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<document id="145D14E68FCD129B1274BEE60D38FB48" ID-DOI="10.1016/j.jcz.2023.05.003" ID-ISSN="1873-2674" IM.bibliography_approvedBy="felipe" IM.illustrations_approvedBy="felipe" IM.materialsCitations_approvedBy="felipe" IM.metadata_approvedBy="felipe" IM.taxonomicNames_approvedBy="felipe" IM.treatments_approvedBy="felipe" checkinTime="1702540870477" checkinUser="felipe" docAuthor="Poddubnaya, Larisa G., Hemmingsen, Willy &amp; MacKenzie, Ken" docDate="2023" docId="03CA3B6FF331FFC4FCD4D55BFB47C5A3" docLanguage="en" docName="ZoolAnz.305.11-22.pdf" docOrigin="Zoologischer Anzeiger 305" docSource="http://dx.doi.org/10.1016/j.jcz.2023.05.003" docStyle="DocumentStyle:32483D6D135DE2FF927BA3F1400E0FFD.1:ZoolAnz.2022-.journal_article" docStyleId="32483D6D135DE2FF927BA3F1400E0FFD" docStyleName="ZoolAnz.2022-.journal_article" docStyleVersion="1" docTitle="Aporocotyle simplex" docType="treatment" docVersion="1" lastPageNumber="19" masterDocId="FFF34317F336FFCCFFE6D47AFFD2C425" masterDocTitle="Digestive system of the marine blood fluke, Aporocotyle simplex (Odhner, 1900) (Digenea: Aporocotylidae) with consideration of the digenean digestive morphology" masterLastPageNumber="22" masterPageNumber="11" pageNumber="18" updateTime="1702541034874" updateUser="GgImagineBatch">
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<mods:title id="5833E0965C3B5AC1469974F9CE77067B">Digestive system of the marine blood fluke, Aporocotyle simplex (Odhner, 1900) (Digenea: Aporocotylidae) with consideration of the digenean digestive morphology</mods:title>
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4.3. Esophageal ultrastructural features of
<taxonomicName id="4C63F1FAF331FFCBFB5ED55BFAC4C511" ID-CoL="67QN2" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[1208,1302,289,308]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="simplex">A. simplex</taxonomicName>
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Thirdly, our ultrastructural data allowed us to subdivide the long esophagus of
<taxonomicName id="4C63F1FAF331FFCBFC48D50FFBDBC5A2" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[942,1033,372,392]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF331FFCBFC48D50FFBDBC5A2" bold="true" box="[942,1033,372,392]" italics="true" pageId="7" pageNumber="18">A. simplex</emphasis>
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into two morphologically different parts - a short anterior and a long middle - posterior esophagus. The anterior esophageal portion resembles that of the body tegument and retains morphologically similar cytoplasmic inclusions and sunken perikarya, but differs in the absence of knob-like surface outgrowths and the presence of irregular, broad and angular cytoplasmic protrusions along the esophageal cytoplasmic lining. However, the middle-posterior esophageal portion retains only the basic tegumental cytoarchitecture and possesses considerable morphological modifications of the syncytial cytoplasmic lining by variations in the shape of long, bunched, extensive cytoplasmic protrusions, forming a reticular network through the esophageal lumen, and the sunken epithelial bodies are modified into well-developed esophageal glands. In
<emphasis id="B917566BF331FFCBFB7BD6B9FAD0C6F3" bold="true" box="[1181,1282,707,726]" italics="true" pageId="7" pageNumber="18">
<taxonomicName id="4C63F1FAF331FFCBFB7BD6B9FB2FC6F3" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[1181,1277,707,726]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="simplex">A. simplex</taxonomicName>
,
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however, based on a light microscopical description by
<bibRefCitation id="EFF2F788F331FFCBFB6DD6A5FAC1C6D7" author="Thulin, J." box="[1163,1299,735,754]" pageId="7" pageNumber="18" pagination="35 - 48" refId="ref12872" refString="Thulin, J., 1980. A redescription of the fish blood-fluke, Aporocotyle simplex Odhner, 1900 (Digenea, anguinicolidae) with comments on its biology. Sarsia 65, 35 - 48." type="journal article" year="1980">Thulin (1980)</bibRefCitation>
, the length of the esophagus in adult specimens is about 1/6 of the total body length and it may be divided into two parts, the first of which occupies 2/3 of the total esophageal length. Our TEM data supported another kind of esophageal morphological division, which corresponds to the data observed for schistosomatid blood flukes. As in
<taxonomicName id="4C63F1FAF331FFCBFB95D711FB02C758" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[1139,1232,874,894]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF331FFCBFB95D711FB02C758" bold="true" box="[1139,1232,874,894]" italics="true" pageId="7" pageNumber="18">A. simplex</emphasis>
