Rhythms of the hippocampal network — TEI-tagged view

Colgin · 2016 · neuroscience · doi:10.1038/nrn.2016.21 · full text at the source · TEI xml · tagged edition · index · concepts · authors
Where this record's text came from. faceted 2026-08 by reading the locally held text (xml); every evidence string verified as an exact substring of that text before merging. Claim relations left EMPTY on purpose: this review cites everything by superscript number (e.g. 'a recent study 51', 'reviewed elsewhere 15 , 16'), so no sentence relates the paper to a NAMED piece of prior work; the only personal name in the body is 'Donald Hebb's cell assembly hypothesis 2', which is background rather than a stated relation of this paper to that work. Rule 7 says leave empty rather than infer from bare citations. Claim type was a genuine choice between 'mechanism' and 'critique': the abstract's 'However, recent findings have challenged traditional views on these behavioral functions.' is critique-shaped, but the body is organised as Mechanisms/Functions sections for each rhythm, so mechanism is the dominant shape. Replication status 'contested' reflects the paper's own reporting of unresolved disagreement, e.g. 'There is less agreement in the field about the functional significance of gamma rhythms 14 .' - it describes the state of the claims reviewed, not an independent replication attempt of this review. 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.
39 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.
This record’s own method is review, and the tags below follow from that. A review’s findings, quantities and statistics are things stated IN this paper, and some of them report results obtained elsewhere. No tag on this page names a different author. Measured across every review in this corpus, 81% of a review’s tags carry no citation at all — most of them are the review speaking. The corpus does not adjudicate the rest: read an unmarked tag as a claim this paper makes, not necessarily one it made first.
Other fields define these differently the same word, defined another way in another discipline
memory 23 fields · novelty 3 fields · frequency 2 fields · terminology 2 fields
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
theta15 gamma9 slow gamma fast gamma2 beta3 rhythm3 frequency16 LFP8 hippocampus29 CA116 CA36 interneuron24 place cell6 memory16 encoding6 retrieval10 replay6 consolidation4 terminology5 band2 HCN novelty7 review319

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 network rhythms— “The hippocampal local field potential (LFP) exhibits three major types of rhythms, theta, sharp wave-ripples and gamma.”
claim-typemechanism— “I will discuss the mechanisms underlying hippocampal rhythm generation and provide an overview of what is known about their mnemonic functions.”
method — evidence— “Here I review our current understanding of the origins and mnemonic functions of hippocampal theta, sharp-wave ripples and gamma rhythms based on findings from rodent studies, and present an updated, synthesized view of their roles and interactions within the hippocampal network.”
object of study — evidence— “The hippocampal local field potential (LFP) exhibits three major types of rhythms, theta, sharp wave-ripples and gamma. These rhythms are defined by their frequencies, have behavioral correlates in several species including rats and humans, and have been proposed to perform distinct functions in hippocampal memory processing. However, recent…
research question — evidence— “The hippocampal local field potential (LFP) exhibits three major types of rhythms, theta, sharp wave-ripples and gamma. These rhythms are defined by their frequencies, have behavioral correlates in several species including rats and humans, and have been proposed to perform distinct functions in hippocampal memory processing. However, recent…
contribution — evidence— “Recently, several studies have begun to address the question of whether slow and fast gamma perform different functions, and a few main hypotheses have emerged from these studies. Evidence for and against each of these hypotheses is discussed below.”
transferfrom cell assembly theory: Hebb's idea that memories are represented by distributed ensembles activated concurrently, used as the reason synchrony should matter to memory— “Such rhythms reflect the synchronized activity of large numbers of neurons because synchronous currents sum together to generate large-amplitude fluctuations in LFP, whereas non-synchronized currents do not sum together and thus remain too small to be detected.”transfers[0]
transferfrom computational modelling: network simulations in which gamma frequency follows excitatory drive and GABA conductance, used to explain why one circuit could produce both subtypes— “Theoretical work has shown that the frequency of gamma rhythms generated by networks of hippocampal interneurons can vary depending on the strength of excitatory and inhibitory inputs 141 . Specifically, in simulations of gamma activity in a model neuron network, gamma frequency was found to increase with increased excitatory drive onto the…transfers[1]
mechanismSeptal GABAergic pacemaker interneurons rhythmically disinhibit hippocampal pyramidal cells, entraining them to theta— “These septal interneurons rhythmically disinhibit hippocampal pyramidal cells and thereby promote their theta rhythmic firing.”mechanisms[0]
mechanismRipple-associated shunting inhibition raises spike threshold so that only strongly potentiated cells fire, selecting which memories are sent to long-term storage— “That is, only the cells that receive sufficient excitation, which are likely to be those with synapses that were potentiated during earlier learning, would be able to overcome this inhibition and send their memory traces to long-term storage.”mechanisms[1]
