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Development of the Primary Auditory Cortex in the Ferret

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Development of the Primary Auditory Cortex in the Ferret UNIVERSITY OF CINCINNATI DATE: August 13, 2003 I, , Bethany A. Adler hereby submit this as part of the requirements for the degree of: Master of Arts in: Communication Sciences and Disorders It is entitled: Development of the Primary Auditory Cortex in the Ferret Approved by: Susan Stanton, Ph.D. Linda Graeter, Ph.D. Gina Montuoro, M.A. Development of the Primary Auditory Cortex in the Ferret A thesis submitted to the Division of Research and Advanced Studies of the University of Cincinnati In partial fulfillment of the Requirements for the degree of MASTER OF ARTS in the Department of Communication Sciences and Disorders of the College of Allied Health Sciences 2003 by Bethany A. Adler B.S., Liberty University, 2001 Committee Chair: Susan Stanton, Ph.D. ABSTRACT The purpose of this study was to examine the development of the primary auditory cortex (AI) in the ferret. This study was conducted using five sable ferrets (Mustela putorius furo) of the following ages: (1) postnatal age 11 (2) postnatal age 18 (3) postnatal age 25 (4) postnatal age 32 and (5) postnatal age 63. Each subject was perfused transcardially with physiological saline, followed by tissue fixation using 4% paraformaldehyde in 0.1 M sodium phosphate buffer (pH 7.4). Each ferret was then decapitated, and the skull was dissected to allow removal of the brain, which was stored in the same fixative. The auditory cortex of each ferret was dissected from the whole brain and subsequently embedded in a gelatin and egg yolk solution. The embedded tissue blocks were then immersed in 4% paraformaldehyde for 72 hours and then sectioned using a vibrotome. Sections cut in 70 µm increments were stained using cresyl violet, and qualitatively analyzed using a light microscope. The results of this study supported previous findings, indicating that the laminar structure of the primary auditory cortex is in place by postnatal day 18 in the ferret. Additional studies investigating the laminar structure in younger ferrets and utilizing cell counts is recommended. ii iii ACKNOWLEDGEMENTS I would like to thank my committee for their guidance, suggestions, and feedback during the production of this thesis. Dr. Susan Stanton, Dr. Linda Graeter, and Gina Montuoro were gracious enough to lend their expertise to this investigation and I greatly appreciated their time. I considered it a privilege to study under Dr. Stanton, who turned out to be even more brilliant than I had garnered during my first year of graduate school, when I attended her lectures. Her vision and direction made this project possible and I owe her a great debt of gratitude. I was especially grateful for the vast quantity of laboratory supplies she allowed me to expend during this learning experience. I would also like to thank my family for their support and encouragement, particularly my sister, who provided late-night chemistry phone support, free of charge. She has always been my academic inspiration, and as usual, she turned out to have the answers I needed. She also served as a part-time counselor as the pressure mounted, which I appreciated more than anything else. Finally, I must thank my classmates, and my favorite practicum supervisor, Dr. Thomas Goldman, who provided invaluable compassion and empathy throughout this project. Their insightful comments helped me maintain a realistic perspective, and their humor helped me keep my sanity. I am sure that God put these special people in my path just when I needed them most. iv TABLE OF CONTENTS Page LIST OF FIGURES AND TABLES............................................................................................ 2 LIST OF ABBREVIATIONS ...................................................................................................... 3 CHAPTER 1.................................................................................................................................. 4 INTRODUCTION ............................................................................................................................ 4 CHAPTER 2.................................................................................................................................. 6 LITERATURE REVIEW................................................................................................................... 6 Anatomy of Neocortex............................................................................................................. 6 Laminar Structure of Neocortex ............................................................................................. 9 Specific Layers of Neocortex ................................................................................................ 10 Development of the Neocortical Layers in Humans ............................................................. 12 Summary of the Neural Migration Process in Adult Mammals............................................ 14 Development of the Human Primary Auditory Cortex ......................................................... 17 Development of the Ferret Sensory Cortex........................................................................... 19 Visual Cortex .................................................................................................................................................................. 19 Somatosensory Cortex .................................................................................................................................................... 21 Auditory Cortex .............................................................................................................................................................. 21 Histological Techniques........................................................................................................ 23 CHAPTER 3................................................................................................................................ 25 MATERIALS AND METHODS ....................................................................................................... 25 Subjects ................................................................................................................................. 25 Materials............................................................................................................................... 25 Procedures ............................................................................................................................ 25 Tissue Preparation................................................................................................................ 25 Qualitative Observation........................................................................................................ 26 CHAPTER 4................................................................................................................................ 28 RESULTS .................................................................................................................................... 28 Development of the Neocortical Layers................................................................................ 28 CHAPTER 5................................................................................................................................ 33 DISCUSSION, LIMITATIONS, AND CONCLUSIONS ........................................................................ 33 Discussion............................................................................................................................. 33 Limitations ............................................................................................................................ 34 Conclusions........................................................................................................................... 35 REFERENCES............................................................................................................................ 37 1 LIST OF FIGURES AND TABLES FIGURES Figure 1: The six layers of neocortex ............................................................................................ 6 Figure 2: The lobes of neocortex ................................................................................................... 8 Figure 3: Brodmann’s map of human cerebral cortex ................................................................... 8 Figure 4: Tonotopic organization of human primary auditory cortex ........................................... 9 Figure 5: Migration of neurons to create new layers of cells in human cortex ........................... 16 Figure 6: Neural migration from the ventricular zone to the CP in primates .............................. 17 Figure 7: Lateral view of the auditory cortex in the ferret........................................................... 22 Figure 8: P18, Section 25............................................................................................................. 29 Figure 9: P25, Section 64............................................................................................................. 29 Figure 10: P32, Section 35........................................................................................................... 30 Figure 11: P32, Section 35........................................................................................................... 31 Figure 12: P63, Section 44........................................................................................................... 32 TABLES Table 1: Ferrets vs. humans:
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