Mapping of a non-spatial dimension by the hippocampal–entorhinal circuit — TEI-tagged view

Aronov, Nevers, Tank · 2017 · neuroscience · doi:10.1038/nature21692 · 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. The paper is a Nature report whose citations are all numeric, so no prior work is named in the running text; the single claim_relation recorded is one where the relation is stated explicitly ('A conceptual framework... We tested this idea'), not inferred from a bare citation. The paper also positions itself against the view that physical space is primary for this circuit ('...rather than a primacy of physical space for this network 9 , 10 , 18 – 21 .'), but that target is a position rather than a named study, so no 'contradicts' relation was recorded. Claim type could arguably be 'mechanism' given the closing claim that a common circuit mechanism is shared across tasks ('Critically, spatial and non-spatial representations are produced by the same neuronal population, suggesting a common circuit mechanism...'), but the load-bearing result is the demonstration that frequency fields exist, hence 'existence'. 2026-08-13 and 2026-08-14 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.
14 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
agency 33 fields · salience 21 fields · reward 16 fields · principal component analysis 5 fields · coefficient of variation 4 fields · information 4 fields · and 9 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
place cell6 grid cell2 border cell head direction2 time cell2 interneuron24 firing field CA116 entorhinal6 hippocampal6 SMT random foraging passive playback linear track task13 frequency16 location5 spatial4 non-spatial navigation3 cognitive map3 continuous3 mechanism4 population15 modulated information7 spatial information3 grid score shuffled3 percentile2 environment2 module3 theta15 speed2 SWR joystick press2 release trial3 duration5 octave2 reward12 salience8 agency15 behavioral context2 model22 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
phenomenonhippocampal-entorhinal coding of a non-spatial continuous dimension— “We tested this idea by recording hippocampal and entorhinal neurons in a task that required rats to use a joystick to manipulate sound along a continuous frequency axis.”
claim-typeexistence— “We found neural representation of the entire behavioral task, including activity that formed discrete firing fields at particular sound frequencies.”
method — evidence— “We recorded 2208 units in the dorsal CA1 of 5 rats and 1164 units in the dorsal MEC of 9 rats ( Extended Data Fig. 2 ).”
object of study — evidence— “Our results therefore suggest that the well-known spatial patterns in the hippocampal/entorhinal circuit may be a consequence of the continuous nature of the relevant task variables (e.g., location), rather than a primacy of physical space for this network”
research question — evidence— “However, the hippocampal/entorhinal circuit is involved not only in spatial navigation, but in a variety of memory-guided behaviors 6 . The relationship between this general function and the specialized spatial activity patterns is unclear. A conceptual framework reconciling these views is that spatial representation is just one example of a…
contribution — evidence— “We tested this idea by recording hippocampal and entorhinal neurons in a task that required rats to use a joystick to manipulate sound along a continuous frequency axis. We found neural representation of the entire behavioral task, including activity that formed discrete firing fields at particular sound frequencies. Neurons involved in this…
transferfrom spatial navigation electrophysiology: place cells, grid cells, border cells and head direction cells, carried into a task with no spatial dimension— “SMT firing fields of both place cells and grid cells were similar to those of other cells and spanned the entire task, including periods when the rat was immobile ( Extended Data Fig. 10 ). Other spatial cell types, including border cells and head direction cells, were also SMT-modulated (e.g. cell 9 in Fig. 4b ; Extended Data Fig. 10 ).…transfers[0]
transferfrom spatial navigation electrophysiology: the spatial information measure, applied unchanged to time bins instead of spatial bins— “To detect place cells, we calculated “spatial information” 34 – the mutual information rate between spikes and location – using the same formula as above ( Eqn. 2 ), but using the 1600 spatial bins instead of time bins. Place cells were defined as cells for which the information rate exceeded 99% of the values for the shuffled samples.”transfers[1]
transferfrom auditory psychophysics: the perceptually uniform logarithmic frequency axis, used as the continuous dimension to be traversed— “To test this idea, we designed a “sound manipulation task” (SMT), in which rats changed the frequency of sound in their environment ( Fig. 1a ). Animals deflected a joystick to activate a pure tone produced by a sound speaker. They continued deflecting in order to increase frequency along a perceptually uniform logarithmic axis 22 . Rewards…transfers[2]
transferfrom human fMRI of cognitive spaces: the claim that continuous non-spatial dimensions are mapped by this circuit, brought down to single units— “However, the hippocampal/entorhinal circuit is involved not only in spatial navigation, but in a variety of memory-guided behaviors 6 . The relationship between this general function and the specialized spatial activity patterns is unclear. A conceptual framework reconciling these views is that spatial representation is just one example of a…transfers[3]