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, the anterior portion of the esophagus in
<taxonomicName id="4C63F1FAF331FFCBFC54D7FCFBC1C7BC" authorityName="Sambon" authorityYear="1907" box="[946,1043,902,921]" class="Trematoda" family="Schistosomatidae" genus="Schistosoma" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="mansoni">
<emphasis id="B917566BF331FFCBFC54D7FCFBC1C7BC" bold="true" box="[946,1043,902,921]" italics="true" pageId="7" pageNumber="18">S. mansoni</emphasis>
</taxonomicName>
has tegumental inclusions identical to those of the external tegument, while in the middle and posterior portions the folded projections contain other cytoplasmic inclusions and the esophageal nucleated cell bodies form so-called esophageal glands (
<bibRefCitation id="EFF2F788F331FFCBFA8CD7A0FC08C02C" author="Morris, G. P. &amp; Threadgold, L. T." pageId="7" pageNumber="18" pagination="15 - 27" refId="ref11561" refString="Morris, G. P., Threadgold, L. T., 1968. Ultrastructure of the tegument of adult Schistosoma mansoni. J. Parasitol. 54, 15 - 27." type="journal article" year="1968">Morris and Threadgold, 1968</bibRefCitation>
;
<bibRefCitation id="EFF2F788F331FFCBFC01D78DFB87C02C" author="Ernst" box="[999,1109,1014,1034]" pageId="7" pageNumber="18" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst, 1975</bibRefCitation>
). Besides, in the freshwater adult aporocotylid blood fluke,
<taxonomicName id="4C63F1FAF331FFCBFC1BD068FB87C000" box="[1021,1109,1042,1061]" class="Trematoda" family="Aporocotylidae" genus="Sanguinicola" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="inermis">
<emphasis id="B917566BF331FFCBFC1BD068FB87C000" bold="true" box="[1021,1109,1042,1061]" italics="true" pageId="7" pageNumber="18">S. inermis</emphasis>
</taxonomicName>
, the esophagus is divided into three regions, distinguished by the morphology and arrangement of the apical cytoplasmic lining (
<bibRefCitation id="EFF2F788F331FFCBFC14D030FAF8C078" author="McMichael-Phillips, D. F. &amp; Lewis, J. W. &amp; Thorndyke, M. C." box="[1010,1322,1098,1117]" pageId="7" pageNumber="18" pagination="149 - 154" refId="ref11342" refString="McMichael-Phillips, D. F., Lewis, J. W., Thorndyke, M. C., 1994 a. Ultrastructure of the digestive system of adult Sanguinicola inermis. J. Helminthol. 68, 149 - 154." type="journal article" year="1994">McMichael-Phillips et al., 1994a</bibRefCitation>
, b). In the three abovementioned blood flukes, the esophageal epithelial cytoplasm consists of a number of morphological characters, which are unique for each studied species (
<bibRefCitation id="EFF2F788F331FFCBFBF3D0E4FA93C094" author="Morris, G. P. &amp; Threadgold, L. T." box="[1045,1345,1182,1201]" pageId="7" pageNumber="18" pagination="15 - 27" refId="ref11561" refString="Morris, G. P., Threadgold, L. T., 1968. Ultrastructure of the tegument of adult Schistosoma mansoni. J. Parasitol. 54, 15 - 27." type="journal article" year="1968">Morris and Threadgold, 1968</bibRefCitation>
,
<bibRefCitation id="EFF2F788F331FFCBFAB3D0E4FA1BC095" author="Ernst" box="[1365,1481,1181,1201]" pageId="7" pageNumber="18" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst, 1975</bibRefCitation>
;
<bibRefCitation id="EFF2F788F331FFCBFCD4D0C3FBA6C0E8" author="McMichael-Phillips, D. F. &amp; Lewis, J. W. &amp; Thorndyke, M. C." box="[818,1140,1209,1229]" pageId="7" pageNumber="18" pagination="149 - 154" refId="ref11342" refString="McMichael-Phillips, D. F., Lewis, J. W., Thorndyke, M. C., 1994 a. Ultrastructure of the digestive system of adult Sanguinicola inermis. J. Helminthol. 68, 149 - 154." type="journal article" year="1994">McMichael-Phillips et al., 1994a</bibRefCitation>
; Present study). Generally, in the
<taxonomicName id="4C63F1FAF331FFCBFCD4D0AFFC4AC0CD" authorityName="Rudolphi" authorityYear="1808" box="[818,920,1237,1256]" class="Trematoda" kingdom="Animalia" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="class">Trematoda</taxonomicName>
the esophagus varies in the fine morphology from group to group. In the trematode
<emphasis id="B917566BF331FFCBFBF2D08BFB3EC121" bold="true" box="[1044,1260,1265,1284]" italics="true" pageId="7" pageNumber="18">
<taxonomicName id="4C63F1FAF331FFCBFBF2D08BFB3AC121" authorityName="von Baer" authorityYear="1827" box="[1044,1256,1265,1284]" class="Trematoda" family="Aspidogastridae" genus="Aspidogaster" kingdom="Animalia" order="Aspidogastrida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="conchicola">Aspidogaster conchicola</taxonomicName>
,
</emphasis>
the short esophagus has tegumental traits with a number of uniciliate sensory endings (
