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<mods:title id="28E8BBF0DE86B69D758F992729A451D1">Neurogenesis in directly and indirectly developing enteropneusts: of nets and cords</mods:title>
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<mods:namePart id="3CCCABFE589AF45478CE0F5BD55FB19D">Wanninger, Sabrina Kaul-Strehlow Makoto Urata Takuya Minokawa Thomas Stach Andreas</mods:namePart>
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<mods:title id="F22E924A930BA5174DA2C48B1D1D5B1E">Organisms Diversity & Evolution</mods:title>
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<treatment id="AB59ED17564AFFB6FCAEFB58FE63FED8" LSID="urn:lsid:plazi:treatment:AB59ED17564AFFB6FCAEFB58FE63FED8" httpUri="http://treatment.plazi.org/id/AB59ED17564AFFB6FCAEFB58FE63FED8" lastPageId="11" lastPageNumber="416" pageId="7" pageNumber="412">
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<subSubSection id="6BEA0F8A564AFFBAFCAEFB58FBDDF9CE" pageId="7" pageNumber="412" type="nomenclature">
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<paragraph id="234F5C01564AFFBAFCAEFB58FB1EFB38" blockId="7.[816,1272,1189,1212]" box="[816,1272,1189,1212]" pageId="7" pageNumber="412">
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<heading id="7807EB6D564AFFBAFCAEFB58FB1EFB38" box="[816,1272,1189,1212]" fontSize="10" level="2" pageId="7" pageNumber="412" reason="7">
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<taxonomicName id="E4F02782564AFFBAFCAEFB58FBDCFB38" ID-CoL="4TX3B" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[816,1082,1189,1212]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="7" pageNumber="412" phylum="Hemichordata" rank="species" species="kowalevskii">
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<emphasis id="11848013564AFFBAFCAEFB58FBDCFB38" box="[816,1082,1189,1212]" italics="true" pageId="7" pageNumber="412">Saccoglossus kowalevskii</emphasis>
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</taxonomicName>
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: dorsal kink stage
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</heading>
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</paragraph>
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<paragraph id="234F5C01564AFFBAFCAEFB1AFBDDF9CE" blockId="7.[816,1452,1255,1909]" pageId="7" pageNumber="412">
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Embryos at this stage are of elongate shape measuring approximately 700 μm in maximum length (
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<figureCitation id="BBCB4084564AFFBAFB59FAF5FAF4FA9B" box="[1223,1298,1288,1311]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 6a</figureCitation>
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). The anterior proboscis region is of ovoid shape with a tapering anterior tip and is subdivided from the posterior body regions by a deep circular constriction (
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<figureCitation id="BBCB4084564AFFBAFB92FA91FB89FA07" box="[1036,1135,1388,1411]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 6a, d</figureCitation>
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). The proboscis region is covered by short cilia, whereas at the anterior tip, a ciliary tuft composed of long cilia is present (
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<figureCitation id="BBCB4084564AFFBAFB0EFA53FAEFFA41" box="[1168,1289,1454,1477]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 6a, c, d</figureCitation>
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). A second, yet shallow circular groove is discernable within the prospective collar region (
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<figureCitation id="BBCB4084564AFFBAFC5EFA0CFBECF98C" box="[960,1034,1521,1544]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 6d</figureCitation>
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). The opisthotroch is composed of compound cilia of multiciliary cells and demarcates the posterior perianal field (
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<figureCitation id="BBCB4084564AFFBAFC56F9CEFBCDF9CE" box="[968,1067,1587,1610]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 6a, d</figureCitation>
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).
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</paragraph>
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</subSubSection>
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<subSubSection id="6BEA0F8A564AFFB6FCD1F9A9FE63FED8" lastPageId="11" lastPageNumber="416" pageId="7" pageNumber="412" type="description">
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<paragraph id="234F5C01564AFFB5FCD1F9A9FD4BF8F1" blockId="7.[816,1452,1255,1909]" lastBlockId="8.[136,771,1687,1909]" lastPageId="8" lastPageNumber="413" pageId="7" pageNumber="412">
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Serotonin-LIR reveals the presence of several bipolar neurons that are located circumferentially around the midlevel of the proboscis region (
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<figureCitation id="BBCB4084564AFFBAFB32F96AFB1DF92A" box="[1196,1275,1687,1710]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 6a</figureCitation>
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). These bipolar neurons project a proximal neurite into a basiepidermal plexus of the proboscis region. A distal neurite connects to the apical cell surface where a single cilium is present. α- Tubulin-LIR reveals an apical ciliary tuft at the anterior tip; yet, no serotonin-LIR somata are detectable within the apical region close to the apical tuft (
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<figureCitation id="BBCB40845645FFB5FF0EF96AFF1BF92A" box="[144,253,1687,1710]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="8" pageNumber="413">Fig. 6a, c</figureCitation>
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). Throughout the prospective collar and trunk region, basiepidermal serotonin-LIR neurites form a loose network that thins out posteriorly into individual neurites within the perianal field (
|
||||
<figureCitation id="BBCB40845645FFB5FD94F907FDBAF895" box="[522,604,1786,1809]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="8" pageNumber="413">Fig. 6b</figureCitation>
|
||||
). No traces of a serotonin-LIR opisthotroch neurite bundle are present within the trunk region, although tubulin staining reveals the existence of an opisthotroch (
|
||||
<figureCitation id="BBCB40845645FFB5FDD2F8A3FD7AF8F1" box="[588,668,1886,1909]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="8" pageNumber="413">Fig. 6a</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015645FFB5FCAEF96AFB71F92A" blockId="8.[816,1175,1687,1710]" box="[816,1175,1687,1710]" pageId="8" pageNumber="413">
|
||||
<taxonomicName id="E4F027825645FFB5FCAEF96AFC25F92A" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[816,963,1687,1710]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="8" pageNumber="413" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="118480135645FFB5FCAEF96AFC25F92A" box="[816,963,1687,1710]" italics="true" pageId="8" pageNumber="413">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
: 1-gill-slit hatchling
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015645FFB4FCAEF924FE7CFE9E" blockId="8.[816,1452,1753,1909]" lastBlockId="9.[136,771,159,1611]" lastPageId="9" lastPageNumber="414" pageId="8" pageNumber="413">
|
||||
The 1-gill-slit hatchling measures between 800 μm and
|
||||
<quantity id="E408F1E45645FFB5FCAEF907FC95F895" box="[816,883,1786,1809]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="8" pageNumber="413" unit="mm" value="1.0">1 mm</quantity>
|
||||
in length when fully expanded (
|
||||
<figureCitation id="BBCB40845645FFB5FB6DF907FA83F895" box="[1267,1381,1786,1809]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="8" pageNumber="413">Fig. 6e, h</figureCitation>
|
||||
). The animals exhibit a conical proboscis with an apical tuft composed of long cilia at the anterior tip (
|
||||
<figureCitation id="BBCB40845645FFB5FAB5F8C0FA7DF8D0" box="[1323,1435,1853,1876]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="8" pageNumber="413">Fig. 6f, h</figureCitation>
|
||||
). The opisthotroch is present within the trunk region, just posterior to the position of the first pair of dorsolateral gill pores (
|
||||
<figureCitation id="BBCB40845644FFB4FE9DFF3DFEB5FF53" box="[259,339,192,215]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h</figureCitation>
|
||||
). The posterior trunk region including the perianal field is elongated ventrally to a total length of 150 μm (
|
||||
<figureCitation id="BBCB40845644FFB4FE86FEFEFE6CFE9E" box="[280,394,259,282]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6e, h</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<caption id="778F0C895644FFB4FCAEFF5EFAB5FCB1" pageId="9" pageNumber="414" startId="9.[816,848,163,183]" targetBox="[136,1452,144,1651]" targetPageId="10">
|
||||
<paragraph id="234F5C015644FFB4FCAEFF5EFA4AFF68" blockId="9.[816,1478,158,821]" pageId="9" pageNumber="414">
|
||||
<emphasis id="118480135644FFB4FCAEFF5EFC8EFF33" bold="true" box="[816,872,163,183]" pageId="9" pageNumber="414">Fig. 5</emphasis>
|
||||
Neurogenesis in the 2-gill slit juvenile of
|
||||
<taxonomicName id="E4F027825644FFB4FB69FF59FA9EFF33" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[1271,1400,163,183]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="118480135644FFB4FB69FF59FA9EFF33" box="[1271,1400,163,183]" italics="true" pageId="9" pageNumber="414">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
.
|
||||
<emphasis id="118480135644FFB4FA18FF5EFA4DFF33" bold="true" box="[1414,1451,163,183]" pageId="9" pageNumber="414">a, b</emphasis>
|
||||
Scanning electron micrographs:
|
||||
<emphasis id="118480135644FFB4FBE1FF43FB7AFF56" bold="true" box="[1151,1180,190,210]" pageId="9" pageNumber="414">c–i</emphasis>
|
||||
Z-projections of confocal microscopy image stacks. Anterior is to the right.
|
||||
<emphasis id="118480135644FFB4FB64FF25FAE3FF68" bold="true" box="[1274,1285,216,236]" pageId="9" pageNumber="414">a</emphasis>
|
||||
Dorsolateral view
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB4FCAEFF0EFA4DFE08" blockId="9.[816,1478,158,821]" pageId="9" pageNumber="414">
|
||||
of a 3-day-old settled juvenile.
|
||||
<emphasis id="118480135644FFB4FBD0FF0EFBBCFE83" bold="true" box="[1102,1114,243,263]" pageId="9" pageNumber="414">b</emphasis>
|
||||
Higher magnification of the two gill slits and the dorsal tongue bars.
|
||||
<emphasis id="118480135644FFB4FBD9FEF0FBB7FEA5" bold="true" box="[1095,1105,269,289]" pageId="9" pageNumber="414">c</emphasis>
|
||||
Overview of the 5-HT+ NS of a 3-day-old juvenile.
|
||||
<emphasis id="118480135644FFB4FC33FED5FC5FFEB8" bold="true" box="[941,953,296,316]" pageId="9" pageNumber="414">d</emphasis>
|
||||
The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord.
|
||||
<emphasis id="118480135644FFB4FC04FEA0FC42FEF5" bold="true" box="[922,932,349,369]" pageId="9" pageNumber="414">e</emphasis>
|
||||
Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons.
|
||||
<emphasis id="118480135644FFB4FAF4FE85FA97FE08" bold="true" box="[1386,1393,376,396]" pageId="9" pageNumber="414">f</emphasis>
|
||||
Detail
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB4FF38FED9FEB2FA07" blockId="9.[136,771,159,1611]" pageId="9" pageNumber="414">
|
||||
The serotonin-LIR nervous system comprises a welldeveloped basiepidermal plexus that extends throughout all three body regions (
|
||||
<figureCitation id="BBCB40845644FFB4FE0FFE9AFDE6FEFA" box="[401,512,359,382]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h, g</figureCitation>
|
||||
). Numerous serotoninLIR epithelial neurons are present throughout the epidermis of the proboscis. These bipolar neurons are evenly distributed except for the most anterior area around the apical ciliary tuft. Within a radius of approximately 65 μm from the central apical tuft, no serotonin-LIR somata are present (
|
||||
<figureCitation id="BBCB40845644FFB4FD8AFDD3FD64FDC1" box="[532,642,558,581]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6f, h</figureCitation>
|
||||
). A higher density of serotonin-LIR neurites characterizes a part of the proboscis plexus dorsally at the base of the proboscis (
|
||||
<figureCitation id="BBCB40845644FFB4FF27FD6CFEEDFD2C" box="[185,267,657,680]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h</figureCitation>
|
||||
). Within the collar epidermis, serotonin-LIR somata are concentrated into two multi-rowed rings, encircling the anterior and middle collar region (
|
||||
<figureCitation id="BBCB40845644FFB4FF0EFD08FEE6FC88" box="[144,256,757,780]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h, i</figureCitation>
|
||||
). The soma is located within the apical part of the epidermis, and each bipolar neuron sends a long proximal neurite into the basiepidermal plexus of the collar region. Detection of serotonin-LIR in the collar region reveals a neurite bundle that passes through the neurulating collar cord (
|
||||
<figureCitation id="BBCB40845644FFB4FE32FC66FE1BFC36" box="[428,509,923,946]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6i</figureCitation>
|
||||
). While the posterior part is already clearly in a subepidermal position, the more anterior part shows dorsal connections to the basiepidermal plexus within the collar epidermis (
|
||||
<figureCitation id="BBCB40845644FFB4FF0EFBDDFF39FBB3" box="[144,223,1056,1079]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6i</figureCitation>
|
||||
). The collar cord in
|
||||
<taxonomicName id="E4F027825644FFB4FE7BFBDDFD60FBB3" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[485,646,1056,1079]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="118480135644FFB4FE7BFBDDFD60FBB3" box="[485,646,1056,1079]" italics="true" pageId="9" pageNumber="414">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
neurulates successively from posterior to anterior, leaving just a neuropore at the posterior and anterior margin of the collar region (for more details, see
|
||||
<bibRefCitation id="476121F05644FFB4FDD9FB7EFF58FB38" author="Kaul, S. & Stach, T." pageId="9" pageNumber="414" pagination="1240 - 1259" refId="ref13149" refString="Kaul, S., & Stach, T. (2010). Ontogeny of the collar cord: Neurulation in the hemichordate Saccoglossus kowalevskii. Journal of Morphology, 271, 1240 - 1259." type="journal article" year="2010">Kaul and Stach 2010</bibRefCitation>
|
||||
). Tubulin-LIR reveals the opisthotroch within the trunk region that propels the living hatchlings through the water (
|
||||
<figureCitation id="BBCB40845644FFB4FE8CFB1AFE8AFB7A" box="[274,364,1255,1278]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h</figureCitation>
|
||||
). No serotonin-LIR opisthotroch neurite ring is present (
|
||||
<figureCitation id="BBCB40845644FFB4FE38FAF5FDC7FA9B" box="[422,545,1288,1311]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h, g</figureCitation>
|
||||
). Serotonin-LIR at the ventroposterior pole of the trunk region shows the presence of approximately 10 to 15 epithelial bipolar neurons (
|
||||
<figureCitation id="BBCB40845644FFB4FF6CFA91FEA2FA07" box="[242,324,1388,1411]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6h</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB4FCAEFE6FFAB5FCB1" blockId="9.[816,1478,158,821]" pageId="9" pageNumber="414">
|
||||
of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in
|
||||
<emphasis id="118480135644FFB4FBAFFE3AFBDAFE5F" bold="true" box="[1073,1084,455,475]" pageId="9" pageNumber="414">g</emphasis>
|
||||
, see also
|
||||
<emphasis id="118480135644FFB4FB0AFE3AFCB1FE71" italics="true" pageId="9" pageNumber="414">double arrowheads and dashed area</emphasis>
|
||||
.
|
||||
<emphasis id="118480135644FFB4FCFCFE1FFC8BFE72" bold="true" box="[866,877,482,502]" pageId="9" pageNumber="414">g</emphasis>
|
||||
The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see
|
||||
<emphasis id="118480135644FFB4FC62FDEAFADAFDAE" box="[1020,1340,535,555]" italics="true" pageId="9" pageNumber="414">double arrowheads and dashed area</emphasis>
|
||||
.
|
||||
<emphasis id="118480135644FFB4FAD9FDEAFAB5FDAF" bold="true" box="[1351,1363,535,555]" pageId="9" pageNumber="414">h</emphasis>
|
||||
Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive.
