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        <title>Auditory Thalamocortical Synaptic Transmission In Vitro</title>
        <author>Cruikshank</author>
        <author>Rose</author>
        <author>Metherate</author>
      </titleStmt>
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        <bibl>
          <title>Auditory Thalamocortical Synaptic Transmission In Vitro</title>
          <date>2002</date>
          <publisher>Journal of Neurophysiology</publisher>
          <idno type="DOI">10.1152/jn.00549.2001</idno>
          
          <idno type="record">Cruikshank2002-auditory-thalamocortical-synaptic-transmissi</idno>
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          <note type="text-sha1">9138ba54e36ec737aa320fb26a7b3340d8e02dbe</note>
          <note type="rights">not-redistributable</note>
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        <person xml:id="paper"><p>the paper's own words</p></person>
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      <span xml:id="t0" type="claim_type" ana="#claim_type" resp="#paper">
        <label>method</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t1" type="phenomenon" ana="#phenomenon" resp="#paper">
        <label>auditory thalamocortical synaptic transmission</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t2" type="method" ana="#method" resp="#paper">
        <label>electrophysiology</label>
        <term>electrophysiology</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t3" type="mechanisms" ana="#mechanisms[0]" resp="#tagger">
        <label>mechanisms</label>
        <term>Thalamic glutamatergic axons excite cortical inhibitory interneurons, which then synapse on the recorded cell, producing the disynaptic fast IPSP</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t4" type="mechanisms" ana="#mechanisms[1]" resp="#tagger">
        <label>mechanisms</label>
        <term>Antidromic activation of layer 5 corticotectal cells whose axons cross the shell stimulation site, feeding back through their intracortical collaterals to generate the layer 1 sink</term>
        <note>status: the paper&#x27;s own words: it is possible that</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t5" type="limitations" ana="#limitations[0]" resp="#tagger">
        <label>The alternative explanation for the shell slice layer 1 sink (antidromically activated layer 5 corticotectal cells) cannot be excluded</label>
        <note>what this limits: the layer 1 sink interpretation: IT DOES NOT PROVE THAT THIS HYPOTHETICAL MODE OF ACTIVATION ACTUALLY OCCURS OR THAT DIRECT INPUT FROM THE NONPRIMARY AUDITORY THALAMUS HAD NO ROLE</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t6" type="limitations" ana="#limitations[1]" resp="#tagger">
        <label>Because of that unresolved alternative, the authors advise caution in using the shell preparation</label>
        <note>what this limits: any use of the shell preparation while the alternative is unresolved: THUS AT PRESENT IT SHOULD BE USED WITH CAUTION</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t7" type="limitations" ana="#limitations[2]" resp="#tagger">
        <label>The pharmacological design cannot isolate the AMPA/KA contribution, since APV was never given alone</label>
        <note>what this limits: any AMPA/KA conclusion from the pharmacology: THIS SUBTRACTIVE LOGIC CANNOT BE USED TO IMPLICATE AMPA/KA RECEPTORS BECAUSE APV WAS NEVER APPLIED BY ITSELF</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t8" type="limitations" ana="#limitations[3]" resp="#tagger">
        <label>The synaptic order of the early response is unresolved</label>
        <note>what this limits: the synaptic order of the early response: EXACTLY HOW MANY LINKS ARE INVOLVED IN THE EARLY RESPONSE REMAINS TO BE DETERMINED</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t9" type="limitations" ana="#limitations[4]" resp="#tagger" cert="1.0">
        <label>A measured quantity is stated not to be comparable across studies, because its visibility depends on method and criteria.</label>
        <note>what this limits: any comparison of IPSP amplitudes between this study and another: the authors state</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t10" type="limitations" ana="#limitations[5]" resp="#tagger" cert="1.0">
        <label>The conventional stimulation method is stated to be possibly unreliable for the purpose it is used for.</label>
        <note>what this limits: any result attributing a response to thalamocortical input on the basis of columnar stimulation: the authors state their MG-evoked profiles differ from those produced by conventional columnar stimulation, which may not be a reliable means of selectively activating thalamocortical inputs</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t11" type="limitations" ana="#limitations[6]" resp="#tagger" cert="1.0">
        <label>Two stimulation sites are reported not to activate the same synapses, with different laminar profiles.</label>
        <note>what this limits: any treatment of on-beam and MG stimulation as interchangeable probes: they are stated not to activate the same sets of synapses in cortex, and to evoke different laminar profiles</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t12" type="limitations" ana="#limitations[7]" resp="#tagger" cert="1.0">
        <label>The AMPA/KA conclusion is conditional on an assumption the authors state is difficult to prove.</label>
        <note>what this limits: the claim that AMPA/KA receptors mediate transmission at these synapses: it follows if one assumes the fast EPSP in the middle layers is a monosynaptic event, and the authors state it is difficult to prove that a response is monosynaptic</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t13" type="limitations" ana="#limitations[8]" resp="#tagger" cert="1.0">
        <label>Whether the preparation&#x27;s evoked activation resembles the intact system is stated to be undetermined.</label>
        <note>what this limits: any inference from these evoked responses to sound-driven cortical activity: the authors state it remains to be determined whether the activation produced by MG stimulation represents a natural pattern similar to that produced by acoustic stimulation in vivo</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t14" type="limitations" ana="#limitations[9]" resp="#tagger" cert="1.0">
        <label>Inhibition may be incompletely developed at the age used, indicated by the marker staining in this material.</label>
        <note>what this limits: any absolute inhibitory strength taken from these recordings: cortical inhibitory systems are known not to be fully developed in juvenile rodents, and the modest cortical PV labeling observed here is stated to indicate it</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t15" type="controls" ana="#controls[0]" resp="#tagger" cert="0.95">
        <label>on-beam and MG-evoked IPSPs compared DIRECTLY WITHIN THE SAME SLICES rather than across the preparation</label>
        <note>eliminates: preparation-dependent differences - slice health, cutting angle, anything that varies between preparations and would otherwise be read as a difference between the two pathways · outcome (stated by the tagger, no quotation): held</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t16" type="controls" ana="#controls[1]" resp="#tagger" cert="0.95">
        <label>a re-analysis restricted to the SEVEN CELLS in which MG-evoked EPSPs were at least as strong as on-beam EPSPs</label>
        <note>eliminates: weaker MG excitation as the explanation of weaker MG inhibition - if the thalamic pathway simply drove the slice less hard, smaller IPSPs would follow with no difference in inhibition itself · outcome (stated by the tagger, no quotation): held</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t17" type="concepts" ana="#concepts[0]" resp="#paper">
        <label>columnar stimulation</label>
        <term>columnar stimulation</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t18" type="concepts" ana="#concepts[1]" resp="#paper">
        <label>thalamocortical inputs</label>
        <term>thalamocortical inputs</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t19" type="concepts" ana="#concepts[2]" resp="#paper">
        <label>MG stimuli</label>
        <term>MG stimuli</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t20" type="concepts" ana="#concepts[3]" resp="#paper">
        <label>on-beam stimuli</label>
        <term>on-beam stimuli</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t21" type="concepts" ana="#concepts[4]" resp="#paper">
        <label>IPSPs</label>
        <term>IPSPs</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t22" type="concepts" ana="#concepts[5]" resp="#paper">
        <label>excitatory thalamic projections</label>
        <term>excitatory thalamic projections</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t23" type="concepts" ana="#concepts[6]" resp="#paper">
        <label>cortical inhibitory interneurons</label>
        <term>cortical inhibitory interneurons</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t24" type="senses" ana="#senses[0]" resp="#tagger" cert="1.0">
        <label>an IPSP amplitude, across papers - NOT A COMPARABLE QUANTITY: whether an IPSP can be observed at all, and how large it is, depends heavily on methodology and criteria, so the same named measurement in two studies is two measurements and the authors state the comparison is difficult</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t25" type="senses" ana="#senses[1]" resp="#tagger" cert="1.0">
        <label>columnar (on-beam) stimulation - NOT A SELECTIVE THALAMOCORTICAL PROBE: it is used to activate thalamocortical inputs, and the authors state it may not be a reliable means of doing so selectively, because MG and on-beam stimuli do not activate the same sets of synapses</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t26" type="senses" ana="#senses[2]" resp="#tagger" cert="1.0">
        <label>the AMPA/KA finding - HELD UP BY AN ASSUMPTION, NOT A CONTROL: the subtractive logic could not be used because APV was never applied alone, so the conclusion follows only if the fast EPSP is a monosynaptic event, which the authors state is difficult to prove</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t27" type="senses" ana="#senses[3]" resp="#tagger" cert="0.8">
        <label>the early IPSPs - CORTICAL IN ROUTE, INFERRED FROM A BLOCKER RATHER THAN FROM THE RESPONSE: a direct inhibitory thalamic projection is ruled out because an AMPA/KA antagonist blocked them, and the remaining account is that excitatory thalamic projections drove cortical interneurons which then synapsed on the recorded cells; the paper states this as likely, not as shown</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t28" type="claim_relations" ana="#claim_relations[0]" resp="#tagger">
        <label>builds-on → Agmon and Connors (1991) somatosensory thalamocortical slice preparation</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t29" type="claim_relations" ana="#claim_relations[1]" resp="#tagger">
        <label>extends → Metherate and Cruikshank (1999), the authors&#x27; own initial feasibility study in rat and mouse slices</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t30" type="claim_relations" ana="#claim_relations[2]" resp="#tagger">
        <label>builds-on → Barth and MacDonald (1996) finding that nonlemniscal stimulation evokes cortical gamma band activity</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t31" type="concept_edges" ana="#concept_edges[0]" resp="#tagger">
        <label>MG stimuli —contrasted-with→ on-beam stimuli</label>
        <term>MG stimuli</term>
        <term>on-beam stimuli</term>
        <note>differs in: which synapses they activate, and the laminar profile they evoke · strength of the relation: 1</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t32" type="concept_edges" ana="#concept_edges[1]" resp="#tagger">
        <label>cortical inhibitory interneurons —causes→ IPSPs</label>
        <term>cortical inhibitory interneurons</term>
        <term>IPSPs</term>