mechanismGamma frequency is set by the balance of excitatory drive onto and GABA conductance within the interneuron network— “Specifically, in simulations of gamma activity in a model neuron network, gamma frequency was found to increase with increased excitatory drive onto the interneurons, and was found to decrease with increased GABA conductances in the interneurons.”mechanisms[2]
mechanismSharp-wave excitation drives place cells outside their place fields, enabling representation of distant locations— “Thus, sharp waves are likely to provide the excitation that allows place cells to fire outside of their place fields, during waking behaviors that require representations of distant locations.”mechanisms[3]
mechanismWhich gamma subtype is nested in the theta cycle switches the hippocampus between externally driven encoding and internally driven retrieval— “The switch between extrinsically and intrinsically driven network operations may depend on which type of gamma rhythms are nested within the theta cycle, with fast gamma promoting extrinsic inputs from MEC and slow gamma promoting intrinsic signals from CA3.”mechanisms[4]
mechanismLFP rhythms are visible only because synchronous currents summate while non-synchronous ones cancel— “Such rhythms reflect the synchronized activity of large numbers of neurons because synchronous currents sum together to generate large-amplitude fluctuations in LFP, whereas non-synchronized currents do not sum together and thus remain too small to be detected.”mechanisms[5]
mechanismgamma-frequency spiking of FAST-SPIKING INTERNEURONS produces the observed gamma rhythms - interneuron spikes CONSISTENTLY PHASE-LOCK to gamma while pyramidal cell spikes DO NOT— “There is much evidence to indicate that inhibitory interneurons are critical for gamma generation 14 , 132 . Gamma rhythms in the LFP of rats have been shown to reflect inhibitory events recorded intracellularly in CA3 and CA1 pyramidal cells 133 and in dentate gyrus granule cells 40 . In addition, interneuron spikes consistently phase-lock…mechanisms[6]
mechanismand the frequency itself is set by the balance of drive - in simulations gamma frequency INCREASED WITH INCREASED EXCITATORY DRIVE onto the interneurons and DECREASED WITH INCREASED GABA CONDUCTANCES— “Theoretical work has shown that the frequency of gamma rhythms generated by networks of hippocampal interneurons can vary depending on the strength of excitatory and inhibitory inputs 141 . Specifically, in simulations of gamma activity in a model neuron network, gamma frequency was found to increase with increased excitatory drive onto the…mechanisms[7]
mechanismwith different interneuron classes firing at different theta phases, so DIFFERENT TYPES OF INTERNEURONS SERVE DIFFERENT FUNCTIONS AT DISTINCT PHASES of the theta cycle— “A recent study of hippocampal interneurons in awake, behaving mice found that PV+ basket cells preferentially fire at an earlier theta phase (when place cells preferentially fire) than do somatostatin-positive (SOM+) oriens lacunosum-moleculare (OLM) interneurons 33 . These findings suggest that different types of interneurons in the…mechanisms[8]
critiques the traditional assignment of one memory function to each rhythm, which recent findings have challenged— “The hippocampal local field potential (LFP) exhibits three major types of rhythms, theta, sharp wave-ripples and gamma. These rhythms are defined by their frequencies, have behavioral correlates in several species including rats and humans, and have been proposed to perform distinct functions in hippocampal memory processing. However, recent…claim_relations[0]
contradicts the theory that theta is required for forming spatial memory representations, since place fields form without it and bats have place cells with little theta— “Place fields emerge as animals gain experience with new environments 68 - 70 , and it is often assumed that the development of place cell representations corresponds to spatial learning. It was therefore surprising that this study 51 showed that, in rats, place fields…claim_relations[1]
qualifies the memory-encoding hypothesis for fast gamma, which one study links instead to working memoryclaim_relations[2]
qualifies the memory-retrieval hypothesis for slow gamma, which novelty-related increases do not fit
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claim_relations[3]
critiques the medial septum as sole pacemaker, since theta occurs in a preparation with no septal input
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claim_relations[4]
predicts that whether different gamma frequencies mean different circuits or the same circuit differently driven is the question to settle next
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claim_relations[5]
senseTHE OBJECT IS DEFINED BY FREQUENCY AND JUDGED BY FUNCTION: these rhythms ARE DEFINED BY THEIR FREQUENCIES, have behavioral correlates, and HAVE BEEN PROPOSED TO PERFORM DISTINCT FUNCTIONS - HOWEVER, RECENT FINDINGS HAVE CHALLENGED TRADITIONAL VIEWS ON THESE BEHAVIORAL FUNCTIONS
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senses[0]
senseand the assignment is stated as an expectation rather than a result: each rhythm is observed during particular behaviors and generated by specific mechanisms, so IT IS THEREFORE NOT SURPRISING THAT THESE DIFFERENT RHYTHMS ARE THOUGHT TO PERFORM DISTINCT FUNCTIONS - with THERE IS LESS AGREEMENT IN THE FIELD ABOUT THE FUNCTIONAL SIGNIFICANCE OF GAMMA
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senses[1]
senseA LIVE NAMING CONFLICT SETTLED IN A PARENTHESIS: the same activity HAS BEEN TERMED FAST GAMMA IN SOME STUDIES AND MIDFREQUENCY GAMMA IN OTHERS - (THE FORMER TERMINOLOGY WILL BE USED HERE). One review, one clause, and the field's vocabulary is decided for the reader
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senses[2]
senseAND THE EARLIER LITERATURE COULD NOT HAVE TOLD THEM APART: MOST OF THE STUDIES DESCRIBED ABOVE DID NOT DIFFERENTIATE BETWEEN DIFFERENT FREQUENCY VARIANTS OF GAMMA, and IT IS POSSIBLE THAT SLOW AND FAST GAMMA ARE GENERATED BY DIFFERENT MECHANISMS - a whole mechanism literature written with one word for two things
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senses[3]
senseA RESULT THAT CANNOT BE ASSIGNED TO A CATEGORY BECAUSE THE CATEGORIES OVERLAP: the observed ~20-40 Hz activity OVERLAPPED WITH BOTH BETA AND SLOW GAMMA FREQUENCY BANDS AND THUS MAY REFLECT EITHER BETA, SLOW GAMMA, OR A COMBINATION OF BOTH
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senses[4]
sensewith the same term carrying two ranges inside one review - slow gamma introduced as ~25-55 Hz here, and reported as ~30-80 Hz three paragraphs later, against ~60-100 Hz for the fast one
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senses[5]
senseand the second range printed where a layer difference is being claimed: SLOW GAMMA (~30-80 Hz) ACTIVITY DOMINATED IN STRATUM RADIATUM, WHEREAS ~60-100 Hz GAMMA DOMINATED IN STRATUM LACUNOSUM-MOLECULARE - the two definitions overlap by 20 Hz
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senses[6]
sensewith the measurement problem named as a property of the object: THE HIGH VARIABILITY OF GAMMA FREQUENCY ALSO COMPLICATES ITS MEASUREMENT IN BEHAVING ANIMALS
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senses[7]
senseA FOUNDING THEORY CALLED INTO QUESTION BY ONE STUDY: in the 1970s the extent to which theta was present PREDICTED HOW QUICKLY ANIMALS LEARNED, which LED TO THE THEORY that theta plays an essential role in learning and memory - HOWEVER, THE THEORY HAS BEEN CALLED INTO QUESTION
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senses[8]
senseand the evidence is two facts that no amount of theta-learning correlation can absorb: place fields WERE FORMED in novel environments DESPITE THETA BEING BLOCKED by septal inactivation, and FLYING BATS HAVE PLACE CELLS EVEN THOUGH these cells exhibit LITTLE TO NO THETA RHYTHMICITY - so THETA RHYTHMICITY IS NOT REQUIRED FOR THE FORMATION OF PLACE FIELDS
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senses[9]
senseTHE REVIEW'S STRUCTURE IS THE ARGUMENT: several studies have begun to address whether slow and fast gamma perform different functions, A FEW MAIN HYPOTHESES HAVE EMERGED, and EVIDENCE FOR AND AGAINST EACH OF THESE HYPOTHESES IS DISCUSSED BELOW
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senses[10]
senseand both hypotheses survive with their counter-evidence attached: HOWEVER, OTHER RESULTS DO NOT SUPPORT THE HYPOTHESIS THAT FAST GAMMA HAS A MEMORY ENCODING FUNCTION - one study links it to WORKING MEMORY, BUT NOT MEMORY ENCODING
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senses[11]
senseand for the other: NOT ALL FINDINGS FROM STUDIES OF SLOW GAMMA FIT WITH THIS MEMORY RETRIEVAL HYPOTHESIS - THESE APPARENT INCONGRUITIES ACROSS STUDIES HIGHLIGHT THE NEED FOR FURTHER INVESTIGATION
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senses[12]
senseA MECHANISM THAT CONTRADICTS ITSELF IN TWO CITATIONS: HCN1 IS THOUGHT TO FACILITATE theta rhythmic firing and its blockade REDUCED theta entrainment - HOWEVER, THE PRECISE ROLE OF HCN CHANNELS REMAINS UNCLEAR BECAUSE IT HAS ALSO BEEN REPORTED THAT ENHANCED THETA RHYTHMS OCCUR IN HCN1 KNOCKOUT MICE
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senses[13]
senseand a pacemaker that turns out not to be necessary in vitro but necessary in vivo: THETA RHYTHMS CAN OCCUR IN THE ABSENCE OF MS PACEMAKER INPUTS in a whole hippocampal preparation, suggesting theta CAN ALSO EMERGE FROM LOCAL CIRCUIT INTERACTIONS - NONETHELESS, IT IS CLEAR THAT MS INPUTS ARE IMPORTANT for driving theta in behaving animals
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senses[14]
sensewith the clean two-circuit story qualified by the cells themselves: A SUBSTANTIAL PROPORTION OF CA1 INTERNEURONS PHASE-LOCK TO BOTH SLOW AND FAST GAMMA, and MAY BE PART OF A NETWORK THAT CAN GENERATE EITHER, DEPENDING ON THE STATE OF THE NETWORK
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senses[15]
senseand the frequency is a consequence of drive rather than an identity: gamma frequency INCREASED WITH INCREASED EXCITATORY DRIVE onto the interneurons and DECREASED WITH INCREASED GABA CONDUCTANCES - which means the label follows the input, not the circuit
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senses[16]
sensewith the mechanism section closing on its own status: HOWEVER, THESE HYPOTHESES ARE TENTATIVE, AND MUCH WORK REMAINS TO BE DONE
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senses[17]
senseTHE OBJECT IS VISIBLE ONLY WHERE IT IS SYNCHRONOUS: rhythms reflect synchronized activity BECAUSE SYNCHRONOUS CURRENTS SUM TOGETHER to generate large-amplitude fluctuations, WHEREAS NON-SYNCHRONIZED CURRENTS DO NOT SUM AND THUS REMAIN TOO SMALL TO BE DETECTED - the measurement selects what it can see
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senses[18]
senseand the region was chosen for the same reason: the hippocampus contains DENSELY PACKED NEURONS THAT GENERATE LARGE LFPs, especially CA1 because THE PYRAMIDAL CELL DENDRITES ARE ALIGNED IN PARALLEL - ACCORDINGLY, MANY STUDIES of the role of brain rhythms in memory HAVE BEEN CONDUCTED IN THE HIPPOCAMPUS
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senses[19]
senseand the species too: many findings are from LOWER MAMMALS, PARTICULARLY RODENTS, IN PART BECAUSE THE MEASUREMENT of brain rhythms in healthy human subjects WITHIN A DEEP-LYING STRUCTURE such as the hippocampus IS DIFFICULT
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senses[20]
senseTHE OPEN QUESTIONS ARE LISTED AS OPEN: what exactly IS the functional significance of variations in theta frequency, and WHY WOULD MEMORIES BE STORED OR CONSOLIDATED IN REVERSE ORDER? DO FORWARD AND REVERSE REPLAY SERVE DIFFERENT FUNCTIONS?
