419 lines
91 KiB
XML
419 lines
91 KiB
XML
<document ID-DOI="10.1126/science.1108397" ID-GBIF-Dataset="cd2ea154-a1f5-4aeb-914c-fa2ea6fe1dd4" ID-GBIF-Taxon="163433813" ID-Zenodo-Dep="3739828" checkinTime="1586002569485" checkinUser="jeremy" docAuthor="Mary H. Schweitzer, Jennifer L. Wittmeyer, John R. Horner & Jan K. Toporskrif" docDate="2005" docId="03E28E29FFD6190DFC2EF4006FE5F91E" docLanguage="en" docName="Schweitzeretal2005SoftTissueVesselsABBYY2.pdf.imf" docOrigin="Science 307" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex Osborn 1905" docType="treatment" docVersion="12" lastPageId="4" lastPageNumber="1955" masterDocId="FFDBF651FFD71909FF81FFB66A60FF99" masterDocTitle="Soft-Tissue Vessels and Cellular Preservation in Tyrannosaurus rex" masterLastPageNumber="1955" masterPageNumber="1952" pageId="1" pageNumber="1952" updateTime="1632324545270" updateUser="ExternalLinkService">
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<mods:mods xmlns:mods="http://www.loc.gov/mods/v3">
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<mods:titleInfo>
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<mods:title>Soft-Tissue Vessels and Cellular Preservation in Tyrannosaurus rex</mods:title>
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</mods:titleInfo>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Mary H. Schweitzer</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Jennifer L. Wittmeyer</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>John R. Horner</mods:namePart>
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</mods:name>
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<mods:name type="personal">
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<mods:role>
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<mods:roleTerm>Author</mods:roleTerm>
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</mods:role>
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<mods:namePart>Jan K. Toporskrif</mods:namePart>
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</mods:name>
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<mods:typeOfResource>text</mods:typeOfResource>
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<mods:relatedItem type="host">
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<mods:titleInfo>
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<mods:title>Science</mods:title>
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</mods:titleInfo>
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<mods:part>
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<mods:date>2005</mods:date>
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<mods:detail type="volume">
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<mods:number>307</mods:number>
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</mods:detail>
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<mods:extent unit="page">
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<mods:start>1952</mods:start>
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<mods:end>1955</mods:end>
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</mods:extent>
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</mods:part>
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</mods:relatedItem>
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<mods:classification>journal article</mods:classification>
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<mods:identifier type="DOI">10.1126/science.1108397</mods:identifier>
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<mods:identifier type="GBIF-Dataset">cd2ea154-a1f5-4aeb-914c-fa2ea6fe1dd4</mods:identifier>
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<mods:identifier type="Zenodo-Dep">3739828</mods:identifier>
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</mods:mods>
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<treatment ID-GBIF-Taxon="163433813" LSID="urn:lsid:plazi:treatment:03E28E29FFD6190DFC2EF4006FE5F91E" httpUri="http://treatment.plazi.org/id/03E28E29FFD6190DFC2EF4006FE5F91E" lastPageId="4" lastPageNumber="1955" pageId="1" pageNumber="1952">
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<subSubSection lastPageId="2" lastPageNumber="1953" pageId="1" pageNumber="1952" type="nomenclature">
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<paragraph blockId="1.[942,1623,2996,3117]" lastBlockId="2.[189,869,284,405]" lastPageId="2" lastPageNumber="1953" pageId="1" pageNumber="1952">
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A newly discovered specimen of
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<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1952" phylum="Chordata" rank="species" species="rex">
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<emphasis italics="true" pageId="1" pageNumber="1952">Tyrannosaurus rex</emphasis>
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</taxonomicName>
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[
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<materialsCitation ID-GBIF-Occurrence="2598703895" collectionCode="MOR" formation="Hell Creek Formation, 8 m above the Fox Hills Sandstone" pageId="1" pageNumber="1952" specimenCode="MOR 1125">Museum of the Rockies (MOR) specimen 1125] was found at the base of the Hell Creek Formation, 8 m above the Fox Hills Sandstone</materialsCitation>
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, as an association of disarticulated elements. The specimen was
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<heading box="[189,868,288,316]" centered="true" fontSize="10" level="5" pageId="2" pageNumber="1953" reason="1">incorporated within a soft, well-sorted sandstone that</heading>
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</paragraph>
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</subSubSection>
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<subSubSection lastPageId="4" lastPageNumber="1955" pageId="2" pageNumber="1953" type="description">
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<paragraph blockId="2.[189,869,284,405]" lastBlockId="2.[928,1608,283,404]" pageId="2" pageNumber="1953">
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<heading box="[190,867,332,360]" centered="true" fontSize="10" level="5" pageId="2" pageNumber="1953" reason="1">
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was interpreted as estuarine in origin.
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<heading box="[190,869,375,403]" centered="true" fontSize="10" level="5" pageId="2" pageNumber="1953" reason="1">Although some bones are slightly deformed or</heading>
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</heading>
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<heading box="[928,1608,288,316]" fontSize="10" level="4" pageId="2" pageNumber="1953" reason="1">
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crushed, preservation is excellent.
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<materialsCitation ID-GBIF-Occurrence="2595781661" collectionCode="MOR" pageId="2" pageNumber="1953" specimenCode="MOR 1125">
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MOR
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<paragraph blockId="2.[928,1608,283,404]" lastBlockId="2.[1667,2351,288,883]" pageId="2" pageNumber="1953">
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<heading box="[932,1607,331,359]" fontSize="10" level="4" pageId="2" pageNumber="1953" reason="1">1125 represents a relatively small individual</heading>
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<heading box="[928,1607,375,403]" fontSize="10" level="4" pageId="2" pageNumber="1953" reason="1">
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of
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<emphasis box="[971,1062,375,403]" italics="true" pageId="2" pageNumber="1953">
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[971,1056,375,403]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1953" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
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,
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</emphasis>
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with a femoral length of 107 cm, as
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</heading>
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compared to the Field Museum (Chicago) specimen (
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<materialsCitation ID-GBIF-Occurrence="2598703896" box="[1836,2083,332,360]" collectionCode="FMNH" pageId="2" pageNumber="1953" specimenCode="FMNH PR2081">
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FMNH
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<emphasis box="[1962,2010,332,360]" italics="true" pageId="2" pageNumber="1953">PR</emphasis>
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2081
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</materialsCitation>
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) that has a femoral length of approximately 131 cm. On the basis of calculated lines of arrested growth (LAG), we estimated that this animal was 18 + 2 years old at death (
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<bibRefCitation author="M. Wuttke" box="[1988,2007,506,534]" editor="S. Schaal & W. Ziegler" journalOrPublisher="Verlag Waldemar Kramer, Frankfurt am Main, Germany" pageId="2" pageNumber="1953" pagination="265 - 274" refId="ref3328" refString="7. M. Wuttke, in Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens, S. Schaal, W. Ziegler, Eds. (Verlag Waldemar Kramer, Frankfurt am Main, Germany, 1988), pp. 265 - 274." type="book chapter" volumeTitle="Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens" year="1988">7</bibRefCitation>
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).