|
||||
<emphasis id="118480135644FFB4FBA0FD9AFBA2FDFF" bold="true" box="[1086,1092,615,635]" pageId="9" pageNumber="414">i</emphasis>
|
||||
Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α- tub, acetylated α- tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="234F5C015644FFB4FF38FA70FE05F9CE" blockId="9.[136,771,159,1611]" pageId="9" pageNumber="414">
|
||||
In transmission electron micrographs, numerous basiepidermal neurites of different diameters, exhibiting oblique to oval profiles, are present in 1-gill-slit hatchlings of
|
||||
<taxonomicName id="E4F027825644FFB4FF36FA0CFED1F98C" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[168,311,1521,1544]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="species" species="kowalevkii">
|
||||
<emphasis id="118480135644FFB4FF36FA0CFED1F98C" box="[168,311,1521,1544]" italics="true" pageId="9" pageNumber="414">S. kowalevkii</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
<figureCitation id="BBCB40845644FFB4FED7FA0CFE56F98C" box="[329,432,1520,1544]" captionStart="Fig" captionStartId="13.[136,170,1423,1443]" captionTargetBox="[137,1452,146,1399]" captionTargetId="figure-155@13.[136,1452,144,1404]" captionTargetPageId="13" captionText="Fig. 7 Ultrastructural details of the developing nervous system in Saccoglossus kowalevskii. a Sagittal section of a 1-gill-slit hatchling of S. kowalevskii. b Ultrastructural detail of the red-marked box in a showing a continuous layer of neurites of 2–4-μm thickness. c Colorcoded image of the micrograph shown in b. d Sagittal section of a 2- gill-slit juvenile of S. kowalevskii. e Ultrastructural detail of the redmarked box in c showing only single and scattered, small neurite" figureDoi="http://doi.org/10.5281/zenodo.12764150" httpUri="https://zenodo.org/record/12764150/files/figure.png" pageId="9" pageNumber="414">Fig. 7a–c</figureCitation>
|
||||
). The high number of neurites composes a continuous and circumferential neurite layer of 2- to 4-μm thickness (
|
||||
<figureCitation id="BBCB40845644FFB4FEF1F9CEFE35F9CE" box="[367,467,1587,1610]" captionStart="Fig" captionStartId="13.[136,170,1423,1443]" captionTargetBox="[137,1452,146,1399]" captionTargetId="figure-155@13.[136,1452,144,1404]" captionTargetPageId="13" captionText="Fig. 7 Ultrastructural details of the developing nervous system in Saccoglossus kowalevskii. a Sagittal section of a 1-gill-slit hatchling of S. kowalevskii. b Ultrastructural detail of the red-marked box in a showing a continuous layer of neurites of 2–4-μm thickness. c Colorcoded image of the micrograph shown in b. d Sagittal section of a 2- gill-slit juvenile of S. kowalevskii. e Ultrastructural detail of the redmarked box in c showing only single and scattered, small neurite" figureDoi="http://doi.org/10.5281/zenodo.12764150" httpUri="https://zenodo.org/record/12764150/files/figure.png" pageId="9" pageNumber="414">Fig. 7b, c</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB4FF16F98BFE07F909" blockId="9.[136,481,1654,1677]" box="[136,481,1654,1677]" pageId="9" pageNumber="414">
|
||||
<taxonomicName id="E4F027825644FFB4FF16F98BFEFDF909" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[136,283,1654,1677]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="118480135644FFB4FF16F98BFEFDF909" box="[136,283,1654,1677]" italics="true" pageId="9" pageNumber="414">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
: 3-gill-slit juvenile
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB4FF16F945FCE5F8F1" blockId="9.[136,771,1720,1909]" pageId="9" pageNumber="414">
|
||||
The juveniles at this stage of development resemble the adult acorn worms in many aspects (
|
||||
<figureCitation id="BBCB40845644FFB4FE5DF924FDC6F974" box="[451,544,1753,1776]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6k, l</figureCitation>
|
||||
). Obvious differences are a lower number of gill slits and the presence of the postanal tail, a feature only present in juvenile acorn worms of the family
|
||||
<taxonomicName id="E4F027825644FFB4FF42F8C0FE9BF8D0" authorityName="Spengel" authorityYear="1901" box="[220,381,1853,1876]" class="Enteropneusta" family="Harrimaniidae" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="family">Harrimaniidae</taxonomicName>
|
||||
(
|
||||
<bibRefCitation id="476121F05644FFB4FE0FF8C0FDDBF8D0" author="Cameron, C. B." box="[401,573,1853,1876]" pageId="9" pageNumber="414" pagination="196 - 215" refId="ref11902" refString="Cameron, C. B. (2005). A phylogeny of the hemichordates based on morphological characters. Canadian Journal of Zoology, 83, 196 - 215." type="journal article" year="2005">Cameron 2005</bibRefCitation>
|
||||
;
|
||||
<bibRefCitation id="476121F05644FFB4FDCEF8C0FF5AF8F1" author="Stach, T. & Kaul, S." pageId="9" pageNumber="414" pagination="150 - 160" refId="ref14441" refString="Stach, T., & Kaul, S. (2012). The postanal tail of the enteropneust Saccoglossus kowalevskii is a ciliarly creeping organ without distinct similarities to the chordate tail. A Zool, 92, 150 - 160." type="journal article" year="2012">Stach and Kaul 2012</bibRefCitation>
|
||||
). Scanning electron micrographs of the anterior tip of
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB4FCAEFC87FB44FC57" blockId="9.[816,1452,890,1909]" pageId="9" pageNumber="414">
|
||||
the proboscis show that the former ciliary tuft is degraded (
|
||||
<figureCitation id="BBCB40845644FFB4FCA6FC66FC73FC36" box="[824,917,923,946]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6k, l</figureCitation>
|
||||
). The opisthotroch is partially altered and has transformed into a ventral creeping sole.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015644FFB7FCD1FC20FABFF8F0" blockId="9.[816,1452,890,1909]" lastBlockId="10.[816,1452,1719,1908]" lastPageId="10" lastPageNumber="415" pageId="9" pageNumber="414">
|
||||
Within the proboscis and collar region, a serotonin-LIR basiepidermal plexus is present. The concentration of neurites is denser at the dorsoposterior base of the proboscis, constituting the proboscis plexus (
|
||||
<figureCitation id="BBCB40845644FFB4FA85FBBCFA7EFBDC" box="[1307,1432,1089,1112]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6k, o</figureCitation>
|
||||
). Serotonin-LIR bipolar neurons are distributed throughout the epidermis of the proboscis with the exception of the most anterior region. Each soma has a bulbous central part containing the nucleus and a slender distal neurite reaching the apical cell surface, where a single cilium is present (
|
||||
<figureCitation id="BBCB40845644FFB4FCA6FAF5FC61FA9B" box="[824,903,1288,1311]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6n</figureCitation>
|
||||
). Basally from the bulbous part, each soma sends a proximal neurite into the basiepidermal plexus. The dorsal proboscis stem sends two serotonin-LIR neurite bundles posteriorly into the subepidermal collar cord, thereby passing the proboscis neck region. At the posterior end of the collar region, the serotonin-LIR becomes superficial again and continues into a dorsal neurite bundle (
|
||||
<figureCitation id="BBCB40845644FFB4FCA6FA0CFC49F98C" box="[824,943,1521,1544]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">
|
||||
Fig.
|
||||
<quantity id="E408F1E45644FFB4FCF1FA0CFC75F98C" box="[879,915,1521,1544]" metricMagnitude="0" metricUnit="m" metricValue="6.0" pageId="9" pageNumber="414" unit="m" value="6.0">6m</quantity>
|
||||
, o
|
||||
</figureCitation>
|
||||
). Unlike
|
||||
<taxonomicName id="E4F027825644FFB4FB80FA0CFB5DF98C" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[1054,1211,1521,1544]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="118480135644FFB4FB80FA0CFB5DF98C" box="[1054,1211,1521,1544]" italics="true" pageId="9" pageNumber="414">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
,
|
||||
<taxonomicName id="E4F027825644FFB4FB53FA0CFA8FF98C" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[1229,1385,1521,1544]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="9" pageNumber="414" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="118480135644FFB4FB53FA0CFA8FF98C" box="[1229,1385,1521,1544]" italics="true" pageId="9" pageNumber="414">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
lacks mesocoelic pores as well as a prebranchial nerve ring at this stage of development. A vast number of serotonin-LIR bipolar neurons are present within the epidermis of the anterior half of the collar region (
|
||||
<figureCitation id="BBCB40845644FFB4FB05F98BFAE3F909" box="[1179,1285,1654,1677]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6k, o</figureCitation>
|
||||
). The architecture of the serotonin-LIR nervous system within the trunk region differs considerably from that described for the preceding hatchling stage. Serotonin-LIR is present in a longitudinal ventral as well as dorsal neurite bundle along the trunk region (
|
||||
<figureCitation id="BBCB40845644FFB4FC56F8E1FBDAF8B7" box="[968,1084,1820,1843]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="9" pageNumber="414">Fig. 6k, m</figureCitation>
|
||||
). The ventral bundle is more extensive and extends to the posterior end of the postanal tail. The dorsal serotonin-LIR neurite bundle is less voluminous and is composed of few neurites only. It runs along the dorsal midline to the anus, with an interruption of the serotonin-LIR signal in the middle region of the trunk (
|
||||
<figureCitation id="BBCB40845647FFB7FF0EF8E6FEEFF8B6" box="[144,265,1819,1842]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="10" pageNumber="415">Fig. 6k, m</figureCitation>
|
||||
). Several serotonin-LIR bipolar neurons are present within the trunk region (
|
||||
<figureCitation id="BBCB40845647FFB7FE74F8C1FDA6F8D7" box="[490,576,1852,1875]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="10" pageNumber="415">
|
||||
Fig.
|
||||
<quantity id="E408F1E45647FFB7FD80F8C1FDA6F8D7" box="[542,576,1852,1875]" metricMagnitude="0" metricUnit="m" metricValue="6.0" pageId="10" pageNumber="415" unit="m" value="6.0">6m</quantity>
|
||||
</figureCitation>
|
||||
). The majority of the somata are located dorsolaterally, sending a proximal neurite into the ventral nerve cord (
|
||||
<figureCitation id="BBCB40845647FFB7FB26F94AFAE9F94A" box="[1208,1295,1719,1742]" captionStart="Fig" captionStartId="11.[816,848,163,183]" captionTargetId="figure-71@12.[136,1452,144,1650]" captionTargetPageId="12" captionText="Fig. 6 Neurogenesis in S. kowalevskii. a–C, f–k, m–o Z-projections of confocal microscopy image stacks. d, e, l Scanning electron micrographs (SEM). Anterior is to the right and ventral to the bottom in all images. a Overview of a dorsal kink stage. Serotonin-LIR neurons are present in the proboscis epidermis. Note the larval apical ciliary tuft. b Detail showing the developing nervous plexus in the trunk region of a dorsal kink stage. c Partial Z-projection of the anterior tip of a dorsal kink stage highlighting the absence of serotonin-LIR somata from the apical plate. d SEM of dorsal kink stage. e SEM of 1-gill-slit stage. f Partial Z-projection of the anterior tip of a 1-gill-slit stage. g Detail showing an elaborate 5-HT+ nervous plexus in the trunk region of the 1-gill-slit stage. h Overview of a 1-gill-slit stage showing the entire serotonin-LIR nervous system. i Close-up of the dorsal collar region demonstrating the 5-HT+ part in the neurulating collar cord. Double arrowheads point to dorsal connections of the anterior portion of the collar cord that is still in contact with the epidermis. k Overview of the serotonin-LIR nervous system of a 3-gill-slit juvenile. l SEM of a 3-gill-slit juvenile in lateral view. m Detail of the anterior trunk region of a 3-gill-slit juvenile showing scattered serotoninLIR bipolar neurons projecting into the ventral neurite bundle. n Detail of two biploar neurons within the proboscis epidermis. o Detail of the collar region showing 5-HT+ neurites within the subepidermal collar cord. 5- HT, serotonin; ac-α-tub, acetylated α-tubulin; at, apical tuft; cn, circumferential neurite; cc, collar cord; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; ms, mesocoel; nn, nervous plexus; ot, opisthotroch; pat, postanal tail; pc, protocoel; pf, perianal field; ph, pharynx; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="10" pageNumber="415">
|
||||
Fig.
|
||||
<quantity id="E408F1E45647FFB7FB72F94AFAE9F94A" box="[1260,1295,1719,1742]" metricMagnitude="0" metricUnit="m" metricValue="6.0" pageId="10" pageNumber="415" unit="m" value="6.0">6m</quantity>
|
||||
</figureCitation>
|
||||
). The number of serotonin-LIR somata in the trunk region is particularly low compared to the collar and proboscis region. Between the gill pores, a higher density of serotonin-LIR neurons is present. Notably, serotonin-LIR does not show a plexus-like pattern in the trunk region as in earlier stages.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015646FFB6FF38FF62FE63FED8" blockId="11.[136,771,159,348]" pageId="11" pageNumber="416">
|
||||
Ultrastructural investigations of transmission electron micrographs of a 2-gill-slit juvenile show that only scattered neurites are present below the one-layered epidermis (
|
||||
<figureCitation id="BBCB40845646FFB6FF78FEFEFEB6FE9E" box="[230,336,258,282]" captionStart="Fig" captionStartId="13.[136,170,1423,1443]" captionTargetBox="[137,1452,146,1399]" captionTargetId="figure-155@13.[136,1452,144,1404]" captionTargetPageId="13" captionText="Fig. 7 Ultrastructural details of the developing nervous system in Saccoglossus kowalevskii. a Sagittal section of a 1-gill-slit hatchling of S. kowalevskii. b Ultrastructural detail of the red-marked box in a showing a continuous layer of neurites of 2–4-μm thickness. c Colorcoded image of the micrograph shown in b. d Sagittal section of a 2- gill-slit juvenile of S. kowalevskii. e Ultrastructural detail of the redmarked box in c showing only single and scattered, small neurite" figureDoi="http://doi.org/10.5281/zenodo.12764150" httpUri="https://zenodo.org/record/12764150/files/figure.png" pageId="11" pageNumber="416">Fig. 7d–f</figureCitation>
|
||||
). The majority of neurites is concentrated within the dorsal and ventral nerve cord (read also
|
||||
<bibRefCitation id="476121F05646FFB6FF16FEB8FE93FED8" author="Kaul, S. & Stach, T." box="[136,373,325,348]" pageId="11" pageNumber="416" pagination="1240 - 1259" refId="ref13149" refString="Kaul, S., & Stach, T. (2010). Ontogeny of the collar cord: Neurulation in the hemichordate Saccoglossus kowalevskii. Journal of Morphology, 271, 1240 - 1259." type="journal article" year="2010">Kaul and Stach 2010</bibRefCitation>
|
||||
).
