Emile Van Handel, a renowned mosquito biochemist with an illustrious career at the Florida Medical Entomology Laboratory in Vero Beach, passed away on August 8, 2013, at the age of 95. Emile's first job was in 1946 as a food chemist, with the task of creating many of the raw materials that were unavailable during the war, such as marzipan. Another research area was the purity analysis of commercial egg powders used by bakers. Because pure egg powders were expensive, bakers often suspected their adulteration with imported egg powders. Emile examined and improved existing methods by investigating the phosphatides, the major components of avian eggs, and was able to distinguish them from vegetable phosphatides. He learned from anyone who had something to teach, but he had already become the only recognized authority with reproducible experimental results. Emile was a methodical researcher, relying on careful measurement and observation in meticulous experimental designs. His results on phosphatide chemistry ultimately led to analytical improvements in the differentiation of egg and vegetable phosphatides. In 1954 he was awarded a Ph.D. in physical organic chemistry from the University of Amsterdam, based on his thesis, “The Chemistry of Phosphoaminolipids, (with special reference to the phosphatides of the avian egg and commercial vegetable “lecithins”).” This research led to an invitation by Donald B. Zilversmit, a physiologist well-known for his solutions of differential equations used for the determination of turnover rates in metabolism and the first Editor of the Journal of Lipid Research, to a position as Assistant Professor of biomedical research at the medical school of the University of Memphis in 1955. In 1957, at the Division of Physiology in Memphis and at the Public Health Laboratories in Stuart, Florida, Van Handel published a paper with Zilversmit in the Journal of Laboratory Clinical Medicine that was cited so often that it was recognized as a Current Contents Citation Classic in 1977. In the “Micromethod for the direct determination of serum triglycerides,” he and Zilversmit described and established procedures that were recognized and followed worldwide, even after an “improved” method was published in the journal Clinical Chemistry in 1961. In 1958 he moved to the Entomological Research Laboratory in Vero Beach, Florida, which was founded by Maurice Provost of the Florida State Board of Health in 1950. Emile joked that he would have remained in Memphis for longer than the three years that he did if “…Memphis offered an ocean beach.” With a grant from NIH, Emile created and ran his research laboratory there. When the laboratory was reorganized and renamed the Florida Medical Entomology Laboratory (FMEL), Emile was appointed as a Professor at the University of Florida. The FMEL, a major center of mosquito science, celebrated its 50th anniversary in 2006, and Emile Van Handel was an essential part of it until his retirement. With his background in lipid research, Emile quickly initiated his groundbreaking investigations on the physiology of mosquitoes. He published a series of fundamental contributions to the methodology for energetic and metabolic determinations of lipid and carbohydrate utilization, all in relation to feeding, reproduction, and flight activities. He established new and simplified biochemical methods at the extremely small scale required for investigating the various physiological aspects of these relatively small organisms. One of his standard classics appeared in 1965, “The obese mosquito,” which summarized his initial results and also stimulated future research in many other laboratories and in many other directions. The clever title reflected his whimsical nature, but he later regretted using it because he thought it was often miscategorized by abstracting services. In 1958, Arden O. Lea joined the FMEL after returning from Guatemala, where he had been conducting an onchocerciasis project for NIH, and Emile met a physiologist of equal stature and common interests. Through long discussions, frequently during lunchtime on the roof of the laboratory, they speculated that hormones were as important in the metabolic regulations of mosquitoes as they were in vertebrates. Consequently, Lea took a sabbatical with Ellen Thomsen in Copenhagen, Denmark, to become proficient in microsurgery on the brains of flies. Afterwards, a productive collaboration between Van Handel and Lea, with Lea performing the brain surgery and Van Handel analyzing its metabolic effects, elucidated the neurosecretory and hormonal control of carbohydrate and lipid metabolism in mosquitoes. Again, doors were opened for further research in expanding areas of mosquito physiology and biology. Emile's highest priority was the investigation of the biochemical bases of energy metabolism. He also examined the role and relationships of various dietary sugar components in considerable detail, supported by analyses of the enzymatic mechanisms involved, and the synthesis and utilization of lipid. For several years, his mind was occupied with the question of how fat was converted to carbohydrate through the operation of the glyoxylate cycle, as found in plants and microorganisms. A thorough search for the two crucial enzymes, maltase and isocitrate lyase, was carried out. Although the former was detected in the mosquito and cockroach, the latter was found to be missing in insects. As more scientists joined FMEL, the interests there also shifted toward fieldwork, ecological questions, and problems with various vector species. Again, Emile developed simple and reliable quantification methods for biologists working in the field, such as the determination of caloric reserves, nectar analyses, sulfite determinations, fecal uric acid, and suspended protein in wastewater. Occasionally, Emile turned his work to other insects besides mosquitoes, including cockroaches, moths, honey bees, love bugs, and mealworms. Several post-docs and other collaborators were attracted to the FMEL from the U.S. and Europe. In later years, Emile taught laboratory courses for international students, which was easy for him because of his fluency in Spanish, Dutch, and German. Occasionally, he also offered courses in turnover rate calculations of metabolites for advanced students in Dutch universities, because, as Emile had mentioned once, this delicate matter is often incompletely or erroneously applied. Emile was known for his unique personality. His quick, honest, and open-minded intellect gave rise to an often sharp, but creative, criticism. A manuscript reviewed by Emile was often feared and returned full of comments and concerns, but the comments were always extremely helpful. He was guided only by scientific truth, and results were only considered valid when the experiments and the controls were repeated. “If you don't have relevant data, don't publish” was one of his cautioning remarks. But in addition to his admirable, truly scientific conduct, he had a refreshing and ironic sense of humor. He had an impressive knowledge of European history, particularly Spanish and Dutch culture, together with an admiration of the respective arts. Furthermore, Emile was a competent chess player, preferably with his son Marius, and equally accomplished on the piano. In his later years he still enjoyed playing Beethoven's Pathetic Sonata, certainly another source of his tremendous scientific creativity. After his retirement from FMEL in 2008, Emile enjoyed activities of daily thinking while walking in the sand along the beach “to keep my body busy,” playing chess “to keep my brain busy,” and playing piano every day “to keep my soul busy.” Emile was also fond of his achievements. His scientific papers have been cited over 5,500 times, and as he proudly wrote in 2007, “Many of the methods I published 30–50 years ago are still widely used today.” In November, 2012, Emile fell at home, resulting in his hospitalization. He returned home on New Years Eve, where his beloved Henriette cared for him until he died peacefully later that summer. Van Handel, E. 1955. Atherosclerosis: Overnutrition or malnutrition? Am. J. Digest. Dis. 22: 206–211. Van Handel, E. and D.B. Zilversmit. 1957. Micromethod for the direct determination of serum triglycerides. J. Lab. Clin. Med. 50: 152–157. Van Handel, E. and D.B. Zilversmit. 1958. Limitation of radioiodine as a label for fat. J. Lab. Clin. Med. 52: 831–839. Van Handel, E. 1959. Separation and chemical assay of lipid classes. J. Am. Oil Chem. Soc. 36: 294–297. Van Handel, E. 1961. Suggested modifications of the micro determination of triglycerides. Clin. Chem. 7: 249–251. Van Handel, E. and P.T.M. Lum. 1961. Sex as regulator of triglyceride metabolism in the mosquito. Science 134: 1979–1980. Van Handel, E. 1962. Coronary thrombosis and insect bites. Lancet 280: 886–887. Van Handel, E. 1965. The obese mosquito. J. Physiol. 181: 478–486. Van Handel, E. 1965. Estimation of glycogen in small amounts of tissue. Analyt. Biochem. 11: 256–265. Van Handel, E. 1965. Microseparation of glycogen, sugars and lipids. Analyt. Biochem. 11: 266–271. Van Handel, E. and A.O. Lea. 1965. Medial neurosecretory cells as regulators of glycogen and triglyceride synthesis. Science 149: 298–300. Van Handel, E. 1966. Temperature independence of the composition of triglyceride fatty acids synthesized de novo by the mosquito. J. Lipid Res. 7: 112–115. Van Handel, E. 1967. The thermal dependence of the rate of glycogen and triglyceride synthesis in the mosquito. J. Exp. Biol. 46: 523–528. Van Handel, E. 1967. Determination of fructose and fructose-yielding carbohydrates with cold anthrone. Analyt. Biochem. 19: 193–194. Van Handel, E. 1967. Non-dependence of the saturation of depot fat on temperature and photoperiod in a hibernating mosquito. J. Exp. Biol. 46: 487–490. Van Handel, E. 1968. Direct microdetermination of sucrose. Analyt. Biochem. 22: 280–283. Van Handel, E. 1968. Utilization of injected maltose and sucrose by insects: evidence for non-intestinal oligosaccharide. Comp. Biochem. Physiol. 24: 537–541. Van Handel, E. 1968. Trehalase and maltase in the serum of vertebrates. Comp. Biochem. Physiol. 26: 561–566. Van Handel, E. 1969. Do trehalose and trehalase function in renal glucose transport? Science 163: 1075–1076. Van Handel, E. 1969. Metabolism of hexoses in the intact mosquito: Exclusion of glucose and trehalose as intermediates. Comp. Biochem. Physiol. 29: 413–421. Van Handel, E. 1969. The equilibrium reaction sorbitol –– fructose in the intact mosquito. Comp. Biochem. Physiol. 29: 1023–1030. Van Handel, E. 1970. Serum trehalase: assay and normal values. Clin. Chim. Acta 29: 349–353. Van Handel, E. and A.O. Lea. 1970. Suppression of glycogen synthesis in the mosquito by a hormone from the medial neurosecretory cells. J. Insect Physiol. 16: 319–321. Van Handel, E. and A.O. Lea. 1970. Control of glycogen and fat metabolism in the mosquito. Gen. Comp. Endocrinol. 14: 381–384. Nayar, J.K. and E. Van Handel. 1971. Flight performance and metabolism of moth Spodoptera frugiperda. J. Insect Physiol. 17: 2475–2479. O'Meara G.F. and E. Van Handel. 1971. Triglyceride metabolism in thermally-feminized males of Aedes aegypti. J. Insect Physiol. 17: 1411–1413. Van Handel, E. 1971. Mannose metabolism: a comparison between the honeybee and the mosquito. Comp. Biochem. Physiol. 38B: 141–145. Nayar, J.K. and E. Van Handel. 1971. The fuel for sustained mosquito flight. J. Insect Physiol. 17: 471–481. Nayar, J.K. and E. Van Handel. 1972. Utilization of injected glucose by tsetse fly (Glossina) and stable fly (Stomoxys). J. Insect Physiol. 18: 105–107. Van Handel, E. 1972. The detection of nectar in mosquitoes. Mosq. News 32: 458. Van Handel, E., J.S. Haeger, and C.W. Hansen. 1972. The sugars of some Florida nectars. Am. J. Bot. 59: 1030–1032. Van Handel, E. 1972. Simple biological and chemical methods to determine the caloric reserves of mosquitoes. Mosq. News 32: 589–591. Van Handel, E. and J.K. Nayar. 1972. Turn-over of diglycerides during flight and rest in the moth Spodoptera frugiperda. Insect Biochem. 2: 8–12. Van Handel, E. and J.K. Nayar. 1972. Direct use of carbohydrates during sustained flight on the moth Spodoptera frugiperda. Insect Biochem. 2: 203–208. Van Handel, E. 1973. Temperature dependence of caloric expenditure and mortality in the starving mosquito. Comp. Biochem. Physiol. 44A: 1321–1323. Van Handel, E. 1974. Lipid utilization during sustained flight of moths. J. Insect Physiol. 20: 2329–2332. Skye G.E. and E. Van Handel. 1974. Malate synthase in insects. Comp. Biochem. Physiol. 49B: 83–86. Van Handel, E. 1975. Direct Determination of uric acid in fecal material. Biochem. Med. 12: 92–93. Van Handel, E. 1976. The chemistry of egg maturation in the unfed mosquito Aedes atropalpus. J. Insect Physiol. 22: 521–522. Van Handel, E. 1976. Metabolism of lovebug Plecia nearctica (Diptera, Bibionidae). Ann. Entomol. Soc. Am. 69: 215–216. Borovsky, D. and E. Van Handel. 1977. Specific radioimmunoassay for vitellogenesis in mosquitoes. J. Insect Physiol. 23: 655–658. Van Handel, E. 1978. Trehalose turnover and the coexistence of trehalose and active trehalase in cockroach haemolymph. J. Insect Physiol. 24: 151–153. Vanderhorst, D.J., A.M.C. Baljet, A.M.T. Beenakkers, and E. Van Handel. 1978. Turnover of locust hemolymph diglycerides during flight and rest. Insect Biochem. 8: 369–373. Borovsky, D. and E. Van Handel. 1979. Does ovarian ecdysone stimulate mosquitoes to synthesize vitellogenin? J. Insect Physiol. 25: 861–865. Borovsky, D. and E. Van Handel. 1980. Synthesis of ovary-specific proteins in mosquitoes. Int. J. Invert. Reprod. 2: 153–163. Borovsky, D., E. Van Handel, and A.O. Lea. 1980. Feedback inhibition of vitellogenin synthesis in ovariectomized mosquitoes. Am. Zool. 20: 903. Van Handel, E. and A.O. Lea. 1981. On and off signals from the ovary regulating egg maturation in Aedes aegypti. Am. Zool. 21: 10. Lea, A.O. and E. Van Handel. 1981. An EDNH-releasing factor from the ovaries of mosquitos fed blood. Am. Zool. 21: 1027. Lea, A.O. and E. Van Handel. 1982. A neurosecretory hormone-releasing factor from ovaries of mosquitoes fed blood. J. Insect Physiol. 28: 503–508. Van Handel, E. 1984. Metabolism of nutrients in the adult mosquito. Mosq. News 44: 573–579. Van Handel, E. and A.O. Lea. 1984. Vitellogenin synthesis in blood-fed Aedes aegypti in the absence of the head, thorax and ovaries. J. Insect Physiol. 30: 871–875. Van Handel, E. 1985. Rapid determination of glycogen and sugars in mosquitoes. J. Am. Mosq. Contr. Assoc. 1: 299–301. Van Handel, E. 1985. Rapid determination of total lipids in mosquitoes. J. Am. Mosq. Contr. Assoc. 1: 302–304. Van Handel, E. 1986. Determination and significance of suspended protein in wastewater. J. Am. Mosq. Contr. Assoc. 2: 146–149. Day, J.F. and E. Van Handel. 1986. Differences between the nutritional reserves of laboratory-maintained and field-collected adult mosquitoes. J. Am. Mosq. Contr. Assoc. 2: 154–157. Van Handel, E. 1986. Growth of three mosquitoes on two larval diets measured by protein accumulation. J. Am. Mosq. Contr. Assoc. 2: 289–291. Van Handel, E. 1987. A sulfide detection test for field use. J. Am. Mosq. Contr. Assoc. 3: 644–645. Van Handel, E. and W.S. Romoser. 1987. Proteolytic activity in the ectoperitrophic fluid of blood-fed Culex nigripalpus. Med. Vet. Entomol. 1: 251–255. Van Handel, E. 1988. Nutrient accumulation in three mosquito species during larval development and its effect on young adults. J. Am. Mosq. Contr. Assoc. 4: 374–376. Van Handel, E. and J.F. Day. 1988. Assay of lipids, glycogen and sugars in individual mosquitoes: correlations with wing length in field-collected Aedes vexans. J. Am. Mosq. Contr. Assoc. 4: 549–550. Day, J.F. and E. Van Handel. 1988. Differences in carbohydrate reserves between resting and flying Culex nigripalpus collected in the field. J. Am. Mosq. Contr. Assoc. 4: 547–548. Van Handel, E. and J.F. Day. 1989. Correlation between wing length and protein content of mosquitoes. J. Am. Mosq. Contr. Assoc. 5: 180–182. Van Handel, E. and J.F. Day. 1990. Nectar-feeding habits of Aedes taeniorhynchus. J. Am. Mosq. Contr. Assoc. 6: 270–273. Van Handel, E. 1991. Sugar-deprivation following a blood meal does not reduce yolk formation and fertility in Culex quinquefasciatus. J. Am. Mosq. Contr. Assoc. 7: 66–68. Van Handel, E., T.W. Scott, J.F. Day, and P. Reiter. 1992. Sugar-feeding habits of Aedes aegypti females from San Juan, Puerto Rico. J. Am. Mosq. Contr. Assoc. 8: 311. Van Handel, E. 1992. Postvitellogenic metabolism of the mosquito (Culex quinquefasciatus) ovary. J. Insect Physiol. 38: 75–79. Greene, L.E. and E. Van Handel. 1992. Sulfide sensitivity in fishes of the Indian River Lagoon, Florida. J. Am. Mosq. Contr. Assoc. 8: 95–96. Van Handel, E. 19923. The sweet and fat mosquito. Wing Beats 4: 4–6. Van Handel, E. 1993. Fuel metabolism of the mosquito (Culex quinquefasciatus) embryo. J. Insect Physiol. 39: 831–833. Van Handel, E. 1994. Hold that tiger. Wing Beats 5: 16,17,23. Van Handel, E., J.D. Edman, J.F. Day, T.W. Scott, G.G. Clark, P. Reiter, and H.C. Lynn. 1994. Plant-sugar, glycogen and lipid assay of Aedes aegypti collected in urban Puerto Rico and rural Florida. J. Am. Mosq. Contr. Assoc. 10: 149–153. Borovsky, D., D.A. Carlson, R.G. Hancock, H. Rembold, and E. Van Handel. 1994. De novo biosynthesis of juvenile hormone III and I by the accessory glands of the male mosquito. Insect Biochem. Molec. Biol. 24: 437–444. Van Handel, E. and M.J. Klowden. 1996. Defecation by the mosquito Aedes aegypti is controlled by the terminal abdominal ganglion. J. Insect Physiol. 42: 139–142.