Working Memory-Related Hippocampal Deactivation Interferes with Long-Term Memory Formation

Axmacher, Elger, Fell · Journal of Neuroscience · 2009 · neuroscience · 10.1523/jneurosci.5277-08.2009 · record Axmacher2009-working-memory-related-hippocampal-deactivat
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Where this record's text came from. faceted 2026-08 by reading the locally held text (html); every evidence string verified as an exact substring of that text before merging. method is recorded as 'fmri' from the controlled list, but the study is genuinely dual-modality: new fMRI data in 19 healthy subjects plus a reanalysis of intracranial EEG (DC potentials) from 8 epilepsy patients with hippocampal depth electrodes - i.e. 'electrophysiology' applies equally. Claim type is 'mechanism' because the contribution is an account of WHY early-list items are forgotten (failure to recruit hippocampus-dependent WM), not merely that a difference exists. Replication status is 'untested' because the article itself reports no independent replication attempt of its own finding. 2026-08-13, 2026-08-15 and 2026-08-25 schema-0.3 passes: every quote verified as an exact substring of the staged text and anchored at merge time; concept terms verified present; every edge endpoint an existing node.
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Other fields define these differently the same word, defined another way in another discipline
working memory 4 fields · statistical-significance 2 fields · fmri 2 fields

Tags on this paper

Every tag the record carries, each linking to every other paper in the corpus that carries it. A tag appears here whether or not its words occur in the article — the spans below are only the ones a sentence in this paper licenses. Of the 12 concept tags on this paper, none is grounded in a quoted sentence. All carry the tagger's gloss of what the term means in this paper, which is not something the article can license.
working memory long-term memory primacy recency hippocampus deactivation DC potentials BOLD serial position anterior cingulate cortex rote strategy Sternberg Early list items processed by single-item, hippocampus-independent working memory leave the hippocampus deactivated, so no encoding into long-term memory takes place and the item is forgotten When multi-item, hippocampus-dependent working memory is recruited instead at early list positions, the same hippocampal engagement supports long-term encoding Items early in the list are rehearsed more, and later items interfere with earlier ones, which is why the primacy advantage arises and why hippocampal engagement stops predicting memory later in the list intracranial rote epilepsy hippocampal sclerosis free recall encoding medial temporal lobe rehearsal subsequent memory primacy effect linear effects of position effects of position hippocampus-dependent WM hippocampus-independent WM list position group DC slopes negative BOLD response hemisphere slope of the DC potential ANOVA A two-way ANOVA with “position” as repeated measure a one-way ANOVA with position as repeated measure sphericity correction Huynh–Feldt corrected for inhomogeneities of covariance t-test post hoc t tests two-tailed paired t tests second-level one-sample t test regression we fitted a linear regression line on the DC slopes general linear model box-car regressor in a general linear model multiple-comparison correction statistical-significance repeated measures ANOVA with Huynh-Feldt correction linear regression of slopes across positions conditional versus unconditional probability cluster-level significance fmri

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second readingalternativesan independent re-reading of a facet, recorded whether or not it agrees
relation to other workclaim_relationshow this paper stands to another: agrees, contradicts, extends, qualifies
claim typeclaim_typethe shape of the claim: existence, mechanism, method, critique, prediction, null
concept edgeconcept_edgesan asserted relation between two concepts, with the sentence asserting it
content tagcontent_tagsa finding, definition, quantity, statistic or variable named in the paper
contributioncontributionwhat the paper claims to add
controlcontrolsan experiment run to eliminate a rival explanation
limitationlimitationswhat the paper says its evidence cannot support
mechanismmechanismshow the reported effect is said to come about
methodmethodhow the paper produced its evidence
object of studyobject_of_studywhat the paper is about — the thing it investigates
phenomenonphenomenonthe effect or regularity under study
research questionresearch_questionthe question the paper sets itself
sensesensesa distinct use of a term, quoted where the paper fixes it
transfertransfersa method or concept carried in from another field

The paper

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Every tag on this page

alternatives 3 · claim_relations 7 · claim_type 1 · concept_edges 3 · content_tags 56 · contribution 1 · controls 1 · limitations 12 · mechanisms 8 · method 1 · object_of_study 1 · phenomenon 1 · research_question 1 · senses 4 · transfers 1

