Obituary William L Woolverton

Authors: Robert L Balster Michael A Nader Kevin B Freeman

Publish Date: 2013/08/16

Volume: 229, Issue: 2, Pages: 379-381

PDF Link

Abstract

Dr William L Woolverton Billy S Guyton Distinguished Professor and Vice Chair for Research in the Department of Psychiatry and Human Behavior at the University of Mississippi Medical Center died after a brief illness at the age of 62 on June 13 2013 Prof Woolverton was a leading scientist and educator in the behavioral pharmacology of drugs of abuseAn Alabama native Woolverton attended the University of the South in Sewanee Tennessee graduating Phi Beta Kappa in 1972 While at Sewanee he did undergraduate behavioral pharmacology research with Dr Ronald Trost which led him to graduate school at the University of Chicago studying pharmacology under the mentorship of Dr Charles R Schuster and graduating with a PhD in 1977 Although Woolverton received his formal training as a pharmacologist he was strongly influenced by the behavioral sciences conducting his doctoral research in the Psychiatry Department with a psychologist in a behavioral sciences research laboratory Thus many of Woolvertons major scientific contributions were on the behavioral determinants of the abuserelated effects of drugs He did most of his research in nonhuman primate laboratories His predoctoral research was on tolerance to the effects of stimulantsUpon graduation Woolverton was recruited to the Virginia Commonwealth University Drug Abuse Training Program where he was supported under a National Institute on Drug Abuse training grant and individual postdoctoral fellowship for the period of 1977–1980 He did his postdoctoral training primarily in the laboratories of Dr Robert Balster in the Department of Pharmacology and Toxicology headed by Dr Louis Harris This was an enormously productive period as he coauthored 18 scientific journal publications based on those 3 years of work Woolverton wanted to understand better how the reinforcing effects of drugs were altered by the context of their availability This led him to a series of complex choice studies in which rhesus monkeys were given opportunities to select cocaine or food reinforcement and how various manipulations such as concurrent drug treatment altered this choice behavior eg Woolverton and Balster 1981 The results of this line of work convinced Woolverton that drug taking behavior was not “impulsive” and “out of control” as has often been proposed but rather is directly determined by the context of its availability providing a nuanced understanding of drug–behavior interactions In addition to this line of work Woolverton also completed some important basic work on the abuserelated properties of phencyclidinelike drugs as well as other researchIn 1980 Woolverton returned to the University of Chicago as a Research Associate Assistant Professor in the Department of Psychiatry in the research group headed by Charles R Schuster During the early years back in Chicago he was surrounded by likeminded behavioral pharmacologists ChrisEllyn Johanson Lewis Seiden Harriett deWit Larry Chait Mark Kleven and Kathleen Grant When Schuster left the University of Chicago to become Director of the National Institute on Drug Abuse in 1986 Woolverton replaced him as the Director of the Drug Abuse Research Center This too was an extremely productive time in Woolvertons career Woolverton played a key role overseeing several projects which led to some of his most significant findings related to the role of dopamine in cocaine abuse For example Woolverton Goldberg and Ginos 1984 examined the reinforcing effects of several dopamine receptor agonists At the time of this study D1 and D2 receptor subtypes had only recently been proposed and their functional roles were unknown Woolverton showed that only agonists at D2 receptor subtypes functioned as reinforcers Woolverton followed this study by examining D1 and D2like receptor antagonists on the reinforcing and discriminative stimulus effects of cocaine While D1like agonists would not function as reinforcers he would later amend this conclusion by testing higher efficacy D1 agonists D1like antagonists blocked cocaine selfadministration indicating an important role for this receptor subtype in cocaine reinforcement Woolverton extended his initial findings with receptor antagonists by showing that these effects were not selective for cocaine by using a multiple schedule in which responses by monkeys were also maintained by food reinforcement Much of his research focused on the pharmacology of cocaine using novel drugs developed by chemistry colleaguesWhile in Chicago Woolverton continued his development of homologous animal models of drug addiction For example Woolverton and Kleven 1988 reported withdrawal symptoms following high doses of cocaine in monkeys He also continued his studies with drug choice paradigms To determine the reinforcing efficacy of cathinone an alkaloid of the Khat plant that has amphetaminelike effects Woolverton and Johanson 1984 gave monkeys the opportunity to choose between cocaine and cathinone and found that