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        <title>Time scales of memory, learning, and plasticity</title>
        <author>Tetzlaff</author>
        <author>Kolodziejski</author>
        <author>Markelic</author>
        <author>Wörgötter</author>
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          <title>Time scales of memory, learning, and plasticity</title>
          <date>2012</date>
          <publisher>Biological Cybernetics</publisher>
          <idno type="DOI">10.1007/s00422-012-0529-z</idno>
          
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        <label>time scales of memory, learning, and synaptic plasticity</label>
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        <label>mechanisms</label>
        <term>Forgetting on finite, graded storage times acts as an implicit compression algorithm, holding down both the capacity needed and the time to retrieve an item</term>
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        <term>New memories overwrite and interfere with old ones, which lowers the network&#x27;s capacity demand but destroys important items - the cost that multiple time scales exist to avoid</term>
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        <term>Hebbian potentiation feeds back on itself - a strengthened synapse drives the postsynaptic neuron harder, which strengthens it further - producing runaway weight growth unless a counteracting mechanism is present</term>
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        <term>An organism evolves a given memory time scale only if its lifespan and environment exert evolutionary pressure for correlations at that scale</term>
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        <term>Spike-triggered calcium influx into the presynaptic terminal raises release probability (facilitation) while use depletes the vesicle pool (depression), giving synapses a seconds-long memory</term>
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        <label>The review is selective rather than exhaustive; only key findings are discussed for each link.</label>
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        <note>what this limits: the set of links the review treats at all: UP TO THIS POINT WE HAVE ONLY DISCUSSED LINKS FOR WHICH THERE IS ALREADY EXPERIMENTAL SUPPORT OR THEORETICAL MODELING</note>
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        <label>The ordering between the two categories is stated to be unresolved and posed as a question.</label>
        <note>what this limits: any account of memory transfer drawn from this review: the interaction between short- and long-term memory is said to remain unclear, and whether an item must pass through short-term memory first is asked rather than answered</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>A construct used throughout the learning-algorithm literature has no established physiology for its longer form.</label>
        <note>what this limits: any physiological reading of eligibility traces: they are used in many learning-algorithm approaches to bridge time gaps, and the authors state the physiological basis of longer traces is unclear</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <term>eligibility traces</term>
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        <label>time scale</label>
        <term>time scale</term>
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        <label>consolidation → retrieval</label>
        <note>THE TWO CANNOT BE SEPARATED IN THIS STUDY - the DESIGN form of the negative relation / No edge type can express it: all eight assert that a relation HOLDS. Recorded here because a confounding statement BOUNDS WHAT THE PAPER&#x27;S OWN RESULTS CAN MEAN, and it lives in a methods paragraph where nothing downstream will find it. A memory deficit cannot be attributed to either of the two processes the field distinguishes - so the distinction survives in the vocabulary while being unusable on the data.</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>a time scale - THREE DIFFERENT SPANS UNDER ONE EXPRESSION: working memory milliseconds to minutes, short-term memory minutes to days, long-term memory up to what the authors put in scare quotes as &#x27;infinity&#x27;; the phrase that organises the whole review denotes a different quantity in each of its three sections</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>long-term memory&#x27;s upper bound - IN THE AUTHORS&#x27; OWN SCARE QUOTES: given as up to &#x27;infinity&#x27;, with the immediate qualification that it is still under debate how items can be stored for such a long time, so the widest value in the review is the one marked as least settled</label>
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        <label>an eligibility trace - A MODELLING DEVICE WITHOUT ITS PHYSIOLOGY: named in 1943 and used in many learning-algorithm approaches to bridge time gaps, while the physiological basis of the longer traces is stated to be unclear, so the algorithms assume a mechanism the review declines to underwrite</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t23" type="claim_relations" ana="#claim_relations[0]" resp="#tagger">
        <label>builds-on → Martin et al. (2000) four criteria for relating plasticity to learning and memory</label>
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        <label>contradicts → Hebb (1949) proposed link between long-term memory and long-term plasticity</label>
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        <label>working memory —contrasted-with→ short-term memory</label>
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        <note>differs in: the span each covers: milliseconds to minutes, versus minutes to days · strength of the relation: 1</note>
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        <label>Working memory spans milliseconds to minutes, while short-term memory stores items over minutes to days.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>Having different storage time scales could avoid the deletion of important information.</label>
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        <label>Robot experiments show that when different time scales interact, more complex behaviour can be obtained.</label>
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      <span xml:id="t29" type="content_tags" ana="#content_tags.findings[3]" resp="#tagger">
        <label>Caenorhabditis elegans shows synaptic plasticity but no experience-based structural plasticity.</label>
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      <span xml:id="t30" type="content_tags" ana="#content_tags.findings[4]" resp="#tagger">
        <label>Of the proposed memory-physiology links, working memory-to-activity and long-term-memory-to-structural-plasticity have experimental support, whereas the classic long-term-memory-to-long-term-plasticity link is still disputed.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t31" type="content_tags" ana="#content_tags.findings[5]" resp="#tagger">
        <label>So far, mimicry cannot be experimentally induced.</label>
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      <span xml:id="t32" type="content_tags" ana="#content_tags.findings[6]" resp="#tagger">
        <label>Rats in enriched environments build significantly more dendritic spines than controls.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t33" type="content_tags" ana="#content_tags.findings[7]" resp="#tagger">
        <label>Adding synaptic scaling with a quadratic weight dependence to a generic plasticity rule can be shown mathematically to stabilise weight development.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>Structural plasticity is modelled only computationally because no analytical models of network topology change exist yet.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t35" type="content_tags" ana="#content_tags.findings[9]" resp="#tagger">
        <label>Whether the exact human forgetting curve is exponential or a power law remains unsettled.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t36" type="content_tags" ana="#content_tags.definitions[0]" resp="#tagger">
        <label>working memory</label>
        <term>working memory</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t37" type="content_tags" ana="#content_tags.definitions[1]" resp="#tagger">
        <label>long-term memory</label>
        <term>long-term memory</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>chunking</label>
        <term>chunking</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>detectability</label>
        <term>detectability</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t40" type="content_tags" ana="#content_tags.definitions[4]" resp="#tagger">
        <label>retrograde alteration</label>
        <term>retrograde alteration</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t41" type="content_tags" ana="#content_tags.definitions[5]" resp="#tagger">
        <label>structural plasticity</label>
        <term>structural plasticity</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t42" type="content_tags" ana="#content_tags.definitions[6]" resp="#tagger">
        <label>eligibility traces</label>
        <term>eligibility traces</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t43" type="content_tags" ana="#content_tags.definitions[7]" resp="#tagger">
        <label>intrinsic plasticity</label>
        <term>intrinsic plasticity</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t44" type="content_tags" ana="#content_tags.definitions[8]" resp="#tagger">
        <label>synaptic tagging</label>
        <term>synaptic tagging</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t45" type="content_tags" ana="#content_tags.definitions[9]" resp="#tagger">
        <label>reservoir computing</label>
        <term>reservoir computing</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t46" type="content_tags" ana="#content_tags.definitions[10]" resp="#tagger">
        <label>synaptic scaling</label>
        <term>synaptic scaling</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t47" type="content_tags" ana="#content_tags.quantities[0]" resp="#tagger">
        <label>about 10 days — how quickly the nervous system&#x27;s storage capacity would be exhausted if every bit of input were stored</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t48" type="content_tags" ana="#content_tags.quantities[1]" resp="#tagger">
        <label>four and seven — the number of items working memory can hold</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t49" type="content_tags" ana="#content_tags.quantities[2]" resp="#tagger">
        <label>tens of milliseconds — the time scale of the postsynaptic potential, the shortest memory-like physiological trace</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t50" type="content_tags" ana="#content_tags.quantities[3]" resp="#tagger">
        <label>μ = 0.01 corresponds to 100 ms — how the alpha-function time-scale parameter maps onto the filter&#x27;s temporal duration</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t51" type="content_tags" ana="#content_tags.quantities[4]" resp="#tagger">
        <label>four — the number of criteria Martin et al. proposed for establishing that plasticity underlies learning and memory (detectability, mimicry, anterograde and retrograde alteration)</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t52" type="content_tags" ana="#content_tags.quantities[5]" resp="#tagger">
        <label>milliseconds to seconds — the time scale of short-term plasticity</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t53" type="content_tags" ana="#content_tags.quantities[6]" resp="#tagger">
        <label>hours to several days — the time scale on which structural plasticity rewires the network</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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