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        <title>Spatial cognition in birds</title>
        <author>Ben-Tov</author>
        <author>Gutfreund</author>
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        <p>Tag layer of the DHSS Meta-Tagging Project. The article text is reproduced from the
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        <bibl>
          <title>Spatial cognition in birds</title>
          <date>2022</date>
          <publisher>Current Biology</publisher>
          <idno type="DOI">10.1016/j.cub.2022.06.080</idno>
          
          <idno type="record">Ben-Tov2022-spatial-cognition-in-birds</idno>
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          <note type="rights">not-redistributable</note>
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              who="build_paper_edition.py">0 spans located by searching the stored text for the quote's exact words, allowing any whitespace between them</change>
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      <span xml:id="t0" type="claim_type" ana="#claim_type" resp="#paper">
        <label>existence</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t1" type="phenomenon" ana="#phenomenon" resp="#paper">
        <label>spatial cognition in birds</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t2" type="method" ana="#method" resp="#paper">
        <label>review</label>
        <term>review</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t3" type="mechanisms" ana="#mechanisms[0]" resp="#tagger">
        <label>mechanisms</label>
        <term>Species differ in place-cell abundance because the ecological demand for spatial memory differs: caching and homing species that must remember many locations develop robust place coding, species that do not need it show little</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t4" type="mechanisms" ana="#mechanisms[1]" resp="#tagger">
        <label>mechanisms</label>
        <term>Because caching birds must retrieve many hidden items, selection for that behaviour is what produces the enlarged hippocampus and abundant place cells, so place cells are proposed to serve spatial memory rather than merely accompany it</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t5" type="mechanisms" ana="#mechanisms[2]" resp="#tagger">
        <label>mechanisms</label>
        <term>Head direction tuning in a migratory seabird is anchored by the earth&#x27;s magnetic field, the cells encoding head angle relative to magnetic north</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t6" type="mechanisms" ana="#mechanisms[3]" resp="#tagger">
        <label>mechanisms</label>
        <term>Knowing one&#x27;s own place and heading requires converting head- and body-centred sensory input into an environment-fixed frame, a transformation performed in and upstream of the hippocampus</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t7" type="limitations" ana="#limitations[0]" resp="#tagger">
        <label>The neural mechanisms of avian spatial cognition remain poorly understood.</label>
        <note>what this limits: any mechanistic claim in the review: OUR UNDERSTANDING OF THE NEURAL MECHANISMS OF SPATIAL COGNITION IN BIRDS IS STILL POOR</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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        <label>All avian hippocampal recordings so far come from small enclosures, unlike the scales at which birds actually navigate.</label>
        <note>what this limits: every avian hippocampal recording in the literature: TO DATE, ALL THE EXPERIMENTS IN WHICH BRAIN ACTIVITY WAS RECORDED IN THE HIPPOCAMPUS OF BIRDS HAVE BEEN CONDUCTED IN RELATIVELY SMALL ENVIRONMENTS - unlike the scales at which birds navigate</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[2]" resp="#tagger">
        <label>How the earth&#x27;s magnetic field shapes the avian head-direction system is not yet resolved.</label>
        <note>what this limits: the magnetic-field question specifically: FURTHER EXAMINATION OF HEAD DIRECTION CELLS IN STREAKED SHEARWATER AND OTHER BIRDS IS NECESSARY TO RESOLVE HOW THE HEAD DIRECTION SYSTEM IN THE AVIAN HIPPOCAMPUS IS INFLUENCED BY THE EARTH&#x27;S MAGNETIC FIELD</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t10" type="limitations" ana="#limitations[3]" resp="#tagger" cert="1.0">
        <label>The category&#x27;s criterion comes from another species, and 2% of the units tested meet it.</label>
        <note>what this limits: any claim about place cells in birds: of more than 2,000 single units recorded in quail hippocampus, only about 2% passed the criterion for place cells, and most of those did not show the clear single field expected from the classical place cells found in rodents</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t11" type="limitations" ana="#limitations[4]" resp="#tagger" cert="1.0">
        <label>The structure is called a homologue and stated to differ fundamentally in architecture.</label>
        <note>what this limits: any transfer of a mammalian hippocampal result to birds: the avian homologue of the mammalian hippocampus has an architecture that, after three hundred million years of independent evolution, differs fundamentally from the mammalian one</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t12" type="concepts" ana="#concepts[0]" resp="#paper">
        <label>place cells</label>
        <term>place cells</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t13" type="concepts" ana="#concepts[1]" resp="#paper">
        <label>avian homologue</label>
        <term>avian homologue</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t14" type="concepts" ana="#concepts[2]" resp="#paper">
        <label>mammalian hippocampus</label>
        <term>mammalian hippocampus</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t15" type="concepts" ana="#concepts[3]" resp="#paper">
        <label>cognitive maps</label>
        <term>cognitive maps</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t16" type="concepts" ana="#concepts[4]" resp="#paper">
        <label>criterion</label>
        <term>criterion</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <span xml:id="t17" type="senses" ana="#senses[0]" resp="#tagger" cert="1.0">
        <label>a place cell - A RODENT-DERIVED CRITERION APPLIED ACROSS A PHYLUM: about 2% of more than 2,000 quail hippocampal units passed the criterion, and most of those lacked the clear single field of activity the classical rodent place cells show, so the category is being tested in birds by a standard defined on the animal it was discovered in</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t18" type="senses" ana="#senses[1]" resp="#tagger" cert="1.0">