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</paragraph>
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</materialsCitation>
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</heading>
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</paragraph>
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<paragraph blockId="2.[1667,2351,288,883]" lastBlockId="3.[1670,2359,288,3116]" lastPageId="3" lastPageNumber="1954" pageId="2" pageNumber="1953">
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No preservatives were applied to interior fragments of the femur of
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<materialsCitation ID-GBIF-Occurrence="2595781663" box="[2073,2241,593,621]" collectionCode="MOR" pageId="2" pageNumber="1953" specimenCode="MOR 1125">MOR 1125</materialsCitation>
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during preparation, and these fragments were reserved for chemical analyses. In addition to the dense compact bone typical of theropods, this specimen contained regions of unusual bone tissue on the endosteal surface (2). Cortical and endosteal bone tissues were demineralized (3), and after 7 days, several fragments of the lining tissue exhibited unusual characteristics not normally observed in fossil bone. Removal of the mineral phase left a flexible vascular tissue that demonstrated great elasticity and resilience upon manipulation. In some cases, repeated stretching was possible (
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<figureCitation box="[2121,2233,293,321]" captionStart="Fig. 1" captionStartId="2.[947,992,963,986]" captionTargetBox="[910,1502,456,912]" captionTargetPageId="2" captionText="Fig. 1. Demineralized fragments of endosteally derived tissues lining the mar row cavity of the T. rex femur. (A) The demineralized fragment is flexible and resilient and, when stretched (arrow), returns to its original shape. (B) Demineralized bone in (A) after air dry ing. The overall structural and functional characteristics remain after dehydration. (C) Regions of demineralized bone show fibrous character (arrows). Scale bars, 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.4751428" httpUri="https://zenodo.org/record/4751428/files/figure.png" pageId="3" pageNumber="1954">Fig. 1 A</figureCitation>
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, arrow), and small pieces of this demineralized bone tissue could undergo repeated dehydrationrehydration cycles (Fig. IB) and still retain this elastic character. Demineralization also revealed that some regions of the bone were highly fibrous (
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<figureCitation box="[1899,2013,554,582]" captionStart="Fig. 1" captionStartId="2.[947,992,963,986]" captionTargetBox="[910,1502,456,912]" captionTargetPageId="2" captionText="Fig. 1. Demineralized fragments of endosteally derived tissues lining the mar row cavity of the T. rex femur. (A) The demineralized fragment is flexible and resilient and, when stretched (arrow), returns to its original shape. (B) Demineralized bone in (A) after air dry ing. The overall structural and functional characteristics remain after dehydration. (C) Regions of demineralized bone show fibrous character (arrows). Scale bars, 0.5 mm." figureDoi="http://doi.org/10.5281/zenodo.4751428" httpUri="https://zenodo.org/record/4751428/files/figure.png" pageId="3" pageNumber="1954">Fig. 1C</figureCitation>
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, arrows).
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</paragraph>
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<footnote pageId="2" pageNumber="1953">
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<paragraph blockId="2.[1668,2351,927,1396]" pageId="2" pageNumber="1953">department of Marine, Earth, Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695, USA. zNorth Carolina State Museum of Natural Sciences, Raleigh, NC 27601, USA. 3Museum of the Rockies, Montana State University, Bozeman, MT 59717, USA. “Carnegie Institution of Washington, Geophysical Laboratory, 5251 Broad Branch Road N.W., Washington, DC 20018, USA.</paragraph>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.4751428" ID-Zenodo-Dep="4751428" captionStart="Fig. 1" httpUri="https://zenodo.org/record/4751428/files/figure.png" pageId="2" pageNumber="1953" startId="2.[947,992,963,986]" targetBox="[910,1502,456,912]" targetPageId="2">
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<paragraph blockId="2.[946,1504,954,1397]" pageId="2" pageNumber="1953">
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<emphasis bold="true" box="[947,1035,963,986]" pageId="2" pageNumber="1953">
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<emphasis bold="true" box="[947,992,963,986]" italics="true" pageId="2" pageNumber="1953">Fig</emphasis>
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. 1.
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</emphasis>
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Demineralized fragments of endosteally derived tissues lining the mar row cavity of the
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1205,1284,1033,1061]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1953" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[1205,1227,1033,1061]" italics="true" pageId="2" pageNumber="1953">T.</emphasis>
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rex
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</taxonomicName>
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femur. (
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<emphasis box="[1408,1431,1033,1061]" italics="true" pageId="2" pageNumber="1953">A</emphasis>
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) The demineralized fragment is flexible and resilient and, when stretched (arrow), returns to its original shape. (
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<emphasis bold="true" box="[1404,1425,1145,1173]" pageId="2" pageNumber="1953">B</emphasis>
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) Demineralized bone in (A) after air dry
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<emphasis box="[948,990,1220,1248]" italics="true" pageId="2" pageNumber="1953">ing</emphasis>
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. The overall structural and functional characteristics remain after dehydration. (C) Regions of demineralized bone show fibrous character (arrows). Scale bars, 0.5 mm.
