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David Schooley, Ph.D.

Emeritus Professor

David Schooley

Contact Information

  • Email:
  • Phone: (775) 784-4136
  • Fax: (775) 784-1419
  • Office: Anderson Health Sciences Building 160
  • Mail Stop: 0352


  • Ph.D., Organic Chemistry, Stanford University, 1968
  • B.S., Chemistry, New Mexico Highland University, 1963

Research Interests

Insect growth and development differs from that of mammals, and the hormones controlling this development have no counterpart in vertebrates. Ecdysone is a steroid which causes the shedding of the exoskeleton necessary for the insect to molt. Juvenile hormone modulates the action of ecdysone, controlling whether a larva molts to a larva or to a pupa. Juvenile hormone must be absent for a pupa to molt to an adult: this forms the basis of the insecticide methoprene, a synthetic molecule with juvenile hormone activity. I was involved in the development of this product before moving to University of Nevada, Reno.

My group identified four of the six known insect juvenile hormones and a related juvenile hormone from crabs. We have studied how insects create these sesquiterpenoids, four of which are biosynthetically unique with ethyl side branches replacing the usual methyl group. We found that the "extra" carbon atoms in these molecules comes from propionate, which is converted to homologs of normal terpene precursors. The glands producing these hormones vigorously metabolize the amino acids valine and/or isoleucine to a coenzyme A ester of propionate. We showed that the ability of the insect to produce the ethyl branched juvenile hormones is related to its ability to metabolize isoleucine or valine.

I have also done a great deal of research on the identification and biochemistry of insect peptide hormones which control a variety of processes. We identified from cockroach brains two peptides which raise its blood sugar levels, and later identified from Manduca sexta the hormones which stimulate (allatotropin) and stop (allatostatin) juvenile hormone production by the corpora allata, as well as the eclosion hormone, a 62 amino acid peptide which initiates the behavior necessary for an insect to escape from the old exoskeleton after it molts. We also identified a diuretic hormone from Manduca which stimulates ion secretion leading to urine production. This peptide is related to a vertebrate peptide known as CRF. We have done extensive research on this family of peptides, showing how Manduca DH is metabolized, identified seven more of these CRF-like DH from five other species of insects, and are currently trying to determine the intracellular mechanisms by which these peptides stimulate the vacuolar ATPase which powers cation secretion. From a cockroach we identified the first member of another family of diuretic hormones related to calcitonin, a controller of calcium balance in mammals. Using genomic data, we have synthesized diuretic hormones from Drosophila melanogaster and Anopheles gambiae (the malaria mosquito) and studied their physiological effects with Geoff Coast from Birkbeck College. We showed that the Anopheles calcitonin-like DH is the specific form which causes the massive excretion of sodium required when the adult female mosquito takes a blood meal. We are also interested in studying the differences in action of these two different families of peptides, both of which elevate intracellular cyclic AMP but with different outcomes in different species.

Awards, Honors, & Professional Recognition

  • 1990- Baxter, Burdick, and Jackson International Award; Agrochemical Division, American Chemical Society
  • 1993- Nevada Agricultural Foundation Arvin Boerlin Research Award
  • 1993- University of Nevada Foundation Professor
  • 2003- UNR College of Ag., Biotech., and Natural Resources Teacher of the Year
  • 2004- UNR Outstanding Researcher Award
  • 2004- University of Nevada Regents Researcher Award
  • 2006- Ted Hopkins Insect Physiologist Award, Kansas State University
  • 2007- Kenneth A. Spencer Award, Kansas City American Chemical Society Section