mechanismSpatial firing arises because location happens to be a continuous task-relevant variable, not because physical space is privileged for this circuit— “Our results therefore suggest that the well-known spatial patterns in the hippocampal/entorhinal circuit may be a consequence of the continuous nature of the relevant task variables (e.g., location), rather than a primacy of physical space for this network”mechanisms[0]
mechanismPerforming a task drives a sequence of firing fields that are elicited parametrically as behaviour progresses— “In this framework, task performance activates a sequence of neural activity, in which firing fields are elicited parametrically with progress through behavior.”mechanisms[1]
mechanismOverlap between neighbouring fields encodes the order and adjacency of behavioural states, supporting episodic linking and planning— “Neighboring and partially overlapping fields therefore represent the order and adjacency of behavioral states.”mechanisms[2]
mechanismtraversal speed is randomised trial by trial so that sound frequency and elapsed time are uncoupled, which is what makes a frequency field distinguishable from a time field— “To uncouple frequency from the amount of elapsed time, we randomly varied, across trials, the “speed” of frequency traversal. Resulting trials varied in duration by up to a factor of 2, on average in the range of 5–10 s.”mechanisms[3]
mechanisma model tested the strength of alignment of the neural activity to various task events, and each field was found to align best either to particular sound frequencies or to the press or the release of the joystick— “To quantify this frequency locking, we implemented a model that tested the strength of alignment of the neural activity to various task events ( Extended Data Fig. 7 ). Of the fields that occurred during sound presentation, more than half in both CA1 and MEC aligned best to particular sound frequencies, whereas the rest were more strongly…mechanisms[4]
mechanismthe proposed general mechanism is that task performance activates a progression of firing fields, so that neighbouring and overlapping fields represent the order and adjacency of behavioural states— “However, our observations lead to an intriguing conjecture that in more complex tasks (e.g. those containing memory-guided decision points), the hippocampal/entorhinal system might similarly represent arbitrary behavioral states. In this framework, task performance activates a sequence of neural activity, in which firing fields are elicited…mechanisms[5]
mechanismand the absence of field broadening across the trial is attributed to an informative sensory variable being continuously available, unlike in a delay task where the code can drift— “Note that fields in the SMT did not progressively broaden during the delay period, as they typically do in time cells; this may be due to the fact that an informative sensory variable (sound frequency) was always available to the animal, preventing a drift in the neural code.”mechanisms[6]
builds-on the cognitive-map framework (refs 7-10) holding that spatial coding is one case of a general mechanism for continuous task-relevant variables, including recent human fMRI work on cognitive spaces— “A conceptual framework reconciling these views is that spatial representation is just one example of a more general mechanism for encoding continuous, task-relevant variables 7 – 10 .”claim_relations[0]
generalises the cognitive map account, from locations to any continuous task-relevant variable— “However, the hippocampal/entorhinal circuit is involved not only in spatial navigation, but in a variety of memory-guided behaviors 6 . The relationship between this general function and the specialized spatial activity patterns is unclear. A conceptual framework reconciling these views is that spatial representation is just one example of a…claim_relations[1]
reinterprets spatial firing itself, as a consequence of the continuity of the task variable rather than of space— “Our results therefore suggest that the well-known spatial patterns in the hippocampal/entorhinal circuit may be a consequence of the continuous nature of the relevant task variables (e.g., location), rather than a primacy of physical space for this network”claim_relations[2]
extends the non-spatial modulation literature, from discrete stimuli and time points to a continuously traversed dimension— “Consistent with this idea, the firing of hippocampal cells is modulated not only by location, but also by non-spatial variables, including sensory, behavioral and internal parameters 12 , 13 . For…claim_relations[3]
distinguishes-from time cells, whose fields broaden across a delay where these did notclaim_relations[4]
qualifies the reported relationship between theta frequency and running speed, which did not appear when speed was not locomotion
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claim_relations[5]
builds-on the grid module literature, with wide-spacing modules producing wide fields in this task too
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claim_relations[6]
confirms the previously reported dependence of grid cell detection on environment size
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claim_relations[7]
senseA CELL TYPE IS NAMED AFTER THE TASK IT WAS DISCOVERED IN: the well-known spatial patterns may be a consequence of the continuous nature of the task variable, rather than a primacy of physical space for this network
