Dopamine Cells Respond to Predicted Events during Classical Conditioning: Evidence for Eligibility Traces in the Reward-Learning Network

Wei‐Xing Pan, Robert Schmidt, Jeffery R. Wickens, Brian I. Hyland · Journal of Neuroscience · 2005 · neuroscience · 10.1523/jneurosci.1478-05.2005 · record pan2005-dopamine-cells-respond-to-predicte
tag layer · index · concepts · authors · עברית

Where this record's text came from. faceted 2026-08 by reading the locally held text (html); every evidence string verified as an exact substring of that text before merging. The paper is a hybrid of in vivo electrophysiology and computational simulation; the single method facet was set to electrophysiology because the modelling is used to interpret recorded data, and the method_evidence sentence names both components. Claim type is 'mechanism' rather than 'critique' because the paper explicitly declines to treat its mismatch with earlier TD models as a refutation of TD, instead locating the discrepancy in parameter choice. The extraction contains PMC site furniture before the article body, but the body itself is clean prose. 2026-08-13 and 2026-08-25 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.

How to read this page. The article itself is not reproduced here — this paper is not cleared for redistribution, so what follows is the tag layer alone. The evidence sentences are withheld with the article. The table at the foot lists every tag the record carries; each row names the facet it belongs to, and the whose words column says whether the tag records the paper's own wording or the tagger's sentence about the paper. Every tag links to each other paper in the corpus carrying it. The machine-readable edition of the same thing is the TEI file beside this one, where the tags live in standOff, and the article is at its publisher via the link below. 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 WITHDRAWN AND REPLACED — the entry was wrong, and a corrected version of it appears in the entries below.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.

Other fields define these differently the same word, defined another way in another discipline
prediction error 6 fields · standard deviation 3 fields · latency 2 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 21 concept tags on this paper, none is grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.

DA cells prediction error temporal difference eligibility trace learning rate TD(0) TD(1) serial compound state vector discount factor conditioned omission modulation index apomorphine quinpirole substantia nigra pars compacta ventral tegmental area supramammillary nucleus random reward cued-reward licking A long-lasting eligibility trace keeps the weights of an earlier sensory signal open to modification for many time steps, so the reward-driven prediction error can be assigned back to the cue in one step instead of migrating one time step per trial A very low learning rate combined with a high eligibility-trace decay makes each trial change the weights only slightly, so responses to the cue and to the predicted reward coexist for a prolonged period before the reward response is suppressed Sustained firing in reverberating circuits, or biochemical processes at the synapse, could hold the seconds-long eligibility trace the model requires learning rate parameter complete serial compound stimulus TD(1) model TD(0) model oddball sequences decay parameter firing rate intertrial interval control paradigms latency t-test paired t test standard deviation (mean ± SD) standard error of the mean Error bars represent SEM. standard deviation threshold criterion histogram averaged peri-event time histograms SD electrophysiology

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second reading	`alternatives`	an independent re-reading of a facet, recorded whether or not it agrees
relation to other work	`claim_relations`	how this paper stands to another: agrees, contradicts, extends, qualifies
claim type	`claim_type`	the shape of the claim: existence, mechanism, method, critique, prediction, null
concept edge	`concept_edges`	an asserted relation between two concepts, with the sentence asserting it
content tag	`content_tags`	a finding, definition, quantity, statistic or variable named in the paper
contribution	`contribution`	what the paper claims to add
control	`controls`	an experiment run to eliminate a rival explanation
limitation	`limitations`	what the paper says its evidence cannot support
mechanism	`mechanisms`	how the reported effect is said to come about
method	`method`	how the paper produced its evidence
object of study	`object_of_study`	what the paper is about — the thing it investigates
phenomenon	`phenomenon`	the effect or regularity under study
research question	`research_question`	the question the paper sets itself
sense	`senses`	a distinct use of a term, quoted where the paper fixes it
transfer	`transfers`	a method or concept carried in from another field

The paper

The full text is not reproduced here. This article is not cleared for redistribution, so the page carries the tag layer only. Its evidence sentences are withheld; the grounding status remains with each tag, and the article itself is at https://doi.org/10.1523/jneurosci.1478-05.2005. The reading view exists for the papers whose licence permits it.