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
413
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413
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<treatment id="AB59ED17564EFFBAFCAEFE75FAE1FBF3" LSID="urn:lsid:plazi:treatment:AB59ED17564EFFBAFCAEFE75FAE1FBF3" httpUri="http://treatment.plazi.org/id/AB59ED17564EFFBAFCAEFE75FAE1FBF3" lastPageId="7" lastPageNumber="412" pageId="3" pageNumber="408">
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<heading id="7807EB6D564EFFBEFCAEFE75FB05FE1B" box="[816,1251,392,415]" fontSize="10" level="2" pageId="3" pageNumber="408" reason="7">
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<taxonomicName id="E4F02782564EFFBEFCAEFE75FBA0FE1B" ID-CoL="KFVG" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[816,1094,392,415]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="3" pageNumber="408" phylum="Hemichordata" rank="species" species="misakiensis">
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<emphasis id="11848013564EFFBEFCAEFE75FBA0FE1B" box="[816,1094,392,415]" italics="true" pageId="3" pageNumber="408">Balanoglossus misakiensis</emphasis>
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</taxonomicName>
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: Spengel stage
|
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</heading>
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</paragraph>
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</subSubSection>
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<subSubSection id="6BEA0F8A564EFFBAFCAEFE37FAE1FBF3" lastPageId="7" lastPageNumber="412" pageId="3" pageNumber="408" type="description">
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<paragraph id="234F5C01564EFFBEFCAEFE37FBEDFCF4" blockId="3.[816,1453,458,1677]" pageId="3" pageNumber="408">
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After about 13 days postfertilization (pf), the tornaria larvae start transforming into the Spengel stage accompanied by a series of morphological changes. The larvae are approximately 950 μm in length and are subdivided by a deep circular constriction into a slightly conical preoral region forming the future proboscis and a voluminous spherical posterior region bearing the distinct opisthotroch (
|
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<figureCitation id="BBCB4084564EFFBEFB13FD6CFB16FD2C" box="[1165,1264,657,680]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2a, b</figureCitation>
|
||||
). The oral field is almost completely obliterated as the ciliary bands of the neotroch are fused together. The position of the ciliary bands is still indicated by grooves (
|
||||
<figureCitation id="BBCB4084564EFFBEFBFBFD08FB56FC88" box="[1125,1200,757,780]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2a</figureCitation>
|
||||
). A pair of dorsolateral slit-like depressions on the anterior part of the postoral body indicates the position of the prospective first gill pores (
|
||||
<figureCitation id="BBCB4084564EFFBEFCA6FCA4FC67FCF4" box="[824,897,857,880]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2a</figureCitation>
|
||||
, arrowhead).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564EFFBEFCD1FC87FB90F909" blockId="3.[816,1453,458,1677]" pageId="3" pageNumber="408">
|
||||
The entire larva is covered by short cilia. An apical tuft composed of longer cilia is present at the apical tip (
|
||||
<figureCitation id="BBCB4084564EFFBEFACCFC66FA7AFC36" box="[1362,1436,923,946]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2b</figureCitation>
|
||||
). Within the apical epidermis, numerous serotonin-LIR neurons are grouped together to form an anterior and a posterior cluster of cells which are interconnected by a median neuropil (
|
||||
<figureCitation id="BBCB4084564EFFBEFCA6FBDDFC67FBB3" box="[824,897,1056,1079]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2f</figureCitation>
|
||||
). The serotonin-LIR somata are slender, sometimes flask-shaped, cells with a nucleus that is positioned centrally or basally within the cytoplasm. Each soma is connected to the apical cell surface by a narrow elongation, the distal neurite (
|
||||
<figureCitation id="BBCB4084564EFFBEFCA6FB58FC9AFB38" box="[824,892,1189,1212]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2f</figureCitation>
|
||||
). The serotonin-LIR cells are bipolar neurons that send a neurite into the median neuropil of the apical organ. Aside from the cells of the apical organ, numerous serotonin-LIR somata are scattered throughout the proboscis’ epidermis. Each bipolar neuron exhibits a distal neurite connecting to the apical surface and a proximal neurite. The proximal neurites of those bipolar neurons form a basiepidermal plexus-like arrangement throughout the proboscis epidermis. Within the voluminous postoral region, serotonin-LIR somata are only occasionally present. A conspicuous number of neurites are present at the level of the opisthotroch and form a distinct neurite bundle, namely, the opisthotroch neurite ring (
|
||||
<figureCitation id="BBCB4084564EFFBEFCA6F9CEFC7CF9CE" box="[824,922,1587,1610]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2b, e</figureCitation>
|
||||
). Aside from that neurite bundle, the postoral body shows only individual neurites arranged in a loose, sometimes discontinuous pattern (
|
||||
<figureCitation id="BBCB4084564EFFBEFB83F98BFB80F909" box="[1053,1126,1654,1677]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2e</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564EFFBEFCAEF945FB86F94B" blockId="3.[816,1120,1720,1743]" box="[816,1120,1720,1743]" pageId="3" pageNumber="408">
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<taxonomicName id="E4F02782564EFFBEFCAEF945FC23F94B" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[816,965,1720,1743]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="3" pageNumber="408" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564EFFBEFCAEF945FC23F94B" box="[816,965,1720,1743]" italics="true" pageId="3" pageNumber="408">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
: Agassiz stage
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564EFFBEFCAEF907FA4DF8F1" blockId="3.[816,1452,1786,1909]" pageId="3" pageNumber="408">
|
||||
The larvae of the Agassiz stage (14 days pf) are of elongated shape measuring approximately
|
||||
<quantity id="E408F1E4564EFFBEFB1CF8E1FB26F8B7" box="[1154,1216,1820,1843]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="3" pageNumber="408" unit="mm" value="1.0">1 mm</quantity>
|
||||
in length and 650 μm in width. The anterior proboscis is of conical acorn shape (
|
||||
<figureCitation id="BBCB4084564EFFBEFCA6F8A3FC7AF8F1" box="[824,924,1886,1909]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="3" pageNumber="408">Fig. 2c, d</figureCitation>
|
||||
). It is separated from the posterior body by a deep
|
||||
</paragraph>
|
||||
<caption id="778F0C895649FFB9FF16F9C8FCE5F8D6" ID-DOI="http://doi.org/10.5281/zenodo.12764146" ID-Zenodo-Dep="12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="4" pageNumber="409" startId="4.[136,168,1589,1609]" targetBox="[136,1452,144,1569]" targetPageId="4" targetType="figure">
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<paragraph id="234F5C015649FFB9FF16F9C8FCE5F8D6" blockId="4.[136,772,1589,1875]" pageId="4" pageNumber="409">
|
||||
<emphasis id="118480135649FFB9FF16F9C8FF58F9CD" bold="true" box="[136,190,1589,1609]" pageId="4" pageNumber="409">Fig. 2</emphasis>
|
||||
Neurogenesis in metamorphosing stages of
|
||||
<taxonomicName id="E4F027825649FFB9FDCFF9CBFD28F9CD" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[593,718,1589,1609]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="4" pageNumber="409" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="118480135649FFB9FDCFF9CBFD28F9CD" box="[593,718,1589,1609]" italics="true" pageId="4" pageNumber="409">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
.
|
||||
<emphasis id="118480135649FFB9FD45F9C8FD01F9CD" bold="true" box="[731,743,1589,1609]" pageId="4" pageNumber="409">b</emphasis>
|
||||
,
|
||||
<emphasis id="118480135649FFB9FD6AF9C8FD18F9CD" bold="true" box="[756,766,1589,1609]" pageId="4" pageNumber="409">c</emphasis>
|
||||
,
|
||||
<emphasis id="118480135649FFB9FF16F9B2FF4FF9E7" bold="true" box="[136,169,1615,1635]" pageId="4" pageNumber="409">e–k</emphasis>
|
||||
Z-projections of confocal microscopy image stacks.
|
||||
<emphasis id="118480135649FFB9FD1BF9B2FD76F9E7" bold="true" box="[645,656,1615,1635]" pageId="4" pageNumber="409">a</emphasis>
|
||||
,
|
||||
<emphasis id="118480135649FFB9FD00F9B2FD4CF9E7" bold="true" box="[670,682,1615,1635]" pageId="4" pageNumber="409">d</emphasis>
|
||||
Scanning electron micrographs. Anterior is to the
|
||||
<emphasis id="118480135649FFB9FE6FF996FDEBF9F9" box="[497,525,1643,1661]" italics="true" pageId="4" pageNumber="409">top</emphasis>
|
||||
.
|
||||
<emphasis id="118480135649FFB9FD84F997FDC3F9FA" bold="true" box="[538,549,1642,1662]" pageId="4" pageNumber="409">a</emphasis>
|
||||
SEM of early metamorphosing stage (Spengel) from lateral left.
|
||||
<emphasis id="118480135649FFB9FE6CF979FDB2F91C" box="[498,596,1668,1688]" italics="true" pageId="4" pageNumber="409">Arrowhead</emphasis>
|
||||
points to the dorsolateral slit-like depression.
|
||||
<emphasis id="118480135649FFB9FEEFF962FE9BF937" bold="true" box="[369,381,1695,1715]" pageId="4" pageNumber="409">b</emphasis>
|
||||
Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from
|
||||
<emphasis id="118480135649FFB9FE6AF947FDFEF94A" box="[500,536,1722,1742]" italics="true" pageId="4" pageNumber="409">left.</emphasis>
|
||||
Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring.
|
||||
<emphasis id="118480135649FFB9FF69F912FEE7F887" bold="true" box="[247,257,1775,1795]" pageId="4" pageNumber="409">c</emphasis>
|
||||
Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view.
|
||||
<emphasis id="118480135649FFB9FE32F8F4FE5EF899" bold="true" box="[428,440,1801,1821]" pageId="4" pageNumber="409">d</emphasis>
|
||||
Dorsal view of metamorphosing Agassiz stage.
|
||||
<emphasis id="118480135649FFB9FF70F8D9FF1EF8BC" bold="true" box="[238,248,1828,1848]" pageId="4" pageNumber="409">e</emphasis>
|
||||
Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage.
|
||||
<emphasis id="118480135649FFB9FE5BF8C2FE2AF8D7" bold="true" box="[453,460,1855,1875]" pageId="4" pageNumber="409">f</emphasis>
|
||||
Close-up of the 5-HT+ apical organ
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="234F5C015649FFB9FCAEF9C8FABCF8D7" blockId="4.[816,1452,1589,1875]" pageId="4" pageNumber="409">
|
||||
of the Spengel stage in lateral view. Note the anterior and posterior cluster of bipolar neurons connected by a central neuropil.
|
||||
<emphasis id="118480135649FFB9FB6BF9B2FAE6F9E7" bold="true" box="[1269,1280,1615,1635]" pageId="4" pageNumber="409">g</emphasis>
|
||||
Detail showing the elaborated 5-HT+ nervous plexus in the prospective trunk region of an Agassiz stage larva.
|
||||
<emphasis id="118480135649FFB9FC77F979FC13F91C" bold="true" box="[1001,1013,1668,1688]" pageId="4" pageNumber="409">h</emphasis>
|
||||
Detail view of 5-HT+ apical bipolar neurons. A slender distal neurite connects the soma to the apical cell surface.
|
||||
<emphasis id="118480135649FFB9FAF5F962FA97F937" bold="true" box="[1387,1393,1695,1715]" pageId="4" pageNumber="409">i</emphasis>
|
||||
Detail of the neuite bundles passing the neck region, connecting the proboscis to the collar region.
|
||||
<emphasis id="118480135649FFB9FC50F929FC3CF96C" bold="true" box="[974,986,1748,1768]" pageId="4" pageNumber="409">k</emphasis>
|
||||
Close-up of the dorsal collar region showing the 5- HT+ bipolar neurons.5-HT, serotonin; ac-α- tub, acetylated α- tubulin; ao, apical organ; at, apical tuft; ci, cilia; co, collar; ne, neurite; nn, nervous plexus; np, neuropil; onr, opisthotroch nerve ring; ot, opisthotroch; pf, perianal field; pr, proboscis; sn, serotonin-LIR neuron; tr, trunk
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015648FFB8FF16FF62FD10FE1B" blockId="5.[136,771,159,548]" pageId="5" pageNumber="410">
|
||||
circular constriction (
|
||||
<figureCitation id="BBCB40845648FFB8FEF0FF62FE3EFF32" box="[366,472,159,182]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2c, d</figureCitation>
|
||||
). The grooves of the fused ciliary bands are no longer visible externally (compare
|
||||
<figureCitation id="BBCB40845648FFB8FF16FF1FFEFCFF7D" box="[136,282,226,249]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2a and d</figureCitation>
|
||||
). A second circular constriction indicates the posterior margin of the collar region (
|
||||
<figureCitation id="BBCB40845648FFB8FDB5FEFEFD9DFE9E" box="[555,635,259,282]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2d</figureCitation>
|
||||
). The collar region is about 150 μm long. The remaining posterior part of the body constitutes the future trunk region. It measures approximately 400 μm in total length and bears the opisthotroch at the midlevel of the trunk region (
|
||||
<figureCitation id="BBCB40845648FFB8FD1CFE75FD00FE1B" box="[642,742,392,415]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2c, d</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C015648FFB8FF38FE54FA4AFDA0" blockId="5.[136,771,159,548]" lastBlockId="5.[816,1452,159,548]" pageId="5" pageNumber="410">
|
||||
Serotonin-LIR is present at the anterior tip of the proboscis region (
|
||||
<figureCitation id="BBCB40845648FFB8FF45FE37FEC0FE65" box="[219,294,458,481]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2c</figureCitation>
|
||||
) and reveals numerous bipolar neurons with thin and long extensions to the cell surface where each neuron bears a single cilium (
|
||||
<figureCitation id="BBCB40845648FFB8FEF7FDF0FE54FDA0" box="[361,434,525,548]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2h</figureCitation>
|
||||
). The overall morphology of the serotonin-LIR bipolar neurons of the apical organ is comparable to that of the Spengel stage. Further individual serotoninLIR neurons are interspersed throughout the proboscis with a higher density present near the base of the proboscis (
|
||||
<figureCitation id="BBCB40845648FFB8FAC2FEFEFA42FE9E" box="[1372,1444,259,282]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2c</figureCitation>
|
||||
, sn). All serotonin-LIR somata project into a basiepidermal plexus that is present throughout the proboscis. The plexus of the proboscis region is connected to that of the collar region by two laterally condensed neurite bundles that pass through a neck-like transition into the collar region (
|
||||
<figureCitation id="BBCB40845648FFB8FB41FE54FADBFE44" box="[1247,1341,425,448]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2c, i</figureCitation>
|
||||
). A pair of bilateral muscle bundles that run anteriorly into the base of the proboscis accompanies the neurite bundles (
|
||||
<figureCitation id="BBCB40845648FFB8FA9FFE16FAA1FD86" box="[1281,1351,491,514]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="5" pageNumber="410">Fig. 2i</figureCitation>
|
||||
). Several serotonin-LIR bipolar neurons are present within the anterior
|
||||
</paragraph>
|
||||
<caption id="778F0C895648FFB8FF16F93CFCE5F8F0" ID-DOI="http://doi.org/10.5281/zenodo.12764148" ID-Zenodo-Dep="12764148" httpUri="https://zenodo.org/record/12764148/files/figure.png" pageId="5" pageNumber="410" startId="5.[136,171,1729,1749]" targetBox="[136,1452,598,1709]" targetPageId="5" targetType="figure">
|
||||
<paragraph id="234F5C015648FFB8FF16F93CFCE5F8F0" blockId="5.[136,773,1729,1908]" pageId="5" pageNumber="410">
|
||||
<emphasis id="118480135648FFB8FF16F93CFF22F951" bold="true" box="[136,196,1729,1749]" pageId="5" pageNumber="410">Fig. 3</emphasis>
|
||||
Ultrastructural details of the developing nervous system
|
||||
<taxonomicName id="E4F027825648FFB8FF16F921FE95F974" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[136,371,1756,1776]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="5" pageNumber="410" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="118480135648FFB8FF16F921FE95F974" box="[136,371,1756,1776]" italics="true" pageId="5" pageNumber="410">Balanoglossus misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
.
|
||||
<emphasis id="118480135648FFB8FE1EF921FE6DF974" bold="true" box="[384,395,1756,1776]" pageId="5" pageNumber="410">a</emphasis>
|
||||
Sagittal section of an Agassiz stage larva of
|
||||
<taxonomicName id="E4F027825648FFB8FF38F90AFED2F88E" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[166,308,1782,1802]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="5" pageNumber="410" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="118480135648FFB8FF38F90AFED2F88E" box="[166,308,1782,1802]" italics="true" pageId="5" pageNumber="410">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
.
