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On the Structure and Function of the Digestive System of the Nudibranch Mollusc Acanthodoris Pilosa M Patricia Morse

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On the Structure and Function of the Digestive System of the Nudibranch Mollusc Acanthodoris Pilosa M Patricia Morse University of New Hampshire University of New Hampshire Scholars' Repository Doctoral Dissertations Student Scholarship Spring 1966 ON THE STRUCTURE AND FUNCTION OF THE DIGESTIVE SYSTEM OF THE NUDIBRANCH MOLLUSC ACANTHODORIS PILOSA M PATRICIA MORSE Follow this and additional works at: https://scholars.unh.edu/dissertation Recommended Citation MORSE, M PATRICIA, "ON THE STRUCTURE AND FUNCTION OF THE DIGESTIVE SYSTEM OF THE NUDIBRANCH MOLLUSC ACANTHODORIS PILOSA" (1966). Doctoral Dissertations. 837. https://scholars.unh.edu/dissertation/837 This Dissertation is brought to you for free and open access by the Student Scholarship at University of New Hampshire Scholars' Repository. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of University of New Hampshire Scholars' Repository. For more information, please contact [email protected]. This dissertation has been microfilmed exactly as received 67-166 MORSE, M. Patricia,1938- ON THE STRUCTURE AND FUNCTION OF THE DIGESTIVE SYSTEM OF THE NUDIBRANCH MOLLUSC A CA NT HODO RIS PILOSA. University of New Hampshire, Ph.D.,1966 Zoology University Microfilms, Inc., Ann Arbor, Michigan ON THE STRUCTURE AND FUNCTION OF THE DIGESTIVE SYSTEM OF THE NUDIBRANCH MOLLUSC ACANTHODORIS PILOSA BY M. PATRICIA MORSE B.S., Bates College, I960 M.S., University of New Hampshire, 1962 A DISSERTATION Submitted to the University of New Hampshire In Partial Fulfillment of The Requirements for the Degree of Doctor of Philosophy Graduate School Department of Zoology May, 1966 This dissertation has been examined and approved. VAJ//J ‘"M L , /7,/ftr/ I Date i ACKNOWLEDGEMENTS Sincere appreciation Is expressed to Dr. Wilbur L. Bullock who has guided and advised me during the preparation of this dis­ sertation. I am indebted to all the members of my Doctoral Committee for their many helpful suggestions and direction during this Investi­ gation, especially to Dr. Melbourne Carriker for his constructive criticism of thesis content. I am most appreciative for the editorial suggestions of Dr. Arthur C. Borror and for the many kindnesses from the faculty and graduate students at the University of New Hampshire. Thanks are due to the faculty at Northeastern University, especially Dr. Nathan W. Riser for his encouragement toward the completion of .this investigation. lYty sincere appreciation is ex­ tended to Dr. Edmund Smith for his suggestions and advice during the initial stages of this investigation, to Dr. Robert S. Friedman and Dr. Burdette Barrett for their help in preparation of the illustrations, and to Mrs. Allan Pollock for her many hours of typing during the preliminary preparation of the text. Finally I would like to express my deepest appreciation to Dr. Arthur J. West II who has been an inspiration and help not only in preparation of the dissertation but throughout my grad­ uate study. ii TABLE OP CONTENTS LIST OP ILLUSTRATIONS ill SECTION I. INTRODUCTION 1 SECTION II. HISTORY ’ 4 SECTION III. MATERIALS AND METHODS II Collection of Animals 11 Fixation 12 Staining Techniques 15 Ciliary Current Studies 16 Feeding Experiments 17 Natural feeding cycle 17 Experimental feeding cycle 17 SECTION IV. OBSERVATIONS 19 Gross Morphology 19 Histology 23 Buccal mass 23 Salivary glands 26 Esophagus 27 Stanach and hepatic ducts 28 Digestive gland 30 Cecum 36 Intestine 37 Ciliary Currents 38 SECTION V. DISCUSSION 39 Area of Reception 39 Area of Conduction and Storage 4l Region of Distribution, Digestion Absorption 41 Area of Peces Formation 48 SECTION VI. CONCLUSIONS 50 SECTION VII. FIGURES 53 SECTION VIII. LITERATURE CITED 86 SECTION IX. APPENDIX 93 iii LIST OP ILLUSTRATIONS FIGURE PAGE 1. Adult of Acanthodoris pilosa (Mailer) Abildgaard, 1789. 5*1 2. Gross morphology of digestive tract. 55 3. Lateral view of buccal mass. 56 *1. Ventral view of mouth and oral veil. 57 5. Dissected view of inner lip and spined buccal lip. 57 6 . Exposed interior of ventral stomach. 58 7. Photomicrograph of oral veil. 59 8. Photomicrograph of inner lip and buccal lip. 59 9. Buccal lip and rudimentary jaws. 60 10. Photomicrograph of buccal spines. 6l 11. Inner groove of buccal lip. 62 12. Cuticular portion of buccal cavity. 63 13. Photomicrograph of cuticular portion of buccal cavity. 64 14. Photomicrograph of ciliated portion of buccal cavity. 64 15. Photomicrograph of glandular ridge of buccal cavity. 65 16. Photomicrograph of salivary gland and buccal mass. 65 17. Salivary gland of A. pilosa. 66 18. Photomicrograph of salivary gland of A. pilosa. 67 19. Photomicrograph of junction of buccal cavity and crop. 67 20. Photomicrograph of valves in crop region. 