434 lines
103 KiB
XML
434 lines
103 KiB
XML
<document id="4EACB322D5ED94CAE014509E0A2CAC60" ID-DOI="10.1126/science.1112158" ID-GBIF-Dataset="3a6b4cb7-b97f-443e-9aec-f5be22e0d320" ID-Zenodo-Dep="3742896" IM.metadata_requiresApprovalFor="plazi" IM.taxonomicNames_requiresApprovalFor="plazi" checkinTime="1586257997572" checkinUser="jeremy" docAuthor="Mary H. Schweitzer, Jennifer L. Wittmeyer & John R. Horner" docDate="2005" docId="0942513AFFA6FF98FF0133FC2461F837" docLanguage="en" docName="Schweitzeretal2005GenderSpecificTissueABBYY2.pdf" docOrigin="Science 308" docStyle="DocumentStyle{}" docTitle="Tyrannosaurus rex" docType="treatment" docVersion="16" lastPageNumber="1459" masterDocId="F57B2942FFA7FF9CFFA63039216EFFE7" masterDocTitle="Gender-Specific Reproductive Tissue in Ratites and Tyrannosaurus rex" masterLastPageNumber="1460" masterPageNumber="1456" pageNumber="1456" updateTime="1698736073715" updateUser="plazi">
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<mods:titleInfo id="F8FA88E0C6CE0EC733F4E8543FEDEA04">
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<mods:title id="2FC110630F9907C1D64CCC311DC252F6">Gender-Specific Reproductive Tissue in Ratites and Tyrannosaurus rex</mods:title>
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<mods:name id="30B4ED3E9CDB9359A30A90121828CCD7" type="personal">
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<mods:namePart id="454D77C455150822849B5F9DCC6504C5">Mary H. Schweitzer</mods:namePart>
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<mods:affiliation id="668ECB6C48E2A437D27F91DC4BE3FDD3">Department of Marine, Earth, and Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695, USA. North Carolina State Museum of Natural Sciences, Raleigh, NC 27601, USA. Museum of the Rockies, Montana State University, Bozeman, MT 59717, USA.</mods:affiliation>
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<mods:nameIdentifier id="92185C0996DACC89884B92E1D274DCAB" type="email">schweitzer@ncsu.edu</mods:nameIdentifier>
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<mods:namePart id="935270223B3CF7DED5A980F327DBEDC0">Jennifer L. Wittmeyer</mods:namePart>
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<mods:affiliation id="A6F0AC3A6F3C85A2FFDEA3D6D3F340D9">Department of Marine, Earth, and Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695, USA.</mods:affiliation>
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<mods:name id="A85C0D4EB78B1BB945B52A354ABF1B3E" type="personal">
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<mods:roleTerm id="C356BC589A8ED2D68C3B45DDD58FD67A">Author</mods:roleTerm>
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<mods:namePart id="9DC1C57EBC06F8E4C0917C8560818D83">John R. Horner</mods:namePart>
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<mods:affiliation id="C7D7834C0194ECC9F2CF4587B0F2F54B">Museum of the Rockies, Montana State University, Bozeman, MT 59717, USA.</mods:affiliation>
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<mods:typeOfResource id="43FD447BDB2CC2DBA7039BDE4B719FBF">text</mods:typeOfResource>
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<mods:title id="203C714D847D3566506C3CFBC6411B9E">Science</mods:title>
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<mods:part id="81AC5ADA4921912756F0B0AE21692020">
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<mods:date id="FE3225D99BA92BEC870AE5AC7BC5D438">2005</mods:date>
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<mods:detail id="7581B7AA9834FBF9BEFDCE4E98E981AC" type="volume">
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<mods:number id="AF39040A8D19B63BA1CC44ACFC58785D">308</mods:number>
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<mods:classification id="AF444B54938EEFE7B0A446F201997B86">journal article</mods:classification>
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<mods:identifier id="7E707F967CF6B8EC1EB2405C071B23AD" type="DOI">10.1126/science.1112158</mods:identifier>
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<mods:identifier id="9A9F89EB9A6BA03D6714F33319074579" type="GBIF-Dataset">3a6b4cb7-b97f-443e-9aec-f5be22e0d320</mods:identifier>
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<treatment id="0942513AFFA6FF98FF0133FC2461F837" ID-DOI="http://doi.org/10.5281/zenodo.3809972" ID-GBIF-Taxon="163536553" ID-Zenodo-Dep="3809972" LSID="urn:lsid:plazi:treatment:0942513AFFA6FF98FF0133FC2461F837" httpUri="http://treatment.plazi.org/id/0942513AFFA6FF98FF0133FC2461F837" lastPageId="4" lastPageNumber="1459" pageId="1" pageNumber="1456">
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<subSubSection id="C9F1B3A7FFA6FF9DFF0133FC2255FAA6" pageId="1" pageNumber="1456" type="nomenclature">
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<paragraph id="8154E02CFFA6FF9DFF0133FC2255FAA6" blockId="1.[167,849,964,1434]" pageId="1" pageNumber="1456">
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A relatively small (femur length, 107 cm)
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<taxonomicName id="46EB9BAFFFA6FF9DFCB933FE20C8FBF7" authority="Osborn, 1905" class="Reptilia" family="Tyrannosauridae" genus="Tyrannosaurus" kingdom="Animalia" order="Dinosauria" pageId="1" pageNumber="1456" phylum="Chordata" rank="species" species="rex">Tyrannosaurus rex</taxonomicName>
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[
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<materialsCitation id="3183EA71FFA6FF9DFE6333C923CBFBDA" ID-GBIF-Occurrence="2979293303" collectionCode="MOR" pageId="1" pageNumber="1456" specimenCode="MOR 1125">Museum of the Rockies (MOR) specimen number 1125</materialsCitation>
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] was discovered at the base of the Hell Creek Formation (dated at ~70 million years ago) as an association of disarticulated elements with excellent preservation (1). At death,
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<materialsCitation id="3183EA71FFA6FF9DFD5B34F32181FAF1" ID-GBIF-Occurrence="2979293318" collectionCode="MOR" pageId="1" pageNumber="1456" specimenCode="MOR 1125">MOR 1125</materialsCitation>
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was estimated to be 18 T 2 years (2), on the basis of lines of arrested growth (LAG).
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</paragraph>
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</subSubSection>
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<footnote id="E2F0FC22FFA6FF9DF9CB344B2651FAD8" pageId="1" pageNumber="1456">
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<paragraph id="8154E02CFFA6FF9DF9CB344B2651FAD8" blockId="1.[1645,2326,1138,1343]" pageId="1" pageNumber="1456">1Department of Marine, Earth, and Atmospheric Sciences, North Carolina State University, Raleigh, NC 27695, USA. 2North Carolina State Museum of Natural Sciences, Raleigh, NC 27601, USA. 3Museum of the Rockies, Montana State University, Bozeman, MT 59717, USA.</paragraph>
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</footnote>
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<subSubSection id="C9F1B3A7FFA6FF98FF7F35772461F837" lastPageId="4" lastPageNumber="1459" pageId="1" pageNumber="1456" type="description">
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<paragraph id="8154E02CFFA6FF9DFF7F35772452FB0D" blockId="1.[167,849,964,1434]" lastBlockId="1.[905,1587,964,1434]" pageId="1" pageNumber="1456">
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Interior femur fragments from
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<materialsCitation id="3183EA71FFA6FF9DFD0E35772223FA89" ID-GBIF-Occurrence="2979293316" box="[680,845,1358,1390]" collectionCode="MOR" pageId="1" pageNumber="1456" specimenCode="MOR 1125">MOR 1125</materialsCitation>
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were reserved without preservatives for chem ical and molecular analyses. Gross examination revealed a thin layer of bone tissue lining the inner (medullary) surfaces of the bone fragments that was structurally distinct from other described bone types (
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<figureCitation id="19D0FCA9FFA6FF9DFADA344A2483FB74" box="[1404,1517,1139,1171]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="1" pageNumber="1456" targetBox="[526,2320,1508,2848]">Fig. 1D</figureCitation>
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) but possessed characteristics in common with avian medullary bone (MB).
