INFORMATION TO USERS This material was produced from a microfilm copy of the original document. While the most advanced technological means to photograph and reproduce this document have been used, the quality is heavily dependent upon the quality of the original submitted. The following explanation of techniques is provided to help you understand markings or patterns which may appear on this reproduction. 1.The sign or "target" for pages apparently lacking from the document photographed is "Missing Page(s)". If it was possible to obtain the missing page(s) or section, they are spliced into the film along with adjacent pages. This may have necessitated cutting thru an image and duplicating adjacent pages to insure you complete continuity. 2. When an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that die copy may have moved during exposure and thus cause a blurred image. You will find a good image of the page in the adjacent frame. 3. When a map, drawing or chart, etc., was part of the material being photographed the photographer followed a definite method in "sectioning" the material. It is customary to begin photoing at the upper left hand corner of a large sheet and to continue photoing from left to right in equal sections with a small overlap. If necessary, sectioning is continued again — beginning below the first row and continuing on until complete. 4. The majority of users indicate that the textual content is of greatest value, however, a somewhat higher quality reproduction could be made from "photographs" if essential to the understanding of the dissertation. Silver prints of "photographs" may be ordered at additional charge by writing the Order Department, giving the catalog number, title, author and specific pages you wish reproduced. 5. PLEASE NOTE: Some pages may have indistinct print. Filmed as received. Xerox University Microfilms 300 North Zeeb Road Ann Arbor, Michigan 48106 HECTOR, Dwight Harold, 1944- H FUNCTIONAL ANATCMY OF THE RESPIRATORY APPARATUS § OF THE STARLING. The Ohio State University, Ph.D., 1974 Zoology University Microfilms,A XEROX Company, Ann Arbor, Michigan THIS DISSERTATION HAS BEEN MICROFILMED EXACTLY AS RECEIVED. FUNCTIONAL ANATOMY OF THE RESPIRATORY APPARATUS OF THE STARLING DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Dwight H. Hector, B.S, * * * * * The Ohio State University 1974 Reading Committee* Approved By Dr. Ahbot S. Gaunt Dr. Lewis Greenwald Dr. Harold S. Weiss Adviser Department of Zoology ACKNOWLEDGMENTS I wish to extend sincere thanks to Dr. Abbot S, Gaunt, my adviser, who has struggled with me during the past four and one-half years of study and research. He has given valuable advice to me through all phases of this thesis and has helped me to begin to think as a professional zoologist. Not withstanding, Drs, Sheldon I. Lustick, Jerry Downhower, Lewis Greenwald of the Department of Zoology and Harold S. Weiss of the Department of Physiology have been constant sources of expert advice and their critical evaluations of this thesis are greatly appreciated. Special assistance in the area of microtechnique was provided by Drs, Roy Tassava and Jack Bantle, and by Ms. Jeanne Manson, Mr. Bob Lloyd, and Mr. Tony Mescher. Dr. Robert M. Nerem of the Department of Aeronautical Engineering greatly aided me in understanding air flow dynamics, and Dr. Willard C. Myser was helpful in taking initial radiographs. I am indebted to the Ohio State University Department of Radiology for permitting me, with the aid of technicians Jerry Tennery and Mike Fields, to use a cinefluoroscope for radiographic analysis. Much of this work would not have been possible without this assistance. ii And finally to ray wife Rogette, I cannot express enough my appreciation for the constant aid and support she has given me. She has worked many long days and nights side by side with me in the preparation of this manuscript and other smaller undertakings along the way. iii VITA March if, 19*44 B o m - Lima, Ohio 1966 B.S., Findlay College, Findlay, Ohio I966-I969 Medical Student, College of Medicine, Ohio State University, Columbus, Ohio 1970-1974 Teaching Associate, Department of Zoology, The Ohio State University, Columbus, Ohio PUBLICATIONS Hector, Dwight H., and Abbot S. Gaunt. 1972. Functional anatomy of the respiratory system of the starling, Stumus vulgaris. American Zoologist, 12(4), p. 726. Abstr. Gaunt, Abbot S. , Dwight H. Hector and Sandra L. L. Gaunt. 1973. Pressure events, mini-breaths and avian vocalization. American Zoologist 13(^)» P.» 13^6. Abstr. FIELDS OF STUDY Major Field1 Zoology Studies in Avian Respiration. Dr. Abbot S. Gaunt Studies in Avian Vocalization. Dr. Abbot S. Gaunt iv TABLE OP CONTENTS Page ACKNOWLEDGMENTS...................................... ii VITA ................................................ iv LIST OP T A B L E S ................................... vii LIST OP FIGURES............................. viii LIST OP ABBREVIATIONS............................. xi INTRODUCTION ...................................... 1 METHODS AND MATERIALS FOR ANATOMICAL S T U D Y ........... 10 Preparation and Injection of Bird ANATOMY OP THE RESPIRATORY SYSTEM................. 15 Lungs Bronchi Medial Bronchi Dorsal Bronchi Ventral Bronchi Lateral Bronchi MICROSCOPIC STRUCTURE OF THE L U N G ..................... 27 Parabronchial Mantle Epithelium Cartilage Smooth Muscle Bronchial Septae Cervical Air Sac (CS) Interclavicular Air Sac (ICS) Sternal Air Sac (SS) Diverticulae of the Anterior Air Sacs D. Subscapularea D. Axillarae D. Subpectorale posterior v Dd. Subpectorale anterior, Subdeltoideae, and Coracoclaviculare D. Bronchidesmoideus Posterior Thoracic Air Sacs (PTS) EXTRAPULMONARY STRUCTURES ............................ ^7 Pleural Membranes Visceral Structures Thoracoabdominal Wall RESPIRATORY MECHANICS BASED ON ANATOMICAL FEATURES . 53 Mechanics of the Air Sacs Mechanics of the Lungs Air Flow in the Lung RESPIRATION IN THE STARLING* ARM AND CRADLE MECHANISM 65 Basic Design Single System Mode Synchronous Mode Asynchronous Mode METHODS AND MATERIALS FOR FUNCTIONAL MEASUREMENTS . 80 Mechanical Measurements Air Flow Measurements RESULTS OF MEASUREMENTS............................. 87 Awake Birds Anesthetized Birds DISCUSSION ......................................... 99 LITERATURE C I T E D ................................... 108 vi LIST OP TABLES I Table Page 1. Approximate relative volume of lungs and air s a c s .................................... 18 2, Summary of evidence and utility for arm and cradle mechanism .......... ......... 100 vii LIST OF FIGURES Figure Page 1. Diagrammatic cross section of the l u n g ........ 3 2. Scheme of interconnecting air capillaries "between adjacent parabronchi................ 3 3. Scheme of air sacs and their connections to the l u n g ................... 4 If. Bellows movement of the thorax................ 7 5. Apparatus for evacuation of air from respiratory system of birds ................ 11 6. Cast of right lung, dorsal a s p e c t ............ 16 7. Cast of right lung, ventral aspect............ 16 8. Cast of right primary "bronchus with origins of secondary bronchi, lateral view ...... 20 9. Cast of right primary bronchus with origins of secondary bronchi, medial view ...... 20 10. Diagram of right lung as viewed from ventromedial . ............................ 21 11. Photomicrograph of parabronchi and tissue mantle 28 12. Photomicrograph of lung cross section showing cartilagenous plate, septum and smooth muscle separating orifices of Mb 1 and 2 .......... 30 13. Photomicrographs of lung cross section showing relative thickness of musculature of primary bronchus and pulmonary artery ........ 30 1*K Photomicrograph of longitudinal section of lung showing septal valve ................ 32 15. Latex cast of the lungs and air s a c s .......... 35 viii Figure Page 16. Exploded cast of lungs and air s a c s .......... 36 17. Shoulder region of skeleton and air sac preparation........... 39 18. Dorsal view of cast of sternal and interclavicular s a c s ................. 39 19. Cast of lungs and air sacs, aj transverse section of a starling at level indicated by line in a, viewed posteriorly, b .......... 4-1 20. Plastic cast of respiratory system prepared with skeleton and transilluminated ........ 44 21. Ventral view of last cervical (C-14) and thoracic vertebrae (T 1-5).................. 51 22. The thoracic cage as a chain of elements . 55 23. Rotation of the ventilatory lever about the bisector of the ventral surface area (•) of the anterior s a c s ...................... 55 24. Rotation of the ventilatory lever about an axis (o) anterior to the bisector of the ventral surface area (•)................. .. 56 25. Rotation of the ventilatory lever about an axis (o) posterior to the bisector of the ventral surface area ( • ) ........... 56 26. Displacement of the oblique septum resulting from movements of the sternum.............. 59 27. Mechanism of volume change caused by anterior displacement of vertebral r i b s ............ 62 28. Movement of septal valve during ventilation . 62 29. Arm and cradle arrangement of the thoracic c a g e ...................................... 67 30. Schematic representation of the lungs and air sacs of the s t a r l i n g ...................... 67 31. Movements of arm and cradle during single system mode of ventilation of the anterior s a c