges[5]
notecorticostriatal —causes→ choice biases — RE-GROUNDS another row of this record: the relation was always stated, the endpoint names were not in the span.
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concept_edges[6]
weightcorticostriatal —causes→ choice biases — 0.95
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concept_edges[6]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
for which uselimitation — the non-specific activation experiments: THE METHOD OF NON-SPECIFIC ACTIVATION OF DIVERSE NEURONAL POPULATIONS USED IN THESE EXPERIMENTS DOES NOT PROVIDE SUFFICIENT control over the relevant populations
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limitations[0]
for which uselimitation — any session-level claim: THE BIASES WERE NOT SIGNIFICANT ON INDIVIDUAL SESSIONS (p<0.05 ON 1/33 SESSIONS), IN PART BECAUSE THE NUMBER OF STIMULATED TRIALS PER SESSION WAS TYPICALLY SMALL (70+/-24 (s.d.) TRIALS)
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limitations[1]
for which uselimitation — the chronometric result specifically: A SIMILAR TREND WAS REFLECTED IN THE SUBJECTS' CHRONOMETRIC FUNCTIONS BUT DID NOT REACH SIGNIFICANCE
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limitations[2]
for which uselimitation — the cell counts for two named fibres: FOR 2 FIBERS IN ONE OF THE RATS WE COULD NOT LOCATE THE TRACKS IN HISTOLOGICAL SECTIONS AND PRECISELY ESTIMATE CELL COUNTS
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limitations[3]
for which uselimitation — any claim that this pathway is the only one: OUR RESULTS DO NOT EXCLUDE THE PARTICIPATION OF OTHER PARALLEL PATHWAYS
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limitations[4]
for which uselimitation — the same scope as limitations[4] - other parallel pathways remain possible
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limitations[5]
for which uselimitation — the same scope as limitations[1] - individual sessions are underpowered at 70+/-24 trials
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limitations[6]
for which uselimitation — the sites with large choice biases: STIMULATION ALSO HAD A MODEST EFFECT ON THE SUBJECTS' PERFORMANCE AT SITES WITH LARGE CHOICE BIASES, POSSIBLY DUE TO A CEILING EFFECT
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limitations[7]
for which uselimitation — the inactivation result: it BIASED SUBJECTS' CHOICES AWAY FROM THE REWARD PORT ASSOCIATED WITH THE FREQUENCY BAND OF THE INACTIVATION SITE BUT DID NOT AFFECT the chronometric measure
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limitations[8]
for which uselimitation — every PRIOR characterisation this paper builds on: AUDITORY STRIATAL NEURONES HAVE PREVIOUSLY BEEN CHARACTERIZED IN ANAESTHETISED AND PASSIVELY LISTENING ANIMALS, NOT DURING BEHAVIOUR
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limitations[9]
for which uselimitation — sensory-recipient striatal regions as a class: THE FUNCTION OF STRIATAL REGIONS WHICH RECEIVE DIRECT INPUT FROM SENSORY CORTICAL AREAS IS LESS WELL ESTABLISHED
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limitations[10]
replicationuntested
limitationNon-specific cortical activation did not give tuning-predicted biases and lacked adequate control over the relevant projection neurons.
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limitations[0]
limitationThe stimulation biases were not statistically significant within individual sessions.
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limitations[1]
limitationThe inactivation effect on chronometric functions did not reach significance.
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limitations[2]
limitationCell counts could not be estimated for two fibres whose tracks were not found histologically.
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limitations[3]
limitationOther parallel pathways contributing to the decision are not excluded.
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limitations[4]
limitationother parallel pathways are not excluded
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limitations[5]
limitationindividual sessions were underpowered; the effect is visible in aggregate
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limitations[6]
limitationat sites with large biases, performance was also modestly affected, possibly a ceiling effect
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limitations[7]
limitationthe chronometric counterpart of the inactivation result trended but did not reach significance
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limitations[8]
limitationauditory striatal neurons had only ever been characterised in anaesthetised or passively listening animals before this
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limitations[9]
limitationthe function of sensory-recipient striatal regions was poorly established at the outset
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limitations[10]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingOptogenetic activation of auditory corticostriatal neurons shifted rats' choices toward the reward port associated with the frequency the stimulated neurons preferred.