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</paragraph>
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<paragraph id="8154E02CFFA6FF9DFC1B34CF291CFD53" blockId="1.[905,1587,964,1434]" lastBlockId="1.[1644,2328,136,1085]" pageId="1" pageNumber="1456">
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MB is an ephemeral tissue, deposited on the endosteal surface of avian long bones
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<emphasis id="B39F3C3EFFA6FF9DFC2A35772298FA89" box="[908,1014,1358,1390]" italics="true" pageId="1" pageNumber="1456">(3-10).</emphasis>
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Its formation in female birds is triggered by increasing levels of gonadal hor mones produced upon ovulation (
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<emphasis id="B39F3C3EFFA6FF9DF7D330B329E9FF4D" box="[2165,2183,138,170]" italics="true" pageId="1" pageNumber="1456">4</emphasis>
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,
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<emphasis id="B39F3C3EFFA6FF9DF70630B329AAFF4D" box="[2208,2244,138,170]" italics="true" pageId="1" pageNumber="1456">10</emphasis>
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,
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<emphasis id="B39F3C3EFFA6FF9DF77A30B3286FFF4D" box="[2268,2305,138,170]" italics="true" pageId="1" pageNumber="1456">11</emphasis>
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), but it can also be artificially induced in male birds by the administration of estrogen (
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<emphasis id="B39F3C3EFFA6FF9DF9DD313527E3FECB" box="[1659,1677,268,300]" italics="true" pageId="1" pageNumber="1456">3</emphasis>
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,
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<emphasis id="B39F3C3EFFA6FF9DF903313527D6FECB" box="[1701,1720,268,300]" italics="true" pageId="1" pageNumber="1456">4</emphasis>
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,
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<emphasis id="B39F3C3EFFA6FF9DF97531352796FECB" box="[1747,1784,268,300]" italics="true" pageId="1" pageNumber="1456">12</emphasis>
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). Because MB is densely mineralized and extremely well vascularized, it provides an easily mobilized source of calcium necessary for the production of calcareous eggshells (
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<emphasis id="B39F3C3EFFA6FF9DF8B031822654FE3C" box="[1814,1850,443,475]" italics="true" pageId="1" pageNumber="1456">13</emphasis>
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). We compare MB from emu and ostrich (
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<emphasis id="B39F3C3EFFA6FF9DF89831DC260DFDE2" box="[1854,1891,485,517]" italics="true" pageId="1" pageNumber="1456">14</emphasis>
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) at different stages of the laying cycle with newly identified dinosaur tissues, because these basal birds share more primitive features with nonavian dinosaurs than do extant neognaths (
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<emphasis id="B39F3C3EFFA6FF9DF7A532AD2931FD53" box="[2051,2143,660,692]" italics="true" pageId="1" pageNumber="1456">15-18</emphasis>
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).
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</paragraph>
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<paragraph id="8154E02CFFA6FF9EF90632862050FBF7" blockId="1.[1644,2328,136,1085]" lastBlockId="2.[150,834,136,1346]" lastPageId="2" lastPageNumber="1457" pageId="1" pageNumber="1456">
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Our investigations show that ratite MB differs from that seen in better-studied neognaths. We observed substantial variation between emu and ostrich MB tissues and between both ratites and reported neognath tissues (
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<figureCitation id="19D0FCA9FFA6FF9DF94233A02656FC5E" box="[1764,1848,921,953]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="1" pageNumber="1456" targetBox="[526,2320,1508,2848]">Fig. 1</figureCitation>
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and fig. S1). MB (
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<figureCitation id="19D0FCA9FFA6FF9DF7C233A029D2FC5E" box="[2148,2236,921,953]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="1" pageNumber="1456" targetBox="[526,2320,1508,2848]">Fig. 1</figureCitation>
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) may be thick (chicken and ostrich) or quite thin (emu) at midshaft; and it may be separated by a distinct layer of endosteal laminar bone (ELB) as described by Chinsamy et al. (19) (chicken and emu), or not (ostrich). The innermost layer of MB in the ostrich Eadjacent to the endosteal surface of cortical bone (CB)] appears to arise from dense sheets to form tubular structures that parallel the long axis of the bone (
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<figureCitation id="19D0FCA9FFA5FF9EFE8A31B720FEFE49" box="[300,400,398,430]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="2" pageNumber="1457" targetBox="[526,2320,1508,2848]">Fig. 1I</figureCitation>
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). Thin hairlike spicules (
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<figureCitation id="19D0FCA9FFA5FF9EFCA331B721AFFE3C" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="2" pageNumber="1457" targetBox="[526,2320,1508,2848]">Fig. 1C</figureCitation>
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) of mineralized bone protrude from the tubes and may be intimately involved in their formation from the basal layer. Mineralized spicules were also noted arising from emu MB (fig. S2), but the tubelike structures were not so apparent or distinct. The MB tissues are morphologically distinct from overlying CB and are similarly distributed in both dinosaur (
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<figureCitation id="19D0FCA9FFA5FF9EFF05332F207CFCD1" box="[163,274,790,822]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="2" pageNumber="1457" targetBox="[526,2320,1508,2848]">Fig. 1D</figureCitation>
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) and ratite (
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<figureCitation id="19D0FCA9FFA5FF9EFE6F332F234FFCD1" box="[457,545,790,822]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="2" pageNumber="1457" targetBox="[526,2320,1508,2848]">Fig. 1</figureCitation>
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, E and F) samples. Higher magnifications of T. rex (
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<figureCitation id="19D0FCA9FFA5FF9EFD24337B239AFC85" box="[642,756,834,866]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="2" pageNumber="1457" targetBox="[526,2320,1508,2848]">Fig. 1G</figureCitation>
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) and ratite (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFF5C3357203CFC69" box="[250,338,878,910]" captionStart="Fig" captionStartId="1.[167,216,1509,1536]" captionTargetPageId="1" captionText="Fig. 1. Extant avian MB and homologous dinosaurian bone tissues. (A) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (B) Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB. (C) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (D) MB on endosteal surface of MOR 1125 femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (E) Emu and (F) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types. (G) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (H) and ostrich (I) tibia. Ostrich MB is apparently unique in forming longitudinal tubules." figureDoi="http://doi.org/10.5281/zenodo.4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="2" pageNumber="1457" targetBox="[526,2320,1508,2848]">Fig. 1</figureCitation>
|
||
, H and I) tissues show the open, crystalline, and fibrous structure of these highly vascular tissues, in contrast to the denser CB.
|
||
</paragraph>
|
||
<footnote id="E2F0FC22FFA6FF9DF9CB356326B5FA70" pageId="1" pageNumber="1456">
|
||
<paragraph id="8154E02CFFA6FF9DF9CB356326B5FA70" blockId="1.[1645,2325,1370,1431]" pageId="1" pageNumber="1456">*To whom correspondence should be addressed. E-mail: schweitzer@ncsu.edu</paragraph>
|
||
</footnote>
|
||
<caption id="D594B0A4FFA6FF9DFF0135DC2477F46B" ID-DOI="http://doi.org/10.5281/zenodo.4012006" ID-Zenodo-Dep="4012006" httpUri="https://zenodo.org/record/4012006/files/figure.png" pageId="1" pageNumber="1456" startId="1.[167,177,2892,2919]" targetBox="[526,2320,1508,2848]" targetPageId="1">
|
||
<paragraph id="8154E02CFFA6FF9DFF0135DC208DF4A5" blockId="1.[167,485,1509,2882]" pageId="1" pageNumber="1456">
|
||
Fig. 1. Extant avian MB and homologous dinosaurian bone tissues.
