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Xsrox University Microfilms 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. Whan an image on the film is obliterated with a large round black mark, it is an indication that the photographer suspected that the 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 raw 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. Xsrox University Microfilms 300 North Zoob Road Ann Arbor, Michigan 43106 77-2369 CHIANG, Ming-Jen, 1948- THE EFFECTS OF SULFUR DIOXIDE BLOCK OF AVIAN INTRAPULMONARY CHEMORECEPTORS ON THE VENTILATORY REGULATION OF C02. The Ohio State University, Ph.D., 1976 Physiology Xerox University Microfilms ,Ann Arbor, Michigan 48106 THE EFFECTS OF SULFUR DIOXIDE BLOCK OF AVIAN 1NTRAPULMONARY CHEMORLCEFTORS ON THE VENTILATORY REGULATION OF C02 DISSERTATION Presented in Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy in the Graduate School of The Ohio State University By Ming-Jen Chiang, B.S. The Ohio State University 1976 Reading Committeet Professor Albert L. Kunz Professor E. Keith Michal Professor Charles W. Smith Professor Harold S. Weiss Approved by (. k ( L ~ t s , Adviser q Department of Physiology ACKNO.;i NDGUiftTo I wish to express my approei a ticn to my adviser, Dr. Albert L. Kunz for his guidance throughout this research. His advice and stimulation, both scientific arid ph: losophical, led me to a better understand.!ng of the physical mode.lr used in the analysis of biological systems. His instruction was invaluable in helping me to logically develop my own original and abstract thinking* Thai'H;;.-! are also extended to all the faculty, staff and graduate students in the department of Physiolop.y• whose help and stimulation have made this work enjoyable and cor,, .true live. Specifically I am indebted to Dr. Philip f erg or who offered helpful criticisms and suggestions in preparing thi thesis. Finally, I vrculd like to thank Office of Naval Research Cra nt (No. 101~?33) and National Heart and Lung Institute Giant (No. 14870-02) for financial support for this project. VITA Date of Birthi September 2, 19^8 Place of Birth: Taiwan (Formosa), Republic of China Education: B.S. National Taiwan University Department of Zoology Taipe i, Ta iv/an, 19 70 Teaching and Research Associate, Department of Physiology, The Ohio State University Columbus, Ohio 1972-1976 Academic Experience and Training: Biological Control System Dr. Albert L. Kunz Feedback and Control System Dr. Pimmel and Dr. Thurston Analysis Dept, of Electrical Engineering Avian Physiology Dr. Harold S. Weiss Single Unit Neural Dr. Albert L. Kunz and Recording Dr. Philip Berger iii TABUS OF CONTIS NTS PAGE ACKNOWLEDGEMENTS................................ VITA ............................................. iii LIST OF TABLES .......... vii LIST OF FIGURES.................................. viii INTRODUCTION .................................... 1 METHODS t 11 Unidirectional Flow System Preparation. » . H S02 Delivery System ................. 12 Surgical Vagotomy .......................... 15 Cannulation of Wing Vein and Artery .... 15 Experimental Set-Up ....................... ^ Monitoring Device and Calibration ..... ^ Isolation of the Animal .......... 2^ Programming................................ 22 Evaluation of Preparation ......... 2 3 Closed-Loop Computer Algorithms.. ......... 23 RESULTSi 25 A. Effects of Low Doses of SOg * ........ 25 1. Open-Loop COp Pacing Experiments and Statistical Analysis ..... 25 2. Closed-Loop Steady-State C02 Regulation....................... 3® iv TABLE OP’ CONTENTS (cont.) Page 3 . Closed-Loop Transients of C02 Regulation. 32 B. Effects of Large Doses of SO2 ......... 3^ 1. Post Vagotomy-Like Breathing (PVLB).'............................ 34 2. Statistical Analysis............ * . 39 3* Open-loop Ventilatory Response after Large Doses of S02 ................. ^1 C. Effects of Intravenous Infusion of S02 . A9 1. Closed-Loop C02 Regulation........ il9 2. Open-Loop C02 Pacing............... 51 D. pH Analysis.................................. 53 DISCUSSION: 55 Experimental Block of IPCs . .............. 55 SOg Block of IPCs........................... 57 Single Unit Recording....................... 59 Mechanism of S02 Blocking Activity of IPCs . 6l Steady-State and Transient-State Analysis in the Closed-Loop Kiode.............. 65 Models for S02 Effects Upon the Dynamics of C02 regulation............................ 70 Steady-State and Transient Analysis of Open-Loop C02 Forcing....................... 73 PVLB and Further Surgical Vagotomy .......... 7^ pH Analysis.................................. 76 v TABLB OF CONTENTS (cont.) PAGE SUMMARY .......................................... 77 BIBLIOGRAPHY...................................... 80 LIST or TABLES Table Page 1. Break of COp Pacing with Low Doses of SO2 * ............................. 29 2. Post Vagotomy Like Breathing (PVLB) with Large Doses of SO2 ................. 0 3* pH Measurements............................... 5^ vii LIST OF FIGURES Figure Page 1. Error-Actuated Feedback Respiratory Control System .......................................... 2 2. Avian Intrapulnonary CO2 Receptor Response (Prom Osborne) .................................. ? 3 . Schematic of Bird's Respiratory System (From hunz)......................................... 13 Determination of S02 Concentration............... 1^ 5 . Schematic of Experimental Set-Up (From Kunz) . 17 6 . Determination of C02 Concentration................ 18 7 . Break of C02 Pacing with Small Doses of S02. 26 8 . The Continuation Record of Figure 7 Showing the Recovery of COg Pacing ....... 27 9. Closed-Loop C02 Regulation after Small Doses of SO2 in the Steady-State................. 31 10. Closed-Loop C02 Regulation after Small Doses of SO2 in the Transient-State............... 33 11. Controlled Closed-Loop Steady-State C02 Regulation and Subsequent Surgical Vagotomy. 35 12. Onset of Post Vagotomy Like Breathing (PVLB) . * 36 1 3 . PVLB and Subsequent Unilateral and Bilateral Vagotomy ..................... .... 38 L6. V vs. Time Response of Open-Loop C02 Transient . ^2 1 5 . Respiratory Rate vs. Time Response of Open-Loop ^3 C02 Transient. .................................. 16. V (J, vs. Time Response of Open-Loop C02 Transient. viii LIST 01’ FlCUJtSG (cont.) F igures Page 17* V vs. ^ COp Response of Open-Loop C0? Steady-State......................................... 46 18. Period vs. £ C02 Response of Open-Loop C02 Steady -S late......................................... 47 19* V(n vc. ?■ COp Response of Open-Loop C0? Steady-State......................................... 48 20. Intravenous (l.V.) Infusion of S02-Aerated Saline in the Closed-Loop m o d e ....................... • 50 21. I.V. Infusion of SO?-Aerated Saline in C0? Pacing............... 52 22. Single Unit Recording of S02 Blocking IPC and its Spontaneous Recovery (Prom Kunz ct al)....... * ..... 60 23* Three C02 Receptors with their Inherent Delays. 66 24. Possible Respiratory Drive of Multiple Feedback Diagram.................................... 68 2 5 * Threshold for Instability (Sensitivity or Gain X Delay) ......................... 69 26. Analog Simulation Model ... 71 2 7 . Results of Analog Simulation........................ 72 ix INTRODUCTION The primary goal of the entire ventilatory apparatus is to regulate the supply of oxygen to the organism and to eliminate carbon dioxide. For a control system to function in this manner, adequate sensory elements quantitating the concentration of carbon dioxide and oxygen must be present at accessible sampling sites. Grodins (1950) described the respiratory system as a regulated system where arterial PCOg is the controlled quanity. The influence of C02 tension on respiratory drive has long been of interest to investigators. C02 regulation is compatible with either the concept of a direct action of C02 on receptors or an indirect action of CO2 via a pH effect at receptor sites. In the language of control theory, the
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