|
||||
<emphasis id="118480135648FFB8FEDBF90BFEB7F88E" bold="true" box="[325,337,1782,1802]" pageId="5" pageNumber="410">b</emphasis>
|
||||
Sagittal section of a 2-gill-slit juvenile of
|
||||
<emphasis id="118480135648FFB8FF16F8EFFEEDF8A1" box="[136,267,1809,1829]" italics="true" pageId="5" pageNumber="410">
|
||||
<taxonomicName id="E4F027825648FFB8FF16F8EFFEE0F8A1" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[136,262,1809,1829]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="5" pageNumber="410" phylum="Hemichordata" rank="species" species="misakiensis">B. misakiensis</taxonomicName>
|
||||
.
|
||||
</emphasis>
|
||||
<emphasis id="118480135648FFB8FE93F8ECFEF1F8A1" bold="true" box="[269,279,1809,1829]" pageId="5" pageNumber="410">c</emphasis>
|
||||
Ultrastructural detail of the red-marked box in
|
||||
<emphasis id="118480135648FFB8FD37F8ECFD52F8A1" bold="true" box="[681,692,1809,1829]" pageId="5" pageNumber="410">a</emphasis>
|
||||
showing a continuous layer of neurites of 4–6-μm thickness in the Agassiz stage.
|
||||
<emphasis id="118480135648FFB8FF16F8BAFF54F8DD" box="[136,178,1863,1881]" italics="true" pageId="5" pageNumber="410">Inset</emphasis>
|
||||
color-coded image of the micrograph shown in
|
||||
<emphasis id="118480135648FFB8FDC3F8BBFD98F8DE" bold="true" box="[605,638,1862,1882]" pageId="5" pageNumber="410">c. d</emphasis>
|
||||
Ultrastructural detail of the red-marked box in
|
||||
<emphasis id="118480135648FFB8FE03F89DFE4FF8F0" bold="true" box="[413,425,1888,1908]" pageId="5" pageNumber="410">b</emphasis>
|
||||
showing only individual basiepidermal
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="234F5C015648FFB8FCAEF93CFB49F8DE" blockId="5.[816,1452,1729,1882]" pageId="5" pageNumber="410">
|
||||
neurites in the juvenile.
|
||||
<emphasis id="118480135648FFB8FB88F93FFBA4F950" box="[1046,1090,1730,1748]" italics="true" pageId="5" pageNumber="410">Inset</emphasis>
|
||||
Color-coded image of the micrograph shown in
|
||||
<emphasis id="118480135648FFB8FC19F921FC75F974" bold="true" box="[903,915,1756,1776]" pageId="5" pageNumber="410">d</emphasis>
|
||||
. Color code for insets: ectoderm,
|
||||
<emphasis id="118480135648FFB8FB20F921FAF6F974" box="[1214,1296,1756,1776]" italics="true" pageId="5" pageNumber="410">light blue</emphasis>
|
||||
; neurites,
|
||||
<emphasis id="118480135648FFB8FAF2F921FA43F974" box="[1388,1445,1756,1776]" italics="true" pageId="5" pageNumber="410">yellow</emphasis>
|
||||
; basal lamina (ecm),
|
||||
<emphasis id="118480135648FFB8FC7CF90BFBD3F88E" box="[994,1077,1782,1802]" italics="true" pageId="5" pageNumber="410">dark blue</emphasis>
|
||||
; mesoderm,
|
||||
<emphasis id="118480135648FFB8FB36F90BFB22F88E" box="[1192,1220,1782,1802]" italics="true" pageId="5" pageNumber="410">red</emphasis>
|
||||
. bl, blastocoel; ecm, extra cellular matrix; epc, epidermal cell; gp, gill pore; i, intestine; ms, mesocoel; mt, metacoel; mtc, metacoelic cell; myo, myofilaments; ne, neurite; nu, nucleus; pc, protocoel; yo, yolk
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFF16FF62FE6EFDA0" blockId="6.[136,772,159,548]" pageId="6" pageNumber="411">
|
||||
collar region (
|
||||
<figureCitation id="BBCB4084564BFFBBFE82FF62FE9BFF32" box="[284,381,159,182]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="6" pageNumber="411">Fig. 2i, k</figureCitation>
|
||||
), whereas only scattered neurons are present within the prospective trunk region. The number of neurites within the collar and trunk region has considerably increased compared to the Spengel stage, and the neurites show a distinct plexus-like pattern, particularly in the prospective trunk region (
|
||||
<figureCitation id="BBCB4084564BFFBBFEDFFEB8FE6DFED8" box="[321,395,325,348]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="6" pageNumber="411">
|
||||
Fig.
|
||||
<quantity id="E408F1E4564BFFBBFEEFFEB8FE6DFED8" box="[369,395,325,348]" metricMagnitude="-3" metricUnit="kg" metricValue="2.0" pageId="6" pageNumber="411" unit="g" value="2.0">2g</quantity>
|
||||
</figureCitation>
|
||||
). The opisthotroch nerve ring is still well developed and encircles the larva’ s body in the middle of the prospective trunk region (
|
||||
<figureCitation id="BBCB4084564BFFBBFE2BFE75FE1AFE1B" box="[437,508,392,415]" captionStart="Fig" captionStartId="4.[136,168,1589,1609]" captionTargetBox="[136,1452,144,1569]" captionTargetId="figure-1@4.[136,1452,144,1569]" captionTargetPageId="4" captionText="Fig. 2 Neurogenesis in metamorphosing stages of B. misakiensis. b, c, e–k Z-projections of confocal microscopy image stacks. a, d Scanning electron micrographs. Anterior is to the top. a SEM of early metamorphosing stage (Spengel) from lateral left. Arrowhead points to the dorsolateral slit-like depression. b Overview of the serotonin-LIR nervous system (NS) in Spengel stage, view from left. Numerous serotonin-LIR neurons are part of the apical organ. Note the serotonin-LIR opisthotroch neurite ring. c Overview of the serotonin-LIR NS of a late metamorphosing stage (Agassiz), dorsal view. d Dorsal view of metamorphosing Agassiz stage. e Detail showing the developing 5-HT+ nervous plexus in the postoral region of the Spengel stage. f Close-up of the 5-HT+ apical organ" figureDoi="http://doi.org/10.5281/zenodo.12764146" httpUri="https://zenodo.org/record/12764146/files/figure.png" pageId="6" pageNumber="411">Fig. 2c</figureCitation>
|
||||
). Ultrastructural analyses of the metamorphosing Agassiz stage of
|
||||
<taxonomicName id="E4F02782564BFFBBFDDBFE54FD3BFE44" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[581,733,425,448]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="6" pageNumber="411" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564BFFBBFDDBFE54FD3BFE44" box="[581,733,425,448]" italics="true" pageId="6" pageNumber="411">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
revealed the presence of numerous neurites that form a 4– 6-μm-thick basiepidermal layer within the prospective trunk region (
|
||||
<figureCitation id="BBCB4084564BFFBBFF47FDF0FEDCFDA0" box="[217,314,525,548]" captionStart="Fig" captionStartId="5.[136,171,1729,1749]" captionTargetBox="[136,1452,598,1709]" captionTargetId="figure-249@5.[136,1452,594,1709]" captionTargetPageId="5" captionText="Fig. 3 Ultrastructural details of the developing nervous system Balanoglossus misakiensis. a Sagittal section of an Agassiz stage larva of B. misakiensis. b Sagittal section of a 2-gill-slit juvenile of B. misakiensis.c Ultrastructural detail of the red-marked box in a showing a continuous layer of neurites of 4–6-μm thickness in the Agassiz stage. Inset color-coded image of the micrograph shown in c. d Ultrastructural detail of the red-marked box in b showing only individual basiepidermal" figureDoi="http://doi.org/10.5281/zenodo.12764148" httpUri="https://zenodo.org/record/12764148/files/figure.png" pageId="6" pageNumber="411">Fig. 3a, c</figureCitation>
|
||||
, inset).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFF16FDB2FDE7FDE2" blockId="6.[136,513,591,614]" box="[136,513,591,614]" pageId="6" pageNumber="411">
|
||||
<taxonomicName id="E4F02782564BFFBBFF16FDB2FEFAFDE2" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[136,284,591,614]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="6" pageNumber="411" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564BFFBBFF16FDB2FEFAFDE2" box="[136,284,591,614]" italics="true" pageId="6" pageNumber="411">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
: early settled juvenile
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFF16FD6CFF0CFC70" blockId="6.[136,772,657,1909]" pageId="6" pageNumber="411">
|
||||
Early settled
|
||||
<taxonomicName id="E4F02782564BFFBBFE92FD6CFE45FD2C" authorityName="delle Chiaje" authorityYear="1829" box="[268,419,657,680]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="6" pageNumber="411" phylum="Hemichordata" rank="genus">
|
||||
<emphasis id="11848013564BFFBBFE92FD6CFE45FD2C" box="[268,419,657,680]" italics="true" pageId="6" pageNumber="411">Balanoglossus</emphasis>
|
||||
</taxonomicName>
|
||||
(12-h postsettlement (ps)) have an elongated body of vermiform shape measuring approximately
|
||||
<quantity id="E408F1E4564BFFBBFF16FD29FF21FD6F" box="[136,199,724,747]" metricMagnitude="-3" metricUnit="m" metricValue="1.0" pageId="6" pageNumber="411" unit="mm" value="1.0">1 mm</quantity>
|
||||
in length (
|
||||
<figureCitation id="BBCB4084564BFFBBFEA3FD29FE42FD6F" box="[317,420,724,747]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4a, b</figureCitation>
|
||||
). The proboscis is pointed at the anterior tip and about 450 μm long. The collar region measures 200 μm in length and is subdivided by a circular constriction into a broader anterior and narrower posterior part. The trunk region has significantly increased in length, now being 430 μm long. One pair of dorsolateral gill pores is developed at the anterior margin just behind the collar region. The opisthotroch is still present and encircles the trunk region (
|
||||
<figureCitation id="BBCB4084564BFFBBFF0EFC20FF3CFC70" box="[144,218,989,1012]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4b</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFF38FC02FBB1FDC1" blockId="6.[136,772,657,1909]" lastBlockId="6.[816,1452,159,581]" pageId="6" pageNumber="411">
|
||||
Numerous serotonin-LIR somata are distributed throughout the epidermis of the proboscis region, and the apical organ is still visible at the anterior tip; yet, the number of serotoninLIR cells is reduced (
|
||||
<figureCitation id="BBCB4084564BFFBBFEF9FB9FFE56FBFD" box="[359,432,1122,1145]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4a</figureCitation>
|
||||
) compared to the Agassiz stage. The serotonin-LIR nervous system of the proboscis region differs from the Agassiz stage in having a neurite-rich plexus at the dorsomedian base of the proboscis (
|
||||
<figureCitation id="BBCB4084564BFFBBFDA7FB3BFD7CFB59" box="[569,666,1222,1245]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4a, e</figureCitation>
|
||||
). This socalled proboscis stem comprises one of the most condensed areas of the adult enteropneust nervous system. Posterior of the proboscis plexus, two serotonin-LIR neurite bundles project into the collar region and pass through the collar on a subepidermal level (
|
||||
<figureCitation id="BBCB4084564BFFBBFEF6FA91FE51FA07" box="[360,439,1388,1411]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4e</figureCitation>
|
||||
). Both serotonin-LIR bundles are part of the neurulated collar cord that forms at this stage. Although
|
||||
<collectionCode id="45E1C4C4564BFFBBFF6AFA53FED7FA41" box="[244,305,1454,1477]" pageId="6" pageNumber="411">DAPI</collectionCode>
|
||||
staining reveals numerous nuclei accompanying the collar cord, no soma stains positively for serotonin within the collar cord. At the posterior end of the collar, the two neurite bundles become basiepidermal again and project laterally (
|
||||
<figureCitation id="BBCB4084564BFFBBFF77F9CEFED6F9CE" box="[233,304,1587,1610]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4e</figureCitation>
|
||||
) into the prebranchial nerve ring (see below). Within the epidermis of the collar region, numerous serotoninLIR somata are intercalated between the epidermis cells. Each serotonin-LIR bipolar neuron sends a proximal neurite into the basiepidermal plexus. The majority of serotonin-LIR neurites within the collar region runs in longitudinal direction (
|
||||
<figureCitation id="BBCB4084564BFFBBFF0EF907FF3AF895" box="[144,220,1786,1809]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4d</figureCitation>
|
||||
) and project into one of the two circular serotoninLIR neurite bundles within the posterior half of the collar region (
|
||||
<figureCitation id="BBCB4084564BFFBBFF47F8C0FEC3F8D0" box="[217,293,1853,1876]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4d</figureCitation>
|
||||
, mcb and 4f, pnr). The more anterior circular neurite bundle runs basiepidermally at the level of the internal margin of the collar coelom (mesocoel) (
|
||||
<figureCitation id="BBCB4084564BFFBBFB50FF62FAF1FF32" box="[1230,1303,159,182]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4d</figureCitation>
|
||||
, mcb), whereas the more posterior ring, the prebranchial nerve ring, is present at the very posterior margin of the collar region. The prebranchial nerve ring forms a distinct network of serotonin-LIR neurites around the paired mesocoelic pores (
|
||||
<figureCitation id="BBCB4084564BFFBBFCA6FEB8FC7AFED8" box="[824,924,325,348]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4f, g</figureCitation>
|
||||
). These mesocoelic pores connect the coeloms of the collar region, the mesocoels, to the opening of the first gill pore, and eventually to the exterior. At this stage of development, a basiepidermal plexus is detectable by serotonin-LIR throughout the entire trunk region (
|
||||
<figureCitation id="BBCB4084564BFFBBFB31FE37FB1DFE65" box="[1199,1275,458,481]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4c</figureCitation>
|
||||
), however, with only few serotonin-LIR somata. The opisthotroch neurite ring is less condensed and comprises only a loose arrangement of circular neurites (
|
||||
<figureCitation id="BBCB4084564BFFBBFC7BFDD3FBA1FDC1" box="[997,1095,558,581]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4a, c</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFCAEFD8DFB6DFD03" blockId="6.[816,1163,624,647]" box="[816,1163,624,647]" pageId="6" pageNumber="411">
|
||||
<taxonomicName id="E4F02782564BFFBBFCAEFD8DFC23FD03" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[816,965,624,647]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="6" pageNumber="411" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564BFFBBFCAEFD8DFC23FD03" box="[816,965,624,647]" italics="true" pageId="6" pageNumber="411">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
: 1-gill-slit juvenile
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFCAEFD4EFAAAFCF4" blockId="6.[816,1452,691,1643]" pageId="6" pageNumber="411">
|
||||
After approximately 1-day postsettlement, the majority of juveniles have lost the opisthotroch (
|
||||
<figureCitation id="BBCB4084564BFFBBFB33FD29FB1DFD6F" box="[1197,1275,724,747]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4h</figureCitation>
|
||||
). The proboscis shape has slightly changed into a blunt, rounded acorn, measuring between 250 and 350 μm in length. The collar region resembles that described for the previous stage and the trunk region is shorter than the previous stage (
|
||||
<figureCitation id="BBCB4084564BFFBBFB43FCA4FADBFCF4" box="[1245,1341,857,880]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4h, l</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFCD1FC87FAD3F9EF" blockId="6.[816,1452,691,1643]" pageId="6" pageNumber="411">
|
||||
Within the proboscis and collar region, serotonin-LIR reveals only few changes of the overall architecture of the nervous system. Along the anterior tip of the proboscis, the number of the serotonin-LIR somata within the former apical organ is further reduced (
|
||||
<figureCitation id="BBCB4084564BFFBBFB81FC02FB91FB92" box="[1055,1143,1023,1046]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4i, l</figureCitation>
|
||||
). Along the anterior-posterior axis of the proboscis region, distinct neurite bundles indicate the position of the former larval ciliary bands (
|
||||
<figureCitation id="BBCB4084564BFFBBFA8CFBBCFAB0FBDC" box="[1298,1366,1089,1112]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4i</figureCitation>
|
||||
, cgn, l). Within the anterior two thirds of the collar region, two rings of numerous serotonin-LIR neurons are present (
|
||||
<figureCitation id="BBCB4084564BFFBBFA93FB79FABEFB1F" box="[1293,1368,1156,1179]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4k</figureCitation>
|
||||
). These cells are long, slender bipolar neurons with the somata placed basally, centrally, or even apically within the epidermis (
|
||||
<figureCitation id="BBCB4084564BFFBBFCA6FB1AFC64FB7A" box="[824,898,1255,1278]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4k</figureCitation>
|
||||
). At the posterior collar margin, a pair of prebranchial nerves runs circumferentially, passes the mesocoelic pores, and eventually connects to the ventral part of the nervous plexus (
|
||||
<figureCitation id="BBCB4084564BFFBBFC1AFAB6FC2DFAE6" box="[900,971,1355,1378]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4l</figureCitation>
|
||||
, pnr). While the arrangement and orientation of the basiepidermal neurites within the proboscis and collar region have not changed much, the situation in the trunk region is considerably different. A plexus-like arrangement of serotonin-LIR neurites is only detectable in the anterior half of the trunk (
|
||||
<figureCitation id="BBCB4084564BFFBBFC3AFA0CFBF7F98C" box="[932,1041,1521,1544]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4l, m</figureCitation>
|
||||
), whereas in the posterior part of the trunk, individual circumferential neurites are present. These circular neurites project into a longitudinal neurite bundle that is present along the ventral midline (
|
||||
<figureCitation id="BBCB4084564BFFBBFB33F9A9FB17F9EF" box="[1197,1265,1620,1643]" captionStart="Fig" captionStartId="7.[816,848,163,183]" captionTargetId="figure-116@8.[136,1452,144,1585]" captionTargetPageId="8" captionText="Fig. 4 Neurogenesis in early settled stages of B. misakiensis. a, c–i, l, m Z-projections of confocal microscopy image stacks. b, k Scanning electron micrographs. a Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left. b Left side view of early settled juvenile. c Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus. d Partial Z-projection showing the longitudinally orientated neurites within the collar region. e Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord. f, g Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in f. h SEM of a settled juvenile in lateral right view. i Close-up of the apical tip of the proboscis. k Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar. l Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view. m Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α-tub, acetylated α-tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 4l</figureCitation>
|
||||
, vnb).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBBFCAEF96AFB6DF92A" blockId="6.[816,1163,1687,1710]" box="[816,1163,1687,1710]" pageId="6" pageNumber="411">
|
||||
<taxonomicName id="E4F02782564BFFBBFCAEF96AFC23F92A" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[816,965,1687,1710]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="6" pageNumber="411" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564BFFBBFCAEF96AFC23F92A" box="[816,965,1687,1710]" italics="true" pageId="6" pageNumber="411">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
: 2-gill-slit juvenile
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564BFFBAFCAEF924FEE7FED8" blockId="6.[816,1452,1753,1909]" lastBlockId="7.[136,772,159,1909]" lastPageId="7" lastPageNumber="412" pageId="6" pageNumber="411">
|
||||
After about 3-day postsettlement, most of the juveniles exhibit two pairs of gill slits (
|
||||
<figureCitation id="BBCB4084564BFFBBFB8BF907FB9FF895" box="[1045,1145,1786,1809]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 5a, b</figureCitation>
|
||||
). Both gill slits are U-shaped and heavily ciliated on the inside (
|
||||
<figureCitation id="BBCB4084564BFFBBFB11F8E1FB3FF8B7" box="[1167,1241,1820,1843]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 5b</figureCitation>
|
||||
). No synapticles are developed. The juveniles measure up to 1.5 mm in total length (
|
||||
<figureCitation id="BBCB4084564BFFBBFCA6F8A3FC79F8F1" box="[824,927,1885,1909]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="6" pageNumber="411">Fig. 5a–c</figureCitation>
|
||||
). The proboscis shows the typical acorn shape, while the mesosome has developed into a 200-μm-long threelobed collar region (
|
||||
<figureCitation id="BBCB4084564AFFBAFEC0FF3DFE4EFF53" box="[350,424,192,215]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5a</figureCitation>
|
||||
). Scanning electron micrographs and acetylated-α- tubulin-LIR reveal that the proboscis and collar region are evenly covered with cilia, whereas on the trunk region, only scattered tufts of cilia are present (
|
||||
<figureCitation id="BBCB4084564AFFBAFF0EFEB8FF17FED8" box="[144,241,325,348]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5a, c</figureCitation>
|
||||
).
|
||||
</paragraph>
|
||||
<caption id="778F0C89564AFFBAFCAEFF5EFBB1FCB1" pageId="7" pageNumber="412" startId="7.[816,848,163,183]" targetBox="[136,1452,144,1585]" targetPageId="8">
|
||||
<paragraph id="234F5C01564AFFBAFCAEFF5EFBB1FCB1" blockId="7.[816,1478,158,821]" pageId="7" pageNumber="412">
|
||||
<emphasis id="11848013564AFFBAFCAEFF5EFC82FF33" bold="true" box="[816,868,163,183]" pageId="7" pageNumber="412">Fig. 4</emphasis>
|
||||
Neurogenesis in early settled stages of
|
||||
<taxonomicName id="E4F02782564AFFBAFB58FF59FAA4FF33" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[1222,1346,163,183]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="7" pageNumber="412" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564AFFBAFB58FF59FAA4FF33" box="[1222,1346,163,183]" italics="true" pageId="7" pageNumber="412">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
.
|
||||
<emphasis id="11848013564AFFBAFAD3FF5EFABEFF33" bold="true" box="[1357,1368,163,183]" pageId="7" pageNumber="412">a</emphasis>
|
||||
,
|
||||
<emphasis id="11848013564AFFBAFAFDFF5EFA9BFF33" bold="true" box="[1379,1405,163,183]" pageId="7" pageNumber="412">c–i</emphasis>
|
||||
,
|
||||
<emphasis id="11848013564AFFBAFA16FF5EFA68FF33" bold="true" box="[1416,1422,163,183]" pageId="7" pageNumber="412">l</emphasis>
|
||||
,
|
||||
<emphasis id="11848013564AFFBAFA07FF5EFA4AFF33" bold="true" box="[1433,1452,163,183]" pageId="7" pageNumber="412">m</emphasis>
|
||||
Z-projections of confocal microscopy image stacks.
|
||||
<emphasis id="11848013564AFFBAFABBFF43FAD7FF56" bold="true" box="[1317,1329,190,210]" pageId="7" pageNumber="412">b</emphasis>
|
||||
,
|
||||
<emphasis id="11848013564AFFBAFADEFF43FAAAFF56" bold="true" box="[1344,1356,190,210]" pageId="7" pageNumber="412">k</emphasis>
|
||||
Scanning electron micrographs.
|
||||
<emphasis id="11848013564AFFBAFC6BFF25FBE6FF68" bold="true" box="[1013,1024,216,236]" pageId="7" pageNumber="412">a</emphasis>
|
||||
Ventrolateral overview of the serotonin-LIR NS of an early settled juvenile. Anterior is to the left.
|
||||
<emphasis id="11848013564AFFBAFB47FF0EFB03FE83" bold="true" box="[1241,1253,243,263]" pageId="7" pageNumber="412">b</emphasis>
|
||||
Left side view of early settled juvenile.
|
||||
<emphasis id="11848013564AFFBAFC57FEF0FC35FEA5" bold="true" box="[969,979,269,289]" pageId="7" pageNumber="412">c</emphasis>
|
||||
Detail of the anterior trunk region of early settled juvenile showing a ubiquitous basiepidermal nervous plexus.
|
||||
<emphasis id="11848013564AFFBAFAC0FED5FA8CFEB8" bold="true" box="[1374,1386,296,316]" pageId="7" pageNumber="412">d</emphasis>
|
||||
Partial Z-projection showing the longitudinally orientated neurites within the collar region.
|
||||
<emphasis id="11848013564AFFBAFC32FEA0FC50FEF5" bold="true" box="[940,950,349,369]" pageId="7" pageNumber="412">e</emphasis>
|
||||
Detail of the 5-HT+ proboscis plexus at the base of the proboscis and the neurite bundles passing through the subepidermal collar cord.
|
||||
<emphasis id="11848013564AFFBAFCFBFE6FFC8AFE22" bold="true" box="[869,876,402,422]" pageId="7" pageNumber="412">f</emphasis>
|
||||
,
|
||||
<emphasis id="11848013564AFFBAFCE5FE6FFC60FE22" bold="true" box="[891,902,402,422]" pageId="7" pageNumber="412">g</emphasis>
|
||||
Close-up of the gill pore and mesocoelic pore showing the prebranchial nerve ring and the nervous plexus entangling the mesocoelic duct in
|
||||
<emphasis id="11848013564AFFBAFC43FE3AFC1AFE5F" bold="true" box="[989,1020,455,475]" pageId="7" pageNumber="412">f. h</emphasis>
|
||||
<collectionCode id="45E1C4C4564AFFBAFB9DFE3AFBD7FE5F" box="[1027,1073,455,475]" pageId="7" pageNumber="412">SEM</collectionCode>
|
||||
of a settled juvenile in lateral right view.
|
||||
<emphasis id="11848013564AFFBAFA3BFE3AFA4DFE5F" bold="true" box="[1445,1451,455,475]" pageId="7" pageNumber="412">i</emphasis>
|
||||
Close-up of the apical tip of the proboscis.
|
||||
<emphasis id="11848013564AFFBAFB27FE1FFB23FE72" bold="true" box="[1209,1221,482,502]" pageId="7" pageNumber="412">k</emphasis>
|
||||
Detail of the dorsolateral collar region showing two clusters of 5-HT+ bipolar neurons that encircle the collar.
|
||||
<emphasis id="11848013564AFFBAFC06FDEAFC78FDAF" bold="true" box="[920,926,535,555]" pageId="7" pageNumber="412">l</emphasis>
|
||||
Overview of the 5-HT+ NS of a settled juvenile 24-h postsettlement, lateral right view.
|
||||
<emphasis id="11848013564AFFBAFBF1FDCFFB64FDC2" bold="true" box="[1135,1154,562,582]" pageId="7" pageNumber="412">m</emphasis>
|
||||
Detail showing circular 5-HT+ neurites within the posterior part of the trunk whereas more anterior a net-like pattern is still present. The apical tuft and cluster of 5-HT+ somata has disappeared. Apical is to the top. 5-HT, serotonin; ac-α- tub, acetylated α- tubulin; a, anus; ao, apical organ; cgn, ciliary groove nerves; cn, circumferential neurite; cc, collar cord; co, collar; gs, gill slit; mcb, middle circular neurite bundle; msp, mesocoel pore; ne, neurite; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tr, trunk; vnb, ventral neurite bundle
|
||||
</paragraph>
|
||||
</caption>
|
||||
<paragraph id="234F5C01564AFFBAFF38FE9AFE2BF8F1" blockId="7.[136,772,159,1909]" pageId="7" pageNumber="412">
|
||||
Throughout the epidermis of the proboscis, numerous serotonin-LIR bipolar neurons are intercalated between the other epidermal cells (
|
||||
<figureCitation id="BBCB4084564AFFBAFEF1FE54FE29FE44" box="[367,463,425,448]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5c, i</figureCitation>
|
||||
). Each soma projects a single proximal neurite into the basiepidermal plexus of the proboscis region (
|
||||
<figureCitation id="BBCB4084564AFFBAFF66FE16FEDDFD86" box="[248,315,491,514]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5i</figureCitation>
|
||||
). The most condensed area of serotonin-LIR neurites is present dorsally at the base of the proboscis, comprising a part of the proboscis plexus (
|
||||
<figureCitation id="BBCB4084564AFFBAFD87FDD3FD9BFDC1" box="[537,637,558,581]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5c, d</figureCitation>
|
||||
). This dense network of neurites measures approximately 200 μm in width and 100 μm in length. From here, three distinct serotonin-LIR neurite bundles leave the proboscis plexus posteriorly to pass through the subepidermal collar cord within the collar region (
|
||||
<figureCitation id="BBCB4084564AFFBAFF0EFD29FF3CFD6F" box="[144,218,724,747]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5d</figureCitation>
|
||||
). These serotonin-LIR neurite bundles are part of the ventral layer of neurites of the collar cord (
|
||||
<figureCitation id="BBCB4084564AFFBAFDA7FD08FD64FC88" box="[569,642,757,780]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5h</figureCitation>
|
||||
). The dorsal area of the collar cord is composed of numerous cells, presumably neurons, which do not show serotonin-LIR in any of the somata (
|
||||
<figureCitation id="BBCB4084564AFFBAFF41FCA4FECEFCF4" box="[223,296,857,880]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5h</figureCitation>
|
||||
). At the posterior pole of the collar region, the three serotonin-LIR neurite bundles become basiepidermal again and connect to the nervous system within the trunk region. The median neurite bundle, composed of 7–10 serotonin-LIR neurites, continues directly into the dorsal midline and is part of the dorsal nerve cord (
|
||||
<figureCitation id="BBCB4084564AFFBAFDA6FC02FD99FB92" box="[568,639,1023,1046]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5f</figureCitation>
|
||||
). SerotoninLIR is detectable until the posterior pole of the animals, with a discontinuity of about 150 μm in the middle of the trunk region (
|
||||
<figureCitation id="BBCB4084564AFFBAFF40FB9FFEA1FBFD" box="[222,327,1122,1145]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5f, g</figureCitation>
|
||||
, double arrowheads). Several serotoninLIR bipolar neurons are located dorsolaterally close to the dorsal midline and project a short neurite into the dorsal neurite bundle (
|
||||
<figureCitation id="BBCB4084564AFFBAFEACFB3BFE9DFB59" box="[306,379,1222,1245]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5f</figureCitation>
|
||||
). The lateral pair of neurite bundles that leaves the collar cord posteriorly runs circumferentially to the ventral side (
|
||||
<figureCitation id="BBCB4084564AFFBAFED4FAF5FE75FA9B" box="[330,403,1288,1311]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5d</figureCitation>
|
||||
). At the level of the dorsolateral gill slits, the bilateral neurite bundles form a distinctive network around the mesocoelic pores as also described for younger stages (
|
||||
<figureCitation id="BBCB4084564AFFBAFF42FA91FECAFA07" box="[220,300,1388,1411]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5d</figureCitation>
|
||||
). The bilateral neurite bundles are part of the prebranchial nerve ring and project into the ventral nerve cord. Numerous serotonin-LIR neurites extend along the entire midline of the trunk region thereby running within the ventral nerve cord that is about 40 μm wide (
|
||||
<figureCitation id="BBCB4084564AFFBAFF53F9EFFED1F9AD" box="[205,311,1554,1577]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5c, e</figureCitation>