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senses[0]
senseand the same neurons produce the same discrete fields on an axis of sound frequency, so spatial and non-spatial representations come from one population
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senses[1]
senseMEMBERSHIP OF A NAMED CLASS BARELY PREDICTS THE SAME CELL IN ANOTHER TASK: the amount of task modulation had a very weak correlation with placeness and gridness, r2 about 0.005, which is read as nearly independent activity patterns
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senses[2]
senseTHE DEFINITION OF THE TYPE IS A THRESHOLD ON THIS DATASET'S OWN SHUFFLES: grid cells are cells whose grid score exceeds the 95th percentile of the shuffled samples, which in this dataset was 0.46
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senses[3]
senseand place cells are cells whose spatial information exceeds 99% of their own shuffled values, so both categories are defined relative to the recording rather than absolutely
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senses[4]
senseAND THE COUNT OF A CELL TYPE DEPENDS ON THE ARENA: fewer grid cells were detected in the smaller environment than the larger one, so every grid comparison was re-verified on the subset recorded in the larger environment
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senses[5]
senseAN EXCLUSION CRITERION THAT DOES NOT EXIST OUTSIDE THE TASK IT WAS BUILT FOR: sharp-wave ripples are normally removed by rejecting low-velocity time points, and with no velocity in this task they had to be detected directly in the field potential
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senses[6]
sensewith the negative disclosure attached: excluding those spikes did not qualitatively change any result, it only raised the in-field to background ratio
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senses[7]
senseTHE SPATIAL CONFOUND IS DESIGNED OUT RATHER THAN CORRECTED FOR: rats moved their heads while deflecting the joystick in the first apparatus, so a nosepoke was added in the second
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senses[8]
senseTHETA DID NOT TRACK SPEED HERE, UNLIKE IN NAVIGATION, which is read as the navigation relationship being dependent on locomotion-related signals
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senses[9]
senseTHE CONTROL IS DECOMPOSED RATHER THAN RUN ONCE: passive playback alone engaged almost no cells, passive playback with a reward engaged a fifth of them with wider fields, so context matters and agency is not strictly required
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senses[10]
senseand a frequency field is called analogous to a place field, naming the new thing after the old one in the same sentence that shows the old name was too narrow
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senses[11]
controlmodifications to the apparatus - a nosepoke aligned with the joystick handle - introduced in the second version — held — rules out the possible SPATIAL CONFOUND of rats moving their heads while deflecting the joystick. It matters because the paper's claim is that these cells map a NON-SPATIAL dimension, so an uncontrolled head movement would make the result the very thing the paper denies
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controls[0]
differs intime cells, whose fields broaden across a delay where these did not — THE AXIS IS WHETHER THE FIELDS BROADEN ACROSS A DELAY, AND THE EXPLANATION IS GIVEN. 'fields in the SMT did not progressively broaden during the delay period, as they typically do in time cells; this may be due to the fact that an informative sensory variable (sound frequency) was always available to the animal, preventing a drift in the neural code.' The broadening is not denied of time cells - it is attributed to the ABSENCE of an external reference, which this task supplies.
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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.
claim_relations[4]
differs inthe reported relationship between theta frequency and running speed, which did not appear when speed was not locomotion — THE AXIS IS THE DOMAIN. 'Unlike in spatial navigation, theta frequency did not correlate to speed; this may imply that the relationship between theta and speed during navigation is' [specific to navigation]. The qualification is that an established correlation is domain-bound - and the evidence for it is a NULL (slope not significantly different from 0, p = 0.70), so the row also carries the polarity gap: see an internal file.
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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.
claim_relations[5]
auditlocation —is-a→ continuous — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span names location as a continuous task variable; is-a is loose.
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concept_edges[1]
notelocation —is-a→ continuous — location is given as an EXAMPLE of a continuous task variable, which is the whole demotion
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concept_edges[1]
auditborder cell —part-of→ SMT — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but border cells were SMT-MODULATED; part-of a task is loose.
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concept_edges[9]
auditplace cell —part-of→ SMT — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but 25.1% of place cells were SMT-MODULATED; part-of a task is loose.
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concept_edges[10]
noteplace cell —part-of→ SMT — a quarter of the place cells and a third of the grid cells were modulated in the sound task
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concept_edges[10]
auditgrid cell —part-of→ SMT — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but 34.3% of grid cells PARTICIPATED in the SMT; part-of a task is loose.
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concept_edges[11]
auditrandom foraging —contrasted-with→ SMT — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but cells were recorded in both the random-foraging task and the SMT and some fired in both; contrasted-with is loose.
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concept_edges[12]
noterandom foraging —contrasted-with→ SMT — the same cells recorded in both, with a mixed representation in CA1 - some cells in one task only, some producing fields in both
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concept_edges[12]
notegrid cell —operationalised-as→ grid score — THE DEFINITION IS A THRESHOLD ON THIS DATASET'S OWN SHUFFLES: the 95th percentile of the shuffled samples, which here was 0.46
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auditgrid score —operationalised-as→ percentile — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the grid score is compared against shuffled percentiles; operationalised-as is loose.
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concept_edges[16]
noteplace cell —operationalised-as→ spatial information — and place cells are those whose information rate exceeds 99% of their own shuffled values
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concept_edges[18]
auditspatial information —operationalised-as→ location — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but spatial information is defined via location; operationalised-as is loose.
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concept_edges[19]
notespatial information —operationalised-as→ location — the same formula as the task-phase information, run over 1600 spatial bins instead of time bins
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concept_edges[19]
auditSWR —part-of→ CA1 — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but some CA1 spikes occurred DURING sharp-wave ripple events; part-of is loose.
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concept_edges[26]
notehead direction —distinct-from→ SMT — and the residual head-direction confound was tested directly, since those cells overlap the modulated population
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concept_edges[29]
edgetime cell —contrasted-with→ SMT
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concept_edges[33]
notetask —contrasted-with→ passive playback — THE CONTROL SEPARATES THE STIMULUS FROM THE TASK: 1.7% of the same CA1 cells responded to the identical sweeps outside the task, against 31.1% inside it
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concept_edges[34]
auditreward —operationalised-as→ salience — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says salience was increased by delivering a reward; the direction of operationalised-as is loose.
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concept_edges[35]
notereward —operationalised-as→ salience — and the control is then run a second time WITH a reward, recovering a fifth of the cells
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concept_edges[35]
notebehavioral context —causes→ firing field — fields in the rewarded playback task were more than twice as wide, so context affected both how many cells were activated and how precise they were
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concept_edges[37]
auditfrequency —causes→ release — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says sound frequency influenced when rats released the joystick; the endpoint 'release' is terse.
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concept_edges[58]
notefrequency —causes→ release — releases were predicted by frequency alone with almost no added influence of elapsed time
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concept_edges[58]
auditfrequency —distinct-from→ duration — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span compares variability of durations and frequencies; distinct-from is loose.
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concept_edges[59]
notefrequency —distinct-from→ duration — trial durations were three times more variable than release frequencies, which is how the strategy is identified
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concept_edges[59]
auditoctave —operationalised-as→ frequency — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span measures frequency in octaves; the direction of operationalised-as is reversed.