|
||
<emphasis id="B39F3C3EFFA6FF9DFF01366C21A7F997" bold="true" box="[167,201,1621,1648]" pageId="1" pageNumber="1456">(A</emphasis>
|
||
) Domestic laying hen, midshaft femur cross section showing the extension of spongy MB deep into the mar row cavity and surrounding preexisting trabeculae (T). (
|
||
<emphasis id="B39F3C3EFFA6FF9DFE623762208AF891" bold="true" box="[452,484,1883,1910]" pageId="1" pageNumber="1456">B)</emphasis>
|
||
Laying emu, midshaft cross section, with a thin layer of MB on the endosteal bone surface, separated from overlying CB by ELB.
|
||
<emphasis id="B39F3C3EFFA6FF9DFF01385921ABF79C" bold="true" box="[167,197,2144,2171]" pageId="1" pageNumber="1456">(C</emphasis>
|
||
) Ostrich MB arising from CB. Convoluted bony projections surround large cavities and form by continued deposition on hairlike spicules of calcified bone (S). (
|
||
<emphasis id="B39F3C3EFFA6FF9DFEEE395F2031F666" bold="true" box="[328,351,2406,2433]" pageId="1" pageNumber="1456">D</emphasis>
|
||
) MB on endosteal surface of
|
||
<materialsCitation id="3183EA71FFA6FF9DFF0139882051F62B" ID-GBIF-Occurrence="2979293317" box="[167,319,2481,2508]" collectionCode="MOR" pageId="1" pageNumber="1456" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
femur fragment delineated from overlying CB by large vascular sinuses and change in color, texture, and density. (
|
||
<emphasis id="B39F3C3EFFA6FF9DFED13A5520E8F560" bold="true" box="[375,390,2668,2695]" pageId="1" pageNumber="1456">E</emphasis>
|
||
) Emu and (
|
||
<emphasis id="B39F3C3EFFA6FF9DFF5D3AA82065F54B" bold="true" box="[251,267,2705,2732]" pageId="1" pageNumber="1456">F</emphasis>
|
||
) ostrich bone taken at same aspect as (D), showing mor phological distinction between bone types.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA6FF9DFF013B752477F46B" blockId="1.[167,2326,2892,2956]" pageId="1" pageNumber="1456">
|
||
<emphasis id="B39F3C3EFFA6FF9DFF013B7521A9F480" bold="true" box="[167,199,2892,2919]" pageId="1" pageNumber="1456">(G</emphasis>
|
||
) Higher magnification of dinosaur femur fragment in oblique view shows dense CB lined with newly described bone tissue, also seen in oblique view of emu (
|
||
<emphasis id="B39F3C3EFFA6FF9DF7553B752864F480" bold="true" box="[2291,2314,2892,2919]" pageId="1" pageNumber="1456">H</emphasis>
|
||
) and ostrich (
|
||
<emphasis id="B39F3C3EFFA6FF9DFEF73B482036F46B" bold="true" box="[337,344,2929,2956]" pageId="1" pageNumber="1456">I</emphasis>
|
||
) tibia. Ostrich MB is apparently unique in forming longitudinal tubules.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8154E02CFFA5FF9EFF61342424ADFDBA" blockId="2.[150,834,136,1346]" lastBlockId="2.[888,1573,136,1346]" pageId="2" pageNumber="1457">
|
||
In a fresh fracture, dense and relatively homogenous dinosaur CB is distinct from the loosely organized and highly vascular MB internal to it (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFECA34A62088FB58" box="[364,486,1183,1215]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2A</figureCitation>
|
||
). A distinct layer corresponding to ELB (19) separates the two bone types. A large erosion room is visible at this boundary, lined with laminar tissue. An emu bone fragment (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFB1830B3245DFF4D" box="[1214,1331,138,170]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2B</figureCitation>
|
||
) in similar orientation shows MB tissues with a distinctive, less organized and Bcrumbly[texture relative to overlying CB. It is interspersed with or laid down between large erosion rooms within the deep cortex and ELB of the tibial shaft. The dense cortex and laminar structure of the ELB are easily distinguished from surrounding MB. The ostrich MB (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFB6F31DC2454FDE2" box="[1225,1338,485,517]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2C</figureCitation>
|
||
) differs in both texture and orientation, with open cavities that are bordered by tubelike bone spicules.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA5FF9EFC0C3251293CFD53" blockId="2.[888,1573,136,1346]" lastBlockId="2.[1627,2309,136,1346]" pageId="2" pageNumber="1457">
|
||
The lacy vascularity of the T. rex tissues (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFC2332AD256EFD53" box="[901,1024,660,692]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2D</figureCitation>
|
||
) is consistent with the larger vascular canals and whorled pattern of the emu (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFC2332D5256DFCEB" box="[901,1027,748,780]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2E</figureCitation>
|
||
) and especially with the ostrich (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFC23332F256EFCD1" box="[901,1024,790,822]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2F</figureCitation>
|
||
), where wide blood-filled sinuses separate forming bone spicules. In ground section (
|
||
<figureCitation id="19D0FCA9FFA5FF9EFC5A3357251BFC69" box="[1020,1141,878,910]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2G</figureCitation>
|
||
),
|
||
<materialsCitation id="3183EA71FFA5FF9EFB343357242EFC69" ID-GBIF-Occurrence="2979293305" box="[1170,1344,878,910]" collectionCode="MOR" pageId="2" pageNumber="1457" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
femur cortical bone is characterized by well-developed multigeneration Haversian systems with obvious and defined cement lines, supporting a mature status for this dinosaur (2). A region of decreased vascularity and laminar structure marks the ELB. In contrast, the medullary tissues arising from the ELB are densely vascularized but show no evidence of Haversian remodeling or cement lines, indicating that this tissue is newly deposited, or younger bone. A fresh cut section of emu bone (
|
||
<figureCitation id="19D0FCA9FFA5FF9EF96E308D2627FF33" box="[1736,1865,180,212]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2H</figureCitation>
|
||
) in comparable orientation shows dense CB and vascular, crystalline, and loosely organized MB, separated by a thin, dense, and less vascular ELB. MB in the ostrich (
|
||
<figureCitation id="19D0FCA9FFA5FF9EF8B6315A2618FE64" box="[1808,1910,355,387]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2I</figureCitation>
|
||
) is more extensive than in the emu samples, most likely because shelling had not yet begun (14). No distinct ELB is visible. MB appears laminar rather than spiculated in
|
||
<figureCitation id="19D0FCA9FFA5FF9EF89D322826C0FDD6" box="[1851,1966,529,561]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2I</figureCitation>
|
||
, because the tubules formed by bony spicules are oriented longitudinally rather than in cross section as in
|
||
<figureCitation id="19D0FCA9FFA5FF9EF9FD32AD27A0FD53" box="[1627,1742,660,692]" captionStart="Fig" captionStartId="2.[150,199,1435,1462]" captionTargetBox="[508,2306,1430,2785]" captionTargetPageId="2" captionText="Fig. 2. Dinosaur and ratite comparative views. (A) Freshly broken fragment of MOR 1125 shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (B) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB. (C) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (D) Higher magnification of MB region of MOR 1125, showing increased porosity and more random orientation of MB than CB or ELB. (E) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (F) Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (G) Ground section of MOR 1125. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (H) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (I) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules." figureDoi="http://doi.org/10.5281/zenodo.4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" targetBox="[508,2306,1430,2785]">Fig. 2F</figureCitation>
|
||
, but it is the same tissue.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA5FF9EF92B3286278EFB74" blockId="2.[1627,2309,136,1346]" pageId="2" pageNumber="1457">
|
||
Additional pattern similarities are seen in demineralized (14) ratite (
|
||
<figureCitation id="19D0FCA9FFA5FF9EF87032D52940FCEB" box="[2006,2094,748,780]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="2" pageNumber="1457" targetBox="[912,2320,134,1544]">Fig. 3</figureCitation>
|
||
, B and C) and T. rex (
|
||
<figureCitation id="19D0FCA9FFA5FF9EF961332F2656FCD1" box="[1735,1848,790,822]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="2" pageNumber="1457" targetBox="[912,2320,134,1544]">Fig. 3A</figureCitation>
|
||
) medullary tissues. In all cases, the matrix is fibrous and randomly organized. The reddish color in extant tissues is due to blood retained in sinuses that separate the bone spicules. The T. rex tissues are similarly pigmented, due either to diagenetic alteration or to close association of bony tissues with blood-producing marrow during the life of the dinosaur.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA5FF9FF92B34A620EAFB0D" blockId="2.[1627,2309,136,1346]" lastBlockId="3.[167,849,136,2959]" lastPageId="3" lastPageNumber="1458" pageId="2" pageNumber="1457">
|
||
In all MB tissues shown, large vascular sinuses are easily discerned (
|
||
<figureCitation id="19D0FCA9FFA5FF9EF84D34F3292FFB0D" box="[2027,2113,1226,1258]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="2" pageNumber="1457" targetBox="[912,2320,134,1544]">Fig. 3</figureCitation>
|
||
, D to F), but in the T. rex, vascular openings are surrounded by circumferentially oriented matrix fibers (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFEA830B320EFFF4D" box="[270,385,138,170]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3D</figureCitation>
|
||
) that are less apparent in extant bone. The ostrich medullary tissues are denser than either the emu (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFDB530D923FCFEE7" box="[531,658,224,256]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3E</figureCitation>
|
||
) or T.