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</paragraph>
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</caption>
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<paragraph blockId="2.[1668,2351,927,1396]" pageId="2" pageNumber="1953">*To whom correspondence should be addressed. E-mail: schweitzer@ncsu.edu</paragraph>
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<paragraph blockId="2.[1668,2351,927,1396]" pageId="2" pageNumber="1953">
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tPresent address: Department of Geosciences, Christian- Albrechts University
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<emphasis box="[1935,1983,1338,1361]" italics="true" pageId="2" pageNumber="1953">Kiel</emphasis>
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, Olshausenstrasse 40, 24098
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<emphasis box="[1670,1717,1373,1396]" italics="true" pageId="2" pageNumber="1953">Kiel</emphasis>
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, Germany.
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</paragraph>
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</footnote>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.4751430" ID-Zenodo-Dep="4751430" captionStart="Fig. 2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="2" pageNumber="1953" startId="2.[1332,1376,2331,2354]" targetBox="[196,2344,1468,2283]" targetPageId="2">
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<paragraph blockId="2.[193,2352,1468,3105]" pageId="2" pageNumber="1953">
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<emphasis bold="true" box="[1332,1422,2331,2354]" pageId="2" pageNumber="1953">Fig. 2.</emphasis>
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Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[2265,2347,2364,2392]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1953" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[2265,2288,2364,2392]" italics="true" pageId="2" pageNumber="1953">T.</emphasis>
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rex
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</taxonomicName>
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cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1693,1783,2551,2579]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1953" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[1693,1716,2551,2579]" italics="true" pageId="2" pageNumber="1953">T.</emphasis>
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rex
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</taxonomicName>
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cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm.
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<emphasis bold="true" box="[587,623,2779,2802]" pageId="2" pageNumber="1953">(E)</emphasis>
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Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible.
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<emphasis bold="true" box="[1798,1807,2854,2877]" pageId="2" pageNumber="1953">(</emphasis>
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F
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<emphasis bold="true" box="[1825,1834,2854,2877]" pageId="2" pageNumber="1953">)</emphasis>
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1850,1933,2850,2878]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1953" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[1850,1872,2850,2878]" italics="true" pageId="2" pageNumber="1953">T.</emphasis>
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rex
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</taxonomicName>
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vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (
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<emphasis bold="true" box="[1495,1518,2888,2916]" pageId="2" pageNumber="1953">C</emphasis>
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) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel.
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<emphasis bold="true" box="[1866,1875,2929,2952]" pageId="2" pageNumber="1953">(</emphasis>
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H
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<emphasis bold="true" box="[1901,1910,2929,2952]" pageId="2" pageNumber="1953">)</emphasis>
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Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I)
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1450,1528,2962,2990]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="2" pageNumber="1953" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[1450,1472,2962,2990]" italics="true" pageId="2" pageNumber="1953">T.</emphasis>
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rex
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</taxonomicName>
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vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed.
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</paragraph>
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</caption>
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<caption ID-DOI="http://doi.org/10.5281/zenodo.3739834" ID-Zenodo-Dep="3739834" captionStart="Fig. 3" captionText="Fig. 3. SEM images of aldehyde-fixed vessels. (A) Isolated vessel from T. rex. (B) Vessel isolated from extant ostrich after demineralization and collagenase digestion (3). (C) Vessel from T. rex, showing internal contents and hollow character. (D) Exploded T. rex vessel showing small round microstructures partially embedded in internal vessel walls. (E) Higher magnification of a portion of T. rex vessel wall, showing hypothesized endothelial nuclei (EN). (F) Similar structures visible on fixed ostrich vessel. Striations are seen in both (E) and (F) that may represent endothelial cell junctions or alternatively may be artifacts of the fixation/dehydration process. Scale bars in (A) and (B), 40 um; in (C) and (D), 10 um; in (E) and (F), 1 um." httpUri="https://zenodo.org/record/3739834/files/figure.png" pageId="3" pageNumber="1954" startId="3.[194,239,485,508]" targetBox="[560,1614,474,1620]" targetPageId="3">
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<paragraph blockId="3.[192,517,477,1670]" pageId="3" pageNumber="1954">
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<emphasis bold="true" box="[194,285,485,508]" pageId="3" pageNumber="1954">Fig. 3.</emphasis>
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SEM images of aldehyde-fixed vessels. (
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<emphasis box="[203,226,555,583]" italics="true" pageId="3" pageNumber="1954">A</emphasis>
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) Isolated vessel from
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<emphasis box="[194,271,592,620]" italics="true" pageId="3" pageNumber="1954">
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[194,267,592,620]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">T. rex</taxonomicName>
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.
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</emphasis>
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(B) Vessel isolated from extant ostrich after demineralization and collagenase digestion (3). (C) Vessel from
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<taxonomicName authorityName="Osborn" authorityYear="1905" box="[194,272,779,807]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">
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<emphasis box="[194,215,779,807]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
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rex
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</taxonomicName>
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, showing internal contents and hollow character.
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<emphasis bold="true" box="[339,380,858,881]" pageId="3" pageNumber="1954">(D)</emphasis>
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Exploded
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[195,282,892,920]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[195,216,892,920]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
|
||
rex
|
||
</taxonomicName>
|
||
vessel showing small round microstructures partially embedded in internal vessel walls.
|
||
<emphasis bold="true" box="[273,307,1045,1068]" pageId="3" pageNumber="1954">(E)</emphasis>
|
||
Higher magnification of a portion of
|
||
<taxonomicName pageId="3" pageNumber="1954">
|
||
<emphasis box="[492,512,1079,1107]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
|
||
rex
|
||
</taxonomicName>
|
||
vessel wall, showing hypothesized endothelial nuclei (
|
||
<emphasis box="[334,378,1191,1219]" italics="true" pageId="3" pageNumber="1954">EN</emphasis>
|
||
).
|
||
<emphasis bold="true" box="[405,413,1195,1218]" pageId="3" pageNumber="1954">(</emphasis>
|
||
F
|
||
<emphasis bold="true" box="[430,438,1195,1218]" pageId="3" pageNumber="1954">)</emphasis>
|
||
Similar structures visible on fixed ostrich vessel. Striations are seen in both (E) and (F) that may represent endothelial cell junctions or alternatively may be artifacts of the fixation/dehydration process. Scale bars in (A) and (B), 40 um; in (C) and (D), 10 um; in (E) and (F), 1 um.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="3.[1670,2359,288,3116]" pageId="3" pageNumber="1954">
|
||
Partial demineralization of the cortical bone revealed parallel-oriented vascular canals that were seen to bifurcate in some areas (
|
||
<figureCitation box="[2232,2347,685,713]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2A</figureCitation>
|
||
, arrows). Occasional fenestrae (marked F) were observed on the surface of the vascular canals, possibly correlating with communicating Volkmann’s canals. Complete demineralization of the cortical bone released thin and transparent soft-tissue vessels from some regions of the matrix (
|
||
<figureCitation box="[2002,2092,991,1019]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2</figureCitation>
|
||
, B and C), which floated freely in the demineralizing solution. Vessels similar in diameter and texture were recovered from extant ostrich bone, when demineralization was followed by digestion with collagenase enzyme (3) to remove densely fibrous collagen matrix (
|
||
<figureCitation box="[2015,2133,1253,1281]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2D</figureCitation>
|
||
). In both dinosaur (
|
||
<figureCitation box="[1754,1867,1296,1324]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2C</figureCitation>
|
||
) and ostrich (
|
||
<figureCitation box="[2074,2192,1296,1324]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2D</figureCitation>
|
||
), remnants of the original organic matrix in which the vessels were embedded can still be visualized under transmitted light microscopy. These vessels are flexible, pliable, and translucent (
|
||
<figureCitation box="[1683,1807,1515,1543]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2E</figureCitation>
|
||
). The vessels branch in a pattern consistent with extant vessels, and many bifurcation points are visible (
|
||
<figureCitation box="[2100,2215,1602,1630]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2E</figureCitation>
|
||
, arrows). Many of the dinosaur vessels contain small round microstructures that vary from deep red to dark brown (
|
||
<figureCitation box="[1910,2002,1733,1761]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2</figureCitation>
|
||
, F and G). The vessels and contents are similar in all respects to blood vessels recovered from extant ostrich bone (
|
||
<figureCitation box="[1770,1894,1864,1892]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2H</figureCitation>
|
||
). Aldehyde-fixed (3) dinosaur vessels (
|
||
<figureCitation box="[1795,1895,1908,1936]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2I</figureCitation>
|
||
) are virtually identical in overall morphology to similarly prepared ostrich vessels (
|
||
<figureCitation box="[1795,1898,1995,2023]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2J</figureCitation>
|
||
), and structures consistent with remnants of nuclei from the original endothelial cells are visible on the exterior of both dinosaur and ostrich specimens (
|
||
<figureCitation box="[2165,2258,2126,2154]" captionStart="Fig. 2" captionStartId="2.[1332,1376,2331,2354]" captionTargetBox="[196,2344,1468,2283]" captionTargetPageId="2" captionText="Fig. 2. Demineralization of cortical bone reveals the presence of soft- tissue structures. (A) Partial demineralization of a fragment of T. rex cortical bone shows an emerging network of vascular canals, some of which are bifurcated (arrows). All are aligned in parallel, consistent with Haversian canals in cortical bone. Small fenestrae (marked F) may indicate invaginations for communicating Volkmann's canals. (B) A second fragment of T. rex cortical bone illustrates transparent vessels (arrows) arising from bone matrix in solution. (C) Complete demineralization reveals transparent flexible vessels in what remains of the cortical bone matrix, represented by a brown amorphous substance (marked M). (D) Ostrich vessel after demineralization of cortical bone and subsequent digestion of fibrous collagenous matrix. Transparent vessels branch and remain associated with small regions of undigested bone matrix, seen here as amorphous, white fibrous material (marked M). Scale bars in (A) to (D), 0.5 mm. (E) Higher magnification of dinosaur vessels shows branching pattern (arrows) and internal contents. Vascular structure is not consistent with fungal hyphae (no septae, and branching pattern is not consistent with fungal morphology) or plant (no cell walls visible, and again branching pattern is not consistent). Round red microstructures within the vessels are clearly visible. (F) T. rex vessel fragment, containing microstructures consistent in size and shape with those seen in the ostrich vessel in (H). (C) Second fragment of dinosaur vessel. Air/fluid interfaces, represented by dark menisci, illustrate the hollow nature of vessels. Microstructure is visible within the vessel. (H) Ostrich vessel digested from demineralized cortical bone. Red blood cells can be seen inside the branching vessel. (I) T. rex vessel fragment showing detail of branching pattern and structures morphologically consistent with endothelial cell nuclei (arrows) in vessel wall. (J) Ostrich blood vessel liberated from demineralized bone after treatment with collagenase shows branching pattern and clearly visible endothelial nuclei. Scale bars in (E) to (J), 50 um. (F), (I), and (J) were subjected to aldehyde fixation (3). The remaining vessels are unfixed." figureDoi="http://doi.org/10.5281/zenodo.4751430" httpUri="https://zenodo.org/record/4751430/files/figure.png" pageId="3" pageNumber="1954">Fig. 2</figureCitation>
|
||
, I and J, arrows).
|
||
</paragraph>
|
||
<caption ID-DOI="http://doi.org/10.5281/zenodo.3739836" ID-Zenodo-Dep="3739836" captionStart="Fig. 4" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="3" pageNumber="1954" startId="3.[196,241,1745,1768]" targetBox="[562,1614,1734,2866]" targetPageId="3">
|
||
<paragraph blockId="3.[195,514,1741,2853]" pageId="3" pageNumber="1954">
|
||
Fig. 4. Cellular features associated with
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[429,513,1778,1806]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[429,451,1778,1806]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
|
||
rex
|
||
</taxonomicName>
|
||
and ostrich tissues. (A) Fragment of demin eralized cortical bone from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[284,371,1927,1955]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[284,305,1927,1955]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
|
||
rex
|
||
</taxonomicName>
|
||
, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher
|
||
<emphasis box="[197,366,2115,2143]" italics="true" pageId="3" pageNumber="1954">magnification</emphasis>
|
||
of one of the microstructures seen embedded in the fibrous material. (
|
||
<emphasis bold="true" box="[354,374,2227,2255]" pageId="3" pageNumber="1954">B</emphasis>
|
||
) Demin eralized and stained (3) ostrich cortical bone, showing
|
||
<emphasis box="[313,399,2339,2367]" italics="true" pageId="3" pageNumber="1954">fibrillar</emphasis>
|
||
, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur
|
||
<emphasis box="[197,255,2750,2778]" italics="true" pageId="3" pageNumber="1954">face</emphasis>
|
||
consistent with filipodia. (C) Isolated microstructure from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[490,511,2825,2853]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
|
||
<paragraph blockId="3.[1670,2359,288,3116]" pageId="3" pageNumber="1954">
|
||
Under scanning electron microscopy (SEM) (
|
||
<figureCitation box="[1684,1769,2257,2285]" captionStart="Fig. 3" captionStartId="3.[194,239,485,508]" captionTargetBox="[560,1614,474,1620]" captionTargetPageId="3" captionText="Fig. 3. SEM images of aldehyde-fixed vessels. (A) Isolated vessel from T. rex. (B) Vessel isolated from extant ostrich after demineralization and collagenase digestion (3). (C) Vessel from T. rex, showing internal contents and hollow character. (D) Exploded T. rex vessel showing small round microstructures partially embedded in internal vessel walls. (E) Higher magnification of a portion of T. rex vessel wall, showing hypothesized endothelial nuclei (EN). (F) Similar structures visible on fixed ostrich vessel. Striations are seen in both (E) and (F) that may represent endothelial cell junctions or alternatively may be artifacts of the fixation/dehydration process. Scale bars in (A) and (B), 40 um; in (C) and (D), 10 um; in (E) and (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739834" httpUri="https://zenodo.org/record/3739834/files/figure.png" pageId="3" pageNumber="1954">Fig. 3</figureCitation>
|
||
), features seen on the external surface of dinosaurian vessels are virtually indistinguishable from those seen in similarly prepared extant ostrich vessels (
|
||
<figureCitation box="[2015,2107,2388,2416]" captionStart="Fig. 3" captionStartId="3.[194,239,485,508]" captionTargetBox="[560,1614,474,1620]" captionTargetPageId="3" captionText="Fig. 3. SEM images of aldehyde-fixed vessels. (A) Isolated vessel from T. rex. (B) Vessel isolated from extant ostrich after demineralization and collagenase digestion (3). (C) Vessel from T. rex, showing internal contents and hollow character. (D) Exploded T. rex vessel showing small round microstructures partially embedded in internal vessel walls. (E) Higher magnification of a portion of T. rex vessel wall, showing hypothesized endothelial nuclei (EN). (F) Similar structures visible on fixed ostrich vessel. Striations are seen in both (E) and (F) that may represent endothelial cell junctions or alternatively may be artifacts of the fixation/dehydration process. Scale bars in (A) and (B), 40 um; in (C) and (D), 10 um; in (E) and (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739834" httpUri="https://zenodo.org/record/3739834/files/figure.png" pageId="3" pageNumber="1954">Fig. 3</figureCitation>
|
||
, B and F), suggesting a common origin. These features include surface striations that may be consistent with endothelial cell junctions, or alternatively may be artifacts of fixation and/or dehydration. In addition, small round to oval features dot the surface of both dinosaur and ostrich vessels, which may be consistent with endothelial cell nuclei (
|
||
<figureCitation box="[1783,1872,2737,2765]" captionStart="Fig. 3" captionStartId="3.[194,239,485,508]" captionTargetBox="[560,1614,474,1620]" captionTargetPageId="3" captionText="Fig. 3. SEM images of aldehyde-fixed vessels. (A) Isolated vessel from T. rex. (B) Vessel isolated from extant ostrich after demineralization and collagenase digestion (3). (C) Vessel from T. rex, showing internal contents and hollow character. (D) Exploded T. rex vessel showing small round microstructures partially embedded in internal vessel walls. (E) Higher magnification of a portion of T. rex vessel wall, showing hypothesized endothelial nuclei (EN). (F) Similar structures visible on fixed ostrich vessel. Striations are seen in both (E) and (F) that may represent endothelial cell junctions or alternatively may be artifacts of the fixation/dehydration process. Scale bars in (A) and (B), 40 um; in (C) and (D), 10 um; in (E) and (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739834" httpUri="https://zenodo.org/record/3739834/files/figure.png" pageId="3" pageNumber="1954">Fig. 3</figureCitation>
|
||
, E and F, arrows).
|
||
</paragraph>
|
||
<paragraph blockId="3.[1670,2359,288,3116]" lastBlockId="4.[190,876,280,3112]" lastPageId="4" lastPageNumber="1955" pageId="3" pageNumber="1954">
|
||
Finally, in those regions of the bone where fibrillar matrix predominated in the demineralized tissues, elongate microstructures could be visualized among the fibers (
|
||
<figureCitation box="[2220,2350,2912,2940]" captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="3" pageNumber="1954">Fig. 4A</figureCitation>
|
||
, inset). These microstructures contain multiple projections on the external surface and are virtually identical in size, location, and overall morphology to osteocytes seen among collagen fibers of demineralized ostrich bone (
|
||
<figureCitation captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="4" pageNumber="1955">Fig. 4B</figureCitation>
|
||
, inset). These cell-like microstructures could be isolated and, when subjected to aldehyde fixation (J), appeared to possess internal contents (
|
||
<figureCitation box="[443,564,458,490]" captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="4" pageNumber="1955">Fig. 4C</figureCitation>
|
||
), including possible nuclei (
|
||
<figureCitation box="[305,423,502,534]" captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="4" pageNumber="1955">Fig. 4C</figureCitation>
|
||
, inset). These microstructures are similar in morphology to fixed ostrich osteocytes, both unstained (
|
||
<figureCitation box="[652,783,589,621]" captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="4" pageNumber="1955">Fig. 4D</figureCitation>
|
||
) and stained
|
||
<emphasis box="[313,354,633,665]" italics="true" pageId="4" pageNumber="1955">(3)</emphasis>
|
||
for better visualization (
|
||
<figureCitation box="[744,866,633,665]" captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="4" pageNumber="1955">Fig. 4D</figureCitation>
|
||
, inset). SEM verifies the presence of the features seen in transmitted light microscopy, and again, projections extending from the surface of the microstructures are clearly visible (
|
||
<figureCitation box="[270,361,852,884]" captionStart="Fig. 4" captionStartId="3.[196,241,1745,1768]" captionTargetBox="[562,1614,1734,2866]" captionTargetPageId="3" captionText="Fig. 4. Cellular features associated with T. rex and ostrich tissues. (A) Fragment of demin eralized cortical bone from T. rex, showing parallel-oriented fibers and cell-like microstruc tures among the fibers. The inset is a higher magnification of one of the microstructures seen embedded in the fibrous material. (B) Demin eralized and stained (3) ostrich cortical bone, showing fibrillar, parallel- oriented collagen matrix with osteocytes embed ded among the fibers. The inset shows a high er magnification of one of the osteocytes. Both inset views show elon gate bodies with multi ple projections arising from the external sur face consistent with filipodia. (C) Isolated microstructure from T. rex after fixation. In addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long filipodia and an internal transparent nucleus-like structure. (D) Fixed ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and filipodia can be seen extending in multiple planes from the cell surface. (E and F) SEM images of aldehyde-fixed (3) microstructures isolated from T. rex cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um." figureDoi="http://doi.org/10.5281/zenodo.3739836" httpUri="https://zenodo.org/record/3739836/files/figure.png" pageId="4" pageNumber="1955">Fig. 4</figureCitation>
|
||
, E and F).