|
||||
). Several serotonin-LIR somata are interspersed throughout the trunk epidermis. The majority of these bipolar neurons project a proximal neurite running circumferential into the ventral nerve cord (
|
||||
<figureCitation id="BBCB4084564AFFBAFDEFF98BFD59F909" box="[625,703,1654,1677]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5e</figureCitation>
|
||||
). The serotonin-LIR somata of the very posterior end send a proximal neurite running radially into the ventral neurite bundle (
|
||||
<figureCitation id="BBCB4084564AFFBAFF7AF924FED4F974" box="[228,306,1753,1776]" captionStart="Fig" captionStartId="9.[816,848,163,183]" captionTargetId="figure-118@10.[136,1452,144,1651]" captionTargetPageId="10" captionText="Fig. 5 Neurogenesis in the 2-gill slit juvenile of B. misakiensis. a, b Scanning electron micrographs: c–i Z-projections of confocal microscopy image stacks. Anterior is to the right. a Dorsolateral view of a 3-day-old settled juvenile. b Higher magnification of the two gill slits and the dorsal tongue bars. c Overview of the 5-HT+ NS of a 3-day-old juvenile. d The collar cord is composed of three 5-HT+ neurite bundles of which the median one projects posteriorly into the dorsal nerve cord. e Detail of the ventral 5-HT+ neurite bundle with numerous incoming circular neurites emerging from lateral bipolar neurons. f Detail of the anterior part of the dorsal 5-HT+ neurite bundle and epidermal collar region. This part is discontinuous with the posterior part of the 5- HT neurite bundle shown in g, see also double arrowheads and dashed area. g The posterior part of the dorsal 5-HT+ neurite bundle contains few serotonin-LIR neurites and is discontinuous with the anterior part of the 5- HT neurite bundle, see double arrowheads and dashed area. h Partial Zprojection of a sagittal scan of the dorsal collar region. The collar cord comprises ventral neurite bundles and a dorsal sheath of somata, which are not serotonin-LIR positive. i Close-up of a part of the proboscis region showing the 5-HT+ bipolar neurons and the basiepidermal nervous plexus. 5-HT, serotonin; a, anus; ac-α-tub, acetylated α-tubulin; cn, circumferential neurite; cc, collar cord; ci, cilia; co, collar; dnb, dorsal neurite bundle; ep, epidermis; gs, gill slit; msp, mesocoel pore; nn, nervous plexus; onr, opisthotroch nerve ring; pnr, prebranchial nerve ring; pr, proboscis; ps, proboscis stem; sn, serotonin-LIR neuron; tb, tongue bar; tm, trunk musculature; vnb, ventral neurite bundle" pageId="7" pageNumber="412">Fig. 5e</figureCitation>
|
||||
). It should be noted that a serotonin-LIR basiepidermal plexus is only present in the epidermis of the proboscis and collar, but completely absent within the trunk region, where instead a condensed dorsal and ventral neurite bundle is present.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564AFFBAFCD1FC85FAE1FBF3" blockId="7.[816,1452,888,1143]" pageId="7" pageNumber="412">
|
||||
The analysis of serial sections for electron microscopy of the trunk epidermis in juvenile
|
||||
<taxonomicName id="E4F02782564AFFBAFB22FC64FABBFC34" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[1212,1373,921,944]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="7" pageNumber="412" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564AFFBAFB22FC64FABBFC34" box="[1212,1373,921,944]" italics="true" pageId="7" pageNumber="412">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
shows comparatively few basiepidermal neurites in the epidermis (
|
||||
<figureCitation id="BBCB4084564AFFBAFCA6FC26FC4EFC76" box="[824,936,987,1010]" captionStart="Fig" captionStartId="5.[136,171,1729,1749]" captionTargetBox="[136,1452,598,1709]" captionTargetId="figure-249@5.[136,1452,594,1709]" captionTargetPageId="5" captionText="Fig. 3 Ultrastructural details of the developing nervous system Balanoglossus misakiensis. a Sagittal section of an Agassiz stage larva of B. misakiensis. b Sagittal section of a 2-gill-slit juvenile of B. misakiensis.c Ultrastructural detail of the red-marked box in a showing a continuous layer of neurites of 4–6-μm thickness in the Agassiz stage. Inset color-coded image of the micrograph shown in c. d Ultrastructural detail of the red-marked box in b showing only individual basiepidermal" figureDoi="http://doi.org/10.5281/zenodo.12764148" httpUri="https://zenodo.org/record/12764148/files/figure.png" pageId="7" pageNumber="412">Fig. 3b, d</figureCitation>
|
||||
and inset). Neurite bundles composed of 4–8 neurites are scattered throughout the lateral areas of the trunk epidermis (
|
||||
<figureCitation id="BBCB4084564AFFBAFC69FBE3FBADFBB1" box="[1015,1099,1054,1077]" captionStart="Fig" captionStartId="5.[136,171,1729,1749]" captionTargetBox="[136,1452,598,1709]" captionTargetId="figure-249@5.[136,1452,594,1709]" captionTargetPageId="5" captionText="Fig. 3 Ultrastructural details of the developing nervous system Balanoglossus misakiensis. a Sagittal section of an Agassiz stage larva of B. misakiensis. b Sagittal section of a 2-gill-slit juvenile of B. misakiensis.c Ultrastructural detail of the red-marked box in a showing a continuous layer of neurites of 4–6-μm thickness in the Agassiz stage. Inset color-coded image of the micrograph shown in c. d Ultrastructural detail of the red-marked box in b showing only individual basiepidermal" figureDoi="http://doi.org/10.5281/zenodo.12764148" httpUri="https://zenodo.org/record/12764148/files/figure.png" pageId="7" pageNumber="412">Fig. 3d</figureCitation>
|
||||
). The most condensed neurite bundles run along the ventral and dorsal midline of the trunk region and comprise the nerve cords.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
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|
|||
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<mods:title id="621D84183749B12630CD62025CBF0916">Neurogenesis in directly and indirectly developing enteropneusts: of nets and cords</mods:title>
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<treatment id="AB59ED17564FFFBEFF16FC30FC9EFF7D" LSID="urn:lsid:plazi:treatment:AB59ED17564FFFBEFF16FC30FC9EFF7D" httpUri="http://treatment.plazi.org/id/AB59ED17564FFFBEFF16FC30FC9EFF7D" lastPageId="3" lastPageNumber="408" pageId="2" pageNumber="407">
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<subSubSection id="6BEA0F8A564FFFBFFF16FC30FDDAFC60" box="[136,572,973,996]" pageId="2" pageNumber="407" type="nomenclature">
|
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<paragraph id="234F5C01564FFFBFFF16FC30FDDAFC60" blockId="2.[136,572,973,996]" box="[136,572,973,996]" pageId="2" pageNumber="407">
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<heading id="7807EB6D564FFFBFFF16FC30FDDAFC60" box="[136,572,973,996]" fontSize="10" level="2" pageId="2" pageNumber="407" reason="7">
|
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<taxonomicName id="E4F02782564FFFBFFF16FC30FDDAFC60" ID-CoL="4TX3B" authority="(Agassiz, 1873)" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[136,572,973,996]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564FFFBFFF16FC30FE77FC60" box="[136,401,973,996]" italics="true" pageId="2" pageNumber="407">Saccoglossus kowalevskii</emphasis>
|
||||
(Agassiz, 1873)
|
||||
</taxonomicName>
|
||||
</heading>
|
||||
</paragraph>
|
||||
</subSubSection>
|
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<subSubSection id="6BEA0F8A564FFFBEFF16FBF2FC9EFF7D" lastPageId="3" lastPageNumber="408" pageId="2" pageNumber="407" type="description">
|
||||
<paragraph id="234F5C01564FFFBFFF16FBF2FE49FA31" blockId="2.[136,771,1039,1461]" pageId="2" pageNumber="407">
|
||||
<materialsCitation id="9398565C564FFFBFFF16FBF2FE4DFA31" collectingDate="2007-09" collectionCode="TEM, SEM" country="United States of America" location="Adult" municipality="Woods Hole" pageId="2" pageNumber="407" specimenCount="1" stateProvince="Massachusetts">
|
||||
<location id="262F0ADA564FFFBFFF16FBF2FF25FBA2" LSID="urn:lsid:plazi:treatment:AB59ED17564FFFBEFF16FC30FC9EFF7D:262F0ADA564FFFBFFF16FBF2FF25FBA2" box="[136,195,1039,1062]" country="United States of America" municipality="Woods Hole" name="Adult" pageId="2" pageNumber="407" stateProvince="Massachusetts">Adult</location>
|
||||
specimens of
|
||||
<taxonomicName id="E4F02782564FFFBFFEC5FBF2FE0BFBA2" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[347,493,1039,1062]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564FFFBFFEC5FBF2FE0BFBA2" box="[347,493,1039,1062]" italics="true" pageId="2" pageNumber="407">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
were collected from intertidal flats near
|
||||
<collectingMunicipality id="C32BC67B564FFFBFFEDFFBCDFE34FBC3" box="[321,466,1072,1095]" pageId="2" pageNumber="407">Woods Hole</collectingMunicipality>
|
||||
(
|
||||
<collectingRegion id="E13492E3564FFFBFFE7BFBCDFD69FBC3" box="[485,655,1072,1095]" country="United States of America" name="Massachusetts" pageId="2" pageNumber="407">Massachusetts</collectingRegion>
|
||||
,
|
||||
<collectingCountry id="5BE71C91564FFFBFFD3EFBCDFD3FFBC3" box="[672,729,1072,1095]" name="United States of America" pageId="2" pageNumber="407">USA</collectingCountry>
|
||||
) in
|
||||
<date id="574E7AC1564FFFBFFF16FBAFFEDEFBED" box="[136,312,1106,1129]" pageId="2" pageNumber="407" value="2007-09">
|
||||
<collectingDate id="470A8329564FFFBFFF16FBAFFEDEFBED" box="[136,312,1106,1129]" pageId="2" pageNumber="407" value="2007-09">September 2007</collectingDate>
|
||||
</date>
|
||||
and 2011. Animals were transported into the laboratory, separated according to sex, and kept individually in finger bowls. Animals were kept at 18 °C on a seawater table. Freshly spawned eggs were mixed with active sperm isolated from a ripe male and diluted in seawater (
|
||||
<bibRefCitation id="476121F0564FFFBFFD5BFB2AFF11FA8B" author="Lowe, C. & Tagawa, K. & Humphreys, T. & Kirschner, M. & Gerhart, J." pageId="2" pageNumber="407" pagination="171 - 194" refId="ref13388" refString="Lowe, C., Tagawa, K., Humphreys, T., Kirschner, M., & Gerhart, J. (2004). Hemichordate embryos: Procurement, culture, and basic methods. Methods in Cell Biology, 74, 171 - 194." type="journal article" year="2004">Lowe et al. 2004</bibRefCitation>
|
||||
). Fertilization envelopes were ruptured using fine forceps, and embryonic and juvenile stages were collected using Pasteur pipettes. Embryos were relaxed in a mix of 7 % MgCl
|
||||
<subScript id="BF745E44564FFFBFFF68FA9BFEE6FAF2" attach="left" box="[246,256,1382,1398]" fontSize="7" pageId="2" pageNumber="407">2</subScript>
|
||||
and seawater (1:1) for 5–10 min prior to fixation for transmission electron microscopy (
|
||||
<collectionCode id="45E1C4C4564FFFBFFDB7FA80FD85FA10" box="[553,611,1405,1428]" pageId="2" pageNumber="407">TEM</collectionCode>
|
||||
) and scanning electron microscopy (
|
||||
<collectionCode id="45E1C4C4564FFFBFFEF7FA63FE45FA31" box="[361,419,1438,1461]" pageId="2" pageNumber="407">SEM</collectionCode>
|
||||
)
|
||||
</materialsCitation>
|
||||
.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564FFFBFFF16FA0CFD93F98C" blockId="2.[136,629,1521,1544]" box="[136,629,1521,1544]" pageId="2" pageNumber="407">Terminology of names of developmental stages</paragraph>
|
||||
<paragraph id="234F5C01564FFFBFFF16F9CEFDB3F8F1" blockId="2.[136,771,1587,1909]" pageId="2" pageNumber="407">
|
||||
For indirectly developing enteropneusts such as
|
||||
<taxonomicName id="E4F02782564FFFBFFF16F9A9FEFBF9EF" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[136,285,1620,1643]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFF16F9A9FEFBF9EF" box="[136,285,1620,1643]" italics="true" pageId="2" pageNumber="407">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
, we follow the terminology sensu
|
||||
<bibRefCitation id="476121F0564FFFBFFD19F9A9FE87F909" author="Nielsen, C. & Hay-Schmidt, A." pageId="2" pageNumber="407" pagination="551 - 570" refId="ref13840" refString="Nielsen, C., & Hay-Schmidt, A. (2007). Development of the enteropneust Ptychodera flava: ciliary bands and nervous system. Journal of Morphology, 268, 551 - 570." type="journal article" year="2007">Nielsen and Hay-Schmidt (2007)</bibRefCitation>
|
||||
. Accordingly, the Spengel stage is characterized by a regression of size of the larva and the fusion of the preoral ciliary feeding band, the neotroch. The Spengel stage is followed by the Agassiz stage. Agassiz larvae have a cone-shaped anterior body region with a smooth surface, the prospective proboscis region. The overall size is further decreased, and the anterior eye spots begin to disappear. The Agassiz larvae are competent for settlement.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564FFFBFFCD1FF62FC51FE9E" blockId="2.[816,1452,159,282]" pageId="2" pageNumber="407">
|
||||
For the direct developer
|
||||
<taxonomicName id="E4F02782564FFFBFFBC7FF62FB08FF32" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[1113,1262,159,182]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564FFFBFFBC7FF62FB08FF32" box="[1113,1262,159,182]" italics="true" pageId="2" pageNumber="407">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
, we followed the terminology of
|
||||
<bibRefCitation id="476121F0564FFFBFFC47FF3DFB7AFF53" author="Lowe, C. J. & Wu, M. & Salic, A. & Evans, L. & Lander, E. & G. C. E & Gerhart, J. & Kirschner, M." box="[985,1180,192,215]" pageId="2" pageNumber="407" pagination="853 - 865" refId="ref13317" refString="Lowe, C. J., Wu, M., Salic, A., Evans, L., Lander, E., Stange-ThomannN, G. C. E., Gerhart, J., & Kirschner, M. (2003). Anteroposterior patterning in hemichordates and the origin of the chordate nervous system. Cell, 113, 853 - 865." type="journal article" year="2003">Lowe et al. (2003</bibRefCitation>
|
||||
,
|
||||
<bibRefCitation id="476121F0564FFFBFFB32FF3DFB07FF53" author="Lowe, C. J. & Terasaki, M. & Wu, M. & Freeman, R. M. & Runft, L. & Kwan, K. & Haigo, S. & Aronowicz, J. & Lander, E. & Gruber, C. & Smith, M. & Kirschner, M. & Gerhart, J." box="[1196,1249,192,215]" pageId="2" pageNumber="407" refId="ref13439" refString="Lowe, C. J., Terasaki, M., Wu, M., Freeman, R. M., Runft, L., Kwan, K., Haigo, S., Aronowicz, J., Lander, E., Gruber, C., Smith, M., Kirschner, M., & Gerhart, J. (2006). Dorsoventral patterning in hemichordates: insights into early chordate evolution. PLoS Biology, 4, e 291." type="journal volume" year="2006">2006</bibRefCitation>
|
||||
). Accordingly, the Kink stage is characterized by a dorsal bending of the future trunk region.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564FFFBFFCAEFEB8FBE1FED8" blockId="2.[816,1031,325,348]" box="[816,1031,325,348]" pageId="2" pageNumber="407">Electron microscopy</paragraph>
|
||||
<paragraph id="234F5C01564FFFBFFCAEFE75FC40F92A" blockId="2.[816,1452,392,1710]" pageId="2" pageNumber="407">
|
||||
<collectionCode id="45E1C4C4564FFFBFFCAEFE75FC81FE1B" box="[816,871,392,415]" pageId="2" pageNumber="407">TEM</collectionCode>
|
||||