|
||
<emphasis id="B39F3C3EFFA4FF9FFC8630DB223EFEE7" box="[800,848,226,256]" italics="true" pageId="3" pageNumber="1458">rex</emphasis>
|
||
samples, particularly closer to the cortex, but the variation in the size and density of vascular sinuses (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFE1E315A2340FE64" box="[440,558,355,387]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3F</figureCitation>
|
||
)
|
||
<emphasis id="B39F3C3EFFA4FF9FFDEF315A230DFE64" box="[585,611,355,387]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
similar to that seen in the T.
|
||
<emphasis id="B39F3C3EFFA4FF9FFE3831A920A0FE49" box="[414,462,400,430]" italics="true" pageId="3" pageNumber="1458">rex</emphasis>
|
||
tissues. In planar view,
|
||
<materialsCitation id="3183EA71FFA4FF9FFF0131822039FE3C" ID-GBIF-Occurrence="2979293308" box="[167,343,443,475]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
undemineralized tissues show a random orientation of fibers, and vascular openings penetrate deep into the tissues (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFCB3322821B6FDBA" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3G</figureCitation>
|
||
, inset) and exhibit an unusual doublet or triplet pattern, where multiple vessels penetrate an osteonlike core (arrows), also seen in the emu (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFE9C328620DAFD38" box="[314,436,703,735]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">
|
||
Fig. 3
|
||
<emphasis id="B39F3C3EFFA4FF9FFE32328620DAFD38" box="[404,436,703,735]" italics="true" pageId="3" pageNumber="1458">H</emphasis>
|
||
</figureCitation>
|
||
, arrows). The ostrich medullary tissues (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFE3032D52366FCEB" box="[406,520,748,780]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3I</figureCitation>
|
||
) are more variable, denser, and less random in appearance than those of the emu, but the morphology changes as the tissues extend into the medullary cavity. Close to the cortex (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFDD333A02389FC5E" box="[629,743,921,953]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3I</figureCitation>
|
||
, inset, and fig. S3), the bone
|
||
<emphasis id="B39F3C3EFFA4FF9FFDA333FC2371FC02" box="[517,543,965,997]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
sheetlike, relatively dense, and punctured by vascular sinuses exhibiting the doublet pattern (arrows) noted above. As tissues extend into the medullary cavity (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFE85344A20F9FB74" box="[291,407,1139,1171]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3I</figureCitation>
|
||
), this pattern becomes obscured. Inset bone has been stained (14) for better contrast.
|
||
</paragraph>
|
||
<caption id="D594B0A4FFA5FF9EFF3035A226A2F46B" ID-DOI="http://doi.org/10.5281/zenodo.4012008" ID-Zenodo-Dep="4012008" httpUri="https://zenodo.org/record/4012008/files/figure.png" pageId="2" pageNumber="1457" startId="2.[150,199,1435,1462]" targetBox="[508,2306,1430,2785]" targetPageId="2">
|
||
<paragraph id="8154E02CFFA5FF9EFF3035A226A2F46B" blockId="2.[150,1205,1435,2956]" lastBlockId="2.[1254,2309,2855,2956]" pageId="2" pageNumber="1457">
|
||
Fig. 2. Dinosaur and ratite comparative views. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFF5D35DC207EF9E7" bold="true" box="[251,272,1509,1536]" pageId="2" pageNumber="1457">A</emphasis>
|
||
) Freshly broken fragment of
|
||
<materialsCitation id="3183EA71FFA5FF9EFE26363221B1F9AC" ID-GBIF-Occurrence="2979293319" collectionCode="MOR" pageId="2" pageNumber="1457" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
shows laminar ELB separating CB and MB. Bone tissues decrease in density internal to the ELB, because of increased vascularity. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFEF237292009F8CC" bold="true" box="[340,359,1808,1835]" pageId="2" pageNumber="1457">B</emphasis>
|
||
) Emu tibia, midshaft section. Erosion rooms extending into ELB are secondarily filled by MB.