|
||
</paragraph>
|
||
<paragraph blockId="3.[196,1615,2862,3115]" pageId="3" pageNumber="1954">
|
||
<heading box="[196,513,2862,2890]" fontSize="10" level="4" pageId="3" pageNumber="1954" reason="1">rex after fixation. In</heading>
|
||
addition to the multiple filipodial-like projections, internal contents can be seen. The inset shows a second structure with long
|
||
<emphasis box="[575,681,2937,2965]" italics="true" pageId="3" pageNumber="1954">filipodia</emphasis>
|
||
and an internal transparent nucleus-like structure.
|
||
<emphasis bold="true" box="[1389,1430,2941,2964]" pageId="3" pageNumber="1954">(D)</emphasis>
|
||
<emphasis box="[1444,1512,2937,2965]" italics="true" pageId="3" pageNumber="1954">Fixed</emphasis>
|
||
ostrich osteocyte; inset, ostrich osteocyte fixed and stained for better visualization. Internal contents are discernible, and
|
||
<emphasis box="[418,524,3011,3039]" italics="true" pageId="3" pageNumber="1954">filipodia</emphasis>
|
||
can be seen extending in multiple planes from the cell surface. (
|
||
<emphasis bold="true" box="[1427,1445,3011,3039]" pageId="3" pageNumber="1954">E</emphasis>
|
||
and F) SEM images of aldehyde-fixed (3) microstructures isolated from
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[1012,1089,3049,3077]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="3" pageNumber="1954" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[1012,1033,3049,3077]" italics="true" pageId="3" pageNumber="1954">T.</emphasis>
|
||
rex
|
||
</taxonomicName>
|
||
cortical bone tissues. Scale bars in (A) and (B), 50 um; in (C) and (D), 20 um; in (E), 10 um; in (F), 1 um.
|
||
</paragraph>
|
||
</taxonomicName>
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph blockId="4.[190,876,280,3112]" pageId="4" pageNumber="1955">
|
||
The fossil record is capable of exceptional preservation, including feathers (
|
||
<bibRefCitation author="X. Xu & X. L. Wang & X. C. Wu" box="[671,693,938,970]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="262" part="401" refId="ref3262" refString="4. X. Xu, X. L. Wang, X. C. Wu, Nature 401, 262 (1999)." type="journal article" year="1999">4</bibRefCitation>
|
||
-
|
||
<bibRefCitation author="M. H. Schweitzer" box="[705,726,938,970]" journalOrPublisher="J. Exp. Zool." pageId="4" pageNumber="1955" pagination="146" part="285" refId="ref3304" refString="6. M. H. Schweitzer et al., J. Exp. Zool. 285, 146 (1999)." type="journal article" year="1999">6</bibRefCitation>
|
||
), hair (
|
||
<bibRefCitation author="M. Wuttke" box="[834,853,938,970]" editor="S. Schaal & W. Ziegler" journalOrPublisher="Verlag Waldemar Kramer, Frankfurt am Main, Germany" pageId="4" pageNumber="1955" pagination="265 - 274" refId="ref3328" refString="7. M. Wuttke, in Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens, S. Schaal, W. Ziegler, Eds. (Verlag Waldemar Kramer, Frankfurt am Main, Germany, 1988), pp. 265 - 274." type="book chapter" volumeTitle="Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens" year="1988">7</bibRefCitation>
|
||
), color or color patterns (
|
||
<bibRefCitation author="M. Wuttke" box="[555,575,983,1015]" editor="S. Schaal & W. Ziegler" journalOrPublisher="Verlag Waldemar Kramer, Frankfurt am Main, Germany" pageId="4" pageNumber="1955" pagination="265 - 274" refId="ref3328" refString="7. M. Wuttke, in Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens, S. Schaal, W. Ziegler, Eds. (Verlag Waldemar Kramer, Frankfurt am Main, Germany, 1988), pp. 265 - 274." type="book chapter" volumeTitle="Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens" year="1988">7</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="D. M. Martill & E. Frey" box="[597,615,983,1015]" journalOrPublisher="N. jb. Geol. Palaont. Mh" pageId="4" pageNumber="1955" pagination="118" part="2" refId="ref3376" refString="8. D. M. Martill, E. Frey, N. jb. Geol. Palaont. Mh. 2, 118 (1995)." type="journal article" year="1995">8</bibRefCitation>
|
||
), embryonic soft tissues (
|
||
<bibRefCitation author="L. M. Chiappe" box="[309,329,1027,1059]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="258" part="396" refId="ref3405" refString="9. L. M. Chiappe et at., Nature 396, 258 (1998)." type="journal article" year="1998">9</bibRefCitation>
|
||
), muscle tissue and/or internal organs
|
||
<emphasis box="[193,317,1070,1102]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="C. Dal Sasso & M. Signore" box="[205,247,1070,1102]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="383" part="392" refId="ref3424" refString="10. C. Dal Sasso, M. Signore, Nature 392, 383 (1998)." type="journal article" year="1998">10</bibRefCitation>
|
||
-
|
||
<bibRefCitation author="D. E. G. Briggs & P. R. Wilby & B. P. Perez-Moreno & J. L. Sanz & M. Fregenal-Martinez" box="[258,299,1070,1102]" journalOrPublisher="J. Geol. Soc. (London)" pageId="4" pageNumber="1955" pagination="587" part="154" refId="ref3491" refString="13. D. E. G. Briggs, P. R. Wilby, B. P. Perez-Moreno, J. L. Sanz, M. Fregenal-Martinez, J. Ceol. Soc. (London) 154, 587 (1997)." type="journal article" year="1997">13</bibRefCitation>
|
||
),
|
||
</emphasis>
|
||
and cellular structure (
|
||