<bibRefCitation id="EFF2F788F331FFCBFA6FD177FCB0C119" author="Halton, D. W." pageId="7" pageNumber="18" pagination="455 - 467" refId="ref10787" refString="Halton, D. W., 1972. Ultrastructure of the alimentary tract of Aspidogaster conchicola (Trematoda: aspidogastrea). J. Parasitol. 58, 455 - 467." type="journal article" year="1972">
<collectingRegion id="49A7449BF331FFCBFA6FD177FA19C105" box="[1417,1483,1293,1312]" country="United Kingdom" name="Halton" pageId="7" pageNumber="18">Halton</collectingRegion>
, 1972
</bibRefCitation>
). In a plagiorchiid digenean of the family
<taxonomicName id="4C63F1FAF331FFCBFAE1D153FA19C119" box="[1287,1483,1321,1340]" class="Trematoda" family="Paramphistomatidae" higherTaxonomySource="GBIF" kingdom="Animalia" order="Plagiorchiida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="family">Paramphistomatidae</taxonomicName>
,
<taxonomicName id="4C63F1FAF331FFCBFCD4D13FFBF1C17D" box="[818,1059,1349,1368]" class="Trematoda" family="Paramphistomidae" genus="Paramphistomum" kingdom="Animalia" order="Plagiorchiida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="epiclitum">
<emphasis id="B917566BF331FFCBFCD4D13FFBF1C17D" bold="true" box="[818,1059,1349,1368]" italics="true" pageId="7" pageNumber="18">Paramphistomum epiclitum</emphasis>
</taxonomicName>
, the esophageal lining has irregular corrugations with flattened ridges and secretory esophageal cytons (
<bibRefCitation id="EFF2F788F331FFCBFA9BD11BFC4FC1B5" author="Mattison, R. G. &amp; Hanna, R. E. B. &amp; Nizami, W. A." pageId="7" pageNumber="18" pagination="1089 - 1101" refId="ref11264" refString="Mattison, R. G., Hanna, R. E. B., Nizami, W. A., 1992. Ultrastructure and histochemistry of the digestive tract of juvenile Paramphistomum epiclitum (Paramphistomidae: Digenea) during migration in Indian ruminants. Int. J. Parasitol. 22, 1089 - 1101. https: // doi. org / 10.1016 / 0020 - 7519 (92) 90029 - k." type="journal article" year="1992">Mattison et al., 1992</bibRefCitation>
). Another plagiorchiid digenean of the family
<taxonomicName id="4C63F1FAF331FFCBFABED107FC87C189" authorityName="Fukui" authorityYear="1929" class="Trematoda" family="Gyliauchenidae" kingdom="Animalia" order="Plagiorchiida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="family">Gyliauchenidae</taxonomicName>
,
<taxonomicName id="4C63F1FAF331FFCBFC85D1E2FBF5C18E" authorityName="Ozaki" authorityYear="1937" box="[867,1063,1432,1451]" class="Trematoda" family="Gyliauchenidae" genus="Gyliauchen" kingdom="Animalia" order="Plagiorchiida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="species" species="nahaensis">
<emphasis id="B917566BF331FFCBFC85D1E2FBF5C18E" bold="true" box="[867,1063,1432,1451]" italics="true" pageId="7" pageNumber="18">Gyliauchen nahaensis</emphasis>
</taxonomicName>
, has three morphologically distinguishable esophageal parts ornamented with rugae-like surface projections and secretory esophageal sunken cytons (
<bibRefCitation id="EFF2F788F331FFCBFB77D1ABFAEAC1C6" author="Jones, M. K. &amp; Hughes-Stamm, S. R. &amp; East, R. M. &amp; Cribb, T. H." box="[1169,1336,1488,1508]" pageId="7" pageNumber="18" pagination="198 - 211" refId="ref10951" refString="Jones, M. K., Hughes-Stamm, S. R., East, R. M., Cribb, T. H., 2000. Ultrastructure of the digestive tract of Gyliauchen nahaensis (Platyhelminthes, Digenea), an inhabitant of the hindgut of herbivorous fishes. J. Morphol. 246, 198 - 211. https: // doi. org / 10.1002 / 1097 - 4687 (200012) 246: 3 &lt;198 :: aid-jmor 4&gt; 3.0. co; 2 - 2." type="journal article" year="2000">Jones et al., 2000</bibRefCitation>
). Aporocotylids have a diet of dissolved matter in the blood fluid surrounding them. Interestingly, in the case of plagiorchiid gyliauchenids and paramphistomids with fluid diet (see
<bibRefCitation id="EFF2F788F331FFCBFB9DD25EFA9BC212" author="Mattison, R. G. &amp; Hanna, R. E. B. &amp; Nizami, W. A." box="[1147,1353,1572,1592]" pageId="7" pageNumber="18" pagination="1089 - 1101" refId="ref11264" refString="Mattison, R. G., Hanna, R. E. B., Nizami, W. A., 1992. Ultrastructure and histochemistry of the digestive tract of juvenile Paramphistomum epiclitum (Paramphistomidae: Digenea) during migration in Indian ruminants. Int. J. Parasitol. 22, 1089 - 1101. https: // doi. org / 10.1016 / 0020 - 7519 (92) 90029 - k." type="journal article" year="1992">Mattison et al., 1992</bibRefCitation>
;
<bibRefCitation id="EFF2F788F331FFCBFABED25FFCB0C276" author="Jones, M. K. &amp; Hughes-Stamm, S. R. &amp; East, R. M. &amp; Cribb, T. H." pageId="7" pageNumber="18" pagination="198 - 211" refId="ref10951" refString="Jones, M. K., Hughes-Stamm, S. R., East, R. M., Cribb, T. H., 2000. Ultrastructure of the digestive tract of Gyliauchen nahaensis (Platyhelminthes, Digenea), an inhabitant of the hindgut of herbivorous fishes. J. Morphol. 246, 198 - 211. https: // doi. org / 10.1002 / 1097 - 4687 (200012) 246: 3 &lt;198 :: aid-jmor 4&gt; 3.0. co; 2 - 2." type="journal article" year="2000">Jones et al., 2000</bibRefCitation>