|
||
<emphasis id="B39F3C3EFFA5FF9EFF3037F221DAF801" bold="true" box="[150,180,1995,2022]" pageId="2" pageNumber="1457">(C</emphasis>
|
||
) Ostrich bone, mid shaft. MB is distinct from CB, but no obvious ELB is visible and several large vascular sinuses are seen. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFE0338BF20A9F746" bold="true" box="[421,455,2182,2209]" pageId="2" pageNumber="1457">D)</emphasis>
|
||
Higher magnification of MB region of
|
||
<materialsCitation id="3183EA71FFA5FF9EFE2638E82188F6F6" ID-GBIF-Occurrence="2979293301" collectionCode="MOR" pageId="2" pageNumber="1457" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, showing increased porosity and more random orientation of MB than CB or ELB. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFF4139B52198F640" bold="true" box="[231,246,2444,2471]" pageId="2" pageNumber="1457">E</emphasis>
|
||
) Emu, stained (14) to distinguish bone from infiltrating marrow fat. MB is more vascular than overlying CB and exhibits a random, whorled pattern. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFE0A3AA820A9F54B" bold="true" box="[428,455,2705,2732]" pageId="2" pageNumber="1457">F)</emphasis>
|
||
Ostrich MB, showing relationship of bony spicules to invading blood sinuses, colored red from remnant blood. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFC9D3B1E223FF4A5" bold="true" box="[827,849,2855,2882]" pageId="2" pageNumber="1457">G</emphasis>
|
||
) Ground section of
|
||
<materialsCitation id="3183EA71FFA5FF9EFBCB3B1E21B1F480" ID-GBIF-Occurrence="2979293307" collectionCode="MOR" pageId="2" pageNumber="1457" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
. Dense cortical Haversian bone shows second- and third-generation remodeling. ELB separates Haversian bone from more vascular MB. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFB8D3B48252DF46B" bold="true" box="[1067,1091,2929,2956]" pageId="2" pageNumber="1457">H</emphasis>
|
||
) Similar orientation of emu femur shows dense CB, distinct ELB, and a thin layer of MB. (
|
||
<emphasis id="B39F3C3EFFA5FF9EFA963B752456F480" bold="true" box="[1328,1336,2892,2919]" pageId="2" pageNumber="1457">I</emphasis>
|
||
) Ostrich MB appears more laminar than in (C) or (F) because of the longitudinal orientation of tubelike medullary spicules.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8154E02CFFA4FF9FFF7F34CF23DAF994" blockId="3.[167,849,136,2959]" pageId="3" pageNumber="1458">
|
||
In regions of
|
||
<materialsCitation id="3183EA71FFA4FF9FFE6F34CF23EEFAF1" ID-GBIF-Occurrence="2979293304" box="[457,640,1270,1302]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
bone where most of the medullary tissues have eroded (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFF143577204EFA89" box="[178,288,1358,1390]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3J</figureCitation>
|
||
), patches of denser CB can be seen, emphasizing the random mazelike pattern and large vascular sinuses of medullary tissues, a pattern also seen in the emu bone (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFF1435C2205FF9FC" box="[178,305,1531,1563]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3K</figureCitation>
|
||
). The ostrich MB shows a similar pattern of bony spicules surrounding large and small blood sinuses (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFD8A366A23CDF994" box="[556,675,1619,1651]" captionStart="Fig. 3" captionStartId="3.[906,955,1586,1613]" captionTargetBox="[912,2320,134,1544]" captionTargetPageId="3" captionText="Fig. 3. Dinosaur and ratite MB. (A) MOR 1125, (B) emu, and (C) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (D) MOR 1125, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (E) emu and (F) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (G) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (I) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of MOR 1125 shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (L) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses." figureDoi="http://doi.org/10.5281/zenodo.3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" targetBox="[912,2320,134,1544]">Fig. 3L</figureCitation>
|
||
).
|
||
</paragraph>
|
||
<caption id="D594B0A4FFA4FF9FFC2C360B2596F79A" ID-DOI="http://doi.org/10.5281/zenodo.3742902" ID-Zenodo-Dep="3742902" httpUri="https://zenodo.org/record/3742902/files/figure.png" pageId="3" pageNumber="1458" startId="3.[906,955,1586,1613]" targetBox="[912,2320,134,1544]" targetPageId="3">
|
||
<paragraph id="8154E02CFFA4FF9FFC2C360B2596F79A" blockId="3.[906,2327,1581,2176]" pageId="3" pageNumber="1458">
|
||
<emphasis id="B39F3C3EFFA4FF9FFC2C360B22B0F9AA" bold="true" box="[906,990,1586,1613]" pageId="3" pageNumber="1458">Fig. 3</emphasis>
|
||
. Dinosaur and ratite MB. (A)
|
||
<materialsCitation id="3183EA71FFA4FF9FFA26360B2771F9AA" ID-GBIF-Occurrence="2979293302" box="[1408,1567,1586,1613]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, (
|
||
<emphasis id="B39F3C3EFFA4FF9FF991360B2724F9AA" bold="true" box="[1591,1610,1586,1613]" pageId="3" pageNumber="1458">B</emphasis>
|
||
) emu, and (
|
||
<emphasis id="B39F3C3EFFA4FF9FF951360B2663F9AA" bold="true" box="[1783,1805,1586,1613]" pageId="3" pageNumber="1458">C</emphasis>
|
||
) ostrich demineralized (14) MB. The coloration in (B) and (C) results from infiltration of tissues with blood sinuses. (
|
||
<emphasis id="B39F3C3EFFA4FF9FF79B366E293AF995" bold="true" box="[2109,2132,1623,1650]" pageId="3" pageNumber="1458">D</emphasis>
|
||
)
|
||
<materialsCitation id="3183EA71FFA4FF9FF7C8366E287FF995" ID-GBIF-Occurrence="2979293309" box="[2158,2321,1623,1650]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, partially demineralized, showing enlarged, randomly arranged vascular openings surrounded by circumferential matrix fibers. Partially demineralized (
|
||
<emphasis id="B39F3C3EFFA4FF9FF908369B27D1F95A" bold="true" box="[1710,1727,1698,1725]" pageId="3" pageNumber="1458">E</emphasis>
|
||
) emu and (
|
||
<emphasis id="B39F3C3EFFA4FF9FF8DF369B26E4F95A" bold="true" box="[1913,1930,1698,1725]" pageId="3" pageNumber="1458">F</emphasis>
|
||
) ostrich medullary tissues show extensive vascular penetration, with randomly spaced and varyingly sized vessel openings. (
|
||
<emphasis id="B39F3C3EFFA4FF9FFC3336D422C3F8EF" box="[917,941,1773,1800]" italics="true" pageId="3" pageNumber="1458">G</emphasis>
|
||
) Plane view of undemineralized dinosaur tissues shows fibrous matrix and an unusual pattern of vascular doublets or triplets within osteonlike structures (arrows). The inset shows variation in depth and diameter of vascular sinuses. (H) Undemineralized emu MB shows similar doublet pattern (arrows) and fibrous matrix. The greater depth of field makes focusing difficult. (
|
||
<emphasis id="B39F3C3EFFA4FF9FF728376429F8F89F" bold="true" box="[2190,2198,1885,1912]" pageId="3" pageNumber="1458">I</emphasis>
|
||
) Ostrich MB is denser closer to the cortex (inset), where the doublet/triplet pattern of vessels (arrows) is evident, but this becomes obscured by the increasing development of bony tubes and spicules as bone extends further into the medullary cavity. (J) Plane view of
|
||
<materialsCitation id="3183EA71FFA4FF9FF8F537F4269AF80F" ID-GBIF-Occurrence="2979293313" box="[1875,2036,1997,2024]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
shows the partially eroded, fibrous MB distributed across the cortex in a mazelike fashion. (K) Emu MB shows white (chloroform-altered) and cream-colored MB in the same mazelike pattern. (
|
||
<emphasis id="B39F3C3EFFA4FF9FF84F382E2696F7D5" bold="true" box="[2025,2040,2071,2098]" pageId="3" pageNumber="1458">L</emphasis>
|
||
) Thicker, randomly oriented tubular spicules of ostrich MB, showing deep penetration and intimate association of blood-containing sinuses.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8154E02CFFA4FF9FFF7F3646223EF7C1" blockId="3.[167,849,136,2959]" pageId="3" pageNumber="1458">
|
||
Scanning electron micrographs (14) reveal the distinctive grainy texture and disorganized morphology of demineralized T.
|
||
<emphasis id="B39F3C3EFFA4FF9FFD2F36EE23D7F912" box="[649,697,1751,1781]" italics="true" pageId="3" pageNumber="1458">rex</emphasis>
|
||
and avian MB (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFF5C37382034F8C6" box="[250,346,1793,1825]" captionStart="Fig" captionStartId="4.[1997,2046,138,165]" captionTargetBox="[1499,1948,134,810]" captionTargetPageId="4" captionText="Fig. 4. Scanning electron microscope images of demineralized MB [(A) to (D)] and CB [(E) to (K)]. Demineralized, aldehyde-fixed (14) MB tissues from (A) MOR 1125, (B) extant laying hen, (C) emu, and (D) ostrich show random, crumbly texture. Organized collagen fiber bundles are not distinct in any sample because of rapid deposition and woven character. Scale bars for (A) and (B), 40 um; for (C) and (D), 20 um. Demineralized fragments of cortical bone from (E) MOR 1125, (F) chicken, (G) emu, and (H) ostrich are shown. A fibrous character dominates all samples. Scale bars for (E), (F), and (H), 30 mm; for (G), 10 mm. Higher magnification of demineralized CB from (I) MOR 1125, (J) emu, and (K) ostrich CB demonstrates the structural similarity between samples, although the MOR 1125 matrix is highly degraded. Scale bars for (I) and (K), 6 um; for (J), 5 um." figureDoi="http://doi.org/10.5281/zenodo.4751441" httpUri="https://zenodo.org/record/4751441/files/figure.png" pageId="3" pageNumber="1458">Fig. 4</figureCitation>
|
||
). This contrasts with the smooth and fibrous texture of demineralized CB from the same specimens (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFE4A376123A2F89F" box="[492,716,1880,1912]" captionStart="Fig" captionStartId="4.[1997,2046,138,165]" captionTargetBox="[1499,1948,134,810]" captionTargetPageId="4" captionText="Fig. 4. Scanning electron microscope images of demineralized MB [(A) to (D)] and CB [(E) to (K)]. Demineralized, aldehyde-fixed (14) MB tissues from (A) MOR 1125, (B) extant laying hen, (C) emu, and (D) ostrich show random, crumbly texture. Organized collagen fiber bundles are not distinct in any sample because of rapid deposition and woven character. Scale bars for (A) and (B), 40 um; for (C) and (D), 20 um. Demineralized fragments of cortical bone from (E) MOR 1125, (F) chicken, (G) emu, and (H) ostrich are shown. A fibrous character dominates all samples. Scale bars for (E), (F), and (H), 30 mm; for (G), 10 mm. Higher magnification of demineralized CB from (I) MOR 1125, (J) emu, and (K) ostrich CB demonstrates the structural similarity between samples, although the MOR 1125 matrix is highly degraded. Scale bars for (I) and (K), 6 um; for (J), 5 um." figureDoi="http://doi.org/10.5281/zenodo.4751441" httpUri="https://zenodo.org/record/4751441/files/figure.png" pageId="3" pageNumber="1458">Fig. 4, E to H</figureCitation>
|
||
). Higher magnifications of demineralized CB (
|
||
<figureCitation id="19D0FCA9FFA4FF9FFD7337BD2184F837" captionStart="Fig" captionStartId="4.[1997,2046,138,165]" captionTargetBox="[1499,1948,134,810]" captionTargetPageId="4" captionText="Fig. 4. Scanning electron microscope images of demineralized MB [(A) to (D)] and CB [(E) to (K)]. Demineralized, aldehyde-fixed (14) MB tissues from (A) MOR 1125, (B) extant laying hen, (C) emu, and (D) ostrich show random, crumbly texture. Organized collagen fiber bundles are not distinct in any sample because of rapid deposition and woven character. Scale bars for (A) and (B), 40 um; for (C) and (D), 20 um. Demineralized fragments of cortical bone from (E) MOR 1125, (F) chicken, (G) emu, and (H) ostrich are shown. A fibrous character dominates all samples. Scale bars for (E), (F), and (H), 30 mm; for (G), 10 mm. Higher magnification of demineralized CB from (I) MOR 1125, (J) emu, and (K) ostrich CB demonstrates the structural similarity between samples, although the MOR 1125 matrix is highly degraded. Scale bars for (I) and (K), 6 um; for (J), 5 um." figureDoi="http://doi.org/10.5281/zenodo.4751441" httpUri="https://zenodo.org/record/4751441/files/figure.png" pageId="3" pageNumber="1458">Fig. 4, I to K</figureCitation>
|
||
) emphasize the smooth, fibrous, and more ordered nature of all specimens, although in
|
||
<materialsCitation id="3183EA71FFA4FF9FFF01383F203FF7C1" ID-GBIF-Occurrence="2979293306" box="[167,337,2054,2086]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
(
|
||
<figureCitation id="19D0FCA9FFA4FF9FFEC2383F20A0F7C1" box="[356,462,2054,2086]" captionStart="Fig" captionStartId="4.[1997,2046,138,165]" captionTargetBox="[1499,1948,134,810]" captionTargetPageId="4" captionText="Fig. 4. Scanning electron microscope images of demineralized MB [(A) to (D)] and CB [(E) to (K)]. Demineralized, aldehyde-fixed (14) MB tissues from (A) MOR 1125, (B) extant laying hen, (C) emu, and (D) ostrich show random, crumbly texture. Organized collagen fiber bundles are not distinct in any sample because of rapid deposition and woven character. Scale bars for (A) and (B), 40 um; for (C) and (D), 20 um. Demineralized fragments of cortical bone from (E) MOR 1125, (F) chicken, (G) emu, and (H) ostrich are shown. A fibrous character dominates all samples. Scale bars for (E), (F), and (H), 30 mm; for (G), 10 mm. Higher magnification of demineralized CB from (I) MOR 1125, (J) emu, and (K) ostrich CB demonstrates the structural similarity between samples, although the MOR 1125 matrix is highly degraded. Scale bars for (I) and (K), 6 um; for (J), 5 um." figureDoi="http://doi.org/10.5281/zenodo.4751441" httpUri="https://zenodo.org/record/4751441/files/figure.png" pageId="3" pageNumber="1458">Fig. 4I</figureCitation>
|
||
), degradation
|
||
<emphasis id="B39F3C3EFFA4FF9FFD05383F23D3F7C1" box="[675,701,2054,2086]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
apparent.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA4FF9FFF7F380B2560F6CB" blockId="3.[167,849,136,2959]" lastBlockId="3.[906,1587,2228,2959]" pageId="3" pageNumber="1458">
|
||
MB occurs naturally only in extant female birds, although it varies in amount and distribution among taxa and with ovulatory phase (5, 20). It
|
||
<emphasis id="B39F3C3EFFA4FF9FFEF9388C2017F732" box="[351,377,2229,2261]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
chemically, functionally, and structurally distinct from both overlying CB and internal trabecular bone (21, 22). Although "medullary" and "trabecular" bone are terms often used interchangeably in the literature, MB has a larger surface area and
|
||
<emphasis id="B39F3C3EFFA4FF9FFC9E39A9223EF657" box="[824,848,2448,2480]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
more vascular than other bone types, allowing rapid calcium mobilization (5). It
|
||
<emphasis id="B39F3C3EFFA4FF9FFD7339DF2380F5E1" box="[725,750,2534,2566]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
more highly mineralized, with a greater apatite-tocollagen ratio (5, 7,
|
||
<emphasis id="B39F3C3EFFA4FF9FFE463A042322F5BA" box="[480,588,2621,2653]" italics="true" pageId="3" pageNumber="1458">20-22),</emphasis>
|
||
and incorporates acidic mucopolysaccharides and glycosaminoglycans that are not present in CB (
|
||
<emphasis id="B39F3C3EFFA4FF9FFD4C3AAD2393F553" box="[746,765,2708,2740]" italics="true" pageId="3" pageNumber="1458">5</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA4FF9FFCB13AAD2252F553" box="[791,828,2708,2740]" italics="true" pageId="3" pageNumber="1458">11</emphasis>
|
||
). Additionally, the matrix of MB
|
||
<emphasis id="B39F3C3EFFA4FF9FFD383AF823DBF506" box="[670,693,2753,2785]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
higher in noncollagenous proteins and lower in collagen, and has a higher collagen III-to-collagen I ratio (
|
||
<emphasis id="B39F3C3EFFA4FF9FFEAB3B7A205FF484" box="[269,305,2883,2915]" italics="true" pageId="3" pageNumber="1458">22</emphasis>
|
||
) relative to other bone types. If preservation allows, these characteristics will be used as part of ongoing research to chemically distinguish the two bone types in this dinosaur.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA4FF9FFC1A3901277CF506" blockId="3.[906,1587,2228,2959]" pageId="3" pageNumber="1458">
|
||
The existence of avian-type MB in dinosaurs has been hypothesized (
|
||
<emphasis id="B39F3C3EFFA4FF9FFAFC395A2401F664" box="[1370,1391,2403,2435]" italics="true" pageId="3" pageNumber="1458">9</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA4FF9FFA20395A24C5F664" box="[1414,1451,2403,2435]" italics="true" pageId="3" pageNumber="1458">23</emphasis>
|
||
) but not identified. In part, this could be because of taphonomic bias, because the death and fossilization of an ovulating dinosaur would be comparatively rare. Additionally, MB in extant birds
|
||
<emphasis id="B39F3C3EFFA4FF9FFC443A042297F5BA" box="[994,1017,2621,2653]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
fragile, the spicules separating easily from the originating layer (fig. S1). Dinosaur MB may separate and be lost from overlying CB in a similar manner during diagenesis.
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA4FF9FFC1A3AD22781F4D0" blockId="3.[906,1587,2228,2959]" lastBlockId="3.[1645,2326,2229,2959]" pageId="3" pageNumber="1458">
|
||
The location, origin, morphology, and microstructure of the new
|
||
<emphasis id="B39F3C3EFFA4FF9FFABC3B2E2418F4D0" box="[1306,1398,2839,2871]" italics="true" pageId="3" pageNumber="1458">T. rex</emphasis>
|
||
tissues support homology with ratite MB. The
|
||
<emphasis id="B39F3C3EFFA4FF9FFA043B7A2494F484" box="[1442,1530,2883,2915]" italics="true" pageId="3" pageNumber="1458">T. rex</emphasis>
|
||
tissues line the medullary cavities of both femora of
|
||
<materialsCitation id="3183EA71FFA4FF9FF8AD388C26D8F732" ID-GBIF-Occurrence="2979293310" box="[1803,1974,2229,2261]" collectionCode="MOR" pageId="3" pageNumber="1458" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, suggesting an organismal response. The tissues are similar in distribution to those of extant ratites, being more extensive in proximal regions of the bone. They are clearly endosteal in origin, and the microstructure with large vascular sinuses
|
||
<emphasis id="B39F3C3EFFA4FF9FF75939A9287AF657" box="[2303,2324,2448,2480]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
consistent with the function of MB as a rapidly deposited and easily mobilized calcium source. The random, woven character indicates rapidly deposited, younger bone. Finally, the robustly supported relationship between theropods and extant birds (
|
||
<emphasis id="B39F3C3EFFA4FF9FF8643AAD294FF553" box="[1986,2081,2708,2740]" italics="true" pageId="3" pageNumber="1458">15-18</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA4FF9FF7953AAD2937F553" box="[2099,2137,2708,2740]" italics="true" pageId="3" pageNumber="1458">24</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA4FF9FF7CD3AAD29FEF553" box="[2155,2192,2708,2740]" italics="true" pageId="3" pageNumber="1458">25</emphasis>
|
||
) permits the application of phylogenetic inference to support the identification of these tissues (
|
||
<emphasis id="B39F3C3EFFA4FF9FF9DC3B2E27CFF4D0" box="[1658,1697,2839,2871]" italics="true" pageId="3" pageNumber="1458">26</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA4FF9FF9133B2E27B5F4D0" box="[1717,1755,2839,2871]" italics="true" pageId="3" pageNumber="1458">27</emphasis>
|
||
).
|
||
</paragraph>
|
||
<paragraph id="8154E02CFFA4FF98F9393B7A20DFF506" blockId="3.[1645,2326,2229,2959]" lastBlockId="4.[150,831,1269,2958]" lastPageId="4" lastPageNumber="1459" pageId="3" pageNumber="1458">
|
||
The morphology of these dinosaur tissues
|
||
<emphasis id="B39F3C3EFFA4FF9FF9C83B5727EBF469" box="[1646,1669,2926,2958]" italics="true" pageId="3" pageNumber="1458">is</emphasis>
|
||
not identical to that of extant neognaths (fig. S1), but is more similar to that seen in ratites. T.
|
||
<emphasis id="B39F3C3EFFA3FF98FE93351D2008FAA5" box="[309,358,1316,1346]" italics="true" pageId="4" pageNumber="1459">rex</emphasis>
|
||
medullary tissues are less extensive than those reported for neognaths, which may be explained by many factors. First, there
|
||
<emphasis id="B39F3C3EFFA3FF98FEEE359D200CFA23" box="[328,354,1444,1476]" italics="true" pageId="4" pageNumber="1459">is</emphasis>
|
||
a wide range of MB morphologies in extant taxa (20), varying with both reproductive phase and the position of the egg within the reproductive tract (6). Medullary tissues become thinner as shelling progresses and disappear completely with deposition of the last egg. If the same was true of dinosaurs,
|
||
<materialsCitation id="3183EA71FFA3FF98FE0F36EC233BF912" ID-GBIF-Occurrence="2979293320" box="[425,597,1749,1781]" collectionCode="MOR" pageId="4" pageNumber="1459" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
may have died toward the end of the laying cycle. Second, MB in extant birds
|
||
<emphasis id="B39F3C3EFFA3FF98FE7237172080F8A9" box="[468,494,1838,1870]" italics="true" pageId="4" pageNumber="1459">is</emphasis>
|
||
hypothesized to provide a buffer against excessive and debilitating bone resorption during shelling (4, 28). It
|
||
<emphasis id="B39F3C3EFFA3FF98FF30378921C1F837" box="[150,175,1968,2000]" italics="true" pageId="4" pageNumber="1459">is</emphasis>
|
||
most extensive in smaller taxa with high reproductive rates, because of the demand for rapid mobilization of skeletal calcium for shelling. Extinct theropods produced hardshelled eggs, as did other dinosaurs and all extant birds (29, 30). Although egg size is not known, eggs were most likely smaller relative to overall body size than in extant birds, resulting in less demand for bone calcium reserves and reducing the need to offset resorption. These factors may also contribute to the smaller ratio of medullary to cortical thickness in theropods than in extant birds. Finally, T.
|
||
<emphasis id="B39F3C3EFFA3FF98FEF139D120E9F5E1" box="[343,391,2536,2566]" italics="true" pageId="4" pageNumber="1459">rex</emphasis>
|
||
, although phylogenetically close to extant birds
|
||
<emphasis id="B39F3C3EFFA3FF98FE553A2B239FF5D5" box="[499,753,2578,2610]" italics="true" pageId="4" pageNumber="1459">(15-18, 24, 25),</emphasis>
|
||
was distinct in size, biomechanical constraints, and, to some degree, physiology (
|
||
<emphasis id="B39F3C3EFFA3FF98FD383A5023AAF56E" box="[670,708,2665,2697]" italics="true" pageId="4" pageNumber="1459">31</emphasis>
|
||
); therefore, slight variations in bone and tissue types would be expected.
|
||
</paragraph>
|
||
<caption id="D594B0A4FFA3FF98F86B30B327D7FB70" ID-DOI="http://doi.org/10.5281/zenodo.4751441" ID-Zenodo-Dep="4751441" httpUri="https://zenodo.org/record/4751441/files/figure.png" pageId="4" pageNumber="1459" startId="4.[1997,2046,138,165]" targetBox="[1499,1948,134,810]" targetPageId="4">
|
||
<paragraph id="8154E02CFFA3FF98F86B30B327D7FB70" blockId="4.[1997,2309,138,838]" lastBlockId="4.[1546,2308,848,1176]" pageId="4" pageNumber="1459">
|
||
<emphasis id="B39F3C3EFFA3FF98F86B30B32690FF42" bold="true" box="[1997,2046,138,165]" pageId="4" pageNumber="1459">Fig</emphasis>
|
||
. 4.
|
||
<emphasis id="B39F3C3EFFA3FF98F79830B329D9FF42" box="[2110,2231,138,165]" italics="true" pageId="4" pageNumber="1459">Scanning</emphasis>
|
||
electron microscope images of demineralized MB [(A) to (D)] and CB [(E) to (K)]. Demineralized, aldehyde-fixed (14) MB tissues from (A)
|
||
<materialsCitation id="3183EA71FFA3FF98F7B731A929D3FE4C" ID-GBIF-Occurrence="2979293314" box="[2065,2237,400,427]" collectionCode="MOR" pageId="4" pageNumber="1459" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, (
|
||
<emphasis id="B39F3C3EFFA3FF98F74331A9286AFE4C" bold="true" box="[2277,2308,400,427]" pageId="4" pageNumber="1459">B)</emphasis>
|
||
extant laying hen, (
|
||
<emphasis id="B39F3C3EFFA3FF98F743318C286AFE37" bold="true" box="[2277,2308,437,464]" pageId="4" pageNumber="1459">C)</emphasis>
|
||
emu, and (
|
||
<emphasis id="B39F3C3EFFA3FF98F7D231E329E4FE12" bold="true" box="[2164,2186,474,501]" pageId="4" pageNumber="1459">D</emphasis>
|
||
) ostrich show random, crumbly texture. Organized collagen fiber bundles are not distinct in any sample because of rapid deposition and woven character. Scale bars for (A) and (B), 40 um; for (C) and (D), 20 um. Demineralized
|
||
<emphasis id="B39F3C3EFFA3FF98F892336926D5FC8C" box="[1844,1979,848,875]" italics="true" pageId="4" pageNumber="1459">fragments</emphasis>
|
||
of cortical bone from
|
||
<emphasis id="B39F3C3EFFA3FF98F74433692987FC8C" bold="true" box="[2274,2281,848,875]" pageId="4" pageNumber="1459">(</emphasis>
|
||
E
|
||
<emphasis id="B39F3C3EFFA3FF98F75D3369286CFC8C" bold="true" box="[2299,2306,848,875]" pageId="4" pageNumber="1459">)</emphasis>
|
||
<materialsCitation id="3183EA71FFA3FF98F9AA334C27CAFC77" ID-GBIF-Occurrence="2979293311" box="[1548,1700,885,912]" collectionCode="MOR" pageId="4" pageNumber="1459" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, (
|
||
<emphasis id="B39F3C3EFFA3FF98F960334C27BAFC77" bold="true" box="[1734,1748,885,912]" pageId="4" pageNumber="1459">F</emphasis>
|
||
) chicken, (
|
||
<emphasis id="B39F3C3EFFA3FF98F8D7334C26E6FC77" box="[1905,1928,885,912]" italics="true" pageId="4" pageNumber="1459">G</emphasis>
|
||
) emu, and (H) ostrich are shown. A fibrous character dominates all samples. Scale bars for (E), (F), and (H), 30
|
||
<emphasis id="B39F3C3EFFA3FF98F836338626CAFC3A" box="[1936,1956,959,989]" italics="true" pageId="4" pageNumber="1459">m</emphasis>
|
||
m; for (G), 10
|
||
<emphasis id="B39F3C3EFFA3FF98F7CA338629EEFC3A" box="[2156,2176,959,989]" italics="true" pageId="4" pageNumber="1459">m</emphasis>
|
||
m. Higher magnification of demineralized CB from (
|
||
<emphasis id="B39F3C3EFFA3FF98F78D33DC295FFBE7" bold="true" box="[2091,2097,997,1024]" pageId="4" pageNumber="1459">I</emphasis>
|
||
)
|
||
<materialsCitation id="3183EA71FFA3FF98F7E133DC29B8FBE7" ID-GBIF-Occurrence="2979293312" box="[2119,2262,997,1024]" collectionCode="MOR" pageId="4" pageNumber="1459" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
, (J) emu, and (K) ostrich CB demonstrates the structural similarity between samples, although the
|
||
<materialsCitation id="3183EA71FFA3FF98F7CF3408286DFBAB" ID-GBIF-Occurrence="2979293315" box="[2153,2307,1073,1100]" collectionCode="MOR" pageId="4" pageNumber="1459" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
matrix is highly degraded. Scale bars for (I) and (K), 6
|
||
<emphasis id="B39F3C3EFFA3FF98F776346D2990FB95" box="[2256,2302,1108,1138]" italics="true" pageId="4" pageNumber="1459">um</emphasis>
|
||
; for (J), 5
|
||
<emphasis id="B39F3C3EFFA3FF98F920344327DBFB7F" box="[1670,1717,1146,1176]" italics="true" pageId="4" pageNumber="1459">um</emphasis>
|
||
.
|
||
</paragraph>
|
||
</caption>
|
||
<paragraph id="8154E02CFFA3FF98FF613AD22461F837" blockId="4.[150,831,1269,2958]" lastBlockId="4.[888,1570,1269,2000]" pageId="4" pageNumber="1459">
|
||
MB most likely first evolved within the lineage in early, small theropods with high productivity. A relatively thicker tyrannosaur bone cortex would reduce the need for MB, and its presence in
|
||
<materialsCitation id="3183EA71FFA3FF98FB1434CF240CFAF1" ID-GBIF-Occurrence="2979293321" box="[1202,1378,1270,1302]" collectionCode="MOR" pageId="4" pageNumber="1459" specimenCode="MOR 1125">MOR 1125</materialsCitation>
|
||
may reflect the retention of a primitive trait. This hypothesis may be tested by examination of the limb bones of the recently reported oviraptor containing eggs in the reproductive tract (
|
||
<emphasis id="B39F3C3EFFA3FF98FA41359D2762FA23" box="[1511,1548,1444,1476]" italics="true" pageId="4" pageNumber="1459">32</emphasis>
|
||
). The existence of MB in crocodiles has been referred to anecdotally (
|
||
<emphasis id="B39F3C3EFFA3FF98FAE335C22437F9FC" box="[1349,1369,1531,1563]" italics="true" pageId="4" pageNumber="1459">3</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA3FF98FAD435C224F8F9FC" box="[1394,1430,1531,1563]" italics="true" pageId="4" pageNumber="1459">21</emphasis>
|
||
), but although they do resorb CB during shelling, experimental evidence suggests that they do not form MB (
|
||
<emphasis id="B39F3C3EFFA3FF98FBFB3646251FF978" box="[1117,1137,1663,1695]" italics="true" pageId="4" pageNumber="1459">6</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA3FF98FB20364625F4F978" box="[1158,1178,1663,1695]" italics="true" pageId="4" pageNumber="1459">9</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA3FF98FB17364625BBF978" box="[1201,1237,1663,1695]" italics="true" pageId="4" pageNumber="1459">11</emphasis>
|
||
,
|
||
<emphasis id="B39F3C3EFFA3FF98FB4A3646247EF978" box="[1260,1296,1663,1695]" italics="true" pageId="4" pageNumber="1459">12</emphasis>
|
||
), even after stimulation with estrogen (
|
||
<emphasis id="B39F3C3EFFA3FF98FB693693259BF92D" box="[1231,1269,1706,1738]" italics="true" pageId="4" pageNumber="1459">33</emphasis>
|
||
). The identification of medullary tissues in dinosaurs supports a closer relationship to birds than to other extant archosaurs, sheds light on reproductive strategies of nonavian theropods, and provides an objective means of gender determination in extinct dinosaurs.
|
||
</paragraph>
|
||
</subSubSection>
|
||
</treatment>
|
||
</document> |