<bibRefCitation author="M. Wuttke" box="[717,737,1070,1102]" editor="S. Schaal & W. Ziegler" journalOrPublisher="Verlag Waldemar Kramer, Frankfurt am Main, Germany" pageId="4" pageNumber="1955" pagination="265 - 274" refId="ref3328" refString="7. M. Wuttke, in Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens, S. Schaal, W. Ziegler, Eds. (Verlag Waldemar Kramer, Frankfurt am Main, Germany, 1988), pp. 265 - 274." type="book chapter" volumeTitle="Messel-Ein Schaufenster in die Geschichte der Erde und des Lebens" year="1988">7</bibRefCitation>
|
||
,
|
||
<emphasis box="[762,871,1070,1102]" italics="true" pageId="4" pageNumber="1955">
|
||
<bibRefCitation author="R. Pawlicki & A. Korbel & H. Kubiak" box="[762,802,1070,1102]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="656" part="211" refId="ref3539" refString="14. R. Pawlicki, A. Korbel, H. Kubiak, Nature 211,656 (1966)." type="book" year="1966">14</bibRefCitation>
|
||
-
|
||
<bibRefCitation author="M. H. Schweitzer & J. R. Horner" box="[812,854,1070,1102]" journalOrPublisher="Ann. Pateontol." pageId="4" pageNumber="1955" pagination="179" part="85" refId="ref3580" refString="16. M. H. Schweitzer, J. R. Horner, Ann. Pateontol. 85, 179 (1999)." type="journal article" year="1999">16</bibRefCitation>
|
||
).
|
||
</emphasis>
|
||
These soft tissues are preserved as carbon films
|
||
<emphasis box="[285,436,1158,1190]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="X. Xu & X. L. Wang & X. C. Wu" box="[297,317,1158,1190]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="262" part="401" refId="ref3262" refString="4. X. Xu, X. L. Wang, X. C. Wu, Nature 401, 262 (1999)." type="journal article" year="1999">4</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="Q. Ji" box="[343,363,1158,1190]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="753" part="393" refId="ref3288" refString="5. Q. Ji et at .. Nature 393, 753 (1998)." type="journal article" year="1998">5</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="C. Dal Sasso & M. Signore" box="[390,426,1158,1190]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="383" part="392" refId="ref3424" refString="10. C. Dal Sasso, M. Signore, Nature 392, 383 (1998)." type="journal article" year="1998">10</bibRefCitation>
|
||
)
|
||
</emphasis>
|
||
or as permineralized threedimensional replications
|
||
<emphasis box="[577,758,1201,1233]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="L. M. Chiappe" box="[589,609,1201,1233]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="258" part="396" refId="ref3405" refString="9. L. M. Chiappe et at., Nature 396, 258 (1998)." type="journal article" year="1998">9</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="A. W. A. Kellner" box="[638,675,1201,1233]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="32" part="379" refId="ref3443" refString="11. A. W. A. Kellner, Nature 379, 32 (1996)." type="journal article" year="1996">11</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="D. E. G. Briggs & P. R. Wilby & B. P. Perez-Moreno & J. L. Sanz & M. Fregenal-Martinez" box="[703,741,1201,1233]" journalOrPublisher="J. Geol. Soc. (London)" pageId="4" pageNumber="1955" pagination="587" part="154" refId="ref3491" refString="13. D. E. G. Briggs, P. R. Wilby, B. P. Perez-Moreno, J. L. Sanz, M. Fregenal-Martinez, J. Ceol. Soc. (London) 154, 587 (1997)." type="journal article" year="1997">13</bibRefCitation>
|
||
),
|
||
</emphasis>
|
||
but in none of these cases are they described as stillsoft, pliable tissues.
|
||
</paragraph>
|
||
<paragraph blockId="4.[190,876,280,3112]" pageId="4" pageNumber="1955">
|
||
Mesozoic fossils, particularly dinosaur fossils, are known to be extremely well preserved histologically and occasionally retain molecular information
|
||
<emphasis box="[438,616,1463,1495]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="M. H. Schweitzer" box="[450,471,1463,1495]" journalOrPublisher="J. Exp. Zool." pageId="4" pageNumber="1955" pagination="146" part="285" refId="ref3304" refString="6. M. H. Schweitzer et al., J. Exp. Zool. 285, 146 (1999)." type="journal article" year="1999">6</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="G. Muyzer" box="[497,535,1463,1495]" journalOrPublisher="Geology" pageId="4" pageNumber="1955" pagination="871" part="20" refId="ref3605" refString="17. G. Muyzer et al .. Geology 20, 871 (1992)." type="journal article" year="1992">17</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="M. H. Schweitzer" box="[561,599,1463,1495]" journalOrPublisher="Proc. Natl. Acad. Sci. U. S. A." pageId="4" pageNumber="1955" pagination="6291" part="94" refId="ref3621" refString="18. M. H. Schweitzer et at., Proc. Natl. Acad. Sci. U. S. A. 94, 6291 (1997)." type="journal article" year="1997">18</bibRefCitation>
|
||
),
|
||
</emphasis>
|
||
the presence of which
|
||
<emphasis box="[307,332,1508,1540]" italics="true" pageId="4" pageNumber="1955">is</emphasis>
|
||
closely linked to morphological preservation
|
||
<emphasis box="[397,466,1550,1582]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="R. E. M. Hedges" box="[409,449,1550,1582]" journalOrPublisher="Archaeometry" pageId="4" pageNumber="1955" pagination="319" part="44" refId="ref3653" refString="19. R. E. M. Hedges, Archaeometry 44, 319 (2002)." type="journal article" year="2002">19</bibRefCitation>
|
||
).
