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Information to Users INFORMATION TO USERS While the most advanced technology has been used to photograph and reproduce this manuscript, the quality of the reproduction is heavily dependent upon the quality of the material submitted. For example: • Manuscript pages may have indistinct print. In such cases, the best available copy has been filmed. • Manuscripts may not always be complete. In such cases, a note will indicate that it is not possible to obtain missing pages. • Copyrighted material may have been removed from the manuscript. In such cases, a note will indicate the deletion. Oversize materials (e.g., maps, drawings, and charts) are photographed by sectioning the original, beginning at the upper left-hand corner and continuing from left to right in equal sections with small overlaps. Each oversize page is also filmed as one exposure and is available, for an additional charge, as a standard 35mm slide or as a 17”x 23” black and white photographic print. Most photographs reproduce acceptably on positive microfilm or microfiche but lack the clarity on xerographic copies made from the microfilm. For an additional charge, 35mm slides of 6”x 9” black and white photographic prints are available for any photographs or illustrations that cannot be reproduced satisfactorily by xerography. Order Number 8726699 The periaqueductal gray: An examination of the distribution of opioid and non-opioid sites, their interaction, and the role of serotonin Nichols, Deborah Sue, Ph.D. The Ohio State University, 1987 U MI 300 N . Zecb Rd. Ann Arbor, M I 48106 PLEASE NOTE: In alt cases this material has been filmed in the best possible way from the available copy. Problems encountered with this document have been identified here with a check mark V 1. Glossy photographs or pages _____ 2. Colored illustrations, paper or print ______ 3. Photographs with dark background _____ 4. Illustrations are poor copy ______ 5. 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THE Ik INTEkAo HuN , AND THE ROLE OF SEROTONIN DISSERTATION rresen teu in Par tia i t u t riiimen t or r ne kequirementa tut the Decree Doctor of Pniiosopny in the uraauate Scnoot of tne Ohio State University By ueoorah Sue Nicnois, B.S., M.Ea. ihe Ohio State U n iversity 1967 u i sser t ation Oomfni t tee ; Approvea Dy b . n. i nor n U.o . ner n t son Aov I ' s d r ..) . i.. nr esriahan uepar t me tit at rsvtnoiuu1. To The Three Men I Love, L.T*, John, &> Mark ACKNOWLEDGEMENTS I woula line to express my sincere appreciation to Dr. Beverly Thorn for her support, enthusiasm, ana guidance throughout this project. A second cnank-you to Dr. Gary Berntson for his advice and assitance tnroughout this project. Thanks also go to Drs. Bresnanan, Bruno, and Stokes for their helpful comments and encouragement. Gratitude is expressed to Orlando Mullins, for his technical assistance, Kathy Moreno, for her clerical assistance, ana Julia Wans and Gary Griggs, for their research assistance. To my husband, L.T., I offer firs t my love and then my thanks for his support, love, ana that gentle push when I needed it most (an additional thank-you tor the Apple 1IC on whicn this has all been w ritten.) To my sons, Jonn and Mark, I thank you for those times you played quietly so i could work, your help with the rats, and your understanding when things got hectic*, I love you both. November, 8, 1955 Born - Dayton, Ohio 1978 ............................................................................B.S. , The Ohio State University, Columbus, Ohio 1982 M.Ed., University of Toledo, To Iedo, Oh i o 1983-Present ....................................................... Graduate Teaching Assistant The Ohio State University Columbus, Ohio FIELDS OF STUDY Major Field: Psychobiology Studies in: Psychophysiology of Mental Retardation and Developmental D isabilities, Gary G. Berntson Pain-medlating Systems, Beverly E. Thorn TABLE OF CONTENTS ACKNOWLEDGEMENTS...................................................................................................in VITA...............................................................................................................................iv LIST OF TABLES........................................................................................................vi LI ST OF FIGURES.............................................................................................................................vii INTRODUCTION............................................................................................................ 1 METHODOLOGY...............................................................................................................29 RESULTS........................................................................................................................36 DISCUSSION.................................................................................................................S3 BIBLIOGRAPHY.............................................................................................................64 v LIST OF TABLES TABLE PAGE 1. Stimulation effects at dorsal ..........s ite s ......................................38 2. Stimulation effects at ventral sites .................................... .39 3. Distribution of effects at anterior-posterior axes for each quadrant .....................................................................................40 4. Results of regression on site specificity for mean difference scores .........................................................................44 5. Results of regression on site specificity for duration...................................... 44 6. Results of regression on site specificity for i ntensi ty .......................................................................................................44 vi LIST OF FIGURES FIGURES PAGE 1. Stimulation effects plotted on rostral-caudal midbrain sections .........................................................................................42 2. Effect of naloxone on stimu I at ion-produced analgesia plotted on rostral-caudal sections ............................ 46 3. Effects of naloxone at dorsal and ventral sites ....................47 4. Cross-tolerance effects at naloxone-reversible and naioxone-non-reversible sites ..............................................................49 5. Effect pf stimulation-produced analgesia, plotted on rostral-caudal sections .....................................................................51 6. Effect of methysergide at dorsal and ventral sites. ............52 vli INTRODUCTION Stimulation of the midbrain periaqueductal gray (PAG) has been reported to e lic it stimuI at ion-produced analgesia (SPA) in humans (Hosobuchi, Adams, & Linchltz, 1977; Richardson & Akll, 1977; & Richardson, 1962) and other animals (Reynolds, 1966; Mayer, Wolfe, Akii, Carder, Uebesklnd, 1971; Balagura A Ralph, 1973). Reynolds (1966) found this analgesia sufficient to allow an experimental laparotomy to be done without need of any chemical anesthetic. Although many areas within the centra) nervous system were found to produce SPA in the rat (le . septal nuclei, dorsomedlal thalamic nucleus, and ventral tegmentum) only stimulation of the mesencephalic central gray matter and periventricular gray matter was found to greatly reduce or totally abolish the responsiveness to all noxious stimuli employed, including radiant heat applied to the tail. This analgesia was reported to be equal to or greater than 10 mg/kg morphine on alI tests (Mayer & Liebesklnd, 1974). Furthermore, this analgesic effect did not appear secondary to general sensory, emotional, or motor mechanisms (Mayer et a l., 1971). Neuroanatomical Pathways Mediating Pain; Small fibers entering the dorsal root of the spinal cord are responsible for the transmission of nociceptive (pain) information; 1 2 these fibers enter the medial aspect of the tract of Lissauer, sending collaterals to both the substantia gelatinosa and the marginal zone, as well as laminae IV and V (Kerr, 1975). Cells responding to nociceptive input are located In laminae I, I I , IV, V, & VI, with the substantia gelatinosa serving as an apparent relay station for the projection of peripheral fibers to these terminal zones (Fields & Basbaum, 1978). Transmission of nociceptive information to higher levels within the central nervous system is by way of multiple ascending pathways: the anterolateral (spinothalamic) system, the spinocervical system, the spinoreticular system, and a dorsal column system (Kerr, 1975; Fields & Basbaum, 1978). The anterolateral system Is the classical pain pathway, lesions of which result In loss of pain contralateral and caudal to the lesion (Spiller & Martin, 1921). This system originates from cells in laminae 1, IV, V, and
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