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senses[21]
sensewith a gap at the center of the leading hypothesis: according to the consolidation hypothesis replay TRANSFERS MEMORIES FROM THE HIPPOCAMPUS TO THE NEOCORTEX, HOWEVER IT REMAINS UNKNOWN WHETHER COORDINATED SEQUENCE REPLAY OCCURS between the hippocampus and its major downstream target
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senses[22]
senseand the question this whole review turns on, asked last: DO DIFFERENT FREQUENCIES OF GAMMA REPRESENT DIFFERENT FUNCTIONAL STATES, AND IF SO, ARE DIFFERENT CIRCUITS INVOLVED IN THEIR GENERATION OR IS THE SAME CIRCUITRY SIMPLY ENTRAINED BY DIFFERENT INPUTS?
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senses[23]
controlseptal inactivation, which removes theta while place fields are being formed — failed — rules out theta as a requirement for spatial representations - place fields FORMED in novel environments DESPITE theta and theta entrainment being BLOCKED
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controls[0]
controlan in vitro whole hippocampal preparation with no septal input at all — partial — rules out the septum as the sole pacemaker - theta OCCURRED IN THE ABSENCE OF MS PACEMAKER INPUTS, which suggests theta can emerge from local circuit interactions
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controls[1]
controlinterneurons checked for phase-locking to both gamma subtypes rather than one — failed — rules out a clean two-circuit account - A SUBSTANTIAL PROPORTION OF CA1 INTERNEURONS PHASE-LOCK TO BOTH slow and fast gamma
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controls[2]
differs inthe traditional assignment of one memory function to each rhythm, which recent findings have challenged — whether the frequency variants within gamma were distinguished at all, since most studies did not
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claim_relations[0]
differs inthe theory that theta is required for forming spatial memory representations, since place fields form without it and bats have place cells with little theta — THE AXIS IS WHETHER PLACE-FIELD DEVELOPMENT TRACKS LEARNING. 'Place fields emerge as animals gain experience with new environments, and it is often assumed that the development of place cell representations corresponds to spatial learning. It was therefore SURPRISING that this study showed...' The contradiction is aimed at the ASSUMED CORRESPONDENCE, which the span states before contradicting it.
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claim_relations[1]
differs inthe memory-encoding hypothesis for fast gamma, which one study links instead to working memory — THE AXIS IS WHICH MEMORY PROCESS. 'a recent study has found that fast gamma is linked to working memory, but not memory encoding, in mice performing a delayed nonmatching-to-place task.' The encoding hypothesis is not denied its association with fast gamma; it is denied THIS process.
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claim_relations[2]
differs inthe memory-retrieval hypothesis for slow gamma, which novelty-related increases do not fit — THE AXIS IS NOVELTY. 'However, not all findings from studies of slow gamma fit with this memory retrieval hypothesis. Specifically, some studies have described novelty-related increases in slow gamma in rats.' The hypothesis is qualified by a condition it does not cover, not replaced.
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claim_relations[3]
notetheta —is-a→ rhythm — IT IS THEREFORE NOT SURPRISING THAT THESE DIFFERENT RHYTHMS ARE THOUGHT TO PERFORM DISTINCT FUNCTIONS - and THERE IS LESS AGREEMENT IN THE FIELD ABOUT THE FUNCTIONAL SIGNIFICANCE OF GAMMA
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concept_edges[1]
noteRETRACTED (REVERSED) · CA3 —causes→ slow gamma — introduced as ~25-55 Hz and DRIVEN BY CA3 - and reported as ~30-80 Hz three paragraphs later — direction corrected 2026-08-20 by the project: the quote puts 'CA3' on the causing side. The edge previously read slow gamma --causes--> CA3, which is the reverse of its own sentence. Nothing else on the row changed; the quote, weight and anchor are untouched.
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concept_edges[3]
retractedRETRACTED (REVERSED) · CA3 —causes→ slow gamma — REVERSED. The span says slow gamma IS DRIVEN BY CA3, so the arrow runs CA3 -> slow gamma. The edge asserts the opposite causal direction.
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concept_edges[3]
noteslow gamma —part-of→ gamma — SLOW GAMMA (~30-80 Hz) DOMINATED IN STRATUM RADIATUM, WHEREAS ~60-100 Hz GAMMA DOMINATED IN STRATUM LACUNOSUM-MOLECULARE - the two definitions overlap by 20 Hz
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concept_edges[4]
auditbeta —contrasted-with→ slow gamma — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span describes 20-40 Hz coupling; the contrast with slow gamma is not stated.
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concept_edges[5]
notebeta —contrasted-with→ slow gamma — A RESULT THAT CANNOT BE ASSIGNED BECAUSE THE CATEGORIES OVERLAP - the ~20-40 Hz activity OVERLAPPED WITH BOTH BANDS and MAY REFLECT EITHER BETA, SLOW GAMMA, OR A COMBINATION OF BOTH
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concept_edges[5]
audittheta —causes→ memory — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says theta predicted learning speed; causes overstates a prediction.
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concept_edges[7]
notetheta —causes→ memory — A FOUNDING THEORY CALLED INTO QUESTION - the 1970s correlation LED TO THE THEORY that theta plays an essential role, and THE THEORY HAS BEEN CALLED INTO QUESTION BY A RECENT STUDY
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concept_edges[7]
auditslow gamma —causes→ retrieval — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span refers to a retrieval hypothesis that some findings do not fit; causes states a hypothesis as a finding.
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concept_edges[10]
noteslow gamma —causes→ retrieval — NOT ALL FINDINGS FIT WITH THIS MEMORY RETRIEVAL HYPOTHESIS - THESE APPARENT INCONGRUITIES ACROSS STUDIES HIGHLIGHT THE NEED FOR FURTHER INVESTIGATION
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concept_edges[10]
effect directionnovelty —causes→ slow gamma — increases
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concept_edges[11]
evidential statusnovelty —causes→ slow gamma — CITED, AND CITED AS A COMPLICATION: the review reports these studies as findings that do NOT fit the memory-retrieval hypothesis it has just set out
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.
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
concept_edges[11]
noteinterneuron —causes→ gamma — INTERNEURON SPIKES CONSISTENTLY PHASE-LOCK TO GAMMA OSCILLATIONS, WHEREAS PYRAMIDAL CELL SPIKES DO NOT
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concept_edges[12]
auditinterneuron —causes→ frequency — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says the frequency of interneuron-generated gamma varies with input strength; the endpoint 'frequency' drops 'gamma'.
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concept_edges[14]
noteinterneuron —causes→ frequency — gamma frequency INCREASED WITH INCREASED EXCITATORY DRIVE onto the interneurons and DECREASED WITH INCREASED GABA CONDUCTANCES - the label follows the input, not the circuit
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
concept_edges[14]
auditHCN —causes→ theta — Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but HCN1 is 'THOUGHT TO facilitate' theta and its role 'remains unclear'; causes is hedged.
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concept_edges[16]
evidential statusHCN —causes→ theta — ATTRIBUTED, WITH ONE CITED EXPERIMENT BEHIND IT AND A QUALIFICATION AFTER IT: 'HCN1 IS THOUGHT TO facilitate theta rhythmic firing in neurons 41, 42, and HCN1 BLOCKADE REDUCED theta entrainment of pyramidal cell spikes 38. HOWEVER, THE PRECISE ROLE OF HCN CHANNELS IN THETA PRODUCTI[ON]...' Enriched rather than retracted because the middle clause is a cited experimental result, not only a belief - but neither the belief nor the experiment is this review's, and the sentence after it reopens the question.
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Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
concept_edges[16]
noteHCN —causes→ theta — A MECHANISM THAT CONTRADICTS ITSELF IN TWO CITATIONS - HCN1 IS THOUGHT TO FACILITATE theta, and ENHANCED THETA RHYTHMS OCCUR IN HCN1 KNOCKOUT MICE
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
concept_edges[16]
auditLFP —operationalised-as→ rhythm — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but rhythms appear as LFP fluctuations; operationalised-as is loose.
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concept_edges[17]
noteLFP —operationalised-as→ rhythm — THE OBJECT IS VISIBLE ONLY WHERE IT IS SYNCHRONOUS - synchronous currents SUM TOGETHER, WHEREAS NON-SYNCHRONIZED CURRENTS DO NOT SUM AND THUS REMAIN TOO SMALL TO BE DETECTED
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concept_edges[17]
audithippocampus —causes→ LFP — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says hippocampal neurons generate large LFPs; 'hippocampus causes LFP' is loose.