        <label>the avian hippocampus - A HOMOLOGUE THAT DIFFERS FUNDAMENTALLY: identified as the avian homologue of the mammalian hippocampus by development, connectivity and cellular markers, and stated in the next sentence to have an architecture fundamentally different after three hundred million years of independent evolution - the correspondence and the difference are asserted together</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t19" type="senses" ana="#senses[2]" resp="#tagger" cert="1.0">
        <label>a cognitive map - A DIFFERENT ORDER OF MAGNITUDE IN BIRDS: the well-studied rodent maps cover small areas while avian maps must represent ranges of hundreds and thousands of kilometres, so the same term names representations whose spatial scale differs by several orders</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t20" type="claim_relations" ana="#claim_relations[0]" resp="#tagger">
        <label>builds-on → Edward Tolman, &#x27;Cognitive maps in rats and men&#x27;</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t21" type="content_tags" ana="#content_tags.findings[0]" resp="#tagger">
        <label>The avian hippocampus is a neuronal sheet on the dorsal surface of the caudal medial pallium, whereas the mammalian hippocampus is a folded structure below the neocortex.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t22" type="content_tags" ana="#content_tags.findings[1]" resp="#tagger">
        <label>The avian hippocampus lacks the granule layer that defines the mammalian structure.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t23" type="content_tags" ana="#content_tags.findings[2]" resp="#tagger">
        <label>Hippocampal-lesioned homing pigeons consistently show impaired homing.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t24" type="content_tags" ana="#content_tags.findings[3]" resp="#tagger">
        <label>Immediate-early gene expression rises in hippocampal neurons of birds searching remembered food locations, a second line of evidence tying the structure to spatial memory.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t25" type="content_tags" ana="#content_tags.findings[4]" resp="#tagger">
        <label>In food-caching species such as titmice, chickadees and jays, the hippocampus is larger than in most non-caching birds.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t26" type="content_tags" ana="#content_tags.findings[5]" resp="#tagger">
        <label>Recordings in Japanese quail found that only about 2% of units passed the place-cell criterion, and most of these lacked a clear single field.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t27" type="content_tags" ana="#content_tags.findings[6]" resp="#tagger">
        <label>The tufted titmouse recordings gave the first clear evidence of place cells in a bird species.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t28" type="content_tags" ana="#content_tags.findings[7]" resp="#tagger">
        <label>In the non-caching zebra finch, place cells were significantly rarer and less robust.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t29" type="content_tags" ana="#content_tags.findings[8]" resp="#tagger">
        <label>Preliminary recordings found hippocampal cells in flying barn owls that fired reliably when the owl flew through a confined region of the flight room.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t30" type="content_tags" ana="#content_tags.findings[9]" resp="#tagger">
        <label>Head direction cells in these birds mostly pointed to or near north.</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t31" type="content_tags" ana="#content_tags.definitions[0]" resp="#tagger">
        <label>spatial cognition</label>
        <term>spatial cognition</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t32" type="content_tags" ana="#content_tags.definitions[1]" resp="#tagger">
        <label>place cell</label>
        <term>place cell</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t33" type="content_tags" ana="#content_tags.definitions[2]" resp="#tagger">
        <label>cognitive map</label>
        <term>cognitive map</term>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t34" type="content_tags" ana="#content_tags.definitions[4]" resp="#tagger">
        <label>border cells</label>
        <term>border cells</term>
        <note>about this quote: THE QUOTE CONTINUES A SENTENCE THAT THIS COPY CUTS IN TWO - another paragraph, moved between the list and its tail, stands between it and the words it follows, which read &#x27;a whole range of spatial cell types have been discovered&#x27;.</note>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
      </span>
      <span xml:id="t35" type="content_tags" ana="#content_tags.definitions[5]" resp="#tagger">
        <label>central-place foraging</label>
        <term>central-place foraging</term>
        <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.definitions[6]" resp="#tagger">
        <label>place cells</label>
        <term>place cells</term>
        <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.definitions[7]" resp="#tagger">
        <label>head direction cells</label>
        <term>head direction cells</term>
        <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.definitions[8]" resp="#tagger">
        <label>grid cells</label>
        <term>grid cells</term>
        <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.quantities[0]" resp="#tagger">
        <label>~2% — Proportion of recorded quail hippocampal units meeting the place-cell criterion</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.quantities[1]" resp="#tagger">
        <label>about 12% — Proportion of quail hippocampal cells classified as head direction cells</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.quantities[2]" resp="#tagger">
        <label>More than 60% — Proportion of recorded tufted titmouse hippocampal cells classified as place cells</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.quantities[3]" resp="#tagger">
        <label>more than 2,000 — Single units analysed in the quail hippocampus study</label>
        <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.quantities[4]" resp="#tagger">
        <label>around 300 million years — Time since the bird and mammal lineages diverged, the evolutionary distance that makes the comparison informative</label>
        <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.quantities[5]" resp="#tagger">
        <label>200-meter-long — Length of the tunnel in which flying bats showed multifocal place fields, the cited evidence that spatial coding changes with environment scale</label>
        <note type="quotation-withheld">Quotation withheld because this record is not cleared for redistribution.</note>
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      <p><note type="rights">The full text of this article is not redistributable, so it is
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      grounding status remain. The article itself is at https://doi.org/10.1016/j.cub.2022.06.080.</note></p>
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