The Neural Correlates of Endogenously Cued Covert
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THE NEURAL CORRELATES OF ENDOGENOUSLY CUED COVERT VISUOSPATIAL ATTENTIONAL SHIFTING IN THE CUE-TARGET INTERVAL: AN ELECTROENCEPHALOGRAPHIC STUDY by Edward Justin Modestino A Dissertation Submitted to the Faculty of The Charles E. Schmidt College of Science in Partial Fulfillment of the Requirements for the Degree of Doctor of Philosophy Florida Atlantic University Boca Raton, FL December 2009 Copyright © Edward Justin Modestino 2009 ii VITA Edward Justin Modestino, son of Louis Anthony Modestino and Elaine Frances Palaima, was born November 05, 1970, in Norwood, Massachusetts. He graduated from Stoughton High School in Stoughton, MA in June 1989. He married Danielle Jean (Guido) Kradin, daughter of Joseph Francis Guido and Carol Anne Kradin, on June 19, 1996. Edward graduated with a Bachelor’s degree in Psychobiology from Harvard in Cambridge, MA in June 1997. His undergraduate thesis, under advisor Michael Hasselmo, D.Phil., was theoretical; it merged neuroscience with clinical neurology on the topic of attention deficit disorder in childhood predisposing to the subsequent development of narcolepsy in adulthood. Next, he entered graduate school at the PENN in Philadelphia, PA, to further explore the theory with Douglas Frye, Ph.D. He graduated with a Master’s degree in Psychobiology in May 1999, and a post-Master’s degree (Master of Philosophy) in Cognitive Neuroscience in May 2001. The focus of his penultimate degree was an fMRI pilot study designed to test his theory using neuroimaging. In August 2001, he entered the Ph.D. program in Complex Systems and Brain Sciences at FAU. Soon thereafter, he began working with Steven Bressler, Ph.D., studying covert visual attentional shifting using EEG. After completing his Ph.D., he will commence a post-doctoral research fellowship at the University of Virginia conducting EEG/neuroimaging research of cognition with Edward Kelly, Ph.D. iv ACKNOWLEDGEMENTS I wish to express sincere gratefulness to the following people. I am thankful for my wife and mother-in-law for their support and encouragement throughout my extensive education. I am indebted to the following people for tutelage in advanced mathematics and programming in C++ and MATLAB: Drs. Taylor, Vallabha, Almonte, Zanto and Nichols. Additionally, Dr. Winchester went out of her way to help with statistics and tutelage in SPSS. I also am indebted to former DIS student/intern Elise Naimo for assisting in the collection of the data and recruiting subjects in trying and near impossible circumstances. Bill McLean, the Center Engineer with a unique sense of humor, also made it his second job to assist me in testing, fixing, repairing, ordering, and building custom equipment. I am most grateful to Dr. Steven Bressler for being a meticulous scientist and mentoring me to become the same. I am grateful to my dissertation committee members: Dr. Howard Hock for his expertise in psychophysics in relation to my research; Dr. Viktor Jirsa for encouraging me to complete my studies here since the very beginning and his insight into non-linear dynamics and EEG; and Dr. Edward Large for his expertise in EEG and vast knowledge of attentional processes. Finally, I am grateful to my Neuroscience II professor, Dean of the College of Science, Dr. Gary Perry. He has believed in me from the very beginning. His assistance in dealing with red tape, administrative and otherwise, has been invaluable. v ABSTRACT Author: Edward Justin Modestino Title: The Neural Correlates of Endogenously Cued Covert Visuospatial Attentional Shifting in the Cue-Target Interval: An Electroencephalographic Study Institution: Florida Atlantic University Dissertation Advisor: Dr. Steven L. Bressler Degree: Doctor of Philosophy Year: 2009 This study investigated electroencephalographic differences related to cue (central left- or right-directed arrows) in a covert endogenous visual spatial attention task patterned after that of Hopf and Mangun (2000). This was done with the intent of defining the timing of components in relation to cognitive processes within the cue- target interval. Multiple techniques were employed to do this. Event-related potentials (ERPs) were examined using Independent Component Analysis. This revealed a significant N1, between 100:200 ms post-cue, greater contralateral to the cue. Difference wave ERPs, left minus right cue-locked data, divulged significant early directing attention negativity (EDAN) at 200:400 ms post-cue in the right posterior which reversed polarity in the left posterior. Temporal spectral evolution (TSE) analysis of the alpha band revealed three stages, (1) high bilateral alpha precue to 120 ms post-cue, (2) an event related desynchronization (ERD) from approximately 120 ms: vi 500 ms post-cue, and (3) an event related synchronization (ERS) rebound, 500: 900 ms post-cue, where alpha amplitude, a measure of activity, was highest contralateral to the ignored hemifield and lower contralateral to the attended hemifield. Using a combination of all of these components and scientific literature in this field, it is possible to plot out the time course of the cognitive events and their neural correlates. vii DEDICATION This manuscript is dedicated to my wife, Danielle Jean Kradin. In this world of chaos, filled with negative and selfish people, she is the one person upon whom I can rely. She has made my life worth living. Additionally, this is dedicated to my maternal grandmother, Helen Pacewicz Palaima, who died on my 8th birthday. Because of her, I took guitar lessons and studied the French language to an advanced level. Although I did not know her very long, she instilled in me the passion and desire to learn and obtain an education. Without her influence during those early years, my life may have gone in a different direction. Finally, this manuscript is dedicated to one of the best feline friends a man could ever have, Nigel Kradin-Modestino. He died unexpectedly of a horrific stroke on one of the very last days I was working on my dissertation. THE NEURAL CORRELATES OF ENOGENOUSLY CUED COVERT VISUOSPATIAL ATTENTIONAL SHIFTING IN THE CUE-TARGET INTERVAL: AN ELECTROENCEPHALOGRAPHIC STUDY LIST OF TABLES ............................................................................................................ xi LIST OF FIGURES ......................................................................................................... xiii 1.0 Introduction .................................................................................................................. 1 1.1 Motivation ....................................................................................................... 1 1.2 Background ...................................................................................................... 2 1.2a Cognitive Studies of Covert Visual Attention .................................... 2 1.2b Neural Network of Covert Visual Attention ....................................... 7 1.2c Visual ERPS and Covert Attentional Shifting with the Posner Paradigm .......................................................................................... 16 1.3 Hypotheses ..................................................................................................... 26 1.3a Posner Task ....................................................................................... 26 1.3b Cue Contrast ..................................................................................... 27 1.4 Overview ......................................................................................................... 27 1.4a Content of Dissertation ..................................................................... 27 1.4b Outline of Chapters ........................................................................... 27 2.0 Methods ...................................................................................................................... 29 2.1 Participant Screening and Recruitment ........................................................... 29 2.2 Experimental Design and Task ........................................................................ 30 viii 2.3 Behavioral Analysis ......................................................................................... 44 2.3a Behavioral Recording ....................................................................... 44 2.3b Behavioral Data Processing .............................................................. 44 2.3c Behavioral Statistical Analysis ......................................................... 44 2.4 EOG Analysis .................................................................................................. 46 2.4a EOG Processing ................................................................................ 46 2.5 EEG Analysis .................................................................................................. 47 2.5a EEG Recording ................................................................................. 47 2.5b EEG Data Processing ........................................................................ 50 2.5c EEG Statistical Analysis ................................................................... 56 3.0 Results ........................................................................................................................ 57 3.1 Behavioral Data ............................................................................................... 57 3.2 Grand Averaged ERPs ....................................................................................