),
<bibRefCitation id="EFF2F788F331FFCBFC93D23AFBF6C276" author="Jones, M. K. &amp; Hughes-Stamm, S. R. &amp; East, R. M. &amp; Cribb, T. H." box="[885,1060,1600,1619]" pageId="7" pageNumber="18" pagination="198 - 211" refId="ref10951" refString="Jones, M. K., Hughes-Stamm, S. R., East, R. M., Cribb, T. H., 2000. Ultrastructure of the digestive tract of Gyliauchen nahaensis (Platyhelminthes, Digenea), an inhabitant of the hindgut of herbivorous fishes. J. Morphol. 246, 198 - 211. https: // doi. org / 10.1002 / 1097 - 4687 (200012) 246: 3 &lt;198 :: aid-jmor 4&gt; 3.0. co; 2 - 2." type="journal article" year="2000">Jones et al. (2000)</bibRefCitation>
suggested that there has been a succession of innovations in the evolution of the feeding of gyliauchenids. Their oral sucker was lost first as a change to a fluid-based diet occurred, and the esophagus lengthened and adopted a major role in the digestion of the food (
<bibRefCitation id="EFF2F788F331FFCBFC8DD2CAFBC6C2E6" author="Jones, M. K. &amp; Hughes-Stamm, S. R. &amp; East, R. M. &amp; Cribb, T. H." box="[875,1044,1711,1731]" pageId="7" pageNumber="18" pagination="198 - 211" refId="ref10951" refString="Jones, M. K., Hughes-Stamm, S. R., East, R. M., Cribb, T. H., 2000. Ultrastructure of the digestive tract of Gyliauchen nahaensis (Platyhelminthes, Digenea), an inhabitant of the hindgut of herbivorous fishes. J. Morphol. 246, 198 - 211. https: // doi. org / 10.1002 / 1097 - 4687 (200012) 246: 3 &lt;198 :: aid-jmor 4&gt; 3.0. co; 2 - 2." type="journal article" year="2000">Jones et al., 2000</bibRefCitation>
). The above suggestion by
<bibRefCitation id="EFF2F788F331FFCBFAFAD2CAFA1DC2E6" author="Jones, M. K. &amp; Hughes-Stamm, S. R. &amp; East, R. M. &amp; Cribb, T. H." box="[1308,1487,1711,1731]" pageId="7" pageNumber="18" pagination="198 - 211" refId="ref10951" refString="Jones, M. K., Hughes-Stamm, S. R., East, R. M., Cribb, T. H., 2000. Ultrastructure of the digestive tract of Gyliauchen nahaensis (Platyhelminthes, Digenea), an inhabitant of the hindgut of herbivorous fishes. J. Morphol. 246, 198 - 211. https: // doi. org / 10.1002 / 1097 - 4687 (200012) 246: 3 &lt;198 :: aid-jmor 4&gt; 3.0. co; 2 - 2." type="journal article" year="2000">Jones et al. (2000)</bibRefCitation>
may also be true for aporocotylid digeneans. As mentioned above, the absence of an oral sucker is not characteristic of all members of the family
<taxonomicName id="4C63F1FAF331FFCBFC95D379FBD4C333" authorityName="Odhner" authorityYear="1912" box="[883,1030,1795,1814]" class="Trematoda" family="Aporocotylidae" kingdom="Animalia" order="Diplostomida" pageId="7" pageNumber="18" phylum="Platyhelminthes" rank="family">Aporocotylidae</taxonomicName>
. Some adult fish blood flukes retain the cercarial anterior sucker, but seemingly lack a pharynx (
<bibRefCitation id="EFF2F788F331FFCBFB15D365FCB0C36B" author="Bullard, S. A. &amp; Overstreet, R. M." pageId="7" pageNumber="18" pagination="97 - 113" refId="ref10056" refString="Bullard, S. A., Overstreet, R. M., 2003. Elaphrobates euzeti gen. and sp. n. (Digenea: Sanguinicolidae) from snappers (Lutjanidae) in the Gulf of Mexico. In: Tome, I., Combes, C., Jourdane, J. (Eds.), Taxonomie,´ecologie et evolution des m´etazoaires parasites. Taxonomy, ecology and evolution of metazoan parasites. PUP Perpignan, pp. 97 - 113." type="book chapter" year="2003">Bullard and Overstreet, 2003</bibRefCitation>
;
<bibRefCitation id="EFF2F788F331FFCBFC88D341FC4CC36B" author="Bullard, S. A. &amp; Overstreet, R. M." box="[878,926,1851,1870]" pageId="7" pageNumber="18" pagination="128 - 136" refId="ref10146" refString="Bullard, S. A., Overstreet, R. M., 2004. Two new species of cardiocola (Digenea: Sanguinicolidae) in drums (sciaenidae) from Mississippi and Louisiana. J. Partasitol. 90, 128 - 136. https: // doi. org / 10.1645 / ge- 106 r." type="journal article" year="2004">2004</bibRefCitation>
;
<bibRefCitation id="EFF2F788F331FFCBFC4CD341FB8FC36B" author="McVay, M. J. &amp; Bakenhaster, M. D. &amp; Bullard, S. A." box="[938,1117,1851,1870]" pageId="7" pageNumber="18" pagination="291 - 305" refId="ref11439" refString="McVay, M. J., Bakenhaster, M. D., Bullard, S. A., 2011. Cardicola laruei Short, 1953 (Digenea: aporocotylisdae) from heart of seatrout, Cynoscion spp., (Perciformes: sciaenidae) in the Gulf of Mexico and Atlantic Ocean: taxonomic redescription, first observations of egg and miracidium, and comments on geographic distribution and host specificity. Comp. Parasitol. 78, 291 - 305. https: // doi. org / 10.1654 / 4501.1." type="journal article" year="2011">McVay et al., 2011</bibRefCitation>
).
</paragraph>
<paragraph id="8BDC8A79F331FFC4FCB7D32DFD36C173" blockId="7.[818,1488,345,1982]" lastBlockId="8.[100,770,148,1980]" lastPageId="8" lastPageNumber="19" pageId="7" pageNumber="18">
For schistosomatid blood flukes, it has been suggested that the digestion of host erythrocytes commences in the esophageal region of the digestive tract, and that the final phases of digestion occur in the lumen of the intestine (Morris, 1968;
<bibRefCitation id="EFF2F788F331FFCBFB77D3D1FB2FC398" author="Ernst" box="[1169,1277,1962,1982]" pageId="7" pageNumber="18" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst, 1975</bibRefCitation>
). Schistosomes have a glandular esophagus, and secretory cells are modified sunken epithelial bodies, which are a source of digestive enzymes, and digestion of blood cells begins in the esophagus (
<bibRefCitation id="EFF2F788F33EFFC4FE62D4B6FE23C4FB" author="Ernst" box="[388,497,203,223]" pageId="8" pageNumber="19" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst, 1975</bibRefCitation>
;
<bibRefCitation id="EFF2F788F33EFFC4FE1BD4B1FDABC4FA" author="Halton, D. W." box="[509,633,203,223]" pageId="8" pageNumber="19" pagination="693 - 704" refId="ref10860" refString="Halton, D. W., 1997. Nutritional adaptations to parasitism within the Platyhelminthes. Int. J. Parasitol. 27, 693 - 704. https: // doi. org / 10.1016 / s 0020 - 7519 (97) 00011 - 8." type="journal article" year="1997">
<collectingRegion id="49A7449BF33EFFC4FE1BD4B1FDEDC4FB" box="[509,575,203,222]" country="United Kingdom" name="Halton" pageId="8" pageNumber="19">Halton</collectingRegion>
, 1997
</bibRefCitation>
). Present data on the
<taxonomicName id="4C63F1FAF33EFFC4FF4ED49DFED4C4DF" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[168,262,231,250]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FF4ED49DFED4C4DF" bold="true" box="[168,262,231,250]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
esophagus are in complete agreement with those obtained for schistosomes. As shown in our study, the esophageal morphological features of
<taxonomicName id="4C63F1FAF33EFFC4FE41D565FDCCC517" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[423,542,287,306]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FE41D565FDCCC517" bold="true" box="[423,542,287,306]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
might suggest a digestive-absorptive function for the middle-posterior esophageal region surrounded by compact cellular complex, in which sunken glandular perikarya of different development stages and their cytoplasmic processes mix with muscle cells as well as with extensive nerve fibres. In
<taxonomicName id="4C63F1FAF33EFFC4FF82D5D1FF13C598" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[100,193,426,446]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FF82D5D1FF13C598" bold="true" box="[100,193,426,446]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
each esophageal secretory granule has heterogeneous content with a dense thin outer halo and less dense inner core. Such granules are surrounded by the lucent or moderately dense matrix of the vacuole, within which each granule is embedded. Such a glandular appearance shows morphological similarity with digestive vacuoles described in the gastrodermal digestive cells of polyopisthocotylean monogeneans (
<bibRefCitation id="EFF2F788F33EFFC4FF8AD628FE9CC640" author="Poddubnaya, L. G. &amp; Hemmingsen, W. &amp; Reed, C. &amp; Gibson, D. I." box="[108,334,594,613]" pageId="8" pageNumber="19" pagination="2599 - 2610" refId="ref12137" refString="Poddubnaya, L. G., Hemmingsen, W., Reed, C., Gibson, D. I., 2015. Ultrastructural characteristics of the caeca of basal polyopisthocotylean monogeneans of the families Chimaericolidae and Hexabothriidae parasitic on cartilaginous fishes. Parasitol. Res. 114, 2599 - 2610. https: // doi. org / 10.1007 / s 00436 - 015 - 4464 - 5." type="journal article" year="2015">Poddubnaya et al., 2015</bibRefCitation>
;
<bibRefCitation id="EFF2F788F33EFFC4FEBFD628FD80C640" author="Cable, J. &amp; El-Naggar, M. M." box="[345,594,594,613]" pageId="8" pageNumber="19" pagination="3181 - 3193" refId="ref10343" refString="Cable, J., El-Naggar, M. M., 2021. Gastrodermis ultrastructure of the different life stages of the polyopisthocotylean monogenean gill parasite Discocotyle sagittata. Parasitol. Res. 120, 3181 - 3193. https: // doi. org / 10.1007 / s 00436 - 021 - 07286 - 6." type="journal article" year="2021">Cable and El-Naggar, 2021</bibRefCitation>
). In the cercaria of the freshwater aporocotylid,
<taxonomicName id="4C63F1FAF33EFFC4FE9FD614FE03C6A4" box="[377,465,622,641]" class="Trematoda" family="Aporocotylidae" genus="Sanguinicola" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="inermis">
<emphasis id="B917566BF33EFFC4FE9FD614FE03C6A4" bold="true" box="[377,465,622,641]" italics="true" pageId="8" pageNumber="19">S. inermis</emphasis>
</taxonomicName>
, the esophageal secretory granules are similar to those of
<taxonomicName id="4C63F1FAF33EFFC4FEBFD6F0FE66C6B9" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[345,436,649,669]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FEBFD6F0FE66C6B9" bold="true" box="[345,436,649,669]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
, but there is no information on their vacuolar appearance (
<bibRefCitation id="EFF2F788F33EFFC4FED2D6DCFDB1C69C" author="McMichael-Phillips, D. F. &amp; Lewis, J. W. &amp; Thorndyke, M. C." box="[308,611,678,697]" pageId="8" pageNumber="19" pagination="149 - 154" refId="ref11342" refString="McMichael-Phillips, D. F., Lewis, J. W., Thorndyke, M. C., 1994 a. Ultrastructure of the digestive system of adult Sanguinicola inermis. J. Helminthol. 68, 149 - 154." type="journal article" year="1994">McMichael-Phillips et al., 1994a</bibRefCitation>
, b). However, in schistosomatid
<emphasis id="B917566BF33EFFC4FF1DD6BBFEB8C6F1" bold="true" box="[251,362,705,724]" italics="true" pageId="8" pageNumber="19">
<taxonomicName id="4C63F1FAF33EFFC4FF1DD6BBFEB7C6F1" authorityName="Sambon" authorityYear="1907" box="[251,357,705,724]" class="Trematoda" family="Schistosomatidae" genus="Schistosoma" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="mansoni">S. mansoni</taxonomicName>
,
</emphasis>
esophageal secretory granules of varied shape display a highly structured morphology, each granule consisting of a dark substance with crystalline structure (
<bibRefCitation id="EFF2F788F33EFFC4FDC7D683FF46C70D" author="Morris, G. P. &amp; Threadgold, L. T." pageId="8" pageNumber="19" pagination="15 - 27" refId="ref11561" refString="Morris, G. P., Threadgold, L. T., 1968. Ultrastructure of the tegument of adult Schistosoma mansoni. J. Parasitol. 54, 15 - 27." type="journal article" year="1968">Morris and Threadgold, 1968</bibRefCitation>
) or with rays of tubules (
<bibRefCitation id="EFF2F788F33EFFC4FE44D76FFD4FC70D" author="Bogitsh, B. J. &amp; Carter, O. S." box="[418,669,789,808]" pageId="8" pageNumber="19" pagination="681 - 686" refId="ref9713" refString="Bogitsh, B. J., Carter, O. S., 1977. Schistosoma mansoni: ultrastructural studies on the esophageal secretory granules. J. Parasitol. 63, 681 - 686. https: // doi. org / 10.2307 / 3279570." type="journal article" year="1977">Bogitsh and Carter, 1977</bibRefCitation>
). Usually, particular attention has been concentrated on the cytochemical composition of the esophageal secretory product in
<taxonomicName id="4C63F1FAF33EFFC4FDBED737FD69C745" authorityName="Sambon" authorityYear="1907" box="[600,699,845,864]" class="Trematoda" family="Schistosomatidae" genus="Schistosoma" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="mansoni">
<emphasis id="B917566BF33EFFC4FDBED737FD69C745" bold="true" box="[600,699,845,864]" italics="true" pageId="8" pageNumber="19">S. mansoni</emphasis>
</taxonomicName>
(
<bibRefCitation id="EFF2F788F33EFFC4FD2DD737FF46C759" author="Ernst" pageId="8" pageNumber="19" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst, 1975</bibRefCitation>
;
<bibRefCitation id="EFF2F788F33EFFC4FF44D713FE41C759" author="Bogitsh, B. J. &amp; Carter, O. S." box="[162,403,873,892]" pageId="8" pageNumber="19" pagination="681 - 686" refId="ref9713" refString="Bogitsh, B. J., Carter, O. S., 1977. Schistosoma mansoni: ultrastructural studies on the esophageal secretory granules. J. Parasitol. 63, 681 - 686. https: // doi. org / 10.2307 / 3279570." type="journal article" year="1977">Bogitsh and Carter, 1977</bibRefCitation>
).