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concept_edges[18]
notehippocampus —causes→ LFP — DENSELY PACKED NEURONS THAT GENERATE LARGE LFPs, especially CA1 because THE PYRAMIDAL CELL DENDRITES ARE ALIGNED IN PARALLEL - ACCORDINGLY, MANY STUDIES HAVE BEEN CONDUCTED IN THE HIPPOCAMPUS
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concept_edges[18]
auditRETRACTED (A HYPOTHESIS EXPLICITLY FRAMED AS A GAP IN KNOWLEDGE) · replay —causes→ consolidation — Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span states the consolidation hypothesis of replay; causes states a hypothesis as a finding.
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concept_edges[20]
noteRETRACTED (A HYPOTHESIS EXPLICITLY FRAMED AS A GAP IN KNOWLEDGE) · replay —causes→ consolidation — a gap at the center of the leading hypothesis - IT REMAINS UNKNOWN WHETHER COORDINATED SEQUENCE REPLAY OCCURS between the hippocampus and its major downstream cortical target
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concept_edges[20]
retractedRETRACTED (A HYPOTHESIS EXPLICITLY FRAMED AS A GAP IN KNOWLEDGE) · replay —causes→ consolidation — A HYPOTHESIS EXPLICITLY FRAMED AS A GAP IN KNOWLEDGE. The span: 'ANOTHER CRITICAL…' The sentence naming the relation is bracketed by 'critical gap in knowledge' before and 'remains unknown' after. A reader joining on 'X --causes--> Y' takes it as a causal relation this paper reports.
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Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
concept_edges[20]
outcome (stated by the tagger, no quotation)control — failedcontrols[0]
outcome (stated by the tagger, no quotation)control — partialcontrols[1]
outcome (stated by the tagger, no quotation)control — failedcontrols[2]
for which uselimitation — the scope of the review: OTHER BRAIN STATE-DEPENDENT RHYTHMS, SUCH AS SLEEP SPINDLES AND SLOW OSCILLATIONS, ALSO INFLUENCE HIPPOCAMPAL OPERATIONS; THESE RHYTHMS WILL NOT BE DISCUSSED IN DEPTH HERE
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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.
limitations[0]
for which uselimitation — the transfer of every finding to humans: MANY OF THE FINDINGS DESCRIBED HERE HAVE BEEN MADE IN STUDIES CARRIED OUT IN LOWER MAMMALS, PARTICULARLY RODENTS
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limitations[1]
for which uselimitation — the mechanistic account of slow versus fast gamma: THESE HYPOTHESES ARE TENTATIVE, AND MUCH WORK REMAINS TO BE DONE
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[2]
for which uselimitation — the memory-retrieval account of slow gamma: THESE APPARENT INCONGRUITIES ACROSS STUDIES HIGHLIGHT THE NEED FOR FURTHER INVESTIGATION - not all findings fit it
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limitations[3]
for which uselimitation — distinguishing slow from fast gamma, which most of the mechanism literature reviewed here cannot do
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[4]
for which uselimitation — the band assignment of that result, and of the band vocabulary generally: the ~20-40 Hz coupling OVERLAPPED WITH BOTH BETA AND SLOW GAMMA FREQUENCY BANDS
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[5]
for which uselimitation — every gamma measurement in behaving animals: THE HIGH VARIABILITY OF GAMMA FREQUENCY ALSO COMPLICATES ITS MEASUREMENT
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[6]
for which uselimitation — the same scope as limitations[2] - the generation mechanisms are tentative
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
limitations[7]
for which uselimitation — the HCN account of theta: HCN1 IS THOUGHT TO FACILITATE THETA RHYTHMIC FIRING, yet ENHANCED theta rhythms occur in HCN1 knockout mice - the review carries both
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limitations[8]
for which uselimitation — the same scope as limitations[1], with the reason: measuring rhythms in healthy humans WITHIN A DE[ep structure] is hard, so the evidence base is rodent
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limitations[9]
replicationcontested
no type for thisfast gamma → memory encoding — THE RELATION FAILS: the review reports that results do not support a memory-encoding function for fast gamma, and that a study found it linked to working memory BUT NOT memory encoding. No schema 0.3 type can express a relation that does not hold, so recording it at all means recording it here. Replaces the retracted concept_edges[9], which quoted this same sentence as evidence FOR the relation.
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unhoused_relations[0]
no type for thistheta frequency → functional significance — DECLARED OPEN BY THE REVIEW: 'some important questions remain ... what exactly is the functional significance of variations in theta frequency? Specifically, is there any functional significance to the lower frequency of theta that occurs during novel experiences compared to familiar ones?' A relation the field has not established, stated as a question. Every edge type asserts that a relation HOLDS.
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unhoused_relations[1]
limitationThe review deliberately excludes other brain-state-dependent rhythms that also affect the hippocampus
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limitations[0]
limitationThe evidence base is overwhelmingly rodent, because deep human hippocampal rhythm measurement is hard
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limitations[1]
limitationThe proposed mechanisms for slow versus fast gamma generation are explicitly provisional
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limitations[2]
limitationFindings on slow gamma do not all fit the memory-retrieval account the review favours
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limitations[3]
limitationMOST OF THE STUDIES DESCRIBED ABOVE DID NOT DIFFERENTIATE BETWEEN DIFFERENT FREQUENCY VARIANTS OF GAMMA, and IT IS POSSIBLE THAT SLOW AND FAST GAMMA ARE GENERATED BY DIFFERENT MECHANISMS
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limitations[4]
limitationthe observed ~20-40 Hz activity OVERLAPPED WITH BOTH BETA AND SLOW GAMMA FREQUENCY BANDS and thus MAY REFLECT EITHER BETA, SLOW GAMMA, OR A COMBINATION OF BOTH
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limitations[5]
limitationTHE HIGH VARIABILITY OF GAMMA FREQUENCY ALSO COMPLICATES ITS MEASUREMENT IN BEHAVING ANIMALS
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limitations[6]
limitationthese hypotheses are TENTATIVE, and MUCH WORK REMAINS TO BE DONE to uncover mechanisms underlying slow and fast gamma rhythms
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limitations[7]
limitationthe precise role of HCN channels REMAINS UNCLEAR because ENHANCED theta rhythms occur in HCN1 knockout mice, where the channel was thought to FACILITATE theta
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limitations[8]
limitationmany of the findings are from RODENTS, in part because measuring rhythms in healthy human subjects WITHIN A DEEP-LYING STRUCTURE such as the hippocampus IS DIFFICULT
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limitations[9]
Content tags
content-observations: finding · quantity · variable · definition · key-term — grounded in the full text
findingTheta rhythmicity is not necessary for single place cells to form place fields, undercutting the assumption that theta is required for building spatial memory representations at the single-cell level.
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content_tags.findings[0]
findingBlocking theta by MS inactivation impaired a delayed spatial alternation task and disrupted ordered place-cell ensembles.
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content_tags.findings[1]
findingRIPPLES ARE GENERATED LOCALLY IN CA1 by ripple-frequency spiking of PERISOMATICALLY TARGETING BASKET CELL INTERNEURONS
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content_tags.findings[2]
findingSharp waves are excitatory events transmitted from CA3 to CA1.
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content_tags.findings[3]
findingRecent evidence suggests sharp wave-ripples also play a key role in aspects of active spatial navigation.
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content_tags.findings[4]
findingSlow (~25-55 Hz) and fast (~60-100 Hz) gamma appear to have different origins and may serve different functions.
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content_tags.findings[5]
findingthese findings suggest that SLOW GAMMA IS IMPORTANT IN MEMORY RETRIEVAL, since replay is thought to reflect REACTIVATION OF MEMORIES OF EARLIER EXPERIENCES
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content_tags.findings[6]
findingthese rhythms ARE DEFINED BY THEIR FREQUENCIES and HAVE BEEN PROPOSED TO PERFORM DISTINCT FUNCTIONS - HOWEVER, RECENT FINDINGS HAVE CHALLENGED TRADITIONAL VIEWS ON THESE BEHAVIORAL FUNCTIONS
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content_tags.findings[7]
findingTHERE IS LESS AGREEMENT IN THE FIELD ABOUT THE FUNCTIONAL SIGNIFICANCE OF GAMMA RHYTHMS
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content_tags.findings[8]
findingthis type of activity HAS BEEN TERMED FAST GAMMA IN SOME STUDIES AND MIDFREQUENCY GAMMA IN OTHERS - THE FORMER TERMINOLOGY WILL BE USED HERE
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content_tags.findings[9]
findingMOST OF THE STUDIES DESCRIBED ABOVE DID NOT DIFFERENTIATE BETWEEN DIFFERENT FREQUENCY VARIANTS OF GAMMA, AND IT IS POSSIBLE THAT SLOW AND FAST GAMMA ARE GENERATED BY DIFFERENT MECHANISMS
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content_tags.findings[10]
findingthe observed ~20-40 Hz activity OVERLAPPED WITH BOTH BETA AND SLOW GAMMA FREQUENCY BANDS AND THUS MAY REFLECT EITHER BETA, SLOW GAMMA, OR A COMBINATION OF BOTH
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content_tags.findings[11]
findingSLOW GAMMA (~30-80 Hz) ACTIVITY DOMINATED IN STRATUM RADIATUM, WHEREAS ~60-100 Hz GAMMA DOMINATED IN STRATUM LACUNOSUM-MOLECULARE
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content_tags.findings[12]
findingTHE HIGH VARIABILITY OF GAMMA FREQUENCY ALSO COMPLICATES ITS MEASUREMENT IN BEHAVING ANIMALS
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content_tags.findings[13]
findingTHE THEORY HAS BEEN CALLED INTO QUESTION BY A RECENT STUDY involving recordings from place cells
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content_tags.findings[14]
findingTHETA RHYTHMICITY IS NOT REQUIRED FOR THE FORMATION OF PLACE FIELDS, and thus theta is NOT REQUIRED FOR THE FORMATION OF SPATIAL MEMORY REPRESENTATIONS AT THE SINGLE CELL LEVEL
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content_tags.findings[15]
findingHOWEVER, OTHER RESULTS DO NOT SUPPORT THE HYPOTHESIS THAT FAST GAMMA HAS A MEMORY ENCODING FUNCTION
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content_tags.findings[16]
findingNOT ALL FINDINGS FROM STUDIES OF SLOW GAMMA FIT WITH THIS MEMORY RETRIEVAL HYPOTHESIS - THESE APPARENT INCONGRUITIES ACROSS STUDIES HIGHLIGHT THE NEED FOR FURTHER INVESTIGATION
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content_tags.findings[17]
findingTHE PRECISE ROLE OF HCN CHANNELS IN THETA PRODUCTION REMAINS UNCLEAR BECAUSE IT HAS ALSO BEEN REPORTED THAT ENHANCED THETA RHYTHMS OCCUR IN HCN1 KNOCKOUT MICE
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content_tags.findings[18]
findingTHETA RHYTHMS CAN OCCUR IN THE ABSENCE OF MS PACEMAKER INPUTS, in a whole hippocampal in vitro preparation
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content_tags.findings[19]
findingA SUBSTANTIAL PROPORTION OF CA1 INTERNEURONS PHASE-LOCK TO BOTH SLOW AND FAST GAMMA
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content_tags.findings[20]
findinggamma frequency was found to INCREASE WITH INCREASED EXCITATORY DRIVE onto the interneurons, and to DECREASE WITH INCREASED GABA CONDUCTANCES
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content_tags.findings[21]
findingHOWEVER, THESE HYPOTHESES ARE TENTATIVE, AND MUCH WORK REMAINS TO BE DONE to uncover mechanisms underlying slow and fast gamma rhythms
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content_tags.findings[22]
findingsynchronous currents SUM TOGETHER to generate large-amplitude fluctuations in LFP, WHEREAS NON-SYNCHRONIZED CURRENTS DO NOT SUM TOGETHER AND THUS REMAIN TOO SMALL TO BE DETECTED
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content_tags.findings[23]
findingIT REMAINS UNKNOWN WHETHER COORDINATED SEQUENCE REPLAY OCCURS BETWEEN THE HIPPOCAMPUS AND ITS MAJOR DOWNSTREAM CORTICAL TARGET
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content_tags.findings[24]
findingDO DIFFERENT FREQUENCIES OF GAMMA REPRESENT DIFFERENT FUNCTIONAL STATES OF THE HIPPOCAMPAL NETWORK, and if so ARE DIFFERENT CIRCUITS INVOLVED IN THEIR GENERATION OR IS THE SAME CIRCUITRY SIMPLY ENTRAINED BY DIFFERENT INPUTS?
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content_tags.findings[25]
quantity~4-12 Hz — frequency band the review assigns to hippocampal theta
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content_tags.quantities[0]
quantity~25-55 Hz and ~60-100 Hz — frequency bands separating slow gamma from fast gamma
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content_tags.quantities[1]
quantity~30-80 Hz — gamma band dominating in stratum radiatum, versus ~60-100 Hz in stratum lacunosum-moleculare
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content_tags.quantities[2]
quantity~25-55 Hz — slow gamma as this review first defines it - the same term is given as ~30-80 Hz three paragraphs later
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content_tags.quantities[3]
quantity~30-80 Hz — slow gamma as reported in the layer comparison, overlapping the ~60-100 Hz fast range by 20 Hz
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content_tags.quantities[4]
quantity~20-40 Hz — the coupling that cannot be assigned to a band, since it overlaps both beta and slow gamma
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content_tags.quantities[5]
variablefrequency
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content_tags.variables[0]
variabletheta
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content_tags.variables[1]
definitionbrain rhythms
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content_tags.definitions[0]
definitionplace cells
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content_tags.definitions[1]
definitionreplay
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content_tags.definitions[2]
definitiontheta phase precession
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content_tags.definitions[3]
definitionneuronal ensembles
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content_tags.definitions[5]
definitionrhythm
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content_tags.definitions[6]
definitionslow gamma
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content_tags.definitions[7]
definitionfast gamma
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.definitions[8]
key-termstheta, gamma, slow gamma, fast gamma, beta, rhythm, frequency, LFP, hippocampus, CA1, CA3, interneuron, place cell, memory, encoding, retrieval, replay, consolidation, terminology, band, HCN, novelty
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.
conceptslow gamma — not defined in this paper's own words; one of them, with two different ranges in one review
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[2]
conceptCA3 — not defined in this paper's own words; the source of the slow one
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[10]
conceptencoding — not defined in this paper's own words; the function proposed for fast gamma
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[14]
conceptnovelty — not defined in this paper's own words; what does not fit the retrieval hypothesis
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[21]
Second readings
a facet read a second time, independently, from its own sentence — and whether the two readings agree
claim typecritique — a second reading DIFFERING from the recorded value; recorded: mechanism
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.claim_type
methodreview of the generation mechanisms and the functional evidence for each rhythm, with the incongruities left standing rather than resolved — a second reading DIFFERING from the recorded value; recorded: review
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
phenomenonperiodically fluctuating waves of neuronal activity in the local field potential, visible only because synchronous currents sum and non-synchronous ones do not — a second reading DIFFERING from the recorded value; recorded: hippocampal network rhythms
Quotation withheld — this paper is not ours to reproduce, and the page has already quoted as much of it as the rights layer allows. The tag, its field and its confidence stand; the sentence is at the publisher.
alternatives.phenomenon

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

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