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facettagfieldconf.whose words
object_of_studywhether working-memory-related hippocampal activity patterns during a word list predict long-term encoding, and explain the primacy effect
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object_of_studythe paper's words
research_questionwhich form of working memory supports the items at the start of a list
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research_questionthe paper's words
contributionshows hippocampal deactivation at the first list position predicts that the item will be forgotten, linking a working-memory signature to long-term encoding failure
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contributionthe paper's words
claim_typemechanism
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claim_typethe paper's words
phenomenonworking memory-long-term memory interaction in the hippocampus
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phenomenonthe paper's words
methodfmri
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also in: fmri
methodthe paper's words
mechanismsmechanisms
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mechanisms[0]the tagger's words
mechanismsmechanisms
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mechanisms[1]the tagger's words
mechanismsmechanisms
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status: the paper's own words: we suggest that
mechanisms[2]the tagger's words
mechanismsworking memory is divided by neural signature: hippocampal activation for multiple items, deactivation for individual items
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mechanisms[3]1.0the tagger's words
mechanismspositive DC slopes in the iEEG are taken to correspond to BOLD deactivations, which is what lets the two datasets be read together
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mechanisms[4]1.0the tagger's words
mechanismsitems at the start of a list are proposed to be rehearsed more, linking the primacy effect to working memory
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mechanisms[5]1.0the tagger's words
mechanismssuccessful early items are encoded in runs: encoding the first raises the conditional probability of encoding the second and third
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mechanisms[6]1.0the tagger's words
mechanismsonly the hemisphere contralateral to the seizure focus was analysed, in patients with confirmed unilateral sclerosis
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mechanisms[7]1.0the tagger's words
limitationsNo behavioural measure of working-memory performance was collected, so it is unknown which items were actually held in WM.
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what this limits: every claim about what was held in working memory: THERE IS NO BEHAVIORAL MONITORING OF WM PERFORMANCE IN THE CURRENT EXPERIMENT
limitations[0]the tagger's words
limitationsThe link between the observed activity and actual long-term memory consequences of WM performance cannot be established.
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what this limits: any claim about what was held in mind: WE DO NOT KNOW FOR SURE HOW LTM IS ACTUALLY AFFECTED BY THE WM PERFORMANCE FOR SPECIFIC ITEMS, I.E., IT IS UNKNOWN WHICH ITEMS ARE ACTUALLY MAINTAINED IN WM
limitations[1]the tagger's words
limitationsThe proposed link to persistent activity recorded in vitro is acknowledged as speculative.
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what this limits: the link to the in vitro literature: THIS ANALOGY BETWEEN ELECTROPHYSIOLOGICAL DATA IN VITRO AND THE NEURAL PATTERNS DURING WM MAINTENANCE MUST REMAIN SPECULATIVE AT THE CURRENT STAGE
limitations[2]the tagger's words
limitationsPosition-specific subsequent-memory analyses were statistically underpowered.
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what this limits: the position-specific null: THIS LACK OF AN EFFECT IS RELATED TO THE REDUCED STATISTICAL POWER FOR POSITION-SPECIFIC ANALYSES OF SUBSEQUENT MEMORY EFFECTS
limitations[3]the tagger's words
limitationsthe two populations differ in recall performance despite a shorter recall phase for the fMRI group
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what this limits: any comparison between the two populations: Quotation withheld
limitations[4]1.0the tagger's words
limitationsthe paradigms differ between the two versions in presentation time, interstimulus interval, block count and recall duration
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what this limits: any comparison between the fMRI and intracranial versions: they differ in presentation time (1000 ms, since goggles REQUIRED LONGER PRESENTATION TIMES), interstimulus interval and block count
limitations[5]1.0the tagger's words
limitationspatients showed both primacy and recency; the fMRI subjects showed no recency at all
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what this limits: any serial-position claim carried across the two groups: NO RECENCY EFFECTS AT THE END of the list were found in the fMRI subjects, against primacy AND recency in the patients
limitations[6]1.0the tagger's words
limitationsthe whole-brain effects did not survive correction for multiple comparisons and are reported at the cluster level instead
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what this limits: the whole-brain effects: THESE EFFECTS DID NOT SURVIVE CORRECTION FOR MULTIPLE COMPARISONS, BUT DID HAVE A LARGE EXTENSION OF 60 VOXELS FOR THE SERIAL POSITION CONTRAST OF SUBSEQUENTLY FORGOTTEN WORDS