Every tag on this page

alternatives 3 · claim_relations 10 · claim_type 1 · concept_edges 6 · content_tags 68 · contribution 1 · controls 1 · limitations 7 · mechanisms 8 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 5 · transfers 4

Relations on this page carry a quotation for the claim. Where a relation was separately evidenced — and so tested against the vocabulary for its type — the row says so; none of 6 relations here carry that mark. The rest are the tagger’s reading of the sentence quoted beside them.

facet	tag	field	conf.	whose words
object_of_study	how the dopamine response moves from reward to predictive cue during learning, and whether the temporal difference algorithm can produce that time course Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`object_of_study`		the paper's words
research_question	whether dopamine cells show the stepwise latency migration that published TD models predict, and if not, which parameter settings would reproduce what they do show Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`research_question`		the paper's words
contribution	shows cue and reward responses coexist rather than trading off, then finds the pattern is reproduced only by long eligibility traces and a low learning rate - turning physiology into a constraint on the algorithm's parameters Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`contribution`		the paper's words
claim_type	mechanism Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_type`		the paper's words
phenomenon	dopamine reward-prediction-error signalling during classical conditioning Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`phenomenon`		the paper's words
method	electrophysiology Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: electrophysiology	`method`		the paper's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: A long-lasting eligibility trace keeps the weights of an earlier sensory signal open to modification for many time steps, so the reward-driven prediction error can be assigned back to the cue in one step instead of migrating one time step per trial	`mechanisms[0]`		the tagger's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: A very low learning rate combined with a high eligibility-trace decay makes each trial change the weights only slightly, so responses to the cue and to the predicted reward coexist for a prolonged period before the reward response is suppressed	`mechanisms[1]`		the tagger's words
mechanisms	mechanisms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: Sustained firing in reverberating circuits, or biochemical processes at the synapse, could hold the seconds-long eligibility trace the model requires	`mechanisms[2]`		the tagger's words
mechanisms	an eligibility trace keeps earlier time steps modifiable, so a cue several steps back can be updated in one trial instead of waiting for the error to migrate Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[3]`	1.0	the tagger's words
mechanisms	with a low learning rate each trial moves the weights only slightly, which is what lets cue and reward responses coexist for hundreds of trials Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[4]`	1.0	the tagger's words
mechanisms	under TD(0) the error must arrive one step earlier per trial, so reaching a cue k steps back takes at least k trials Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[5]`	1.0	the tagger's words
mechanisms	candidate substrates for a seconds-long trace are reverberating circuits and synaptic biochemistry Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[6]`	0.6	the tagger's words
mechanisms	long traces with slow learning is the combination machine learning already found to give early but stable learning, which may be why biology uses it Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`mechanisms[7]`	0.6	the tagger's words
limitations	The fitted parameter values are not claimed to be fixed neural constants, since a different model structure could yield different best-fitting values. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the fitted parameter values: OUR RESULTS SHOULD NOT BE UNDERSTOOD AS PROVIDING EXPLICIT FIXED VALUES FOR NEURAL TD LEARNING PARAMETERS, BECAUSE THE SPECIFIC VALUES ... MAY BE DIFFERENT IF DETAILS OF THE MODEL STRUCTURE WERE CHANGED	`limitations[0]`		the tagger's words
limitations	The stimulus representation assumed by the model (serial compound) is not known to be how the brain represents stimuli for reward learning. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the timing predictions in particular: IT IS NOT KNOWN HOW SENSORY STIMULI ARE REPRESENTED IN THE BRAIN FOR REWARD LEARNING	`limitations[1]`		the tagger's words
limitations	the paper refuses the strong reading of its own result: these are not fixed values for neural parameters, because a different model structure would shift them Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the same scope as limitations[0] - the values are a property of this model structure	`limitations[2]`	1.0	the tagger's words
limitations	how the brain represents stimuli in time is unknown; the serial compound representation is a convention inherited from earlier models Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the same scope as limitations[1], with the convention named: WE FOLLOWED MANY PREVIOUS STUDIES IN USING A SERIAL COMPOUND REPRESENTATION OF CONDITIONED STIMULI	`limitations[3]`	1.0	the tagger's words
limitations	the animals had already had hundreds of solenoid-reward pairings before recording, so the click was not a neutral event at the start of conditioning Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: any reading of the click as a neutral cue: ANIMALS WERE THEREFORE EXPOSED TO HUNDREDS OF SOLENOID-REWARD PAIRINGS BY THE TIME THE RECORDINGS WERE MADE	`limitations[4]`	1.0	the tagger's words
limitations	the negative prediction error signal is small and hard to measure against an already low baseline rate Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: the negative prediction-error signal: THE AMPLITUDE OF INHIBITIONS WAS SMALL, CONSISTENT WITH THE FACT THAT INHIBITORY RESPONSES CAN BE DIFFICULT TO DETECT AND QUANTIFY IN NEURONS THAT ALREADY HAVE A LOW BASELINE RATE	`limitations[5]`	1.0	the tagger's words
limitations	21 of 45 electrophysiologically plausible cells failed the pharmacological test and were discarded Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. what this limits: what a 'dopamine neuron' means here, and in every study that skips the drug test: FROM 45 NEURONS THAT WERE POSSIBLY DOPAMINERGIC ON ELECTROPHYSIOLOGICAL CRITERIA, 24 ... WERE CONFIRMED DA NEURONS BASED ON HAVING A CLEAR INHIBITORY RESPONSE TO DOPAMINE AGONIST DRUGS	`limitations[6]`	1.0	the tagger's words
transfers	The temporal difference algorithm and its prediction-error signal, taken from machine learning, are used as the model of midbrain dopamine cell activity. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[0]`		the tagger's words
transfers	the temporal difference algorithm, imported as a candidate account of what dopamine cells compute Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[1]`	1.0	the tagger's words
transfers	the century-old requirement that some trace of the cue survive until the reward arrives, here identified with the eligibility trace Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[2]`	1.0	the tagger's words
transfers	the known trade-off that high lambda with low alpha buys stable early learning, used here as an adaptive argument Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`transfers[3]`	0.9	the tagger's words
controls	electrode track positions confirmed histologically after the experiments were completed Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. eliminates: attributing the responses to dopamine cells that were not where the recordings were thought to be - the confirmation comes after, from the tissue, not from the coordinates · outcome (stated by the tagger, no quotation): held	`controls[0]`	0.85	the tagger's words
senses	presumed DA cells are a doubly filtered category: rate below 10 Hz and spike at least 1 ms, THEN more than 50% inhibition by a dopamine agonist. Cell identity is pharmacological here, not genetic or anatomical Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[0]`	1.0	the tagger's words
senses	TD(0), TD(1) and TD(lambda) are three different models, not one - the value of lambda changes which weights an error can touch, and the published DA models were TD(0) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[1]`	1.0	the tagger's words
senses	prediction error here is the scalar difference between actual reward and the temporal difference of successive predictions, with the NEGATIVE side deliberately clipped to match dopamine cells' floor Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[2]`	1.0	the tagger's words
senses	early and late training are defined by block count - 6 or fewer against 10 or more - and the paper's central result holds only early Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[3]`	0.9	the tagger's words
senses	the recorded population pools substantia nigra pars compacta, ventral tegmental area and supramammillary nucleus, justified by a null on regional differences Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`senses[4]`	1.0	the tagger's words
alternatives	claim_type: mechanism Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. a second reading reaching the same value. Recorded: mechanism — note: a second reading reaching the same value from a different sentence. Not applied - a person decides.	`alternatives.claim_type`		the tagger's words
alternatives	method: extracellular single-unit recording from pharmacologically identified midbrain dopamine cells during first acquisition, paired with a systematic sweep of the TD algorithm's parameter space Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. a second reading DIFFERING from the recorded value. Recorded: electrophysiology — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.	`alternatives.method`		the tagger's words
alternatives	phenomenon: the shift of dopamine cell firing from unexpected reward to the cue that predicts it Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. a second reading DIFFERING from the recorded value. Recorded: dopamine reward-prediction-error signalling during classical conditioning — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.	`alternatives.phenomenon`		the tagger's words
claim_relations	builds-on → Montague et al. (1996) temporal-difference model of dopamine activity Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[0]`		the tagger's words
claim_relations	contradicts → previous TD models predicting a gradual stepwise latency shift (Montague et al., 1996; Schultz et al., 1997) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: the time course of the latency shift: previous TD models PREDICT A GRADUAL STEPWISE SHIFT IN LATENCY WITH RESPONSES TO REWARDS LOST BEFORE RESPONSES DEVELOP TO THE CONDITIONED CUE	`claim_relations[1]`		the tagger's words
claim_relations	extends → Miller et al., 1981; Kosobud et al., 1994; Kiyatkin and Rebec, 2001; Hyland et al., 2002 (rat DA cell responses to reward-predicting cues) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[2]`		the tagger's words
claim_relations	contradicts → the stepwise latency migration predicted by TD(0) models of dopamine activity Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: the latency profile of the new response: NEW CONDITIONED RESPONSES MADE THEIR FIRST APPEARANCE AT A CONSTANT, SHORT LATENCY AFTER PREDICTIVE CUES, and DEVELOPED RAPIDLY, WITHIN A FEW TRIALS - not the gradual migration the models expect	`claim_relations[3]`	1.0	the tagger's words
claim_relations	contradicts → the prediction that a redundant intermediate cue draws no response Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[4]`	1.0	the tagger's words
claim_relations	contradicts → the prediction that reward responses vanish before cue responses appear Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[5]`	1.0	the tagger's words
claim_relations	reconciles → the temporal difference framework with the recorded time course, by relocating the disagreement to parameter choice Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: the model's free parameters: THESE APPARENT DISPARITIES BETWEEN TD MODEL PREDICTION ERRORS AND DA CELL ACTIVITY ARE ATTRIBUTABLE TO THE PARTICULAR CHOICE OF PARAMETERS USED IN THE MODEL - the settings for ALPHA and LAMBDA, not the framework	`claim_relations[6]`	1.0	the tagger's words
claim_relations	critiques → published primate datasets, which on re-inspection never supported gradual migration either Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. differs in: gradual migration against simultaneous responding, and the stage of learning: the existing primate data PROVIDES NO SUPPORT FOR THE GRADUAL MIGRATION OF A RESPONSE DURING LEARNING AND SUGGESTS THAT DA CELLS SHOW SIMULTANEOUS RESPONDING TO BOTH CUE AND REWARD, AT LEAST EARLY IN LEARNING	`claim_relations[7]`	1.0	the tagger's words
claim_relations	confirms → earlier reports that reward responses are diminished in highly trained animals Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[8]`	0.9	the tagger's words
claim_relations	extends → rat studies showing dopamine responses to reward-predicting cues, by showing they arise during acquisition Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`claim_relations[9]`	1.0	the tagger's words
concept_edges	prediction error —operationalised-as→ DA cells Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 · note: the identification is asymmetric by construction: firing can rise to 100 Hz but only fall to zero from a 5 Hz baseline Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span says DA cells' baseline spiking can be equated to zero prediction error; operationalised-as is loose. also in: prediction error · DA cells	`concept_edges[0]`	1.0	the tagger's words
concept_edges	TD(0) —is-a→ temporal difference Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'TD(0) model'; 'temporal difference' is an abbreviation variant. also in: TD(0) · temporal difference	`concept_edges[2]`	1.0	the tagger's words
concept_edges	TD(1) —is-a→ temporal difference Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span writes 'TD(1) model'; 'temporal difference' is an abbreviation variant. also in: TD(1) · temporal difference	`concept_edges[3]`	1.0	the tagger's words
concept_edges	apomorphine —operationalised-as→ DA cells Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 · note: direction is test-to-category: more than 50% inhibition by the agonist is what makes a recorded cell count as dopaminergic Audit 2026-09-28: DOUBTFUL - read against its span and kept, but apomorphine inhibition was an inclusion criterion for DA cells; operationalised-as is loose. also in: apomorphine · DA cells	`concept_edges[5]`	1.0	the tagger's words
concept_edges	serial compound —operationalised-as→ state vector Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 1 also in: serial compound · state vector	`concept_edges[8]`	1.0	the tagger's words
concept_edges	omission —operationalised-as→ prediction error Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. strength of the relation: 0.8 · note: the negative side of the signal is measured by withholding what was predicted Audit 2026-09-28: DOUBTFUL - read against its span and kept, but responses to cue omission were analysed; operationalised-as 'prediction error' is interpretive. also in: omission · prediction error	`concept_edges[9]`	0.9	the tagger's words
content_tags	t-test Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: t-test	`content_tags.statistics[0]`		the tagger's words
content_tags	standard deviation Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: standard deviation	`content_tags.statistics[1]`		the tagger's words
content_tags	standard error of the mean Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: standard error of the mean	`content_tags.statistics[2]`		the tagger's words
content_tags	standard deviation threshold criterion Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: standard deviation threshold criterion	`content_tags.statistics[3]`		the tagger's words
content_tags	histogram Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: histogram	`content_tags.statistics[4]`		the tagger's words
content_tags	paired t test Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: paired t test	`content_tags.statistics[5]`	1.0	the tagger's words
content_tags	modulation index Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: modulation index	`content_tags.statistics[6]`	0.8	the tagger's words
content_tags	SD Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: SD	`content_tags.statistics[7]`	0.9	the tagger's words
content_tags	Dopamine cells kept responding to reward long after they developed responses to the predictive cue. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[0]`		the tagger's words
content_tags	New conditioned responses appeared straight away at their final short latency after the cue, with no gradual backward drift of latency from reward to cue. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[1]`		the tagger's words
content_tags	Every cell trained with two cues in sequence responded clearly to the second, seemingly redundant cue, and that response persisted for the whole experiment. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[2]`		the tagger's words
content_tags	Cells recorded after many sessions of cue-reward pairing no longer responded to a predictable reward, unlike the cells recorded early in training. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[3]`		the tagger's words
content_tags	Acquiring a cue response and losing the reward response run on different time courses rather than being two sides of one shift. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[4]`		the tagger's words
content_tags	Omitting an expected reward significantly reduced firing rate. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[5]`		the tagger's words
content_tags	Omitting the second cue restored reward responses to the level seen with random reward. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[6]`		the tagger's words
content_tags	With the chosen parameter values, the TD model's prediction error reproduced key elements of the recorded dopamine pattern. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[7]`		the tagger's words
content_tags	A high eligibility-trace decay parameter was the one setting that let the model learn cue significance without stepwise trial-by-trial migration of the response from reward to cue. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[8]`		the tagger's words
content_tags	Once the eligibility-trace decay fell below about 0.6, stepwise migration of the prediction error reappeared no matter what learning rate was used. Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[9]`		the tagger's words
content_tags	conditioned responses appeared at their final short latency within a few trials, with no gradual migration Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[10]`	1.0	the tagger's words
content_tags	the redundant intermediate cue drew a clear response in every two-cue cell, which persisted Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[11]`	1.0	the tagger's words
content_tags	responses to predicted reward persisted after cue responses had developed, in every cell recorded early in training Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[12]`	1.0	the tagger's words
content_tags	responses to predicted reward are lost completely only after training beyond what first establishes cue responses, so the two changes have different time courses Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[13]`	1.0	the tagger's words
content_tags	those late-training cells still responded robustly to reward delivered without cues Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[14]`	1.0	the tagger's words
content_tags	omitting the second cue restored the reward response to its uncued level, so the redundant cue was doing predictive work Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[15]`	1.0	the tagger's words
content_tags	TD with eligibility zero and a fast learning rate reproduced the published pattern, with no second-cue response and no overlap Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[16]`	1.0	the tagger's words
content_tags	TD with lambda 0.9 and alpha 0.005 reproduced the recorded pattern, including the coexistence and the intermediate cue response Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[17]`	1.0	the tagger's words
content_tags	lambda below about 0.6 brought stepwise migration back for every value of alpha, making it the critical parameter Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[18]`	1.0	the tagger's words
content_tags	omitted rewards produced a significant firing decrease but omitted cues did not Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[19]`	1.0	the tagger's words
content_tags	CONTROL NULL: no regional difference in response pattern, which is what licensed pooling the three midbrain sites Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[20]`	1.0	the tagger's words
content_tags	BUILT IN, NOT PREDICTED: the model's small negative errors follow from the clipping imposed to match dopamine cells' floor, as the authors state Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[21]`	1.0	the tagger's words
content_tags	re-inspection of published primate datasets found no support for gradual migration there either Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.findings[22]`	1.0	the tagger's words
content_tags	eligibility trace Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: eligibility trace	`content_tags.definitions[0]`		the tagger's words
content_tags	temporal difference Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: temporal difference	`content_tags.definitions[1]`		the tagger's words
content_tags	learning rate parameter Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: learning rate parameter	`content_tags.definitions[3]`		the tagger's words
content_tags	complete serial compound stimulus Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: complete serial compound stimulus	`content_tags.definitions[6]`		the tagger's words
content_tags	temporal difference Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: temporal difference	`content_tags.definitions[7]`	1.0	the tagger's words
content_tags	prediction error Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: prediction error	`content_tags.definitions[8]`	1.0	the tagger's words
content_tags	eligibility trace Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: eligibility trace	`content_tags.definitions[9]`	1.0	the tagger's words
content_tags	serial compound Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: serial compound	`content_tags.definitions[10]`	1.0	the tagger's words
content_tags	modulation index Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: modulation index	`content_tags.definitions[11]`	1.0	the tagger's words
content_tags	TD(0) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: TD(0)	`content_tags.definitions[12]`	1.0	the tagger's words
content_tags	TD(1) model Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: TD(1) model	`content_tags.definitions[13]`		the tagger's words
content_tags	TD(0) model Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: TD(0) model	`content_tags.definitions[14]`		the tagger's words
content_tags	oddball sequences Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: oddball sequences	`content_tags.variables[1]`		the tagger's words
content_tags	decay parameter Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: decay parameter	`content_tags.variables[2]`		the tagger's words
content_tags	firing rate Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: firing rate	`content_tags.variables[4]`		the tagger's words
content_tags	discount factor Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: discount factor	`content_tags.variables[5]`		the tagger's words
content_tags	intertrial interval Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: intertrial interval	`content_tags.variables[6]`		the tagger's words
content_tags	control paradigms Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: control paradigms	`content_tags.variables[7]`		the tagger's words
content_tags	learning rate Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: learning rate	`content_tags.variables[9]`	1.0	the tagger's words
content_tags	eligibility trace Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: eligibility trace	`content_tags.variables[10]`	1.0	the tagger's words
content_tags	latency Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: latency	`content_tags.variables[11]`	1.0	the tagger's words
content_tags	omission Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above. also in: omission	`content_tags.variables[12]`	1.0	the tagger's words
content_tags	24 of 45, from 11 rats — cells that passed pharmacological confirmation as dopamine neurons out of those meeting the electrophysiological criteria Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[0]`		the tagger's words
content_tags	5.7 ± 3.5 Hz — average baseline firing rate of the confirmed dopamine cells Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[1]`		the tagger's words
content_tags	8 of 11 — cells tested in which conditioned responses to the cue developed during recording Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[2]`		the tagger's words
content_tags	74 ± 33 ms; 53 ± 9 ms — latencies of the newly conditioned excitatory and inhibitory responses to the cue Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[3]`		the tagger's words
content_tags	-0.12 ± 0.17 — average modulation index at the time of an omitted second cue, a nonsignificant departure from baseline Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[4]`		the tagger's words
content_tags	-0.21 ± 0.20; p < 0.01; n = 11 — average modulation index when an expected reward was omitted, a significant reduction from baseline Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[5]`		the tagger's words
content_tags	∼0.6 — threshold value of the eligibility-trace decay parameter below which stepwise migration of the prediction error reappears Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[6]`		the tagger's words
content_tags	-0.05 — ceiling imposed on the model's negative prediction error, to match the limited range over which dopamine cells can be inhibited Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[7]`		the tagger's words
content_tags	0.9 and 0.005 — the lambda and alpha that reproduced the recorded dopamine pattern Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[8]`	0.9	the tagger's words
content_tags	about 0.6 — the lambda threshold below which stepwise migration reappears for every alpha Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[9]`	1.0	the tagger's words
content_tags	-0.05 — the imposed floor on negative prediction error, matching dopamine cells' limited inhibitory range Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[10]`	1.0	the tagger's words
content_tags	24 of 45 — electrophysiologically plausible cells that passed the pharmacological identification Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[11]`	1.0	the tagger's words
content_tags	5.7 Hz, 1.5 ms — mean firing rate and spike duration of the accepted dopamine cells Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[12]`	1.0	the tagger's words
content_tags	-0.21 vs -0.12 — modulation index for omitted reward (significant) against omitted cue (not) Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[13]`	1.0	the tagger's words
content_tags	74 ms and 53 ms — latency of the new conditioned excitations and inhibitions Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.	`content_tags.quantities[14]`	1.0	the tagger's words

Interpretive choices

Tags that could not be placed in the text:

none — every tag was located in the text

What was deliberately not tagged, and why, is recorded per paper by the tagger. Where a paper has no such record, this page says so rather than implying that nothing was declined.