        <note>direction on the measure: increases · strength of the relation: 0.8</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
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        <label>t-test</label>
        <term>t-test</term>
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        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t34" type="content_tags" ana="#content_tags.statistics[1]" resp="#tagger">
        <label>t-test</label>
        <term>t-test</term>
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        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>mean</label>
        <term>mean</term>
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        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t36" type="content_tags" ana="#content_tags.findings[0]" resp="#tagger">
        <label>An in vitro mouse brain preparation can preserve functionally intact thalamocortical connections from the MGv to auditory cortex.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t37" type="content_tags" ana="#content_tags.findings[1]" resp="#tagger">
        <label>Auditory thalamocortical transmission is carried by both AMPA/kainate and NMDA glutamate receptors.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t38" type="content_tags" ana="#content_tags.findings[2]" resp="#tagger">
        <label>Every primary slice analysed showed a dominant current sink in cortical layers 3-4 after medial geniculate stimulation.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t39" type="content_tags" ana="#content_tags.findings[3]" resp="#tagger">
        <label>Thalamus-evoked responses differ from those of conventional columnar stimulation, indicating columnar stimulation may not reliably activate thalamocortical inputs selectively.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t40" type="content_tags" ana="#content_tags.findings[4]" resp="#tagger">
        <label>A second &#x27;shell&#x27; slice is presented in which subcortical stimulation appears to selectively activate synapses in superficial layer 1 of temporal cortex.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t41" type="content_tags" ana="#content_tags.findings[5]" resp="#tagger">
        <label>Tracing confirmed that layer 5 corticotectal axons pass through the shell stimulation region on their way to the inferior colliculus.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t42" type="content_tags" ana="#content_tags.definitions[1]" resp="#tagger">
        <label>on-beam stimulation</label>
        <term>on-beam stimulation</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t43" type="content_tags" ana="#content_tags.definitions[2]" resp="#tagger">
        <label>early response</label>
        <term>early response</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t44" type="content_tags" ana="#content_tags.definitions[3]" resp="#tagger">
        <label>late responses</label>
        <term>late responses</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t45" type="content_tags" ana="#content_tags.definitions[4]" resp="#tagger">
        <label>primary slice</label>
        <term>primary slice</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t46" type="content_tags" ana="#content_tags.definitions[5]" resp="#tagger">
        <label>shell slice</label>
        <term>shell slice</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t47" type="content_tags" ana="#content_tags.variables[3]" resp="#tagger">
        <label>stimulus intensity</label>
        <term>stimulus intensity</term>
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        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t48" type="content_tags" ana="#content_tags.variables[4]" resp="#tagger">
        <label>current source density</label>
        <term>current source density</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t49" type="content_tags" ana="#content_tags.variables[5]" resp="#tagger">
        <label>amplitudes of the early responses</label>
        <term>amplitudes of the early responses</term>
        <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[0]" resp="#tagger">
        <label>66 cells / 24 slices — number of intracellularly recorded layer 3-4 neurons and the number of primary slices they came from</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[1]" resp="#tagger">
        <label>4.0 ± 0.1 ms — mean onset latency of the intracellular early response, statistically indistinguishable from the middle-layer CSD sink latency</label>
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        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t52" type="content_tags" ana="#content_tags.quantities[2]" resp="#tagger">
        <label>16/16 — proportion of histologically checked primary slices whose top contained at least part of the ventral medial geniculate</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[3]" resp="#tagger">
        <label>15/20 — fraction of tested cells that displayed an inhibitory postsynaptic potential following the initial EPSP</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t54" type="content_tags" ana="#content_tags.quantities[4]" resp="#tagger">
        <label>6.0 ± 0.2 ms — mean onset latency of the shell-slice layer 1 surface sink, longer than the infragranular sink latency</label>
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        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t55" type="content_tags" ana="#content_tags.quantities[5]" resp="#tagger">
        <label>22 — number of primary slices with laminar CSD analysis, all of which showed a dominant middle-layer sink</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
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    <body>
      <p><note type="rights">The full text of this article is not redistributable, so it is
      not included here. The evidence sentences are withheld with it; the tag layer and its
      grounding status remain. The article itself is at https://doi.org/10.1152/jn.00549.2001.</note></p>
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