Selected Publications

  • Signal transduction for Schistocerca gregaria ion transport peptide is mediated via both cyclic AMP and cyclic GMP. Audsley, N., Jensen, D., Schooley, D.A., Peptides, 41, 74-80 (2013).
  • Christie AE, Stevens JS, Bowers MR, Chapline MC, Jensen DA, Schegg KM, Goldwaser J, Kwiatkowski MA, Pleasant TK Jr, Shoenfeld L, Tempest LK, Williams CR, Wiwatpanit T, Smith CM, Beale KM, Towle DW, Schooley DA, Dickinson PS. 2012, Identification of a calcitonin-like diuretic hormone that functions as an intrinsic modulator of the American lobster, Homarus americanus, cardiac neuromuscular system., J Exp Biol. 2010 Jan 1;213(1):118-27. doi: 10.1242/jeb.037077.
  • Victoria Te Brugge, Jean-Paul Paluzzi, David A. Schooley and Ian Orchard 2011, Identification of the elusive peptidergic diuretic hormone in the blood-feeding bug Rhodnius prolixus: a CRF-related peptide, J. Exp. Biol., 214, 371-381
  • Geoffrey M. Coast and David A. Schooley 2011, Toward a consensus nomenclature for insect neuropeptides and peptide hormones, Peptides/Elsevier, 32, 620-631
  • Hull, J. J., Copley, K. S., Schegg, K. M., Quilici, D. R., Schooley, D. A., Welch, W. H. 2009, De Novo molecular modeling and biophysical characterization of Manduca sexta eclosion hormone., Biochemistry, 48(38), 9047-9060.

Book or Chapter(s) in Books

  • Schooley, D.A.; Horodyski, F.M. and Coast, Geoffrey M. 2011, Hormones controlling homeostasis in insects, Gilbert, L. (ed.) Insect Endocrinology. Elsevier, pp. 366-429. ISBN 9780123847492

Older Publications

  • Isolation and characterization of CRF-related diuretic hormones from the whitelined sphinx moth Hyles lineata, Furuya, K., Harper, M., Schegg, K. M., and Schooley, D. A., Insect Biochem. Mol. Biol., 30, 127-133 (2000).
  • Cockroach diuretic hormones: characterization of a calcitonin-like peptide in insects, Furuya, K., Milchak, R. J., Schegg, K. M., Zhang, J., Tobe, S. S., Coast, G. M., and Schooley, D. A., Proc. Natl. Acad. Sci. USA, 97, 6469-6474 (2000).
  • Antidiuretic effects of a factor in brain/corpora cardiaca/corpora allata extract on fluid reabsorption across the cryptonephric complex of Manduca sexta, Liao, S., Audsley, N., and Schooley, D. A., J. Exp. Biol., 203, 605-615 (2000).
  • The Drosophila melanogaster homologue of an insect calcitonin-like diuretic peptide stimulates V-ATPase activity in fruit fly Malpighian tubules, Coast, G. M., Webster, S. G., Schegg, K. M., Tobe, S. S., and Schooley, D. A., J. Exp. Biol., 204, 1795-1804 (2001).
  • Isolation and identification of a diuretic hormone from Zootermopsis nevadensis, Baldwin, D., Schegg, K. M., Furuya, K., Lehmberg, E., and Schooley, D. A., Peptides, 22, 147-152 (2001).
  • The distribution of a kinin-like peptide and its co-localization with a CRF-like peptide in the blood-feeding bug, Rhodnius prolixus, Te Brugge, V. A., Nässel, D. R., Coast, G. M., Schooley, D. A., and Orchard, I., Peptides, 22, 161-173 (2001).
  • Antagonistic control of fluid secretion by the Malpighian tubules of Tenebrio molitor: Effects of diuretic and antidiuretic peptides and their second messengers, Wiehart, U. I. M., Nicolson, S. W., Eigenheer, R. A., and Schooley, D. A., J. Exp. Biol., 205, 493-501 (2002).
  • Identification of a potent antidiuretic factor acting on beetle Malpighian tubules, Eigenheer, R. A., Nicolson, S. W., Schegg, K. M., Hull, J. J., and Schooley, D. A., Proc. Natl. Acad. Sci. USA, 99, 84-89 (2002).
  • Simultaneous preparation of both enantiomers of juvenile hormone labeled at C-10 with tritium at high specific activity, Maxwell, R. A., Anderson, R. J., and Schooley, D. A., Anal. Biochem., 305, 40-48 (2002).
  • JH diol kinase: part I- Purification, characterization and substrate specificity of juvenile hormone selective diol kinase from Manduca sexta, Maxwell, R. A., Welch, W. H., and Schooley, D. A., J. Biol. Chem., 277, 21874-21881 (2002).
  • JH diol kinase: part II- Sequencing, cloning, and molecular modeling of juvenile hormone-selective diol kinase from Manduca sexta, Maxwell, R. A., Welch, W. H., Horodyski, F. M., Schegg, K. M., and Schooley, D. A., J. Biol. Chem., 277, 21882-21890 (2002).
  • Insect diuretic and antidiuretic hormones, Coast, G. M., Orchard, I., Phillips, J. E., and Schooley, D. A., Adv. Insect Physiol., 29, 279-409 (2002). The biological activity of diuretic factors in Rhodnius prolixus, Te Brugge, V. A., Schooley, D. A., and Orchard, I., Peptides, 23, 671-681 (2002).
  • Isolation, identification and localization of a second beetle antidiuretic peptide, Eigenheer, R. A., Wiehart, U. M., Nicolson, S. W., Schoofs, L., Schegg, K. M., Hull, J. J., and Schooley, D. A., Peptides, 24, 27-34 (2003).
  • The mechanism of action of the antidiuretic peptide Tenmo ADFa in Malpighian tubules of Aedes aegypti, Massaro, R. C., Lee, L. W., Patel, A. B., Wu, D. S., Yu, M. J., Scott, B. N., Schooley, D. A., Schegg, K. M., and Beyenbach, K. W., J. Exp. Biol., 207, 2877-2888. (2004).
  • A method for selective conjugation of an analyte to enzymes without unwanted enzyme-enzyme cross-linking, Lombardi, V. C., and Schooley, D. A., Anal. Biochem., 331, 40-45 (2004).
  • Presence and activity of a Dippu-DH31-like peptide in the blood-feeding bug, Rhodnius prolixus, Te Brugge, V. A., Lombardi, V. C., Schooley, D. A., and Orchard, I., Peptides, 26, 29-42 (2005).
  • A study of signal transduction for the two diuretic peptides of Diploptera punctata, Tobe, S. S., Zhang, J. R., Schooley, D. A., and Coast, G. M., Peptides, 26, 89-98 (2005).
  • A novel diuretic hormone receptor in Drosophila: Evidence for conservation of CGRP signaling, Johnson, E. C., Shafer, O. T., Trigg, J. S., Schooley, D. A., Dow, J. A., and Taghert, P. H., J. Exp. Biol., 208, 1239-1246 (2005).
  • Hormones controlling homeostasis in insects, Schooley, D. A., Horodyski, F. M., and Coast, G. M., in Comprehensive Molecular Insect Science, Vol. 3: Endocrinology, Gilbert,L. I., Iatrou, K., and Gill,S. S., Eds., Elsevier Pergamon, Oxford, 493-550 (2005).
  • JH8: Recent progress in juvenile hormone research, Schooley, D. A., Tobe, S. S., and Yagi, K. J., Journal of Insect Physiology, 51, 343 (2005).
  • Mosquito natriuretic peptide identified as a calcitonin-like diuretic hormone in Anopheles gambiae (Giles), Coast, G. M., Garside, C. S., Webster, S. G., Schegg, K. M., and Schooley, D. A., J. Exp. Biol., 208, 3281-3291 (2005).
  • Juvenile hormone diol kinase, a calcium-binding protein with kinase activity, from the silkworm, Bombyx mori, Li, S., Zhang, Q. R., Xu, W. H., and Schooley, D. A., Insect Biochem Mol. Biol., 35, 1235-1248 (2005).

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