412 Florida Entomologist 87(3) September 2004

BATTLING WEST NILE VIRUS AND OTHER MOSQUITO-BORNE DISEASES: A TRIBUTE TO DR. MAURICE W. PROVOST

JOHN D. EDMAN Center for Vectorborne Diseases, Department of Entomology, University of California, Davis One Shields Ave., Davis, CA 95616-8584

This pioneer lecture honors the scientific ca- longer have the benefit of Dr. Provost’s keen in- reer of Dr. Maurice W. Provost. Dr. Provost’s dis- sight in planning our defense as we did when SLE tinguished career with the Florida State Board invaded. However, he left us clear directions of (later Division) of Health was devoted to freeing how we should respond to this newest vector Floridians from the scourge of mosquitoes and borne disease challenge. sand flies, and preventing mosquito-borne dis- Dr. Provost was unpretentious and so I will eases such as malaria and St. Louis encephalitis henceforth refer to him, simply as Maury. Despite (SLE). The unprecedented invasion of West Nile his distinguished appearance and sophisticated virus (WNV) into North America in 1999 and its tastes, Maury was at heart a naturalist who en- transcontinental spread from the Northeast over couraged informality. He donned a coat and tie for the past four years has added a new mosquito- the annual staff picture on the front steps of the borne disease to the Florida landscape. WNV is a FMEL (Fig. 1) and for the annual Christmas close relative of SLE virus and has a similar party (Fig. 2) but otherwise he dressed casually in maintenance and amplification cycle involving loose, brightly colored shirts. wild birds and Culex mosquitoes. SLE virus in- Fated with a strong genetic predisposition for vaded Florida in 1959-63, just prior to when I coronary heart disease, Maury witnessed the un- joined Dr. Provost, then director of the Florida timely death from heart failure of his parents and Medical Entomology Laboratory (FMEL) in Vero all his brothers and sisters, except for one Beach. Now, 40 years later, history has been re- younger brother. He too experienced early symp- peated, only this time the invader is an African vi- toms of heart disease and as a biologist Maury rus. WNV moved southward into Florida in 2001 understood inheritance and statistics. He real- bringing human and equine illness and death to ized his own longevity might be abbreviated de- many wild birds. It is unfortunate that we no spite his long daily walks, sensible life style, and

Fig. 1. Dr. Provost (front, center) with Senior Staff at the Florida Medical Entomology Laboratory in 1967.

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Fig. 2. Dr. Provost talking to the cook, Dr. Arden Lea, and hungry staff, Les Bourinot and Larry Webber, at the annual Florida Medical Entomology Laboratory Christmas Party in about 1962.

positive attitude. This gave focus and a quiet not the best way to recruit those who harbored sense of urgency to his life. He was one of famous less passionate feelings toward mother nature cardiologist Denton Cooley’s early coronary by- than he did—and there were many who didn’t pass patients, and this hope-filled trip to Texas during the “drain and develop era” following WW undoubtedly bought Maury some extra time. He II. Maury, the consummate gentleman, found used it unselfishly. Despite all that he contributed such displays of activism to be way outside of his and left us there was still much more that he comfort zone. wanted to give to his profession and to the preser- Maury had a special fondness for waterfowl vation of the wild side of Florida for which he and wetlands, and all their various inhabitants, cared so deeply. Those who knew him and worked except perhaps their mosquitoes, sand flies, and with him realized that Maury represented some- black flies. He must have donated considerable one special. This made his death at just 63 all the blood to these vexers of nature on those summer more tragic. Some reparation can be found in re- sojourns through the humid boreal terrain of New calling some of the positive things that I person- England. However, they did not stimulate an ally learned from Maury, lessons that displayed early calling to medical entomology. That was the depth of his wisdom and demonstrate how something Maury happened into without the slow we as a nation and public health community medical entomology background and training one have been to take them to heart and turn them might expect. Although initially a student of ver- into positive action. tebrate ecology, he ended up becoming a leader Born to French Canadian emigrants, Maury and innovative force in the early development of grew up hiking the mountains and wading the mosquito research and control efforts in Florida. clear, cold streams of rural New England. Early in World War II, malaria and chance all played a life, he developed the deep love of nature that so role in taking him in this fortunate direction. Per- permeated his interests and values as an adult. haps it was this unique background that allowed His quiet, often stoic approach to controversial him to step back and take a fresh and broader environmental issues belied the visceral feelings view of mosquito biology and control possibilities. that were his constant motivator. He was a de- Maury’s education included undergraduate voted environmentalist before the term was studies at St. Anselm College in New Hampshire coined and popularized. Maury was a pragmatist and graduate work at the University of Califor- who knew that marching and flag waving were nia, Berkeley, and Iowa State University, plus

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considerable practical experience. The nesting to generate local support for any control effort, ecology of waterfowl associated with Iowa’s many and Maury had the backing of his politically as- cattail-ringed potholes was the topic of Maury’s tute Bureau Chief, John Mulrennan Sr. Ph.D. dissertation. This came after many sum- Land was donated by Indian River County, and mers of fish and waterfowl surveys for the New a special state appropriation funded construction Hampshire Dept. of Fish and Game while a stu- of the initial facilities that opened in 1957 (Fig. 3). dent at Saint Anselm College and following grad- Maury knew it would take a long-term commit- uation. He talked fondly about the year he spent ment and more resources and patience than he in graduate school at U.C. Berkeley before return- could expect from a non-research oriented health ing to New Hampshire to complete his Master’s in agency. Thus, he launched his strategy of apply- Vertebrate Zoology on his native turf. He then be- ing for NIH grants to support basic research and gan doctoral training at Iowa State in 1940 but collaboration with local mosquito control districts was interrupted by service to his country (1942- for the conduct of more short-term applied re- 45) with the U.S. Public Health Service assigned search. This strategy was highly successful, and to the ‘Malaria Control in War Areas’ program in research funding and staff grew rapidly. The lab Florida. This assignment provided Maury with had four Sections: Ethology, Ecology, and Physiol- his first public health challenge, controlling ma- ogy to conduct basic research, and a Control sec- laria vectors. Paris Green and swamp drainage tion to address operational issues of more were the methods of choice, and seeing the envi- immediate concern. When the Control section was ronmental affects of both convinced Maury that surreptitiously removed to form a separate labo- there must be better ways to do battle with mos- ratory at Panama City in 1964, it disrupted quitoes. When the war ended he stayed on for a Maury’s master plan for a balance between basic year as State Entomologist before returning to and applied research. He quietly accepted the po- Iowa to finish his doctoral studies. These Florida litical reality and replaced the Control Section experiences and connections led Maury to return with a Biochemistry Section, an entomological to accept a position in the Bureau of Entomology field just coming into its own. He also consoli- of the Florida State Board of Health, heading up dated the small chironomid midge substation at research on mosquito biology and control. Winter Haven and used the associated positions Malaria was no longer a significant problem in and resources to develop a stronger salt marsh Florida, but land and economic development management effort within the Ecology section. along the coastline, while expanding, were being The ecology research group, which included inhibited by the hordes of mosquitoes that laid Maury and the mosquito control district directors claim to Florida’s pristine salt marshes and man- in Indian River and Brevard counties, developed grove swamps. This seemed like an almost insur- and fine-tuned salt marsh impoundments to pre- mountable task and tested the strength of vent mosquito egg deposition. This control strat- Maury’s vision and leadership. Salt marsh mos- egy effectively freed large areas of Florida’s East quitoes were thought to disperse great distances, Coast from the traditional hordes of salt marsh but no one knew how far they moved from their mosquitoes. Importantly to Maury, it did so with- estuarine sources and, therefore, how far control out the extensive use of persistent insecticides. efforts needed to be extended to protect the grow- Along with the expanding research capability ing populace. Living in old lighthouse quarters on at Vero Beach came the first major epidemic of St. the tip of Sanibel Island (no bridge in those days), Louis encephalitis (SLE) in Florida, near Tampa Maury and his small team of thick-skinned scien- and St. Petersburg (1959-63). This provided the tists, including Jim Haeger and Bill Bidling- 3rd major mosquito research challenge in Maury’s mayer and their devoted wives, began to study career, after malaria and salt marsh mosquito the migratory behavior of Aedes taeniorhynchus. control. The SLE epidemic led to creation of the This was indeed a challenge and one that contin- Encephalitis Research Laboratory in Tampa. ued to occupy the rest of Maury’s scientific career. Maury forged close ties with this new laboratory Maury encountered a Danish scientist, Erik and enlarged the research focus in Vero Beach to Nielsen, who was studying migration of the com- include studies on birds and Culex nigripalpus. mon salt marsh butterfly in Florida, and this in- This mosquito was quickly shown to be the major teraction stimulated and broadened Maury’s vector transmitting SLE virus in Florida, both thinking about mosquito migration and dispersal. among birds and to dead-end human hosts. It led to a brief but productive research collabora- I arrived at the FMEL during this period and tion between the two. Maury realized that ad- began studying the blood-feeding behavior of dressing the flood-water mosquito problems Florida’s mosquitoes. While waiting for the funds associate with Florida’s many salt marshes, pas- from Maury’s new NIH grant, I became involved tures, glades, and irrigated citrus would require a in several ongoing research projects and, despite major research commitment, and so he launched my youth and naiveté about the subtropics, I plans for the creation of a research laboratory in quickly grew to appreciate how the Provost phi- Vero Beach. Mosquitoes were so thick it was easy losophy had permeated the way research was ap-

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Fig. 3. The Florida Medical Entomology Laboratory at Vero Beach, Florida, as dedicated in 1957.

proached at the laboratory. When I interviewed during the day and engaged in a mass exodus at with Maury, he explained that research ideas at first twilight. Much swarming and mating behav- the FMEL were usually generated from field ob- ior preceded their departure, perhaps because servations, then brought into the laboratory for they would never find each other again following experimentation and clarification under con- distant migration. At the same time, Arden Lea trolled conditions, and finally taken back to the and colleagues at the FMEL were engaged in lab- field and verified again whenever possible. He oratory experiments on the endocrine control of also explained that most U.S. university research- behavior, including mating. They noted that Aedes ers did not have the benefit of Florida’s long sub- aegypti and other mosquitoes, including a colony tropical field season or the opportunity to engage of Ae. taeniorhynchus, would mate soon after in large, multi-year field studies utilizing the emergence but successful sperm transfer did not pooled talents of many different scientists all fo- take place unless females were 30-40 hr old. This cusing on the same insect. Maury was a propo- raised doubts about whether the Ae. taeniorhyn- nent of team research long before current chus observed mating prior to their exodus from biological complexity and rapid technological the marsh were actually being inseminated. A change forced this realization on the entire re- large field experiment with newly-emerged, dye- search community. He felt that the key to under- marked Ae. taeniorhynchus was subsequently con- standing the biology of insects was to be able to ducted on an inland. Indeed, observations alone bridge field and laboratory based science, an ap- can sometimes be misleading, for no fertilized fe- proach that is now gaining many new proponents males <30 hrs old were recovered in the field. in medical entomology. For example, genetically The advent of West Nile virus (WNV) in the modified mosquitoes can be created through novel U.S. has revealed how little is still known about technology but their population dynamics must many of our most important vector species. The be understood before they can be used for mos- current epidemic also exposed a general deteriora- quito control. tion of the public health infrastructure in most I was personally involved in research at Vero states. When WNV arrived, New York City no Beach that exemplified Maury’s approach. In his longer had a mosquito control program, anyone many field experiments on the movement of salt who could identify mosquitoes, or equipment for marsh mosquitoes, Ae. taeniorhynychus, Maury surveillance and operational mosquito control. An and colleagues observed that dense adult popula- emergency response was mounted with the assis- tions emerged synchronously in the salt marsh tance of the Centers for Disease Control and Pre-

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vention (CDC), neighboring states, and private uted a summary of the life history of Cx. nigripal- contractors. New York City was forced to use 30- pis in a monograph on St. Louis encephalitis year-old control technology that had not been published by the Florida State Board of Health properly evaluated for effectiveness in a disease (Provost 1969). Much has been added to our control emergency. New, innovative technologies knowledge about mosquitoes since Maury’s pio- are needed for detecting pathogens, predicting neering work, but he was responsible for provid- risk, and controlling vectors. Until we better un- ing the basis for conducting meaningful research derstand mosquito vectors and the cycles of dis- on mosquito biology. eases they transmit, we will continue to be vulnerable to natural, accidental, and intention- ally caused epidemics. The ongoing WNV epidemic REFERENCES CITED in the U.S., with over 4,000 cases and nearly 300 deaths in 2002, provides a strong argument for a NAYAR, J. K. (Ed). 1985. Bionomics and physiology of dramatic increase in the kind of research advo- Aedes taeniorhynchus and Aedes sollicitans, the salt cated and conducted by Maury Provost. marsh mosquitoes of Florida. Agr. Expt. Sta. Bulle- tin No. 852, University of Florida, Gainesville. The large body of research on salt marsh Aedes PROVOST, M. W. 1969. The natural history of Culex ni- generated by Maury and the staff at the FMEL gripalpus, pp. 46-62 In St. Louis Encephalitis in was published after his death in a monograph Florida. Monograph No. 12, Florida State Board of dedicated to him (Nayar 1985). Maury contrib- Health, Jacksonville.