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concept_edges[62]
noteoctave —operationalised-as→ frequency — and the unit is RELATIVE - octaves above a 2 kHz start, so the number does not join without its reference
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concept_edges[62]
notegrid cell —part-of→ module — and the module a cell belongs to predicts its field width in the SOUND task as well as in the arena
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concept_edges[68]
outcome (stated by the tagger, no quotation)control — heldcontrols[0]
for which uselimitation — the hexagonal-lattice interpretation only: THE SMALL NUMBER OF FIELDS PRODUCED BY GRID CELLS in the sound task makes the 1-dimensional-slice reading untestable here
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limitations[0]
for which uselimitation — every functional claim about the acoustic representation: IN THE SMT, RATS DID NOT ACTUALLY NEED TO REPRESENT THE STRUCTURE OF THE ENTIRE ACOUSTIC SPACE - the representation is not necessary for the behaviour
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limitations[1]
for which uselimitation — any comparison of grid-cell yield between environments: WE DETECTED FEWER GRID CELLS IN THE SMALLER ENVIRONMENT THAN IN THE LARGER ONE
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limitations[2]
for which uselimitation — the three-way classification of firing fields: THE THREE CLUSTERS WERE NOT PERFECTLY SEPARATED ... SOME FIRING FIELDS HAD SIGNIFICANTLY NON-ZERO REGRESSION COEFFICIENTS FOR MORE THAN ONE TASK PARAMETER: 14% IN CA1 - the classes overlap, and by how much is measured
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limitations[3]
for which uselimitation — the same scope as limitations[0] - too few fields to test the lattice reading
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limitations[4]
for which uselimitation — the same scope as limitations[1] - one frequency would suffice to solve the task
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limitations[5]
for which uselimitation — the same scope as limitations[2] - grid comparisons are confounded with arena size
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limitations[6]
for which uselimitation — the field-narrowing result near press and release: it may reflect A HIGHER DENSITY of sampling at those moments rather than a property of the fields
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limitations[7]
for which uselimitation — the attribution of the modulation to the sound dimension: HEAD DIRECTION (HD) CELLS OVERLAP WITH SMT-MODULATED NEURONS, and a separate analysis was needed to exclude them
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limitations[8]
replicationuntested
no type for thisSMT → linear track — AN ANALOGY, BOUNDED BY ITS AUTHORS: the sound-manipulation task evolves along a continuous axis (sound frequency) and is therefore ANALOGOUS to spatial navigation on a linear track, SHARING SOME KEY FEATURES of neural representation - discrete firing fields tiling the task, multiple fields in MEC cells, clustering at salient features, and context-dependence. 'Some key features' is the authors' own bound and it is the part an equivalence edge deletes.
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unhoused_relations[0]
limitationToo few fields per grid cell in the sound task to test whether its firing corresponds to a one-dimensional slice through a hexagonal grid lattice.
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limitations[0]
limitationThe task itself did not require the animals to represent the whole acoustic dimension, so the functional necessity of the representation is not established.
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limitations[1]
limitationGrid-cell yield depended on arena size, fewer being detected in the smaller environment.
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limitations[2]
limitationThe three alignment classes (press-, release-, frequency-aligned) were not cleanly separable, with some fields loading on more than one task parameter.
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limitations[3]
limitationthe hexagonal-lattice reading of the grid cell results could not be tested, because grid cells produced too few fields in this task
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limitations[4]
limitationthe rats did not actually need to represent the whole acoustic space to solve the task, since responding to one frequency would have sufficed
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limitations[5]
limitationfewer grid cells were detected in the smaller environment than the larger one, so all grid comparisons had to be verified on the subset recorded in the larger environment
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limitations[6]
limitationa tightening of firing fields in the vicinity of joystick presses and releases may be due to a higher density of available sensory cues, or alternatively may result from the stronger salience of these events
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limitations[7]
limitationand a head-direction confound had to be excluded by a separate analysis, since head direction cells overlap the modulated population
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limitations[8]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingRats solved the task by tracking sound frequency rather than elapsed time, releasing the joystick later when frequency advanced more slowly.
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content_tags.findings[0]
findingA large fraction of both CA1 and entorhinal cells were significantly task-modulated during the non-spatial sound task.
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content_tags.findings[1]
findingTask-modulated activity took the form of discrete firing fields resembling those seen during spatial navigation.
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content_tags.findings[2]
findingMost fields occurring during sound presentation were locked to sound frequency rather than to the motor events bounding the trial.
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content_tags.findings[3]
findingAlmost no CA1 cells, including the task-modulated ones, responded to passive playback of the same sounds (1.7% against 31.1% during the task).
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content_tags.findings[4]
findingAdding a reward at the end of passive sweeps produced task-modulated activity, but in fewer cells than the task itself (20.2%).
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content_tags.findings[5]
findingFields in the rewarded passive-playback task were about twice as wide as SMT fields, i.e. temporally less precise.
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content_tags.findings[6]
findingA quarter of identified place cells also carried the non-spatial sound representation.
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content_tags.findings[7]
findingGrid cells likewise participated in the non-spatial task.
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content_tags.findings[8]
findingHow spatially tuned a neuron was barely predicted how strongly it was modulated in the sound task, indicating the two codes are near-independent within the same cells.
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content_tags.findings[9]
findingGrid module identity carried over to the sound task: modules with wider spatial fields produced wider sound fields.
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content_tags.findings[10]
findingthe reconciling framework is stated in advance - spatial representation as one example of a general mechanism for encoding continuous task-relevant variables
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content_tags.findings[11]
findingneurons formed discrete firing fields at particular sound frequencies, and overlapped with the known spatial cell types
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content_tags.findings[12]
findingcommon circuit mechanisms are proposed for representations of diverse behavioural tasks
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content_tags.findings[13]
findingthe cognitive map frame already allowed the related entities to be distinct stimuli or abstract concepts, not only locations
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content_tags.findings[14]
findinghippocampal firing was already known to be modulated by non-spatial variables, discrete stimuli and time points in delay tasks
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content_tags.findings[15]
findingTRAVERSAL SPEED WAS RANDOMISED SO THAT FREQUENCY AND ELAPSED TIME WERE UNCOUPLED, which is what makes a frequency field distinguishable from a time field
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content_tags.findings[16]
findingjoystick releases were predicted by sound frequency alone with almost no added influence of elapsed time
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content_tags.findings[17]
findingand trial durations were about three times more variable than release frequencies, which is how the strategy was identified
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content_tags.findings[18]
finding40.0% of CA1 units and 51.3% of MEC units were significantly modulated during the task
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content_tags.findings[19]
findingactivity was largely confined to discrete firing fields akin to those of spatial navigation, clustered at trial boundaries but spanning the entire task
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content_tags.findings[20]
findingAND THE NEW THING IS NAMED AFTER THE OLD ONE: frequency fields, called analogous to place fields
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content_tags.findings[21]
findingmore than half of the fields during sound presentation aligned best to frequency rather than to the press or the release
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content_tags.findings[22]
findingTHE CONTROL SEPARATES THE STIMULUS FROM THE TASK: 1.7% of the same cells responded to identical sweeps outside the task, against 31.1% inside it
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content_tags.findings[23]
findingand the control was then run again WITH a reward, recovering a fifth of the cells
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content_tags.findings[24]
findingwith fields more than twice as wide, so behavioural context affected both how many cells were activated and how precise they were, while agency was not strictly required
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content_tags.findings[25]
findinga quarter of place cells and a third of grid cells were modulated in the sound task, with a mixed representation in CA1
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content_tags.findings[26]
findingMEMBERSHIP OF THE NAMED CLASS BARELY PREDICTED THE SAME CELL IN THE OTHER TASK: r2 of about 0.005 against placeness and gridness
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content_tags.findings[27]
findingso the two sets were not distinct subclasses of the circuit but a shared use of the same neural population between tasks
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content_tags.findings[28]
findingand the grid module a cell belongs to predicted its field width in the sound task as well as in the arena
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content_tags.findings[29]
findingTHE ANALYSIS THAT WOULD HAVE DECIDED THE LATTICE READING COULD NOT BE RUN, for want of fields per grid cell
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content_tags.findings[30]
findingthe sound task is declared analogous to a linear track, with the shared properties then listed one by one
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content_tags.findings[31]
findingspatial and non-spatial representations are produced by the same neuronal population
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content_tags.findings[32]
findingA CELL TYPE IS NAMED AFTER THE TASK IT WAS DISCOVERED IN: the spatial patterns may be a consequence of the continuity of the task variable rather than a primacy of physical space
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content_tags.findings[33]
findingthe rats did not need to represent the whole acoustic space; responding to one particular frequency would have sufficed
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content_tags.findings[34]
findingthe conjecture is that task performance activates a progression of fields carrying the order and adjacency of behavioural states
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content_tags.findings[35]
findingwith place and grid codes as a manifestation of that general mechanism rather than a spatial faculty
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content_tags.findings[36]
findingbecause rats sometimes moved their heads while deflecting the joystick in version 1 of the apparatus, version 2 was modified to remove this possible spatial confound
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content_tags.findings[37]
findingfrequency is reported in octaves RELATIVE to a 2 kHz start, so the number does not join without its reference
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content_tags.findings[38]
findingand deflections from stepping or leaning on the joystick were removed by a 3 s minimum, so not every deflection is a trial
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content_tags.findings[39]
findingAN EXCLUSION CRITERION THAT DOES NOT EXIST OUTSIDE ITS OWN TASK: ripples are normally removed by rejecting low-velocity time points, and with no velocity here they had to be detected directly in the field potential
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content_tags.findings[40]
findingwith the negative disclosure attached - excluding those spikes changed no result, it only raised the in-field to background ratio
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content_tags.findings[41]
findingthe reported information is a RATIO TO CHANCE, real rate over the mean of 100 shuffles
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content_tags.findings[42]
findingand the information rate alone was rejected as the criterion because background differences inflate it, with the test placed on the peak rate instead
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content_tags.findings[43]
findingputative interneurons are defined by a bare rate threshold and excluded
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content_tags.findings[44]
findingplace cells are defined as cells whose spatial information exceeds 99% of their own shuffled values
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content_tags.findings[45]
findingTHE DEFINITION OF A GRID CELL IS A THRESHOLD ON THIS DATASET'S OWN SHUFFLES - the 95th percentile, which here was 0.46
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content_tags.findings[46]
findingAND THE COUNT OF A CELL TYPE DEPENDS ON THE ARENA: fewer grid cells in the smaller environment, so every grid comparison was re-verified on the subset recorded in the larger environment
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content_tags.findings[47]
findingfields with more than one significant alignment coefficient are labelled mixed rather than forced into one class
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content_tags.findings[48]
findingTHETA FREQUENCY DID NOT TRACK SPEED HERE, unlike in navigation, which is read as that relationship depending on locomotion-related signals
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content_tags.findings[49]
findingand fields did not broaden across the trial as time cells do, attributed to a sensory variable being continuously available
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content_tags.findings[50]
findinga head-direction confound was tested directly, since those cells overlap the modulated population
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content_tags.findings[51]
findingand two simultaneously recorded cells with similar head direction tuning had dissimilar sound-task firing, which is what rules it out
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content_tags.findings[52]
findingfield tightening at the press and release is left with two rival explanations, denser sensory cues or greater salience
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content_tags.findings[53]
findingeach interval is linearly warped to its median duration before any average is taken
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content_tags.findings[54]
quantity2208 / 1164 — units recorded in dorsal CA1 (5 rats) and dorsal MEC (9 rats)
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content_tags.quantities[0]
quantity40.0% and 51.3% — proportion of CA1 and MEC cells significantly modulated by the sound task
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content_tags.quantities[1]
quantity70.8±2.6% — proportion of trials on which rats released the joystick inside the reward target zone
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content_tags.quantities[2]
quantity1.7% compared to 31.1% — CA1 cells responding during passive playback versus during the active task
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content_tags.quantities[3]
quantity25.1% of 295 — share of place cells that were also modulated by the sound task
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content_tags.quantities[4]
quantity34.3% of 105 — share of grid cells that were also modulated by the sound task
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content_tags.quantities[5]
quantityr 2 =0.0054 and r 2 =0.0053 — variance in sound-task modulation explained by placeness and by gridness
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content_tags.quantities[6]
quantity2.49±0.29 s and 1.16±0.05 s — mean field width in the rewarded passive-playback task versus the active sound task
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content_tags.quantities[7]
quantity26%, 23% and 51% — share of CA1 sound-period fields aligned to joystick press, to release, and to sound frequency
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content_tags.quantities[8]
quantity0.78±0.02 s and 1.36±0.05 s — sound-task field widths for grid module 1 versus modules 2/3
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content_tags.quantities[9]
quantity0.46 — the grid score above which a cell is a grid cell - the 95th percentile of THIS dataset's own shuffles, not a constant
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content_tags.quantities[10]
quantityr 2 =0.0054 — a very weak correlation between the amount of SMT modulation and placeness, against 0.0053 for gridness, suggesting nearly independent activity patterns in the two tasks
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content_tags.quantities[11]
quantity1.7% — cells responding to the identical sweeps outside the task, against 31.1% inside it
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content_tags.quantities[12]
quantity20.2% — cells responding once a reward was added to the same passive sweeps
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content_tags.quantities[13]
quantity25.1% — of 295 place cells modulated in the sound task, against 34.3% of 105 grid cells
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content_tags.quantities[14]
quantity0.146±0.002 — coefficient of variation of trial duration against 0.046±0.005 for release frequency - the ratio that identifies the strategy
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content_tags.quantities[15]
variablemutual information rate (dependent)
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content_tags.variables[2]
variablefiring field width (dependent)
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content_tags.variables[4]
variableputative interneurons (control)
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content_tags.variables[5]
variablebrief trials (control)
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content_tags.variables[8]
variablefrequency
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content_tags.variables[9]
variableduration
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content_tags.variables[10]
variablegrid score
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content_tags.variables[11]
variablespatial information
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content_tags.variables[12]
variablespeed
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content_tags.variables[13]
variableenvironment
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content_tags.variables[14]
definitionsound manipulation task
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content_tags.definitions[0]
definitioncognitive map
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content_tags.definitions[1]
definitionfrequency fields
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content_tags.definitions[2]
definitionpress
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content_tags.definitions[3]
definitiongrid score
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content_tags.definitions[4]
definitionspatial information
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content_tags.definitions[5]
definitioncognitive map
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content_tags.definitions[6]
definitionplace cell
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content_tags.definitions[7]
definitiongrid cell
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content_tags.definitions[8]
definitioninterneuron
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content_tags.definitions[9]
definitionmodulated
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content_tags.definitions[10]
definitionspatial information
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content_tags.definitions[11]
key-termsplace cell, grid cell, border cell, head direction, time cell, interneuron, firing field, CA1, entorhinal, hippocampal, SMT, random foraging, passive playback, linear track, task, sound frequency, location, spatial, non-spatial, navigation, cognitive map, continuous, mechanism, population, modulated, information, spatial information, grid score, shuffled, percentile, environment, module, theta, speed, velocity, SWR, joystick, press, release, trial, duration, octave, reward, salience, agency, behavioral context, model
Concepts
terms this paper carries, each with the sentence it was cut from — the layer that joins one paper to another. Terms with no quote yet are on CONCEPTS.html.
conceptgrid cell — not defined in this paper's own words; the entorhinal counterpart, also modulated in a task with no space in it
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concepts[1]
conceptmechanism — not defined in this paper's own words; the common circuit operation claimed to underlie both
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.
concepts[22]
conceptinformation — not defined in this paper's own words; mutual information rate between spikes and the task phase
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concepts[25]
conceptenvironment — not defined in this paper's own words; the arena, whose size changes how many grid cells exist
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concepts[30]
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.
t-test p as written: p<0.001, t-test
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content_tags.statistics[0]
coefficient of variation as written: coefficient of variation of 0.146±0.002 and 0.046±0.005
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content_tags.statistics[1]
chi-squared test χ 2 as written: p<0.001, χ 2 test
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content_tags.statistics[2]
Wilcoxon rank-sum test as written: p<0.001, Wilcoxon rank-sum test
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content_tags.statistics[3]
shuffle (permutation) test p as written: p<0.01, shuffle test
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content_tags.statistics[4]
mutual information H as written: mutual information rate between spikes and the phase of the task
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content_tags.statistics[5]
linear regression as written: linear regression was fit to the relationship between theta phase and the warped time
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content_tags.statistics[6]
bootstrap as written: bootstrap analysis on the individual trials using 100 bootstrapped samples
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content_tags.statistics[7]
correlation as written: the Pearson correlation is measured between the PSTHs
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content_tags.statistics[8]
mean squared error as written: mean squared error between the joystick release times simulated by the model and the actual release times
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content_tags.statistics[9]
standard error of the mean s.e.m. as written: mean±s.e.m. here and elsewhere
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content_tags.statistics[10]
principal component analysis as written: the first three principal components of the extracted waveforms
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content_tags.statistics[11]
confidence interval as written: 95% multinomial confidence intervals
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content_tags.statistics[12]
evidence status noteconfidence interval — This quote ends with the stray word 'The', the first word of the next sentence. It is not trimmed, because the quote without it occurs more than once in the paper's text and could not be tied to one place. The claim is carried by the sentence before 'The'.content_tags.statistics[12]
coefficient of determination r 2 as written: r 2 =0.0054 and r 2 =0.0053
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.statistics[13]
shuffle test
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content_tags.statistics[14]
mutual information 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.
content_tags.statistics[15]
Wilcoxon rank-sum test
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.statistics[16]
coefficient of variation
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.statistics[17]
bootstrap
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.statistics[18]
linear regression
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content_tags.statistics[19]
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
methoda sound manipulation task in which a joystick drives frequency along a logarithmic axis at a randomly varied speed, so that frequency is uncoupled from elapsed time, with the same cells recorded again during random foraging — a second reading DIFFERING from the recorded value; recorded: electrophysiology
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.method
phenomenondiscrete firing fields that tile a continuous behavioural variable, and the overlap of that population with the known spatial cell types — a second reading DIFFERING from the recorded value; recorded: hippocampal-entorhinal coding of a non-spatial continuous dimension
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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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