68 21. Photomicrograph of crop in collapsed state. 68 22. Photomicrograph of ciliated epithelium of crop. 69 23. Tubular esophagus of A. pilosa. 70 24. Photanicrograph of glandular area of ventral stomach. 71 25. Photomicrograph of glandular area enlarged. 71 iv FIGURE PAGE 26. Fold of hepatic duct of A. pilosa. 72 27. Photomicrograph of folds of hepatic duct. 73 28. Photanicrograph of minor hepatic duct. 73 29. Photomicrograph of dorsal stomach. 74 30. Photanicrograph of mucosal folds of dorsal stanach. 74 31. Photomicrograph of ciliated epithelium of dorsal stanach. 75 32. Fold of digestive gland tubule. 76 33. Photanicrograph of crypt cell of digestive gland. 77 3k. Photomicrograph of fold of digestive gland tubule. 77 35. Digestive gland tubule with fragmentation phagocytes. 78 36. Tubule cells of digestive gland. 79 37. Photomicrograph of cecum at stomach and intestinal junction. 80 38. Photanicrograph of cecum. 80 39. Ciliated columnar epithelium of cecum. 8l 40. Photanicrograph of cecal epithelium. 82 kl. Photanicrograph of intestine. 82 42. Photomicrograph of anal region. 83 43. Photanicrograph of intestinal wandering cell. 83 44. Ciliated epithellun of intestine. 84 45. Ciliated epithelium of anal region. 85 SECTION I INTRODUCTION One should study both structure and function in order to deter­ mine the nature of any system. As Yonge (1926b) wrote, "...it is to the physiological assumptions of the morphologist and the morphological assumptions of the physiologist that the misconceptions of the past are due." In relation to general invertebrate anatcmy and physiology Delaunay (1931) reflected, "On peut distinguer, schematiquement, deux periodes dans l'histoire de cette question, la periode anatcmique (1800-1900) et la periode physiologique (1900-1930)." In the late 1920's C. M. Yonge established the correlation between anatomical and physiological studies on molluscan digestion. Since then there have been excellent accounts of the functional mor­ phology of the digestive tract, i.e., of prosobranchs (Graham 1932, 1939, Fretter and Graham 1962), opisthobranchs (Millott 1937a, Graham 1938, Fretter 1938-39, Howells 19^2, Forrest 1953) and pulmonates (Carriker 19^6a, 19^6b, Carriker and Bilstad 19^6). These investigators examined gastropods by careful dissection, reconstruction of serial sections, observation of ciliary currents and experimental feeding cycles. The authors1 descriptions combined gross morphology and histol­ ogy of the digestive tract with a functional correlation of its parts. The results of these individual investigations have led to a broader understanding of gastropod digestion. This is reflected in the discussions of form and function of the molluscan stomach by Graham (19^9), the gastropod stanach by Morton (1953), the section dealing with molluscan digestion in Morton's book (1958) and in the monograph 1 of prosobranchs by Fretter and Graham (1962). The major publications on the functional anatomy of the nudi- branch alimentary canal are Graham's (1938) account of eolid digestion and Millott's (1937a) investigation of a sponge-eating dorid Jorunna tcmentosa. Dr. Edmund Smith (personal ccnmunication, 1964) Indicated the need for further basic studies on the functional anatcmy of the nudibranch digestive system. Consequently, the qpisthobranch mollusc Acanthodoris pilosa (Abildgaard, 1789) was chosen for study because there are few accounts of dorid nudibranch digestion and this particular nudibranch is available in a relatively stable inter-tidal population on the northern coast of Maine (Fig. 1). This research was undertaken to determine the functional morphology and histology of the digestive tract of A. pilosa. Gross structure was correlated with the entire organism by dissection and serial section reconstruction. Microscopic anatcmy and function were studied by utilizing a variety of histological fixatives, stains and histochemical reactions. In accordance with the investigations of Yonge (1926a, 1926b), Fretter (1937), Graham (1938) and Millott (1937a), iron saccharate was employed for experimental indication of intracellular digestion. The observations and results of this Investigation have been utilized to correlate form and function in the digestive tract of A. pilosa. Where possible, terminology parallels that used by authors describing closely related animals. However, the characteristic out- pocketings of the molluscan stcxnach have been given a variety of names by numerous investigators. Yonge (1926b) working with lamellibranchs related the problem as follows: 3 Because of the superficial resemblance of the digestive diverticula of the Lamellibranchs, and of many other Invertebrates, to the liver of the Vertebrates, and the discovery in them of glycogen by Bernard (1855), they became known as the "liver." Weber (1880) later intro­ duced the name hepatopancreas as a result of his discovery of the secretory
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