<bibRefCitation id="EFF2F788F33EFFC4FE4FD713FDF3C759" author="Ernst" box="[425,545,873,892]" pageId="8" pageNumber="19" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst (1975)</bibRefCitation>
indicated that the posterior esophageal portion of
<taxonomicName id="4C63F1FAF33EFFC4FE93D7FFFE05C7BD" authorityName="Sambon" authorityYear="1907" box="[373,471,901,920]" class="Trematoda" family="Schistosomatidae" genus="Schistosoma" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="mansoni">
<emphasis id="B917566BF33EFFC4FE93D7FFFE05C7BD" bold="true" box="[373,471,901,920]" italics="true" pageId="8" pageNumber="19">S. mansoni</emphasis>
</taxonomicName>
contains substantial acid phosphatase activity both in the lining of the esophageal lumen and in the sunken perikarya connected to this lining. The reaction product in this region occurs in some membrane-bound vesicles and in the basal infoldings of the cytoplasmic lining. These acid-phosphatase-positive vesicles are found in the esophageal cell bodies and in the Golgi complexes and granular endoplasmic reticulum, suggesting that these proteins are being synthesized and packaged in the cell bodies and then transported to the esophageal cytoplasmic lining. Also, the cytochemical results of
<bibRefCitation id="EFF2F788F33EFFC4FF27D0FAFE6AC0B6" author="Bogitsh, B. J. &amp; Carter, O. S." box="[193,440,1152,1171]" pageId="8" pageNumber="19" pagination="681 - 686" refId="ref9713" refString="Bogitsh, B. J., Carter, O. S., 1977. Schistosoma mansoni: ultrastructural studies on the esophageal secretory granules. J. Parasitol. 63, 681 - 686. https: // doi. org / 10.2307 / 3279570." type="journal article" year="1977">Bogitsh and Carter (1977)</bibRefCitation>
have shown that esophageal granules contain carbohydrate-containing endopeptidase, which is capable of digesting hemoglobin. Biochemical and molecular data indicated proteolytic pathways of haemoglobin digestion in schistosomes (
<bibRefCitation id="EFF2F788F33EFFC4FD5AD0AEFF46C126" author="Halton, D. W." pageId="8" pageNumber="19" pagination="693 - 704" refId="ref10860" refString="Halton, D. W., 1997. Nutritional adaptations to parasitism within the Platyhelminthes. Int. J. Parasitol. 27, 693 - 704. https: // doi. org / 10.1016 / s 0020 - 7519 (97) 00011 - 8." type="journal article" year="1997">
<collectingRegion id="49A7449BF33EFFC4FD5AD0AEFD2CC0C2" box="[700,766,1236,1255]" country="United Kingdom" name="Halton" pageId="8" pageNumber="19">Halton</collectingRegion>
, 1997
</bibRefCitation>
). Immunocytochemistry has shown that the complex of hydrolytic proteinases (cysteine endopeptodases) is involved in the sequential degradation of haemoglobin to readily absorbable peptides, all of which are expressed in the schistosome gastrodermis (
<bibRefCitation id="EFF2F788F33EFFC4FDC0D139FD04C173" author="Dalton, J. P. &amp; Smith, A. M. &amp; Clough, K. A. &amp; Brindley, P. J." box="[550,726,1347,1367]" pageId="8" pageNumber="19" pagination="299 - 303" refId="ref10532" refString="Dalton, J. P., Smith, A. M., Clough, K. A., Brindley, P. J., 1995. Digestion a haemoglobin by schistosomes: 35 years on. Parasitol. Today 11, 299 - 303. https: // doi. org / 10.1016 / 0169 - 4758 (95) 80045 - x." type="journal article" year="1995">Dalton et al., 1995</bibRefCitation>
).
</paragraph>
<paragraph id="8BDC8A79F33EFFC4FF62D125FE1FC210" blockId="8.[100,770,148,1980]" pageId="8" pageNumber="19">
The abovementioned cytochemical data for
<taxonomicName id="4C63F1FAF33EFFC4FDC3D125FD55C157" authorityName="Sambon" authorityYear="1907" box="[549,647,1375,1394]" class="Trematoda" family="Schistosomatidae" genus="Schistosoma" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="mansoni">
<emphasis id="B917566BF33EFFC4FDC3D125FD55C157" bold="true" box="[549,647,1375,1394]" italics="true" pageId="8" pageNumber="19">S. mansoni</emphasis>
</taxonomicName>
is supported by our morphological data on the
<taxonomicName id="4C63F1FAF33EFFC4FE24D101FDF6C1AB" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[450,548,1403,1422]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FE24D101FDF6C1AB" bold="true" box="[450,548,1403,1422]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
esophagus. In studied specimens of
<taxonomicName id="4C63F1FAF33EFFC4FF0ED1EDFE9AC18F" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[232,328,1431,1450]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FF0ED1EDFE9AC18F" bold="true" box="[232,328,1431,1450]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
, clots of host blood cells in various stages of decay were found in close proximity to epithelial cytoplasmic protrusions within the lumen of the posterior esophagus. As a result of blood cell disintegration, the conglomerations of moderately dense substance, in which small vesicles and dense formless inclusions are embedded, may be seen in the esophageal lumen.