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content_tags.findings[0]
findingArch-mediated silencing of corticostriatal neurons pushed choices away from the port matching the inactivated site's frequency band while leaving psychometric slope (sensitivity) unchanged.
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content_tags.findings[1]
findingNon-specific (untargeted) activation of auditory cortex produced biases whose direction did not follow the frequency tuning of neurons near the fiber.
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content_tags.findings[2]
findingNon-specific cortical activation also impaired task performance, flattening the psychometric curve.
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content_tags.findings[3]
findingStimulating corticostriatal axons still biased choices after the auditory cortex was pharmacologically silenced, showing the effect travels through the long-range striatal projection rather than through recurrent cortical circuitry.
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content_tags.findings[4]
findingSilencing only about one third to eight tenths of one percent of the local neurons (roughly 1,000-2,400 Arch-expressing cells) was enough to produce robust choice biases.
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content_tags.findings[5]
findingCorticostriatal stimulation also shifted response-time (chronometric) curves exactly as adding real acoustic evidence for the site's preferred frequency would.
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content_tags.findings[6]
findingIdeal-observer analysis showed that cortical and striatal neurons had similar stimulus selectivity.
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content_tags.findings[7]
findingactivating corticostriatal neurons biased choices toward the port associated with the stimulation site's preferred frequency
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.
content_tags.findings[8]
findinginactivating them biased choices away from that port, as predicted
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content_tags.findings[9]
findingnon-specific cortical activation produced biases whose direction the tuning near the fiber did not predict
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content_tags.findings[10]
findingstimulating corticostriatal axons in the striatum reproduced the effect from the other end of the pathway
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content_tags.findings[11]
findingthe bias survived pharmacological silencing of the cortex, so it is carried by the long-range output rather than recurrent cortical activity
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content_tags.findings[12]
findingstimulation shifted chronometric curves as if acoustic evidence favouring that choice had been added
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content_tags.findings[13]
finding84% of targeted neurons were in layer V
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content_tags.findings[14]
findingrobust biases appeared at sites with 1040-2444 Arch-expressing cells, 0.35-0.83% of the local population
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content_tags.findings[15]
findingsilencing that minute fraction changed behaviour, suggesting a privileged role for these cells
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content_tags.findings[16]
findinginactivation efficiency decayed with a space constant of about 564 micrometres
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content_tags.findings[17]
findingcortical and striatal neurons showed similar stimulus selectivity, so the striatum preserves the information needed for the task
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content_tags.findings[18]
findingtargeted stimulation left psychophysical sensitivity intact, unlike non-specific stimulation
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content_tags.findings[19]
findingno biases appeared in uninjected animals given identical light
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content_tags.findings[20]
findingexciting corticostriatal neurons suppressed local cortical activity overall
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content_tags.findings[21]
quantity84% — share of ChR2-expressing neurons that lay in cortical layer V after the HSV1-Cre/AAV targeting
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content_tags.quantities[0]
quantity0.35–0.83% — fraction of neurons in the illuminated volume that had to be silenced to yield robust choice biases
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content_tags.quantities[1]
quantity~564 μ m — space constant of the decay of inactivation efficiency with distance from the fiber
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.
content_tags.quantities[2]
quantity36/57 sites, p = 0.015 — proportion of non-specific cortical stimulation sites producing contralateral biases, and its significance
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content_tags.quantities[3]
quantityp = 0.016 — significance of the choice bias from corticostriatal axon stimulation while the cortex was pharmacologically inactivated
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content_tags.quantities[4]
quantity20–50 ms — size of the response-time increase on difficult trials
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content_tags.quantities[5]
quantity70±24 (s.d.) trials — typical number of stimulated trials per session, explaining why single-session biases rarely reached significance
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content_tags.quantities[6]
quantity4 of 201 — cells that responded to blue-light pulses on more than half of trials, i.e. the identified light-driven corticostriatal population
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content_tags.quantities[7]
quantity25% — proportion of trials on which photostimulation was delivered, capped to limit behavioural adaptation
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content_tags.quantities[8]
quantity5-40 kHz over three octaves — the stimulus range, split into low and high target octaves
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content_tags.quantities[9]
quantity84% — targeted ChR2-expressing neurons located in layer V
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content_tags.quantities[10]
quantity1040-2444 neurones, 0.35-0.83% — Arch-expressing cells within 1 mm producing robust biases, as a fraction of the local population
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content_tags.quantities[11]
quantity~564 μ m — space constant of inactivation efficiency
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.