limitations[7]1.0the tagger's words
limitationsseveral supporting results are trends at p between 0.05 and 0.07, labelled as such
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what this limits: the subsequently-forgotten results, with their exact values: WE FOUND A TREND FOR AN EFFECT OF POSITION (F (2,36) = 2.943; p = 0.070) AND FOR A LINEAR EFFECT OF POSITION (F (1,18) = 4.249; p = 0.054)
limitations[8]1.0the tagger's words
limitationsa position-specific null is attributed to reduced statistical power
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duplicate of: limitations[3] · what this limits: the same scope as limitations[3] - the position-specific analyses, underpowered
limitations[9]1.0the tagger's words
limitationseffects appear in the left hippocampus and not the right, without an account of why
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what this limits: any bilateral hippocampal claim: FOR THE RIGHT HIPPOCAMPUS, A THREE-WAY ANOVA DID NOT REVEAL AN EFFECT OF, OR INTERACTION WITH, THE FACTOR POSITION (ALL p > 0.3), BUT A SIGNIFICANT MAIN EFFECT OF MEMORY (F (1,18) = 8.898; p < 0.01)
limitations[10]0.9the tagger's words
limitationsthe required rote strategy is verified only by asking participants afterwards
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what this limits: the rote-strategy assumption: ALL SUBJECTS WERE REQUIRED TO USE A ROTE STRATEGY ... AVOIDING MEMORY AIDS - instructed, and verified only by asking afterwards
limitations[11]1.0the tagger's words
transfersserial position curves and the primacy effect, imported as the behavioural handle on a neural question
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transfers[0]1.0the tagger's words
controlsa group of healthy control subjects investigated with fMRI alongside the patient recordings
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eliminates: the hippocampal working-memory effect being a property of the epileptic brains the depth electrodes are implanted in · outcome (stated by the tagger, no quotation): held
controls[0]0.85the tagger's words
sensesworking memory is subdivided here by NEURAL SIGNATURE rather than by task or capacity: hippocampus-dependent when multiple items are held, hippocampus-independent when one is
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senses[0]1.0the tagger's words
sensesa deactivation is used as a positive finding - the absence of hippocampal engagement is the marker, not a null
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senses[1]1.0the tagger's words
senseseffects of position and LINEAR effects of position are distinguished explicitly, with the a priori assumption of the second stated: that if there are effects across positions, these should be linear
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senses[2]1.0the tagger's words
sensesthe DC potential and the BOLD signal are treated as two measures of one underlying quantity, on the strength of a cited correspondence rather than a within-subject comparison
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senses[3]1.0the tagger's words
alternativesclaim_type: mechanism
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a second reading reaching the same value. Recorded: mechanism — note: a second reading reaching the same value from a different sentence. Not applied - a person decides.
alternatives.claim_typethe tagger's words
alternativesmethod: intracranial EEG DC potentials in 8 epilepsy patients reanalysed from a 1999 study, plus new fMRI in 19 healthy subjects on the same word-list paradigm
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a second reading DIFFERING from the recorded value. Recorded: fmri — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.methodthe tagger's words
alternativesphenomenon: the primacy effect and its hippocampal correlates
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a second reading DIFFERING from the recorded value. Recorded: working memory-long-term memory interaction in the hippocampus — note: a second reading, verbatim-verified, disagreeing with the value on the record. Not applied - a person decides.
alternatives.phenomenonthe tagger's words
claim_relationsbuilds-on → Fernández et al., 1999
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claim_relations[0]the tagger's words
claim_relationsextends → Axmacher et al., 2007
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claim_relations[1]the tagger's words
claim_relationsbuilds-on → Atkinson and Shiffrin (1968)
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claim_relations[2]the tagger's words
claim_relationsextends → the authors' own Sternberg study distinguishing hippocampus-dependent from hippocampus-independent working memory
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claim_relations[3]1.0the tagger's words
claim_relationsextends → Fernandez et al. 1999, whose iEEG data are reanalysed here for a different question
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claim_relations[4]1.0the tagger's words
claim_relationssupports → the rehearsal account of the primacy effect
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claim_relations[5]0.85the tagger's words
claim_relationssupports → fMRI findings that working-memory maintenance facilitates long-term encoding
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claim_relations[6]0.8the tagger's words
concept_edgesdeactivation —causes→ long-term memory
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strength of the relation: 0.8 · note: predicts FAILURE of encoding; the sign matters and the schema records only the pair Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span says hippocampal deactivation predicts failure of encoding; the edge carries no direction.
concept_edges[0]0.85the tagger's words
concept_edgesworking memory —causes→ long-term memory
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status: ATTRIBUTED TO NAMED WORK AND OFFERED AS AN IDEA: 'IT HAS BEEN PROPOSED THAT the primacy effect is due to the fact that items at early list positions undergo more extensive rehearsal than items presented later (Rundus, 1971; Brodie and Murdock, 1977). THIS IDEA SUGGESTS a link between LTM formation and rehearsal processes'. The relation is real and this paper builds on it; it belongs to two papers from the 1970s. · strength of the relation: 0.8 · note: via rehearsal, which is the proposed account of the primacy effect Audit 2026-09-28: DOUBTFUL - read against its full span and kept, but the span says the idea 'SUGGESTS A LINK' between LTM formation and rehearsal attributed to working memory; causes is hedged.
concept_edges[2]0.9the tagger's words
concept_edgesprimacy —contrasted-with→ recency
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strength of the relation: 1 Audit 2026-09-28: DOUBTFUL - read against its span and kept, but the span reports only the recency effect; the contrast with primacy is not in it.
also in: primacy · recency
concept_edges[3]1.0the tagger's words
content_tagsANOVA
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also in: ANOVA
content_tags.statistics[0]the tagger's words
content_tagsANOVA
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also in: ANOVA
content_tags.statistics[1]the tagger's words
content_tagssphericity correction
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content_tags.statistics[2]the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[3]the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[4]the tagger's words
content_tagst-test
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also in: t-test
content_tags.statistics[5]the tagger's words
content_tagsregression
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also in: regression
content_tags.statistics[6]the tagger's words
content_tagsgeneral linear model
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content_tags.statistics[7]the tagger's words
content_tagsmultiple-comparison correction
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content_tags.statistics[8]the tagger's words
content_tagsstatistical-significance
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content_tags.statistics[9]the tagger's words
content_tagsrepeated measures ANOVA with Huynh-Feldt correction
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content_tags.statistics[10]0.85the tagger's words
content_tagslinear regression of slopes across positions
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content_tags.statistics[11]1.0the tagger's words
content_tagsconditional versus unconditional probability
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content_tags.statistics[12]1.0the tagger's words
content_tagscluster-level significance
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content_tags.statistics[13]1.0the tagger's words
content_tagsAs expected, more items were successfully encoded at the beginning of the list than at later positions.
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content_tags.findings[0]the tagger's words
content_tagsEarly-list words that were later forgotten were accompanied by hippocampal deactivation signatures in both fMRI and intracranial EEG.
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content_tags.findings[1]the tagger's words
content_tagsNo such deactivation appeared for later list items or for items that were successfully remembered.
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content_tags.findings[2]the tagger's words
content_tagsIn the intracranial data the linear decline of DC slope across list positions was significant only for subsequently forgotten items.
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content_tags.findings[3]the tagger's words
content_tagsA second, independent general linear model reproduced the effect: left-hippocampal negative BOLD for forgotten items at position one was absent at position seven.
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content_tags.findings[4]the tagger's words
content_tagsThe position effect was left-lateralised: the right hippocampus showed a main effect of subsequent memory but no position effect.
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content_tags.findings[5]the tagger's words
content_tagsFor both later-remembered and later-forgotten items, activity was most pronounced in the anterior cingulate cortex.
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content_tags.findings[6]the tagger's words
content_tagsSuccessful encoding at the start of a list came in runs: encoding the first word raised the probability of encoding the second.
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content_tags.findings[7]the tagger's words
content_tagsThe authors conclude that failing to engage hippocampus-dependent working memory early in a list causes the later encoding failure.
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content_tags.findings[8]the tagger's words
content_tagshippocampal deactivation at the first list position occurred for items that were later forgotten
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content_tags.findings[9]1.0the tagger's words
content_tagsno such deactivation appeared for items that were later remembered
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content_tags.findings[10]1.0the tagger's words
content_tagsDC slopes were positive early in the list for subsequently forgotten items and flattened later
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content_tags.findings[11]1.0the tagger's words
content_tagsonly the first position differed significantly from zero
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content_tags.findings[12]1.0the tagger's words
content_tagsin the iEEG subjects, encoding the first item raised the conditional probability of encoding the second (0.35 to 0.51)
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content_tags.findings[13]1.0the tagger's words
content_tagsthe patients showed a recency effect at the end of the list
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content_tags.findings[14]1.0the tagger's words
content_tagsthe healthy fMRI group showed primacy but no recency at all
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content_tags.findings[15]1.0the tagger's words
content_tagsa second general linear model comparing the first and seventh positions reproduced the result
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content_tags.findings[17]1.0the tagger's words
content_tagsprimacy effect
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content_tags.definitions[1]the tagger's words
content_tagslinear effects of position
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content_tags.definitions[2]the tagger's words
content_tagseffects of position
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content_tags.definitions[4]1.0the tagger's words
content_tagshippocampus-dependent WM
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content_tags.definitions[5]1.0the tagger's words
content_tagshippocampus-independent WM
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content_tags.definitions[6]1.0the tagger's words
content_tagslist position
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also in: list position
content_tags.variables[0]the tagger's words
content_tagsgroup
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also in: group
content_tags.variables[2]the tagger's words
content_tagsDC slopes
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also in: DC slopes
content_tags.variables[3]the tagger's words
content_tagsnegative BOLD response
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content_tags.variables[4]the tagger's words
content_tagshemisphere
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also in: hemisphere
content_tags.variables[6]the tagger's words
content_tagsserial position
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content_tags.variables[7]1.0the tagger's words
content_tagssubsequent memory
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[8]1.0the tagger's words
content_tagsslope of the DC potential
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.variables[9]1.0the tagger's words
content_tagsEight — epilepsy patients contributing intracranial EEG data
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[0]the tagger's words
content_tagsNineteen — healthy participants in the fMRI experiment
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[1]the tagger's words
content_tags4.9% — proportion of EEG trials rejected as artefacts before analysis
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[2]the tagger's words
content_tags0.35 / 0.51 — unconditional versus conditional probability of encoding the second list item in the iEEG patients
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[3]the tagger's words
content_tagssmaller than 0.19 — maximum collinearity between the fMRI model regressors for adjacent list items (0 = orthogonal, 1 = collinear)
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[4]the tagger's words
content_tagsF (11,198) = 21.156; p < 10 −16 — effect of serial position on recall performance in the fMRI participants
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[5]the tagger's words
content_tags8 epilepsy patients and 19 healthy subjects — the two samples
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[6]0.9the tagger's words
content_tags0.35 rising to 0.51 for the second item given the first — conditional encoding probability in patients
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[7]1.0the tagger's words
content_tags4.9% of trials excluded for artefacts — iEEG data rejection
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[8]1.0the tagger's words
content_tagsMNI 30/-15/-18 and -30/-15/-18 — the anatomically defined hippocampal regions of interest
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[9]1.0the tagger's words
content_tagsrecall phase 90 s for iEEG versus 60 s for fMRI, presentation 400 ms versus 1000 ms — the paradigm differences between the two versions
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
value source note: THE VALUE DRAWS ON THE PAPER BEYOND THIS QUOTE: part of it is printed elsewhere in the paper, not in this quote ('presented in uppercase letters for a duration of 400 ms.'). The 90 s is the 60 s of this quote plus the 30 s it says the other version lasted longer.
content_tags.quantities[10]0.9the tagger's words
content_tagsp = 0.054, p = 0.070, p = 0.054 — the trends reported as consistent with the iEEG result
Quotation withheld — this record is not cleared for redistribution. The sentence is in the paper at the link above.
content_tags.quantities[11]0.9the tagger's words

Interpretive choices

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© 2026 Shir Sivroni · ORCID 0009-0000-2597-2824