</paragraph>
<paragraph id="8BDC8A79F33EFFC4FF62D244FD21C3A1" blockId="8.[100,770,148,1980]" pageId="8" pageNumber="19">
Additional data by
<bibRefCitation id="EFF2F788F33EFFC4FEDBD245FE67C274" author="Ernst" box="[317,437,1598,1618]" pageId="8" pageNumber="19" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst (1975)</bibRefCitation>
shows that, like the tegument, the anterior portion of the esophagus contains negligible cytochemically demonstrable acid phosphatase activity. Contrary data by
<bibRefCitation id="EFF2F788F33EFFC4FD65D20CFCD0C2AC" author="Cesari, I. M." box="[643,770,1654,1673]" pageId="8" pageNumber="19" pagination="405 - 414" refId="ref10401" refString="Cesari, I. M., 1974. Schistosoma mansoni: distribution and characteristics of alkaline and acid phosphatase. Exp. Parasitol. 36, 405 - 414." type="journal article" year="1974">Cesari (1974)</bibRefCitation>
show that most of the acid phosphatase activity in
<taxonomicName id="4C63F1FAF33EFFC4FDAAD2E8FD7CC280" authorityName="Sambon" authorityYear="1907" box="[588,686,1682,1701]" class="Trematoda" family="Schistosomatidae" genus="Schistosoma" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="mansoni">
<emphasis id="B917566BF33EFFC4FDAAD2E8FD7CC280" bold="true" box="[588,686,1682,1701]" italics="true" pageId="8" pageNumber="19">S. mansoni</emphasis>
</taxonomicName>
males is located in the epidermis, but in the opinion of
<bibRefCitation id="EFF2F788F33EFFC4FDCBD2D4FD77C2E4" author="Ernst" box="[557,677,1710,1729]" pageId="8" pageNumber="19" pagination="633 - 647" refId="ref10734" refString="Ernst, S. C., 1975. Biochemical and cytochemical studies of digestive-absorptive functions of esophagus, caecum and tegument in Schistosoma mansoni: acid phosphatase and tracer studies. J. Parasitol. 61, 633 - 647. https: // doi. org / 10.2307 / 3279456." type="journal article" year="1975">Ernst (1975)</bibRefCitation>
the techniques employed by
<bibRefCitation id="EFF2F788F33EFFC4FEC5D2B0FE70C2F8" author="Cesari, I. M." box="[291,418,1738,1757]" pageId="8" pageNumber="19" pagination="405 - 414" refId="ref10401" refString="Cesari, I. M., 1974. Schistosoma mansoni: distribution and characteristics of alkaline and acid phosphatase. Exp. Parasitol. 36, 405 - 414." type="journal article" year="1974">Cesari (1974)</bibRefCitation>
do not allow the determination of the original cytological site of the soluble enzyme activity measured in the supernatant fractions. Also,
<bibRefCitation id="EFF2F788F33EFFC4FE8AD378FE26C330" author="Pappas, P. W." box="[364,500,1794,1813]" pageId="8" pageNumber="19" pagination="105 - 121" refId="ref11960" refString="Pappas, P. W., 1988. The relative roles of the intestines and external surfaces in the nutrition of monogeneans, digeneans and nematodes. Parasitology 96, 105 - 121. https: // doi. org / 10.1017 / s 0031182000086005." type="journal article" year="1988">Pappas (1988)</bibRefCitation>
suggested that it is basically correct that the tegument is the primary source of nutrients for adult digeneans during short-term
<emphasis id="B917566BF33EFFC4FE9FD343FE69C369" bold="true" box="[377,443,1849,1868]" italics="true" pageId="8" pageNumber="19">in vitro</emphasis>
incubations, but that “trematodes do not readily feed under artificial (
<emphasis id="B917566BF33EFFC4FE5CD32FFE28C34D" bold="true" box="[442,506,1877,1896]" italics="true" pageId="8" pageNumber="19">in vitro</emphasis>
) conditions, so the value of such experiments remains questionable” (see
<bibRefCitation id="EFF2F788F33EFFC4FDF4D308FD42C3A1" author="Pappas, P. W." box="[530,656,1905,1925]" pageId="8" pageNumber="19" pagination="105 - 121" refId="ref11960" refString="Pappas, P. W., 1988. The relative roles of the intestines and external surfaces in the nutrition of monogeneans, digeneans and nematodes. Parasitology 96, 105 - 121. https: // doi. org / 10.1017 / s 0031182000086005." type="journal article" year="1988">Pappas, 1988</bibRefCitation>
, p. S110).
</paragraph>
<paragraph id="8BDC8A79F33EFFC4FF62D3F7FB47C5A3" blockId="8.[100,770,148,1980]" lastBlockId="8.[818,1488,148,390]" pageId="8" pageNumber="19">
Additionally, besides differentiated esophageal secretory perikarya, there are undifferentiated cells, cells at the beginning stage of their differentiation and disintegrated cells in the compact cellular masses surrounding the middle and posterior esophagus of
<taxonomicName id="4C63F1FAF33EFFC4FAC5D4CAFA53C4E7" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[1315,1409,175,195]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FAC5D4CAFA53C4E7" bold="true" box="[1315,1409,175,195]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
. Taking into account that esophageal glandular cells are considered to be an initial site of blood processing, with blood cells being lysed as they pass through the esophagus, the new esophageal glandular cells should be produced prior to the initiation of blood feeding and be accompanied by apoptosis of utilized esophageal glands. The presence of extensive nerve fibres supporting the foregut of
<taxonomicName id="4C63F1FAF33EFFC4FB87D52DFB6DC54F" baseAuthorityName="Odhner" baseAuthorityYear="1900" box="[1121,1215,343,362]" class="Trematoda" family="Aporocotylidae" genus="Aporocotyle" kingdom="Animalia" order="Diplostomida" pageId="8" pageNumber="19" phylum="Platyhelminthes" rank="species" species="simplex">
<emphasis id="B917566BF33EFFC4FB87D52DFB6DC54F" bold="true" box="[1121,1215,343,362]" italics="true" pageId="8" pageNumber="19">A. simplex</emphasis>
</taxonomicName>
suggests that such processes are controlled by the nervous system.
</paragraph>
</subSubSection>
</treatment>
</document>