content_tags.quantities[12]
quantity25% — share of trials carrying photostimulation, capped to prevent adaptation
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content_tags.quantities[13]
quantity70±24 trials — stimulated trials per session, the reason single sessions were underpowered
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content_tags.quantities[14]
quantity45-75 dB (SPL) — the randomised intensity range blocking a loudness strategy
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content_tags.quantities[15]
quantity30 ms tones at 100 Hz — the cloud-of-tones stimulus timing
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content_tags.quantities[16]
variablelight activation (independent)
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content_tags.variables[0]
variabletarget octave (independent)
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content_tags.variables[1]
variablechoice bias (dependent)
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content_tags.variables[2]
variableresponse time (dependent)
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content_tags.variables[3]
variablesound intensity (control)
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content_tags.variables[4]
variablemask LED (control)
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content_tags.variables[5]
variablenumber of Arch-expressing neurones (covariate)
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content_tags.variables[6]
variablestimulus strength
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content_tags.variables[7]
variablechoice biases
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content_tags.variables[8]
variablepreferred frequency
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content_tags.variables[9]
variableresponse times
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content_tags.variables[10]
definitioncloud of tones stimulus
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content_tags.definitions[0]
definitionstimulus strength r
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content_tags.definitions[1]
definitionpreferred frequency
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content_tags.definitions[2]
definitionanti-bias
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content_tags.definitions[3]
definitioncloud of tones
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content_tags.definitions[5]
definitionanti-bias
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content_tags.definitions[6]
definitiontwo alternative choice
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content_tags.definitions[7]
definitionmotor component of the response time
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content_tags.definitions[8]
definitionmagnitude of modulation of response times
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content_tags.definitions[9]
key-termscorticostriatal, auditory cortex, striatum, perceptual decision-making, cloud-of-tones, two alternative choice, tonotopic, Channelrhodopsin-2, archaerhodopsin-3, psychometric curve, chronometric, choice biases, anti-bias, antidromic, compensatory plasticity, logistic regression, ideal observer, space constant, superior colliculus, reinforcement learning, tetrode, optical fiber, layer V
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.
logistic regression β as written: logistic regression model
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content_tags.statistics[0]
Wilcoxon signed-rank test p as written: signed-rank test
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content_tags.statistics[1]
linear regression (ordinary least squares) as written: linear regression using ordinary least squares method
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content_tags.statistics[2]
bootstrap as written: bootstrap resampling
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content_tags.statistics[3]
cross-validation as written: leave-one-out cross-validation
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content_tags.statistics[4]
confidence interval as written: Confidence intervals
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content_tags.statistics[5]
p-value p as written: p-values
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content_tags.statistics[6]
standard error as written: standard errors of regression coefficients
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content_tags.statistics[7]
mean squared error minimisation as written: minimized the mean square error
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content_tags.statistics[8]
logistic regression model
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content_tags.statistics[9]
signed-rank test
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content_tags.statistics[10]
ideal observer
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content_tags.statistics[11]
slope of the psychometric curve
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content_tags.statistics[12]
Second readings
a facet read a second time, independently, from its own sentence — and whether the two readings agree
claim typemechanism — a second reading reaching the same value; recorded: mechanism
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alternatives.claim_type
methodrats performing a two-alternative cloud-of-tones frequency discrimination, with tetrode recording and projection-targeted channelrhodopsin and archaerhodopsin in corticostriatal neurons — a second reading DIFFERING from the recorded value; recorded: electrophysiology
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alternatives.method
phenomenonperceptual decision-making - converting an auditory representation into a motor choice — a second reading DIFFERING from the recorded value; recorded: auditory perceptual decision-making
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alternatives.phenomenon

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