and
|
||||
<collectionCode id="45E1C4C4564FFFBFFC04FE75FC36FE1B" box="[922,976,392,415]" pageId="2" pageNumber="407">SEM</collectionCode>
|
||||
embryos and juveniles of
|
||||
<taxonomicName id="E4F02782564FFFBFFB7FFE75FA95FE1B" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[1249,1395,392,415]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564FFFBFFB7FFE75FA95FE1B" box="[1249,1395,392,415]" italics="true" pageId="2" pageNumber="407">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
were fixed with ice-cold 2.5 % glutaraldehyde (GA) in 0.2 M sodium cacodylate buffer (pH 7.2), adjusted to an osmolarity of approximately 800 mosm with the addition of NaCl (for
|
||||
<taxonomicName id="E4F02782564FFFBFFCAEFDF0FC28FDA0" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[816,974,525,548]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFCAEFDF0FC28FDA0" box="[816,974,525,548]" italics="true" pageId="2" pageNumber="407">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
, 2.5 % GA in 0.05 M phosphate buffer + 0.3 M sodium chloride (
|
||||
<collectionCode id="45E1C4C4564FFFBFFBB3FDD3FBB9FDC1" box="[1069,1119,558,581]" country="United Kingdom" lsid="urn:lsid:biocol.org:col:12782" name="Chambers Institute, Tweeddale Museum" pageId="2" pageNumber="407" type="Herbarium">PBS</collectionCode>
|
||||
)). Primary fixation was stopped after 45 min (30 min,
|
||||
<taxonomicName id="E4F02782564FFFBFFB89FDB2FB4DFDE2" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[1047,1195,591,614]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFB89FDB2FB4DFDE2" box="[1047,1195,591,614]" italics="true" pageId="2" pageNumber="407">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
) with three buffer rinses of 10, 15, and 20 min, respectively (five rinses with PBS for 10 min each for
|
||||
<taxonomicName id="E4F02782564FFFBFFC47FD6CFB8BFD2C" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[985,1133,657,680]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFC47FD6CFB8BFD2C" box="[985,1133,657,680]" italics="true" pageId="2" pageNumber="407">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
). Primary fixation was followed by 30 min of postfixation with 2 % OsO
|
||||
<quantity id="E408F1E4564FFFBFFAB9FD40FAB7FD4E" box="[1319,1361,691,717]" metricMagnitude="-1" metricUnit="m" metricValue="1.016" pageId="2" pageNumber="407" unit="in" value="4.0">
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||||
<subScript id="BF745E44564FFFBFFAB9FD40FAD7FD49" attach="left" box="[1319,1329,701,717]" fontSize="7" pageId="2" pageNumber="407">4</subScript>
|
||||
in
|
||||
</quantity>
|
||||
sodium cacodylate buffer (
|
||||
<collectionCode id="45E1C4C4564FFFBFFC6DFD29FBC0FD6F" box="[1011,1062,724,747]" country="United Kingdom" lsid="urn:lsid:biocol.org:col:12782" name="Chambers Institute, Tweeddale Museum" pageId="2" pageNumber="407" type="Herbarium">PBS</collectionCode>
|
||||
). Postfixation was stopped with three buffer rinses (15, 30, 30 min) followed by two rinses with ddH2O (15, 30 min). After dehydration through a graded series of ethanol, specimens were embedded in Epon resin for
|
||||
<collectionCode id="45E1C4C4564FFFBFFCAEFCA4FC81FCF4" box="[816,871,857,880]" pageId="2" pageNumber="407">TEM</collectionCode>
|
||||
and light microscopy. For
|
||||
<collectionCode id="45E1C4C4564FFFBFFBE4FCA4FB54FCF4" box="[1146,1202,857,880]" pageId="2" pageNumber="407">SEM</collectionCode>
|
||||
, specimens were critical point dried in a CPD 030 (
|
||||
<collectingRegion id="E13492E3564FFFBFFBCCFC87FB47FC15" box="[1106,1185,890,913]" country="Liechtenstein" name="Balzers" pageId="2" pageNumber="407">Balzers</collectingRegion>
|
||||
Union,
|
||||
<collectingCountry id="5BE71C91564FFFBFFB64FC87FA60FC15" box="[1274,1414,890,913]" name="Liechtenstein" pageId="2" pageNumber="407">Liechtenstein</collectingCountry>
|
||||
) or Leica CPD 300 (Leica,
|
||||
<collectingCountry id="5BE71C91564FFFBFFBA5FC66FB45FC36" box="[1083,1187,923,946]" name="Germany" pageId="2" pageNumber="407">Germany</collectingCountry>
|
||||
). Dried specimens were sputter coated with gold in a SCD 040 (
|
||||
<collectingRegion id="E13492E3564FFFBFFA9CFC41FAB3FC57" box="[1282,1365,956,979]" country="Liechtenstein" name="Balzers" pageId="2" pageNumber="407">Balzers</collectingRegion>
|
||||
Union,
|
||||
<collectingCountry id="5BE71C91564FFFBFFCAEFC20FC37FC70" box="[816,977,989,1012]" name="Liechtenstein" pageId="2" pageNumber="407">Liechtenstein</collectingCountry>
|
||||
) sputter coater and viewed with a Fei Quantum 200
|
||||
<collectionCode id="45E1C4C4564FFFBFFC57FC02FC19FB92" box="[969,1023,1023,1046]" pageId="2" pageNumber="407">SEM</collectionCode>
|
||||
at 15 kV (
|
||||
<collectionCode id="45E1C4C4564FFFBFFBEAFC02FB7AFB92" box="[1140,1180,1023,1046]" pageId="2" pageNumber="407">FEI</collectionCode>
|
||||
Co.,
|
||||
<collectingCountry id="5BE71C91564FFFBFFB46FC02FA61FB92" box="[1240,1415,1023,1046]" name="Netherlands" pageId="2" pageNumber="407">The Netherlands</collectingCountry>
|
||||
) or a Philips XL 30
|
||||
<collectionCode id="45E1C4C4564FFFBFFC42FBDDFBC4FBB3" box="[988,1058,1056,1079]" pageId="2" pageNumber="407">ESEM</collectionCode>
|
||||
(Philips,
|
||||
<collectingCountry id="5BE71C91564FFFBFFB1BFBDDFAD0FBB3" box="[1157,1334,1056,1079]" name="Netherlands" pageId="2" pageNumber="407">The Netherlands</collectingCountry>
|
||||
). Complete longitudinal and transverse semithin serial sections (0.5 μm) for light microscopy and ultrathin serial sections (~
|
||||
<quantity id="E408F1E4564FFFBFFAA2FB9FFA66FBFD" box="[1340,1408,1122,1145]" metricMagnitude="-8" metricUnit="m" metricValue="5.5" pageId="2" pageNumber="407" unit="nm" value="55.0">55 nm</quantity>
|
||||
) for
|
||||
<collectionCode id="45E1C4C4564FFFBFFCAEFB79FC81FB1F" box="[816,871,1156,1179]" pageId="2" pageNumber="407">TEM</collectionCode>
|
||||
from three stages of
|
||||
<taxonomicName id="E4F02782564FFFBFFBDAFB79FB31FB1F" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[1092,1239,1156,1179]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564FFFBFFBDAFB79FB31FB1F" box="[1092,1239,1156,1179]" italics="true" pageId="2" pageNumber="407">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
(56, 132, and 432 h pf) were produced on a Leica Ultracut S microtome and from five stages of
|
||||
<taxonomicName id="E4F02782564FFFBFFC55FB3BFB85FB59" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[971,1123,1222,1245]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFC55FB3BFB85FB59" box="[971,1123,1222,1245]" italics="true" pageId="2" pageNumber="407">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
<emphasis id="11848013564FFFBFFBEBFB38FB67FB59" bold="true" box="[1141,1153,1221,1245]" pageId="2" pageNumber="407">B</emphasis>
|
||||
Spengel
|
||||
<emphasis id="11848013564FFFBFFB40FB3BFACEFB59" box="[1246,1320,1221,1245]" italics="true" pageId="2" pageNumber="407">stage^:</emphasis>
|
||||
13 days pf,
|
||||
<emphasis id="11848013564FFFBFFCAEFB1BFCDAFB7A" bold="true" box="[816,828,1254,1278]" pageId="2" pageNumber="407">B</emphasis>
|
||||
Agassiz
|
||||
<emphasis id="11848013564FFFBFFC06FB1AFC07FB7A" box="[920,993,1254,1278]" italics="true" pageId="2" pageNumber="407">stage^:</emphasis>
|
||||
14 days pf, 12-h postsettlement (ps), 1 ay ps and 3 days ps) sectioned on a Leica
|
||||
<collectionCode id="45E1C4C4564FFFBFFB72FAF5FAF7FA9B" box="[1260,1297,1288,1311]" country="USA" name="Upjohn Culture Collection" pageId="2" pageNumber="407">UC</collectionCode>
|
||||
7 microtome. Semithin sections were stained with 1 % toluidin blue for 3 min at 63 °C. Ultrathin sections were stained with 2 % uranylacetate and 2.5 % lead citrate, either in an automatic stainer (NanoWlm Technologie GmbH, Göttingen,
|
||||
<collectingCountry id="5BE71C91564FFFBFFCAEFA53FC75FA41" box="[816,915,1454,1477]" name="Germany" pageId="2" pageNumber="407">Germany</collectingCountry>
|
||||
) or manually. Light microscopic images were recorded with a digital camera (Olympus
|
||||
<collectionCode id="45E1C4C4564FFFBFFB42FA2DFAA6FA63" box="[1244,1344,1488,1511]" pageId="2" pageNumber="407">BX-UCB</collectionCode>
|
||||
) mounted on an Olympus BX51 compound microscope or Nikon Eclipse E800.
|
||||
<collectionCode id="45E1C4C4564FFFBFFC40F9EFFBF1F9AD" box="[990,1047,1554,1577]" pageId="2" pageNumber="407">TEM</collectionCode>
|
||||
pictures were documented with a Philips EM 208 electron microscope at 70 kV equipped with a Nikon digital camera. Images were optimized by using Adobe Photoshop and Illustrator
|
||||
<collectionCode id="45E1C4C4564FFFBFFB39F98BFB2FF909" box="[1191,1225,1654,1677]" country="France" name="Musee des Dinosaures d'Esperaza (Aude)" pageId="2" pageNumber="407">CS</collectionCode>
|
||||
3 (Adobe, San José,
|
||||
<collectionCode id="45E1C4C4564FFFBFFCAEF96AFCB1F92A" box="[816,855,1687,1710]" country="USA" name="Chicago Academy of Sciences" pageId="2" pageNumber="407" type="Museum">CA</collectionCode>
|
||||
,
|
||||
<collectingCountry id="5BE71C91564FFFBFFCFCF96AFC7CF92A" box="[866,922,1687,1710]" name="United States of America" pageId="2" pageNumber="407">USA</collectingCountry>
|
||||
).
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564FFFBFFCAEF924FA64F974" blockId="2.[816,1410,1753,1776]" box="[816,1410,1753,1776]" pageId="2" pageNumber="407">Immunolabeling and confocal laser scanning microscopy</paragraph>
|
||||
<paragraph id="234F5C01564FFFBEFCAEF8E1FD4CFA20" blockId="2.[816,1451,1820,1909]" lastBlockId="3.[136,772,159,1909]" lastPageId="3" lastPageNumber="408" pageId="2" pageNumber="407">
|
||||
For immunocytochemistry embryos and juveniles of
|
||||
<taxonomicName id="E4F02782564FFFBFFCAEF8C0FC21F8D0" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[816,967,1853,1876]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564FFFBFFCAEF8C0FC21F8D0" box="[816,967,1853,1876]" italics="true" pageId="2" pageNumber="407">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
(dorsal kink = 76 hpf, 1-gill-slit hatchling = 156 hpf, 3-gill-slit juvenile = 750 hpf) and larvae and juveniles of
|
||||
<taxonomicName id="E4F02782564EFFBEFF36FF62FEA6FF32" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[168,320,159,182]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="3" pageNumber="408" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564EFFBEFF36FF62FEA6FF32" box="[168,320,159,182]" italics="true" pageId="3" pageNumber="408">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
(
|
||||
<emphasis id="11848013564EFFBEFECCFF62FEB8FF33" bold="true" box="[338,350,159,183]" pageId="3" pageNumber="408">B</emphasis>
|
||||
Spengel
|
||||
<emphasis id="11848013564EFFBEFE22FF62FE18FF33" box="[444,510,159,183]" italics="true" pageId="3" pageNumber="408">stage^</emphasis>
|
||||
= 13 days pf,
|
||||
<emphasis id="11848013564EFFBEFD3BFF62FD57FF33" bold="true" box="[677,689,159,183]" pageId="3" pageNumber="408">B</emphasis>
|
||||
Agassiz
|
||||
<emphasis id="11848013564EFFBEFF16FF3DFF2CFF5C" box="[136,202,192,216]" italics="true" pageId="3" pageNumber="408">stage^</emphasis>
|
||||
= 14 days pf, early settled stage = 12-h postsettlement (ps), 1-gill-slit juvenile = 1 day ps, and 2-gill-slit juvenile = 3 days ps) were fixed with 4 % paraformaldehyde (
|
||||
<collectionCode id="45E1C4C4564EFFBEFD30FEFEFD39FE9E" box="[686,735,259,282]" pageId="3" pageNumber="408">PFA</collectionCode>
|
||||
) in phosphate buffer (
|
||||
<collectionCode id="45E1C4C4564EFFBEFED4FED9FE88FEBF" box="[330,366,292,315]" pageId="3" pageNumber="408">PB</collectionCode>
|
||||
). Specimens were washed three times and permeabilized in PB containing 2 % Triton X-100 (
|
||||
<collectionCode id="45E1C4C4564EFFBEFD5AFEB8FD1FFED8" box="[708,761,325,348]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
). Subsequent blocking was carried out in
|
||||
<collectionCode id="45E1C4C4564EFFBEFDB3FE9AFDBBFEFA" box="[557,605,359,382]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
containing 6 % normal goat serum (Jackson Immuno Research) for 2 h at room temperature (
|
||||
<collectionCode id="45E1C4C4564EFFBEFEC8FE54FE9DFE44" box="[342,379,425,448]" pageId="3" pageNumber="408">RT</collectionCode>
|
||||
). Primary antibodies were diluted in blocking solution in
|
||||
<collectionCode id="45E1C4C4564EFFBEFEC1FE37FE69FE65" box="[351,399,458,481]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
and applied at a final concentration of 1:500 (anti-serotonin, Sigma, Cat. # S5545; anti-acetylated α- tubulin Sigma, Cat. #
|
||||
<collectionCode id="45E1C4C4564EFFBEFE13FDF0FE7BFDA0" box="[397,413,525,548]" name="Tavera, Department of Geology and Geophysics" pageId="3" pageNumber="408">T</collectionCode>
|
||||
6793) overnight at RT. Then, animals were washed six times for 20 min each with
|
||||
<collectionCode id="45E1C4C4564EFFBEFD00FDD3FD28FDC1" box="[670,718,558,581]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
on a rocker table. Secondary antibodies were diluted in blocking solution in
|
||||
<collectionCode id="45E1C4C4564EFFBEFF60FD8DFEC8FD03" box="[254,302,624,647]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
and applied at a final concentration of 1:600 (goat anti-rabbit Alexa Fluor 633, Invitrogen, Cat. # A21070; goat anti-mouse Alexa Fluor 568, Invitrogen, Cat. # A11004) together with a nuclei marker (1:600,
|
||||
<collectionCode id="45E1C4C4564EFFBEFD99FD29FDA1FD6F" box="[519,583,724,747]" pageId="3" pageNumber="408">DAPI</collectionCode>
|
||||
, Invitrogen, Cat. # D1306) and a marker for filamentous actin (1:150, Alexa fluor 488 phalloidin, Invitrogen, Cat. # A12379) for 4 h at RT. Afterward, the samples were rinsed three times for 20 min each in
|
||||
<collectionCode id="45E1C4C4564EFFBEFF43FCA4FEEBFCF4" box="[221,269,857,880]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
and twice in PBS. Subsequently, samples were stepped gradually into 70 % glycerol and eventually mounted in Fluoromount G (SouthernBiotech) on glass slides. The strong signal resulting from the epidermal cilia and mesodermal muscles made it impossible to unambiguously identify neural structures such as neurites in the tubulin and phalloidin stainings; yet, the detected cilia and muscles were used as positional markers. Analysis and digital image acquisition were performed on a Leica TCS SP5 II confocal laser scanning microscope (Leica Microsystems,
|
||||
<collectingCountry id="5BE71C91564EFFBEFDA7FB7EFD46FB1E" box="[569,672,1155,1178]" name="Germany" pageId="3" pageNumber="408">Germany</collectingCountry>
|
||||
). Optical sections were taken at a step size between 0.3 and 0.6 μm in Z-resolution. The resulting image stacks were digitally merged into maximum Z-projections and further processed with the open source image software
|
||||
<collectingCountry id="5BE71C91564EFFBEFDA1FAF5FD83FA9B" box="[575,613,1288,1311]" name="Fiji" pageId="3" pageNumber="408">Fiji</collectingCountry>
|
||||
(Max Planck Society for the Advancement of Science e.