cathinone was equally reinforcing to cocaine This work led Woolverton to a long series of studies that focused on models of reinforcing strength including the systematic evaluation of progressiveratio parameters eg Woolverton 1995 and fooddrug choice studies These latter studies included the evaluation of environmental manipulations in studies with Michael Nader examining changes in the magnitude of alternative reinforcers and the response requirement for each reinforcer eg Nader and Woolverton 1991The Chicago years also included biannual retreats with James Woods and his laboratory at the University of Michigan which featured young scientists presenting their research findings in front of an impressively intimidating audience While at Chicago Bills zest for life included owning a Porsche learning to fly airplanes and a keen interest in Alfred Hitchcock movies Woolverton remained in Chicago until 1993 During that nearly decade and a half he trained 15 students and postdoctoral fellows He also met and married his wife Candy while in ChicagoIn 1993 Woolverton was recruited to the University of Mississippi Medical Center UMMC at the rank of professor and served as Vice Chair for Research in the Department of Psychiatry and Human Behavior until his passing During his time at UMMC Woolverton provided critical leadership for the inception and development of a number of programs that have had a lasting impact on the institution In 1994 he became the first director of the Division of Neurobiology and Behavior Research He also served as the chair of the steering committee for the Program in Neuroscience an interdepartmental graduate program offering a doctoral degree in Neuroscience Woolverton trained a number of researchers while at UMMC including Drs James Rowlett Michael Weed Kristin Wilcox Karen Anderson Sunmee Wee Kevin Freeman and Sally Huskinson In 2010 Woolverton was awarded the Billy S Guyton Distinguished Professorship the highest honor that is bestowed upon researchers at UMMC He was held in the highest regard by the faculty and staff with whom he interacted and his wisdom which he shared selflessly and with humility will continue to shape the academic environment at UMMC for years to come because so many have fashioned their professional behavior after his exampleThe scope of Woolvertons research during his years at UMMC included further work on the reinforcing effects of direct and indirect monoamine agonists the behavioral pharmacology of benzodiazepines the application of behavioral economics to the measurement of drug reinforcement and the investigation of drug interactions as determinants of polydrug abuse However Woolvertons longstanding interest in how environmental circumstances shape the choice to take drugs gave rise to some of his most innovative work at UMMC He investigated the effects of probability of and delay to reinforcement on the choice to selfadminister cocaine when nondrug alternatives were available In 2007 he began a longstanding and productive collaboration with two experts in delay discounting Drs Leonard Green and Joel Myerson which resulted in the first paper to demonstrate hyperbolic discounting of delayed drug injections Woolverton et al 2007 He followed this study with others examining delay discounting of nondrug reinforcers in monkeys and found that monkeys discounted delayed drug injections less steeply than nondrug reinforcers Freeman et al 2012 Woolverton also introduced the use of choice procedures to study drugs as punishers Woolverton 2003 In 2012 he reported hyperbolic discounting of delayed punishment with histamine in monkeys selfadministering cocaine injections which was the first study to examine the effects of delaying punishment on drug selfadministration Woolverton et al 2012 This body of work was ongoing at the time of Woolvertons death as were his ideas on how it should be developed Thus with peaking creativity and no twilight in sight Woolverton pushed the envelope of behavioral pharmacology until the endDr Woolverton was very active in the College on Problems of Drug Dependence serving on the Board of Directors from 1997 to 2001 He also served as Secretary and the President of the International Study Group Investigating Drugs as Reinforcers and President of the Society for the Stimulus Properties of Drugs Reflecting his contributions to behavioral sciences research Woolverton was a Fellow of the American Psychological AssociationWoolverton was much admired by his colleagues and students He had a very inquisitive mind and liked to challenge accepted scientific beliefs He had a gentle way of guiding the development of his trainees and in engaging his research peers in the field He loved to play and sing in the evershifting band of friends and colleagues that he cultivated primarily during his Mississippi years He will be greatly missed He is survived by his wife Candy Woolverton and his son Chris and daughter Lucy Among his last wishes before he died Woolverton characteristically wanted the memorial service to be a “celebration” with the music he loved and poetry by WB Yeats His wishes were honored

Keywords: