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A Tribute to Robert L. Metcalf

Robert L. Metcalf, 1916-1998

Dr. Robert Lee Metcalf died at 11 a.m., Wednesday, November 11, 1998, at his home in Urbana, 2 days before what would been his 82nd birthday. Thus ended one of the most influential lives of 20th century ento-mology. Dr. Metcalf, more than any other single individual, made the goal of environmentally compatible pest management achievable. For over five decades, he consistently worked toward implementing intelligent, rational, and environmentally sustainable pest control; and for many of those years, he was an articulate and courageous spokesperson for a viewpoint that was distinctly unpopular among his less progressive peers.

Dr. Metcalf was born November 13, 1916, at Columbus, Ohio, a son of McClellen Lee and Cleo Esther Fouch Metcalf. As a young child, he moved with his family to Urbana, where his father obtained a faculty position in the Department of Entomology at the University of Illinois. Young Bob enrolled at Illinois and received a bachelor’s degree in 1939 and master’s degree in 1940. For his master’s work with Clyde Kearns, he investigated the toxicity and repellent action of derivatives of picramic acid and of toluanesulfonyl chloride He then departed for Cornell University to conduct his doctoral studies, which swiftly resulted in a Ph.D. degree in 1942; he conducted fluorescence-microscopic studies of the physiology and biochemistry of the Malpighian system of Periplaneta americana (L.), at the same time developing other applications of fluorescence techniques to the field of entomology.

It is all the more remarkable that Metcalf, renowned for alerting the scientific community to the environmental consequences of pesticide abuse, began his career trained as a traditional chemical toxicologist. His first position of employment after graduate school in 1943 was as an assistant entomologist for the Tennessee Valley Authority, where he developed new approaches to chemical control of mosquitoes in impounded waters. After 6 years, he left TVA to join the faculty at University of California-Riverside, where he rose through the ranks rapidly, advancing to full professorship in 1953, serving as department chair from 1953 to 1965 and as vice chancellor for research from 1965 to 1968.

Among his numerous accomplishments during his tenure at Riverside was the development of carbamate insecticides. During this period, Metcalf’s training in organic chemistry and accomplishment in toxicology was to hold him in good stead, as it gave him the expert credentials (and accompanying credibility) to address the limitations of chemical control.

Metcalf began his professional career just as synthetic organic insecticides first came into widespread use and were widely heralded as the ultimate insect control agents. It’s almost impossible to appreciate the level of enthusiasm that permeated the entomological community, and society at large, with respect to the potential utility of these chemicals; Reader’s Digest stories trumpeted the ability of "entire towns" to "abolish flies;" Time Magazine proclaimed in 1947 that "the flies in Iowa can now be counted on the fingers of one hand" as a result of using the new pesticides. Robert Metcalf was among the first few entomologists prescient enough to recognize the dangers of excessive enthusiasm. Most importantly, he was among the first to docu-ment meticulously and incontrovertibly the phenomenon of insecticide resistance. In 1949, he documented DDT and lindane resistance in house flies in southern California. At the same time, he was among the first to document cross-resistance to pesticides; although these flies had never been exposed to dieldrin, a new cyclodiene insecticide, they nonetheless manifested resistance to the chemical when it was first encountered. Metcalf went on to develop a quantitative and repeatable method for assessing the level of resistance, a laboratory bioassay employing a microliter applicator to estimate precisely the LD50, or dose lethal to 50% of a test population, that became the standard in the field. Less than 10 years after the first evidence of resistance to synthetic organic insecticides was obtained, Metcalf authored an article in Scientific American, explaining the phenomenon and its consequences for human health and well-being in clear and simple terms. Metcalf also used his formidable chemical skills to develop sensitive methods for determining environmental fates of pesticides, in particular detecting pesticide residues in food.

In the 1960s, Metcalf focused his attention on the development of insecticide synergists, compounds with no inherent toxicity that can interfere with detoxification of insecticides and thus render greatly reduced quantities of insecticide as toxic as larger amounts. By investigating synergists (biological properties, such as effects on nontarget arthropods and biocontrol agents, as well as chemical properties), he contributed to providing people with an alternative to applying increasingly larger amounts of insecticides to cope with resistant strains. Metcalf was thus instrumental in promoting the reduction of pesticide inputs without compromising efficacy—an approach that was both environmentally less destructive and, in practical terms, acceptable to users.

In 1968, Metcalf left UC-Riverside to return to the University of Illinois, this time as a member of the faculty of the Department of Entomology. From 1969 to 1972 he served as head of Zoology; in 1971, he was named Distinguished Professor of Biology. During this period, Metcalf pioneered what may have been the most effective tool for demonstrating the impact of pesticides on the environment—the model ecosystem. As he stated in 1971, "The threat of environmental catastrophe has demonstrated that we can no longer afford to learn about environmental behavior of pesti-cides through one to two decades of widescale use. We need a realistic laboratory method for screening proposed new pesticides for their environmental fate." Metcalf’s brilliant solution to this problem was to confine a miniature ecosystem—a microcosm—to the laboratory. In a series of enclosed tanks, Metcalf designed self-contained functioning ecosystems—as he described, "an Illinois farm pond in a box." This construct allowed him to document meticulously, precisely, and repeatably the movement of pesticides through the trophic web. Metcalf and his coworkers evaluated over 200 chemicals in this terrestrial-aquatic ecosystem, providing invaluable information on the environmental suitability not only of pesticides but of industrial chemicals such as polychlorinated biphenyls and animal supplements. Information obtained from model ecosystems confirmed information obtained laboriously from decades of field studies, validating the method as a fast, inexpensive, and reliable index of environmental fate. Largely as a result of these studies, and studies inspired by his pioneering efforts, biodegradability has become a pre-requisite for introducing any new pesticide into commerce. The invention of microcosm technology has indisputably made a major contribution to the understanding of the impact of chemicals on environmental quality and, more importantly, provided a tool for minimizing that impact.

In 1975, at UIUC, Dr. Metcalf highlighted in a text-book chapter a promising new approach to pest control that, again, reduces reliance upon synthetic organic insecticides—the use of attractants and repellents, particularly naturally occurring plant compounds, that manipulate insect behavior rather than destroy physiological function. He was involved in one of the firsts tests of the efficacy of kairomones (attractants) as control agents, tests that drew from his groundbreaking studies on the kairomonal activity of methyl eugenol toward the oriental fruit fly Dacus dorsalis. In field tests, as little as a single gram of this material resulted in the mass trapping of over 7000 male flies in a single day. Subsequent work focused on developing attractant baits for corn rootworms, the most economically important pests of corn in the U.S. In studies of importance not only for their practical applicability but also for their theoretical implications for understanding the evolution of hostplant specialization in insects, Metcalf and colleagues documented compulsive feeding behavior on the part of these beetles in response to cucurbitacins, plant constituents restricted to the plant hosts of root-worms. By incorporating cucurbitacins into attractant baits, pesticide applications for corn rootworm control could be reduced from pounds per acre to grams per acre without loss of efficacy. Literally until a few weeks before he died, Dr. Metcalf continued to refine his technique, to minimize further synthetic organic inputs; cucurbitacin baits combined with neem products, as opposed to pesticides, were field-tested in summer 1998.

Metcalf’s contributions to developing environmentally compatible pest controls extend beyond the laboratory. More than the vast majority of his peers, he displayed a deep commitment to public service. He never shirked his public responsibility to serve as an educated and compelling advocate of responsible pest control. While at Riverside, his laboratory functioned as the International Insecticide Reference Center for the World Health Organization. He was a member of the Environmental Protection Agency’s Pesticide Advisory Panel from 1976 to 1982 (a critical period in the establishment of that agency); during his service on the panel he was instrumental in the removal of at least 10 toxic and widely used pesticides. As a member of the National Academy of Sciences, he served on committees evaluating polychlorinated biphenyls in the environment (1977-79), on urban pest management (1978-80), on water quality (1976-78), and on cotton insect control (1980-81). In connection with cotton insect control, he testified before Congress in a highly charged atmosphere on insect control efforts aimed at eradication (efforts that led to the spectacular $55 million failure of the fire ant program, for example). Years later, his view—that eradication is not a feasible goal for all insect introductions—is the prevailing one; his testimony undoubtedly spared the southeastern U.S. from massive pesticide contamination. In 1976, Dr. Metcalf traveled to China in one of the first scientific exchanges with that nation with an aim toward learning more about alternative low-impact control methods.

Aside from influencing public policy through political channels, Metcalf reached out to communicate with citizens in an unprecedented manner. He was frequently consulted by members of the press and willingly and patiently spoke with reporters from a wide range of media. More importantly, he authored several books for use inside and outside classrooms. His Introduction to Insect Pest Management, edited with William Luckmann, is the standard text used in classrooms across the country to instruct the next generation of integrated pest management practitioners. As co-author of Destructive and Useful Insects, a text originally written by Metcalf’s father Clell, a renowned entomologist in his own right, along with William Flint, Robert Metcalf has reached thousands; over 100,000 copies have been published and it is considered an indispensable part of any ento-mologist’s library. The newest edition, published in 1993, places new emphasis on beneficial insects, chemical ecology, and other alternatives to chemical control.

Over his astonishingly productive career, Metcalf authored or co-authored over 450 scientific papers and advised over 80 graduate students. He received many honors for his accomplishments. He was a member of the National Academy of Sciences and a fellow of the American Academy of Arts & Sciences and of the American Association for the Advancement of Science; he was honored with the Order of Cherubini, Pisa, Italy. Past president of the Entomological Society of America, he received its Founder’s Award in 1978. In 1991 he received an honorary doctorate from Ohio State University, and in 1997, he was recognized by the University of Illinois, his intellectual home for over 30 years, with an honorary degree of doctor of science. Dozens of his more than 80 students attended a reception in his honor and spoke movingly of the influence he had on their professional and personal lives.

Dr. Metcalf married Esther Jemina Rutherford on June 22, 1940, at Urbana. She died May 13, 1991. He married Elaine West Reynolds on January 1, 1992, at Albuquerque, NM. She survives. Also surviving are two sons, Robert Alan Metcalf of Bethesda, MD, and Michael Rutherford Metcalf of Raleigh, NC; a daughter, Esther Lee Metcalf Sims of Alexandria, VA; three stepdaughters, Nancy Dotson of Socorro, NM, Peggy Icenogle of Albuquerque, NM, and Jane Crawford of Crediton, England; a stepson, Eric Reynolds of Paradise, CA; a brother, James Richard Metcalf of Mahomet; and four grandchildren. Memorial contributions may be made to the University Foundation; Robert L. Metcalf Fund, Harker Hall, 1305 W. Green St., Urbana, IL 61801. (Modified from Amer. Entomol.) [back to top]

A Tribute to Robert L. Metcalf
by Richard Lampman

Early 20th century entomologists, whether educated in basic or applied research, tended to be well-trained in the bionomics of insects. This emphasis on understanding the ecology of insects and their hosts is why several became pioneers in the management of agricultural and public health pests when problems arose from the exuberant overreliance on broad spectrum synthetic insecticides. What elevated Metcalf above this group of distinguished peers? Scientifically, it was because he was the "cutting edge" for several diverse areas of entomology from the 50s until his death. He had 450+ publications, a phenomenal record, but his impact is best judged by his influence on the research of others.

Robert Metcalf was enamored by the concept of evolution and used it as the unifying theme in his research, publications, lectures, and often in his general conversation. Thus, it seems fitting to look back at his accomplishments in "pseudo- evolutionary" terms. His early training as a biochemist/physiologist preadapted him for the subsequent revolution in insecticide toxicology during the 40s, 50s, and 60s. His trademarks were adaptability and phenotypic plasticity. He saw firsthand the human toxicity of many new synthetic pes-ticides, as well as development of insecticide resistance, resurgences, outbreaks of secondary pests, bioaccumulation, delayed neurotoxicity, and the "pesticide treadmill." He saw the big picture and realized that a complex problem like insect management demanded research at numerous levels. This multidisciplinary approach resulted in major advancements in insecticide toxicology, environmental toxicology, chemical ecology, and integrated pest management.

One thing that distressed Metcalf was his belief that fewer and fewer grad students were being trained in the history of entomology, specifically pest management. He accepted as a truism that "the past is prologue to the future." For that reason, I would like to briefly review his scientific achievements and share a few of our conversations. This is not a chronological or heavily researched history of his accomplishments. It is a personal tribute to an individual I had to pleasure to work with for 15 years and greatly admired. Please forgive any minor inaccuracies.

Metcalf’s accomplishments in the field of insecticide toxi-cology included developing biodegradable DDT analogs, discovering a new class of insecticides (he was properly called the father of carbamate insecticides in one text), elucidating mechanisms of anticholinesterase insecticide activity ("biochemical lesions") and physiological mechanisms of insecticide resistance, exploring mechanisms of delayed neurotoxicity and use of synergists to enhance pesticide activity and study metabolism. He once submitted a paper containing the word "biodegradable" and the editor responded there was no such word in the dictionary and requested it be removed. Metcalf proudly stated that he had trained the brightest and the best in toxicology, starting with his very first, George Georghiou.

Metcalf was a young entomologist in the 40s and early 50s and was one of many responsible for field testing a wide array of newly discovered pesticides (of which they often knew nothing, not even the chemical structure). He was appalled by the non-specific nature of many of these compounds and witnessed first-hand workers (often colleagues) being poisoned during testing. One friend was mixing para-thion and spilled it on his pants. Another drove a truck off the road in the early evening hours after spraying because he could no longer see (his pupils had constricted). Metcalf was determined to develop insecticides that had greater toxicity to the targets (insects) than to the major non-targets (humans). He started using terms like "mammalian selectivity ratio" (and recommended that no pesticide be applied that was equally or more toxic to mammals than to insects). Carbamates were designed to be safer than most organophosphorus insecticides. He regretted that Riverside lost millions of dollars by deciding not to pursue a patent on the carbamates. He said the response from the university research office had been these compounds would be of academic interest only. The carbamates were later the major products of American Cyanamid and included some of the most toxic insecticides to mammals.

His work on acute toxicology focused on understanding the biochemical basis of a compound’s toxicity and how an insect overcomes susceptibility. This was of paramount importance for engineering compounds that would have greater selectivity to the target and, at the same time, slow the development of cross (within class) and multiple (between class) resistance. Metcalf’s laboratory served as a major screening step in WHO’s program to discover ecological and safe public health alternatives for DDT in the 70s. His work with piperonyl butoxide generated a new area of investigation into mixed function oxidases which later had a significant impact on many basic studies of insect-plant interactions. When May Berenbaum first came to UIUC, a group of students asked her to give a graduate seminar on MFOs. Without hesitation, she replied that we should ask Robert Metcalf first, as he was one of the leading authorities. Metcalf made many ripples in the pond.

One outcome of the widespread use of organochlorines and cyclodienes was contamination of urban and agricultural ecosystems and bioaccumulation of these compounds and metabolites in non-target species. Ironically, this generated another new area of study, environmental toxicology, and Metcalf was in the forefront. He preached that you had to systematically evaluate xenobiotics being released in the environment if there was to be any chance of predicting and avoiding potential problems. The model ecosystem (an aquarium with fish and aquatic and terrestrial insects and plants) was a simple solution to complex questions about biomagnification of pesticides and their metabolism in a food chain. Metcalf was a co-founder of the UI’s Institute for Environmental Studies. He was generally considered an environmentalist and frequently asked to lecture or give advice on the phone; however, many groups rapidly dis-covered that his environmentalism was based in science and that he wouldn’t condemn a pesticide indiscriminately.

Metcalf was always quick to apply his skills to various problems that seemed unrelated to others. His work with fruitfly attractants allowed him to use his experience with the chemistry of insecticides and their mode of action and apply it to insect behavior. He felt that structure-activity studies had a dual function: the discovery of new attractants and elucidation of the mode of action (harkening back to his studies of the anticholinesterase "biochemical lesion"). As grad students, we had a brief period before rootworm season when Metcalf and his first wife Esther would travel to Hawaii and work with Wally Mitchell and his wife on Dacus attractants—his concept of a perfect vacation.

Soon after Metcalf came to Urbana from Riverside, he had decided to apply the principles put forth in Science by Chamblis and Jones; namely, that cucurbitacins were specific phytochemicals influencing the feeding behavior of corn rootworms and cucumber beetles. He collaborated with a cucurbit breeding specialist, A.M. "Dusty" Rhodes. While his core group of students, postdocs, and collaborators (including Larry Hansen, Bettina Francis, Joel Coats, Jim Sanborn, Indir Kapoor, Gary Booth, Lena Brattsen, Keith Soloman, and Roy Fukuto, to name a few) continued their studies in toxicology, Metcalf and his wife collaborated with Rhodes on the influence of cucurbitacins on native Diabroticites in Illinois. This research led to what I believe is the best example in entomology of an Old World-New World co-adaptation of insect and plants (Aulacophorites and Diabroticites with the Cucurbitaceae). By 1980, Metcalf was publishing review articles that provided a convincing scenario of coevolution. Working with Jane Ferguson, Dan Fischer, John Andersen, Phil Lewis, Lesley Deem-Dickson, and myself, he explored the chemical ecology of this group of chrysomelids and made it the premier example of the influence of semiochemicals on insect behavior. There is no other group of insects for which so many semiochemicals are chemically defined that elicit such compulsive responses.

Metcalf said that the early breakthrough with Diabrotica was the development of "beetle prints," letting the beetles do the complex chemistry of this triterpenoids for you. The beetles were sensitive to nanogram quantities of cucs and by eluting these chemicals on thin layer chromatographic plates you could rapidly tell what cucs were present by having the beetles eat the silica off the TLC plates at specific spots. Metcalf’s lab produced an abundance of papers on the cucurbitacin chemistry of the Cucurbitaceae, as well as the sensitivity of beetle species to different cucurbitacins, host plant resistance due to cucs in cotyledons of cultivated cucurbits, sequestering of cucs by beetles and the allomonal properties of cucs. Metcalf loved taking his pest management class to the field and asking a volunteer (usually the teaching assistant) to taste a bitter fruit high in cucurbitacins. Students quickly learned the defensive properties of these chemicals. In the early 70s, Metcalf cut open a bitter fruit and dusted it with carbaryl. The next day the fruits were covered with thousands of dead beetles. He immediately saw the utility of cucurbitacins for controlling one of the most important crop pests in the Midwest. From 1978 to 1990, field tests were conducted with numerous collaborators in several states to show the efficacy of cucurbitacin baits. When these baits were finally brought to area-wide testing, he was no longer involved or consulted. In fact, press publications and articles on the internet rewrote history and attributed this groundbreaking research to researchers at other institutions.

In 1980, no one would have believed that there was any-thing as dramatic as cucs left in the chemical ecology of Diabroticites, but, starting with Andersen’s thesis and mine, and the report by T. Ladd of Ohio that the Japanese beetle lure, eugenol, attracted northern corn rootworms, it became evident that corn rootworms also ex-hibited a compulsive flight response to volatile attractants. Once again, Metcalf used his "biochemical lesion" experience with insecticides to study the chemistry of olfaction. By 1988, structure-activity studies resulted in attractants for all native Diabroticites in Illinois, both pest and non-pest species (five species in two genera). This generated over 20 papers and review articles and added a new chapter to the chemical ecology of corn rootworms and cucumber beetles. We were proud of these papers and, particularly, several concepts we felt were obvious but ignored in the literature. For example, attraction has two main components—the release rate of the chemicals (number of molecules impinging on receptors) and the inherent sensitivity of the insect receptor to each chemical (based on enzyme kinetic studies of anticholinesterase insecticides). Others ignored the fact that many of these chemicals differed in release rate by 100-300 times.

Numerous papers were generated on corn rootworm attractants throughout the U.S. What bothered Metcalf was that a significant number seemed to be negatively focused. Rather than experiment with how these lures could be used in IPM, some focused on how they would have little impact. He was quick to point out that endless excuses were made for pesticide failures, but few were willing to even attempt something new. It took almost 20 years to bring the concept of cuc-baits to large-scale field trials and, by 1990, it seemed no one was willing to accept the utility of the attractants.

For most, especially a retired professor with little grant money remaining and only a couple of students left, this would have probably marked the end of a brilliant scientific career. The early 90s were also a period when Metcalf had to deal with two events that all of his scientific prowess could not help—the discovery he had prostate cancer and the death of his wife (which left him rudderless). He concentrated all of his efforts on writing a book about practical aspects of chemical ecology as a tribute to his wife. I had assumed from what he said almost daily that he was giving up his research after the completion of the book, but I had underestimated his resiliency. With his marriage to Elaine Reynolds, the widow of his late friend Hal Reynolds, Metcalf displayed a sudden revitalization. He re-focused on the attractants of corn rootworms and made several advances that showed he was still on the cutting edge of his field. He also began collaborating on projects with Hans Hummel, University of Giessen, and Robert Novak, Center for Economic Entomology. Phil Lewis completed his Ph.D. and Lesley Deem-Dickson returned to complete her degree. Metcalf split his time about equally between Paradise, CA (his second wife’s home) and Urbana. In Paradise he was known as the outstanding clarinetist with an interest in bugs. He returned every summer to Urbana to work with corn rootworms. From 1992 to 1998, his publication record resembles that of a new assistant professor preparing for tenure. He described indole as a synergist in corn and cucur-bit blossoms for phenylpropanoids and terpenoids; he explained why thistles and other native plants always seemed to have certain Diabrotica species feeding in them; and he wrote several reviews on trends in entomology and insecticide research.

Metcalf’s final research summer concentrated on the practical application of corn rootworm attractants for mani-pulating corn rootworm behavior. Once again, he discovered something unexpected—kairomones can inhibit the response of beetles to pheromones—a concept that could have an impact on the mating disruption technology of several important insect pests. He found that by using the attractants you could demonstrate that large numbers of gravid western and northern corn rootworm adults were present in alfalfa and soybean fields. He seemed so revitalized that Hummel returned to Germany and told colleagues of Metcalf’s good health and stamina.

In conversations with his wife, it is evident to me now that he knew 1998 would be his last summer of research. He willed himself to go out to the field almost every day, analyze the data, complete two publications, and leave sections for at least three additional papers. In my last conversation with Dr. Metcalf, he went through a list of things for me to do after he was gone. Afterwards we talked about the summer, the exhilarating joy of discovery of the attractants, and the fun of walking the prairie, corn, and soybean fields of Illinois.

Metcalf was dramatically impacted by Rachel Carson’s Silent Spring. He was aware of the controversy about the accuracy of her examples, but nonetheless he was deeply moved by her spirit. It was one of the few books Metcalf always had out in his living room and was fond of picking up and repeatedly reading. After his funeral, I visited his family at his house, along with many others, and I spotted a well-worn copy of Silent Spring sitting on table.

Although scientifically rewarded and praised for his research, he was often criticized as being "anti-insecticide" and sometimes ostracized by applied entomologists. It per-plexed him that anyone would hold that opinion considering his publications stressed the importance of insecticides and the need to preserve them by taking an ecological approach to insect control. His pest management text with his friend and collaborator William Luckmann brought together all the developing concepts of environmental toxicology, insecticide resistance, and chemical ecology for the integrated management of insect pests and, in my opinion, was, at least partially, a tribute to Rachel Carson. It had also been an attempt to make a bridge between the Department of Entomology and the Center for Economic Entomology by having contributions from both groups. Metcalf had been a champion of both institutions throughout his tenure in Illinois.

Metcalf used to say he had an affinity for using simple tools (topical application, colorimetric assay, model ecosystem, beetle prints, sticky traps, etc.) for answering complex questions. When I asked him what he attributed all of these accomplishments to, he hesitated and said much of it was sudden insight but most of these discoveries were either due to challenges he presented to his students or challenges his students presented to him.

Robert Metcalf was entomology’s Thomas Edison. Both believed in practical application of scientific principles and both believed that a research laboratory should consist of a team of workers systematically investigating a topic. Metcalf was driven to be the best, whether it was science, sports (golf, tennis, and ping-pong), or music (he owned almost every type of clarinet known). Students and faculty drop-ping by his laboratory would sometimes step in gingerly because they thought we were having an argument. We constantly disagreed on how to interpret data or what was the most vital next experiment, yet we were always able to compromise. I once thought he was so exasperated that I apologized for being contentious. At first, it looked like I’d almost physically hit him. He glared at me and said that was the nature of our relationship. Once I learned he loved a good discussion, I seldom agreed with anything, although I quickly learned you had to come prepared and ready to back up your assertions with facts because he did not talk just for the sake of talking. Not all of his students realized that he would say things to spark an argument. It was not meant to belittle, but rather an attempt to generate an honest, although sometimes heated, exchange of ideas. Perhaps the most rewarding aspect for many of us was the next day he would bring up a discussion that totally incorporated your previous arguments into his. Metcalf was famous for being the most demanding questioner of his ownstudents during prelims. It was how you reacted to questions you had to think about that mattered; he had already decided whether you were qualified or not for a doctorate and was willing to vigorously defend his judgment. All of us wished he had told us that before the orals and not afterwards.

Although never a believer in organized religion, Metcalf was a highly moral individual who believed in decorum, honor, and truth. His last words to me were that he was certain that somehow, in some form, he was finally going to be able to understand why beetles did what they did. I only wish he could let me know what he found out. [back to top]


Integrative Biology University of Illinois

Updated 12/08/99