|
||||
<collectionCode id="45E1C4C4564EFFBEFDF8FAD7FD9BFAC5" box="[614,637,1322,1345]" country="Canada" lsid="urn:lsid:biocol.org:col:13946" name="Royal British Columbia Museum - Herbarium" pageId="3" pageNumber="408" type="Museum">V</collectionCode>
|
||||
.,
|
||||
<collectingCountry id="5BE71C91564EFFBEFD0EFAD7FD1EFAC5" box="[656,760,1322,1345]" name="Germany" pageId="3" pageNumber="408">Germany</collectingCountry>
|
||||
), Adobe Photoshop, and Illustrator
|
||||
<collectionCode id="45E1C4C4564EFFBEFE77FAB6FDEDFAE6" box="[489,523,1355,1378]" country="France" name="Musee des Dinosaures d'Esperaza (Aude)" pageId="3" pageNumber="408">CS</collectionCode>
|
||||
3 (San José,
|
||||
<collectionCode id="45E1C4C4564EFFBEFD10FAB6FD53FAE6" box="[654,693,1355,1378]" country="USA" name="Chicago Academy of Sciences" pageId="3" pageNumber="408" type="Museum">CA</collectionCode>
|
||||
,
|
||||
<collectingCountry id="5BE71C91564EFFBEFD5EFAB6FD1EFAE6" box="[704,760,1355,1378]" name="United States of America" pageId="3" pageNumber="408">USA</collectingCountry>
|
||||
). At least ten specimens per stage were stained and scanned and rendered identical results for each staining per stage.
|
||||
</paragraph>
|
||||
<paragraph id="234F5C01564EFFBEFF38FA53FC9EFF7D" blockId="3.[136,772,159,1909]" lastBlockId="3.[816,1452,159,249]" pageId="3" pageNumber="408">
|
||||
Negative controls were performed by excluding either the primary or the secondary antibody. No signal was detected in any specimens of these experiments. In order to test for unspecific binding of the anti-serotonin (5-
|
||||
<collectionCode id="45E1C4C4564EFFBEFF38F9CEFF36F9CE" box="[166,208,1587,1610]" pageId="3" pageNumber="408">HT</collectionCode>
|
||||
) antibody, additional negative controls with preadsorbed antibodies were performed on developmental stages of
|
||||
<taxonomicName id="E4F02782564EFFBEFE82F98BFE5CF909" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[284,442,1654,1677]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="3" pageNumber="408" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564EFFBEFE82F98BFE5CF909" box="[284,442,1654,1677]" italics="true" pageId="3" pageNumber="408">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
and
|
||||
<taxonomicName id="E4F02782564EFFBEFE69F98BFD75F909" baseAuthorityName="Agassiz" baseAuthorityYear="1873" box="[503,659,1654,1677]" class="Enteropneusta" family="Harrimaniidae" genus="Saccoglossus" kingdom="Animalia" pageId="3" pageNumber="408" phylum="Hemichordata" rank="species" species="kowalevskii">
|
||||
<emphasis id="11848013564EFFBEFE69F98BFD75F909" box="[503,659,1654,1677]" italics="true" pageId="3" pageNumber="408">S. kowalevskii</emphasis>
|
||||
</taxonomicName>
|
||||
. For this, the rabbit anti-serotonin (5-
|
||||
<collectionCode id="45E1C4C4564EFFBEFE50F96AFE10F92A" box="[462,502,1687,1710]" pageId="3" pageNumber="408">HT</collectionCode>
|
||||
) antibody (polyclonal; ImmunoStar) was incubated in
|
||||
<collectionCode id="45E1C4C4564EFFBEFE62F945FDCBF94B" box="[508,557,1720,1743]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
blocking solution overnight at 4 °C together with serotonin (5-HT-)-BSA conjugate (ImmunoStar) reconstituted in
|
||||
<collectionCode id="45E1C4C4564EFFBEFDCCF907FD65F895" box="[594,643,1786,1809]" pageId="3" pageNumber="408">PBT</collectionCode>
|
||||
block with a final dilution of the antibody of 1:500 and a final concentration of the serotonin-BSA conjugate of 20 μg/ml. This solution was subsequently used as primary antibody solution according to the protocol described above, and none of the individuals showed any signal.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
96
data/AB/59/ED/AB59ED17564FFFBFFF16FF1FFD5BFC15.xml
Normal file
96
data/AB/59/ED/AB59ED17564FFFBFFF16FF1FFD5BFC15.xml
Normal file
|
|
@ -0,0 +1,96 @@
|
|||
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<mods:titleInfo id="F5394FC08EAE8A1D8B5599E5CF3DA819">
|
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<mods:title id="24B7FC6160311BBA136E8485DE670549">Neurogenesis in directly and indirectly developing enteropneusts: of nets and cords</mods:title>
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</mods:titleInfo>
|
||||
<mods:name id="24CF309E4846B498E0B4E7C95F0ECA22" type="personal">
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||||
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|
||||
<mods:roleTerm id="FBCD385F79AF1338EBE5457516E7E7E8">Author</mods:roleTerm>
|
||||
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|
||||
<mods:namePart id="12A60C51E50BE970466C67F222E9AB85">Wanninger, Sabrina Kaul-Strehlow Makoto Urata Takuya Minokawa Thomas Stach Andreas</mods:namePart>
|
||||
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|
||||
<mods:typeOfResource id="44191CA865E571BC6F2D41D849E4388A">text</mods:typeOfResource>
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|
||||
<mods:title id="25C8B2BD8968B9772B46C44E5C035A78">Organisms Diversity & Evolution</mods:title>
|
||||
</mods:titleInfo>
|
||||
<mods:part id="2ABCEF80BA3DBE265854BD9398605282">
|
||||
<mods:date id="57FF6BED8F1FC9B203584A930E61E2FA">2015</mods:date>
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||||
<mods:detail id="F936F1D38AD3A26B2C6B388CF18C408F" type="pubDate">
|
||||
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|
||||
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|
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|
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|
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|
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|
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|
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|
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|
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|
||||
<treatment id="AB59ED17564FFFBFFF16FF1FFD5BFC15" LSID="urn:lsid:plazi:treatment:AB59ED17564FFFBFFF16FF1FFD5BFC15" httpUri="http://treatment.plazi.org/id/AB59ED17564FFFBFFF16FF1FFD5BFC15" lastPageNumber="407" pageId="2" pageNumber="407">
|
||||
<subSubSection id="6BEA0F8A564FFFBFFF16FF1FFDA8FF7D" box="[136,590,226,249]" pageId="2" pageNumber="407" type="nomenclature">
|
||||
<paragraph id="234F5C01564FFFBFFF16FF1FFDA8FF7D" blockId="2.[136,590,226,249]" box="[136,590,226,249]" pageId="2" pageNumber="407">
|
||||
<heading id="7807EB6D564FFFBFFF16FF1FFDA8FF7D" box="[136,590,226,249]" fontSize="10" level="2" pageId="2" pageNumber="407" reason="7">
|
||||
<taxonomicName id="E4F02782564FFFBFFF16FF1FFDA8FF7D" ID-CoL="KFVG" authority="(Kuwano, 1902)" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[136,590,226,249]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFF16FF1FFE7BFF7D" box="[136,413,226,249]" italics="true" pageId="2" pageNumber="407">Balanoglossus misakiensis</emphasis>
|
||||
(Kuwano, 1902)
|
||||
</taxonomicName>
|
||||
</heading>
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
<subSubSection id="6BEA0F8A564FFFBFFF16FED9FD5BFC15" pageId="2" pageNumber="407" type="description">
|
||||
<paragraph id="234F5C01564FFFBFFF16FED9FD5BFC15" blockId="2.[136,772,292,913]" pageId="2" pageNumber="407">
|
||||
Adult
|
||||
<taxonomicName id="E4F02782564FFFBFFF55FED9FEB9FEBF" baseAuthorityName="Kuwano" baseAuthorityYear="1902" box="[203,351,292,315]" class="Enteropneusta" family="Ptychoderidae" genus="Balanoglossus" kingdom="Animalia" pageId="2" pageNumber="407" phylum="Hemichordata" rank="species" species="misakiensis">
|
||||
<emphasis id="11848013564FFFBFFF55FED9FEB9FEBF" box="[203,351,292,315]" italics="true" pageId="2" pageNumber="407">B. misakiensis</emphasis>
|
||||
</taxonomicName>
|
||||
were collected at a depth of
|
||||
<quantity id="E408F1E4564FFFBFFD0EFED9FD0EFEBF" box="[656,744,292,315]" metricMagnitude="0" metricUnit="m" metricValue="1.5" metricValueMax="2.0" metricValueMin="1.0" pageId="2" pageNumber="407" unit="m" value="1.5" valueMax="2.0" valueMin="1.0">1 to 2 m</quantity>
|
||||
at Sunset beach, Aomori-Bay, Asamushi,
|
||||
<collectingRegion id="E13492E3564FFFBFFDA2FEB8FD74FED8" box="[572,658,325,348]" country="Japan" name="Aomori" pageId="2" pageNumber="407">Aomori</collectingRegion>
|
||||
,
|
||||
<collectingCountry id="5BE71C91564FFFBFFD3EFEB8FD06FED8" box="[672,736,325,348]" name="Japan" pageId="2" pageNumber="407">Japan</collectingCountry>
|
||||
, in
|
||||
<date id="574E7AC1564FFFBFFF16FE9AFF1EFEFA" box="[136,248,359,382]" pageId="2" pageNumber="407" value="2012-06">June 2012</date>
|
||||
. Specimens were maintained in aquaria with running seawater at ambient water temperature (24–26 °C) at the Research Center for Marine Biology Tohoku University in Asamushi. Spawning of gravid females was induced by elevating the temperature to 31 °C, and insemination was carried out artificially (
|
||||
<bibRefCitation id="476121F0564FFFBFFEB2FDF0FDB7FDA0" author="Urata, M. & Yamaguchi, M." box="[300,593,525,548]" pageId="2" pageNumber="407" pagination="533 - 540" refId="ref14555" refString="Urata, M., & Yamaguchi, M. (2004). The Development of the Enteropneust Hemichordate Balanoglossus misakiensis KUWANO. Zoological Science, 21, 533 - 540." type="journal article" year="2004">
|
||||
Urata and
|
||||
<collectingRegion id="E13492E3564FFFBFFE01FDF0FDF2FDA0" box="[415,532,525,548]" country="Japan" name="Yamaguchi" pageId="2" pageNumber="407">Yamaguchi</collectingRegion>
|
||||
2004
|
||||
</bibRefCitation>
|
||||
). Embryos were cultured in membrane-filtered seawater in an incubator at 24 °C in Petri dishes without agitation. Hatched larvae were transferred into 5-l beakers at a concentration of
|
||||
<specimenCount id="35F69788564FFFBFFDF9FD8DFD58FD03" box="[615,702,624,647]" count="2" pageId="2" pageNumber="407" type="larva">2 larvae</specimenCount>
|
||||
/10 ml and were cultured with gentle stirring. From 2-day postfertilization (pf) onward, they were daily fed a mixture of the single-celled algae
|
||||
<taxonomicName id="E4F02782564FFFBFFEF4FD29FDDCFD6F" box="[362,570,724,747]" class="Bacillariophyceae" family="Chaetocerotaceae" genus="Chaetoceros" kingdom="Chromista" order="Chaetocerotales" pageId="2" pageNumber="407" phylum="Ochrophyta" rank="species" species="gracilis">
|
||||
<emphasis id="11848013564FFFBFFEF4FD29FDDCFD6F" box="[362,570,724,747]" italics="true" pageId="2" pageNumber="407">Chaetoceros gracilis</emphasis>
|
||||
</taxonomicName>
|
||||
,
|
||||
<taxonomicName id="E4F02782564FFFBFFDDBFD29FD33FD6F" box="[581,725,724,747]" class="Chlorophyceae" family="Dunaliellaceae" genus="Dunaliella" kingdom="Plantae" order="Chlamydomonadales" pageId="2" pageNumber="407" phylum="Chlorophyta" rank="species" species="undetermined">
|
||||
<emphasis id="11848013564FFFBFFDDBFD29FD57FD6F" box="[581,689,724,747]" italics="true" pageId="2" pageNumber="407">Dunaliella</emphasis>
|
||||
sp.
|
||||
</taxonomicName>
|
||||
, and
|
||||
<taxonomicName id="E4F02782564FFFBFFF16FD08FEF1FC88" box="[136,279,757,780]" class="Pavlovophyceae" family="Pavlovaceae" genus="Pavlova" kingdom="Chromista" order="Pavlovales" pageId="2" pageNumber="407" phylum="Haptophyta" rank="species" species="luteri">
|
||||
<emphasis id="11848013564FFFBFFF16FD08FEF1FC88" box="[136,279,757,780]" italics="true" pageId="2" pageNumber="407">Pavlova luteri</emphasis>
|
||||
</taxonomicName>
|
||||
. Most of the water in the beaker was replaced by fresh membrane-filtered seawater every day, and streptomycin (
|
||||
<quantity id="E408F1E4564FFFBFFF11FCCAFF32FCCA" box="[143,212,823,846]" metricMagnitude="-5" metricUnit="kg" metricValue="5.0" pageId="2" pageNumber="407" unit="mg" value="50.0">50 mg</quantity>
|
||||
/l) was added to prevent fungal growth. Agassiz-stage larvae (14 days pf) were cultured with sand grains from their natural habitat in Petri dishes to induce metamorphosis.
|
||||
</paragraph>
|
||||
</subSubSection>
|
||||
</treatment>
|
||||
</document>
|
||||
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Reference in a new issue