|
||
</emphasis>
|
||
Vascular microstructures that may be derived from original blood materials of Cretaceous organisms have also been reported
|
||
<emphasis box="[325,446,1681,1713]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="R. Pawlicki & A. Korbel & H. Kubiak" box="[337,377,1681,1713]" journalOrPublisher="Nature" pageId="4" pageNumber="1955" pagination="656" part="211" refId="ref3539" refString="14. R. Pawlicki, A. Korbel, H. Kubiak, Nature 211,656 (1966)." type="book" year="1966">14</bibRefCitation>
|
||
-
|
||
<bibRefCitation author="M. H. Schweitzer & J. R. Horner" box="[387,429,1681,1713]" journalOrPublisher="Ann. Pateontol." pageId="4" pageNumber="1955" pagination="179" part="85" refId="ref3580" refString="16. M. H. Schweitzer, J. R. Horner, Ann. Pateontol. 85, 179 (1999)." type="journal article" year="1999">16</bibRefCitation>
|
||
).
|
||
</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="4.[190,876,280,3112]" pageId="4" pageNumber="1955">
|
||
Pawlicki was able to demonstrate osteocytes and vessels obtained from dinosaur bone using an etching and replication technique
|
||
<emphasis box="[286,412,1858,1890]" italics="true" pageId="4" pageNumber="1955">(14,15).</emphasis>
|
||
However, we demonstrate the retention of pliable soft-tissue blood vessels with contents that are capable of being liberated from the bone matrix, while still retaining their flexibility, resilience, original hollow nature, and three-dimensionality. Additionally, we can isolate three-dimensional osteocytes with internal cellular contents and intact, supple filipodia that float freely in solution. This
|
||
<taxonomicName authorityName="Osborn" authorityYear="1905" box="[194,280,2250,2282]" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" higherTaxonomySource="GBIF" kingdom="Animalia" order="Dinosauria" pageId="4" pageNumber="1955" phylum="Chordata" rank="species" species="rex">
|
||
<emphasis box="[194,280,2250,2282]" italics="true" pageId="4" pageNumber="1955">T. rex</emphasis>
|
||
</taxonomicName>
|
||
also contains flexible and fibrillar bone matrices that retain elasticity. The unusual preservation of the originally organic matrix may be due in part to the dense mineralization of dinosaur bone, because a certain portion of the organic matrix within extant bone is intracrystalline and therefore extremely resistant to degradation
|
||
<emphasis box="[375,499,2556,2588]" italics="true" pageId="4" pageNumber="1955">
|
||
(
|
||
<bibRefCitation author="S. Weiner & W. Traub & H. Elster & M. J. DeNiro" box="[386,426,2556,2588]" journalOrPublisher="Appl. Geochem." pageId="4" pageNumber="1955" pagination="231" part="4" refId="ref3671" refString="20. S. Weiner, W. Traub, H. Elster, M. J. DeNiro, Appl. Geochem. 4, 231 (1989)." type="journal article" year="1989">20</bibRefCitation>
|
||
,
|
||
<bibRefCitation author="G. A. Sykes & M. J. Collins & D. I. Walton" box="[443,482,2556,2588]" journalOrPublisher="Org. Geochem" pageId="4" pageNumber="1955" pagination="1059" part="23" refId="ref3702" refString="21. G. A. Sykes, M. J. Collins, D. I. Walton, Org. Geochem. 23, 1059 (1995)." type="journal article" year="1995">21</bibRefCitation>
|
||
).
|
||
</emphasis>
|
||
These factors, combined with as yet undetermined geochemical and environmental factors, presumably also contribute to the preservation of soft-tissue vessels. Because they have not been embedded or subjected to other chemical treatments, the cells and vessels are capable of being analyzed further for the persistence of molecular or other chemical information
|
||
<emphasis box="[733,783,2906,2938]" italics="true" pageId="4" pageNumber="1955">(3).</emphasis>
|
||
</paragraph>
|
||
<paragraph blockId="4.[190,876,280,3112]" lastBlockId="4.[930,1613,282,1671]" pageId="4" pageNumber="1955">Using the methodologies described here, we isolated translucent vessels from two other exceptionally well-preserved tyrannosaurs (figs. S1 and S2) (3), and we isolated microstructures consistent with osteocytes in at least three other dinosaurs: two tyrannosaurs and one hadrosaur (fig. S3). Vessels in these specimens exhibit highly variable preservation, from crystalline morphs to transparent and pliable soft tissues.</paragraph>
|
||
<paragraph blockId="4.[930,1613,282,1671]" pageId="4" pageNumber="1955">
|
||
The elucidation and modeling of processes resulting in soft-tissue preservation may form the basis for an avenue of research into the recovery and characterization of similar structures in other specimens, paving the way for micro- and molecular taphonomic investigations. Whether preservation is strictly morphological and the result of some kind of unknown geochemical replacement process or whether it extends to the subcellular and molecular levels is uncertain. However, we have identified protein fragments in extracted bone samples, some of which retain slight antigenicity
|
||
<emphasis box="[932,982,1115,1147]" italics="true" pageId="4" pageNumber="1955">(3).</emphasis>
|
||
These data indicate that exceptional morphological preservation in some dinosaurian specimens may extend to the cellular level or beyond. If so, in addition to providing
|
||
<emphasis box="[1572,1612,1247,1279]" italics="true" pageId="4" pageNumber="1955">independent</emphasis>
|
||
means of testing phylogenetic hypotheses about dinosaurs, applying molecular and analytical methods to well-preserved dinosaur specimens has important implications for elucidating preservational microenvironments and will contribute to our understanding of biogeochemical interactions at the microscopic and molecular levels that lead to fossilization.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |