Hostname: page-component-78c5997874-j824f Total loading time: 0 Render date: 2024-11-19T16:26:14.593Z Has data issue: false hasContentIssue false
  • English
  • Français

Hormonal control of tick development and reproduction

Published online by Cambridge University Press:  19 April 2005

H. H. REES
H. H. REES
Affiliation:
School of Biological Sciences, University of Liverpool, Biosciences Building, Crown Street, Liverpool, L69 7ZB, UK
Get access Rights & Permissions [Opens in a new window]

Abstract

Ecdysteroids (moulting hormones), juvenoids and neuropeptides in ticks are reviewed but, by far, the emphasis is on the former since this class of hormones has been the subject of most investigations. In immature stages of ticks, ecdysteroids have been shown to regulate moulting and to terminate larval diapause. Although there is a paucity of information on the molecular action of ecdysteroids in ticks, their action appears to be via a heterodimeric ecdysone/ultraspiracle receptor, as in insects. The role of ecdysteroids in sperm maturation in adult males is considered. In females, ecdysteroids function in the regulation of salivary glands, of production of sex pheromones and of oogenesis and oviposition. There is evidence for ecdysteroid production in the integument and pathways of hormone inactivation are similar to those in insects. Ecdysteroids also function in embryogenesis. Although evidence for the occurrence and functioning of juvenile hormones in ticks has been contradictory, in recent thorough work it has not been possible to detect known juvenile hormones in ticks, nor to demonstrate effects of extracts on insects. Factors (neuropeptides) from the synganglion affect physiological processes and limited immunocytochemical studies are reviewed. Sigificantly, a G-protein-coupled receptor has been cloned, expressed, and specifically responds to myokinins.

Keywords

Hormonesendocrine systemecdysteroidsmoulting hormonesjuvenile hormonesneuropeptides
Type
Research Article
Information
Parasitology , Volume 129 , Issue S1 , October 2004 , pp. S127 - S143
Copyright
© 2004 Cambridge University Press

Access options

Get access to the full version of this content by using one of the access options below. (Log in options will check for institutional or personal access. Content may require purchase if you do not have access.)

References

REFERENCES

AESCHLIMANN, A. A. ( 1968). La ponte chez Ornithodoros moubata Murray (Ixodoidea: Argasidae). Revue Suisse Zoologie 75, 1033–1039.Google Scholar
ALLAN, S. A., PHILLIPS, J. S., TAYLOR, D. & SONENSHINE, D. E. ( 1988). Genital sex pheromones of ixodid ticks: evidence for the role of fatty acids from the anterior reproductive tract in mating of Dermacentor variabilis and Dermacentor andersoni. Journal of Insect Physiology 34, 315–323.CrossRefGoogle Scholar
BASSAL, T. T. M. & ROSHDY, M. A. ( 1974). Argas (Persicargas) arboreus: juvenile hormone analog termination of diapause and oviposition control. Experimental Parasitology 36, 34–39.CrossRefGoogle Scholar
BELLÉS, X. ( 1998). Endocrine effectors in insect vitellogenesis. In Recent Advances in Arthropod Endocrinology (ed. Coast, G. M. & Webster, S. G.), pp. 71–90. Cambridge, Cambridge University Press.
BINNINGTON, K. C. ( 1981). Ultrastructural evidence for the endocrine nature of the lateral organs of the cattle tick Boophilus microplus. Tissue and Cell 13, 475–490.CrossRefGoogle Scholar
BINNINGTON, K. C. ( 1987). Histology and ultrastructure of the acarine synganglion. In Arthropod Brain: Its Evolution, Development, Structure, and Functions (ed. Gupta, A. P.), pp. 95–109. Oxford, Pergamon Press.
BINNINGTON, K. C. & OLIVER, J. H. Jr. ( 1982). Structure and function of the circulatory, nervous and neuroendocrine systems of ticks. In Physiology of Ticks (ed. Obenchain, F. D. & Galun, R.), pp. 351–398. Oxford, Pergamon Press.CrossRef
BOUVIER, J., DIEHL, P. A. & MORICI, M. ( 1982). Ecdysone metabolism in the tick Ornithodoros moubata (Argasidae, Ixodoidea). Revue Suisse de Zoologie 89, 967–976.CrossRefGoogle Scholar
CAMPBELL, J. D. & OLIVER, J. H. Jr. ( 1984). Membrane feeding and developmental effects of ingested β-ecdysone on Ornithodoros parkeri (Acari: Argasidae). In Acarology VI, Vol. 1 (ed. Griffiths, D. A. & Bowman, C. E.), pp. 393–399. Chichester, Ellis Horwood.
CHANG, E. S. & KAUFMAN, W. R. ( 2004). Endocrinology of crustaceans and arachnids. In Comprehensive Insect Science – Vol. 3 Endocrinology (ed. Gilbert, L. I., Iatrou, K. & Gill, S.), pp. 805–842. Amsterdam, Elsevier.
CHINZEI, Y. ( 1986). Vitellogenin biosynthesis and processing in a soft tick, Ornithodoros moubata. In Host Regulated Development Mechanisms in Vector Arthropods (ed. Borovsky, D. & Spielman, A.), pp. 18–24. Vero Beach, Florida, University of Florida Press IFAS.
CHINZEI, Y., CHINO, H. & TAKAHASHI, K. ( 1983). Purification and properties of vitellogenin and vitellin from a tick Ornithodoros moubata. Journal of Comparative Physiology 152, 13–21.CrossRefGoogle Scholar
CHINZEI, Y. & TAYLOR, D. ( 1990). Regulation of vitellogenesis induction by engorgement in the soft tick (Ornithodoros moubata). Advances in Invertebrate Reproduction 5, 565–570.Google Scholar
CHINZEI, Y., TAYLOR, D. & ANDO, K. ( 1991). Effects of juvenile hormone and its analogs on vitellogenin synthesis and ovarian development in Ornithodoros moubata (Acari: argasidae). Journal of Medical Entomology 28, 506–513.CrossRefGoogle Scholar
CHINZEI, Y., TAYLOR, D., MIURA, K. & ANDO, K. ( 1992). Vitellogenesis induction by synganglion factor in adult female tick, Ornithodoros moubata (Acari: Argasidae). Journal of the Acarological Society of Japan 1, 15–26.CrossRefGoogle Scholar
CHINZEI, Y. & YANO, I. ( 1985). Fat body is the site of vitellogenin synthesis in the soft tick Ornithodoros moubata. Journal of Comparative Physiology, B 155, 671–678.CrossRefGoogle Scholar
CONNAT, J.-L. ( 1987). Aspects endocrinologiques de la physiologie du dévelopment et de la reproduction chez le tiques. PhD thesis, University of Burgogne.
CONNAT, J.-L., DELBECQUE, J.-P., ALABOUVETTE, J. & PITOIZET, N. ( 1997). Evolution of ecdysteroids and of their apolar conjugates during the post-embryonic development of the tick Ornithodoros moubata. Archives of Insect Biochemistry and Physiology 35, 159–168.3.0.CO;2-A>CrossRefGoogle Scholar
CONNAT, J.-L. & DIEHL, P. A. ( 1986). Probable occurrence of ecdysteroid fatty acid esters in different classes of arthropods. Insect Biochemistry 16, 91–97.CrossRefGoogle Scholar
CONNAT, J.-L., DIEHL, P. A., DUMONT, N., CARMINATI, S. & THOMPSON, M. J. ( 1983 a). Effects of exogenous ecdysteroids on the female tick, Ornithodoros moubata: Induction of supermolting and influence on oogenesis. Zeitschrift für Angewandte Entomologie 96, 520–530.Google Scholar
CONNAT, J.-L., DIEHL, P. A., GFELLER, H. & MORICI, M. ( 1985). Ecdysteroids in females and eggs of the ixodid tick Amblyomma hebraeum. International Journal of Invertebrate Reproduction and Development 8, 103–116.CrossRefGoogle Scholar
CONNAT, J-L., DIEHL, P. A. & MORICI, M. ( 1984). Metabolism of ecdysteroids during the vitellogenesis of the tick Ornithodoros moubata (Ixodoidea: Argasidae): accumulation of apolar metabolites in the eggs. General and Comparative Endocrinology 56, 100–110.CrossRefGoogle Scholar
CONNAT, J.-L., DIEHL, P. A. & THOMPSON, M. J. ( 1986 a). Possible inactivation of ingested ecdysteroids by conjugation with long-chain fatty acids in the female tick Ornithodoros moubata (Acarina: Argasidae). Archives of Insect Biochemistry and Physiology 3, 235–252.Google Scholar
CONNAT, J.-L. & DOTSON, E. M. ( 1988). Comparative investigation of the egg ecdysteroids of ticks using radioimmunoassay and metabolic studies. Journal of Insect Physiology 34, 639–645.CrossRefGoogle Scholar
CONNAT, J.-L., DOTSON, E. M. & DIEHL, P. A. ( 1987). Metabolism of ecdysteroids in the female tick Amblyomma hebraeum (Ixodoidea: Ixodidae): accumulation of free ecdysone and 20-hydroxyecdysone in the eggs. Journal of Comparative Physiology B 157, 689–699.CrossRefGoogle Scholar
CONNAT, J.-L., DOTSON, E. M. & DIEHL, P. A. ( 1988). Apolar conjugates of ecdysteroids are not used as a storage form of molting hormone in the argasid tick Ornithodoros moubata. Archives of Insect Biochemistry and Physiology 9, 221–235.CrossRefGoogle Scholar
CONNAT, J.-L., DUCOMMUN, J. & DIEHL, P. A. ( 1983 b). Juvenile hormone-like substances can induce vitellogenesis in the tick Ornithodoros moubata (Acarina: Argasidae). International Journal of Invertebrate Reproduction 6, 285–294.Google Scholar
CONNAT, J.-L., DUCOMMUN, J., DIEHL, P. A. & AESCHLIMANN, A. ( 1986 b). Some aspects of the control of the gonotrophic cycle in the tick Ornithodoros moubata (Ixodoidea, Argasidae). In Morphology, Physiology, and Behavioral Biology of Ticks (ed. Sauer, J. R. & Hair, J. A.), pp. 194–216. Chichester, Ellis Horwood.
CONNAT, J.-L., LAFONT, R. & DIEHL, P. A. ( 1986 c). Metabolism of [3H]ecdysone by isolated tissues of the female ixodid tick Amblyomma hebraeum (Ixodoidea: Ixodidae). Molecular and Cellular Endocrinology 47, 257–267.Google Scholar
CONNAT, J.-L. & NEPA, M.-C. ( 1990). Effects of different anti-juvenile hormone agents on the fecundity of the female tick Ornithodoros moubata. Pesticide Biochemistry and Physiology 37, 266–274.CrossRefGoogle Scholar
CROSBY, T., EVERSHED, R. P., LEWIS, D., WIGGLESWORTH, K. P. & REES, H. H. ( 1986 a). Identification of ecdysone 22-long chain fatty acyl esters in newly laid eggs of the cattle tick Boophilus microplus. Biochemical Journal 240, 131–138.Google Scholar
CROSBY, T., WIGGLESWORTH, K. P., LEWIS, D. & REES, H. H. ( 1986 b). Moulting hormones in the development of the cattle tick, Boophilus microplus. In Host Regulated Developmental Mechanisms in Vector Arthropods (ed. Borovsky, D. & Spielman, A.), pp. 37–45. Vero Beach, University of Florida-IFAS.
DAVIS, H. H., DOTSON, E. M. & OLIVER, J. H. Jr. ( 1994). Localization of insulin-like immunoreactivity in the synganglion of nymphal and adult Dermacentor variabilis (Acari: Ixodidae). Experimental and Applied Acarology 18, 111–122.CrossRefGoogle Scholar
DEES, W. H., SONENSHINE, D. E. & BREIDLING, E. ( 1984 a). Ecdysteroids in the American dog tick, Dermacentor variabilis (Acari: Ixodidae), during different periods of tick development. Journal of Medical Entomology 21, 514–523.Google Scholar
DEES, W. H., SONENSHINE, D. E. & BREIDLING, E. ( 1984 b). Ecdysteroids in Hyalomma dromedarii and Dermacentor variabilis and their effects on sex pheromone activity. In Acarology VI (ed. Griffiths, D. A. & Bowman, C. E.), pp. 405–413. Chichester, Ellis Horwood.
DEES, W. H., SONENSHINE, D. E. & BRIEDLING, E. ( 1985). Ecdysteroids in the camel tick, Hyalomma dromedarii (Acari: Ixodidae) and comparison with sex pheromone activity. Journal of Medical Entomology 22, 22–27.CrossRefGoogle Scholar
DELBECQUE, J. P., DIEHL, P. A. & O'CONNOR, J. D. ( 1978). Presence of ecdysone and ecdysterone in the tick Amblyomma hebraeum Koch. Experientia 34, 1379–1381.CrossRefGoogle Scholar
DIEHL, P. A., CONNAT, J.-L. & DOTSON, E. M. ( 1986). Chemistry, function, and metabolism of tick ecdysteroids. In Morphology, Physiology and Behavioral Biology of Ticks (ed. Sauer, J. R. & Hair, J. H.), pp. 165–193. Chichester, Ellis Horwood.
DIEHL, P. A., CONNAT, J.-L., GIRAULT, J. P. & LAFONT, R. ( 1985). A new class of apolar ecdysteroid conjugates: esters of 20-hydroxy-ecdysone with long-chain fatty acids in ticks. International Journal of Invertebrate Reproduction and Development 8, 1–13.CrossRefGoogle Scholar
DIEHL, P. A., GERMOND, J. E. & MORICI, M. ( 1982). Correlations between ecdysteroid titres and integument structure in nymphs of the tick, Amblyomma hebraeum Koch (Acarina: Ixodidae). Revue Suisse de Zoologie 89, 859–868.CrossRefGoogle Scholar
DINAN, L. ( 1997). Ecdysteroids in adults and eggs of the house cricket, Acheta domesticus (Orthoptera: Gryllidae). Comparative Biochemistry and Physiology 116B, 129–135.CrossRefGoogle Scholar
DONAHUE, W. A., TEEL, P. D., STREY, O. F. & MEOLA, R. W. ( 1997). Pyriproxyfen effects on newly engorged larvae and nymphs of the lone star tick (Acari: Ixodidae). Journal of Medical Entomology 34, 206–211.CrossRefGoogle Scholar
DOTSON, E. M., CONNAT, J.-L. & DIEHL, P. A. ( 1991). Cuticle deposition and ecdysteroid titres during embryonic and larval development of the argasid tick Ornithodoros moubata. General and Comparative Endocrinology 82, 386–400.CrossRefGoogle Scholar
DOTSON, E. M., CONNAT, J.-L. & DIEHL, P. A. ( 1993). Metabolism of [3H]ecdysone in embryos and larvae of the tick Ornithodoros moubata. Archives of Insect Biochemisty and Physiology 23, 67–78.CrossRefGoogle Scholar
DOTSON, E. M., CONNAT, J.-L. & DIEHL, P. A. ( 1995). Ecdysteroid titre and metabolism and cuticle deposition during embryogenesis of the ixodid tick Amblyomma hebraeum (Koch). Comparative Biochemistry and Physiology 110B, 155–166.CrossRefGoogle Scholar
DUMSER, J. B. & OLIVER, J. H. Jr. ( 1981). Kinetics of spermatogenesis, cell cycle analysis, and testis development in nymphs of the tick Dermacentor variabilis. Journal of Insect Physiology 27, 743–753.CrossRefGoogle Scholar
EL-SAID, A., SWIDERSKI, Z., AESCHLIMANN, A. & DIEHL, P. A. ( 1981). Fine structure of spermiogenesis in the tick Amblyomma hebraeum (Acari: Ixodidae): late stages of differentiation and structure of the mature spermatozoon. Journal of Medical Entomology 18, 464–476.CrossRefGoogle Scholar
FRIESEN, K. J. & KAUFMAN, W. R. ( 2002). Quantification of vitellogenesis and its control by 20-hydroxyecdysone in the ixodid tick, Amblyomma hebraeum. Journal of Insect Physiology 48, 773–782.CrossRefGoogle Scholar
FRIESEN, K. J. & KAUFMAN, W. R. ( 2003). Cypermethrin inhibits egg development in the ixodid tick, Amblyomma hebraeum. Pesticide Biochemistry and Physiology 76, 25–35.CrossRefGoogle Scholar
GÄDE, G., HOFFMANN, K. H. & SPRING, J. ( 1997). Hormonal regulation in insects: facts, gaps, and future directions. Physiological Reviews 77, 963–1032.CrossRefGoogle Scholar
GERMOND, J.-E., DIEHL, P. A. & MORICI, M. ( 1982). Correlations between integument structure and ecdysteroid titres in fifth-stage nymphs of the tick, Ornithodoros moubata. General and Comparative Endocrinology 46, 255–266.CrossRefGoogle Scholar
GILBERT, L. I., IATROU, K. & GILL, S. (Eds.) ( 2004). Comprehensive Insect Science – Vol. 3 Endocrinology. Amsterdam, Elsevier.
GOODMAN, W. & GRANGER, N. ( 2004). The juvenile hormone. In Comprehensive Insect Science – Vol. 3 Endocrinology ( ed. Gilbert, L. I., Iatrou, K. & Gill, S.), pp. 319–408. Amsterdam, Elsevier.
GUO, X., HARMON, M. A., LAUDET, V., MANGELSDORF, D. J. & PALMER, M. J. ( 1997). Isolation of a functional ecdysteroid receptor homologue from the ixodid tick, Amblyomma americanum (L.). Insect Biochemistry and Molecular Biology 27, 945–962.CrossRefGoogle Scholar
GUO, X., XU, Q., HARMON, M. A., JIN, X. LAUDET, V., MANGELSDFORF, D. J. & PALMER, M. J. ( 1998). Isolation of two functional retinoid X receptor subtypes from the Ixodid tick, Amblyomma americanum (L.). Molecular and Cellular Endocrinology 139, 45–60.CrossRefGoogle Scholar
HAMILTON, J. G. C. ( 1992). The role of pheromones in tick biology. Parasitology Today 8, 130–133.CrossRefGoogle Scholar
HAMMOCK, B. D. ( 1985). Regulation of juvenile hormone titer: degradation. In Comprehensive Insect Physiology, Biochemistry and Pharmacology, Vol. 7 ( ed. Kerkut, G. A. & Gilbert, L. I.), pp. 431–472. Oxford, Pergamon.
HARRIS, R. A. & KAUFMAN, W. R. ( 1981). Hormonal control of salivary gland degeneration in the ixodid tick Amblyomma hebraeum. Journal of Insect Physiology 27, 241–243.CrossRefGoogle Scholar
HARRIS, R. A. & KAUFMAN, W. R. ( 1984). Neural involvement in the control of salivary gland degeneration in the ixodid tick. Amblyomma hebraeum. Journal of Experimental Biology 109, 281–290.Google Scholar
HARRIS, R. A. & KAUFMAN, W. R. ( 1985). Ecdysteroids: possible candidates for the hormone which triggers salivary gland degeneration in the ixodid tick Amblyomma hebraeum. Experientia 41, 740–742.CrossRefGoogle Scholar
HAYWARD, D. C., BASTIANI, M. J., TRUEMAN, J. W. H., TRUMAN, J. W., RIDDIFORD, L. M. & BALL, E. E. ( 1999). The sequence of Locusta RXR, homologous to Drosophila Ultraspiracle, and its evolutionary implications. Development Genes and Evolution 209, 564–571.CrossRefGoogle Scholar
HENRICH, V. ( 2004). The ecdysteroid receptor (EcR). In Comprehensive Insect Science – Vol. 3 Endocrinology ( ed. Gilbert, L. I., Iatrou, K. & Gill, S.), pp. 245–285. Amsterdam, Elsevier.
HENRICH, V. C., RYBCZYNSKI, R. & GILBERT, L. I. ( 1999). Peptide hormones, and puffs: mechanisms and models in insect development. Vitamins and Hormones – Advances in Research and Applications 55, 73–125.Google Scholar
HOFFMANN, K. H., MEYERING-VOS, M. & LORENZ, M. W. ( 1999). Allatostatins and allatotropins: is the regulation of corpora allata activity their primary function? European Journal of Entomology 96, 255–266.Google Scholar
HOLMES, S. P., BARHOUMIT, R., NACHMAN, R. J. & PIETRANTONIO, P. V. ( 2003). Functional analysis of a G protein-coupled receptor from the Southern cattle tick Boophilus microplus (Acari: Ixodidae) identifies it as the first arthropod myokinin receptor. Insect Molecular Biology 12, 27–38.CrossRefGoogle Scholar
HOLMES, S. P., HE, H., CHEN, A. C., IVIE, G. W. & PIETRANTONIO, P. V. ( 2000). Cloning and transcriptional expression of a leucokinin-like peptide receptor from the Southern cattle tick, Boophilus microplus (Acari: Ixodidae). Insect Molecular Biology 9, 457–465.CrossRefGoogle Scholar
ISAAC, R. E. & SLINGER, A. J. ( 1989). Storage and excretion of ecdysteroids. In Ecdysone ( ed. Koolman, J.), pp. 250–253. Stuttgart, G. Thieme.
IWAMI, M., KAWAKAMI, A., ISHIZAKI, H., TAKAHASI, S. Y., ADACHI, T., SUZUKI, Y., HAGAWASA, H. & SUZUKI, A. ( 1989). Cloning of a gene encoding bombyxin, an insulin-like brain secretory peptide of the silkmoth Bombyx mori with prothoracicotropic activity. Development Growth and Differentiation 31, 31–37.CrossRefGoogle Scholar
JAFFE, H., HAYES, K. K., SONENSHINE, D. E., DEES, W. H., BEVERIDGE, M. & THOMPSON, M. J. ( 1986). Controlled release reservoirs system for the delivery of insect steroid analogues against ticks. Journal of Medical Entomology 23, 685–691.CrossRefGoogle Scholar
JAMES, A. M. & OLIVER, J. H. Jr. ( 1997). Purification and partial characterization of vitellin from the black-legged tick Ixodes scapularis. Insect Biochemistry and Molecular Biology 27, 639–649.CrossRefGoogle Scholar
JARVIS, T. D., EARLEY, F. G. & REES, H. H. ( 1994). Ecdysteroid biosynthesis in larval testes of Spodoptera littoralis. Insect Biochemstry and Molecular Biology 24, 531–537.CrossRefGoogle Scholar
KAUFMAN, W. R. ( 1989). Tick–host interaction: a synthesis of current concepts. Parasitology Today 5, 47–56.CrossRefGoogle Scholar
KAUFMAN, W. R. ( 1991). Correlation between haemolymph ecdysteroid titre, salivary gland degeneration and ovarian development in the ixodid tick, Amblyomma hebraeum Koch. Journal of Insect Physiology 37, 95–99.CrossRefGoogle Scholar
KHALIL, G. M. ( 1970). Biochemistry and physiological studies on certain ticks (Ixodoidea). Gonad development and gametogenesis in Hyalomma (H.) anatolicum excavatum Koch (Ixodidae). Journal of Parasitology 56, 596–610.Google Scholar
KHALIL, G. M., SONENSHINE, D. E., HANAFY, H. A. & ABDELMONEM, A. E. ( 1984). Juvenile hormone I effects on the camel tick, Hyalomma dromedarii (Acari: Ixodidae). Journal of Medical Entomology 21, 561–566.CrossRefGoogle Scholar
KOZLOVA, T. & THUMMEL, C. S. ( 2000). Steroid regulation of postembryonic development and reproduction in Drosophila. Trends in Endocrinology and Metabolism 11, 276–280.CrossRefGoogle Scholar
KUBO, I., KOMATSU, S., ASAKA, Y. & DE BOER, G. ( 1987). Isolation and identification of apolar metabolites of ingested 20-hydroxyecdysone in frass of Heliothis virescens larvae. Journal of Chemical Ecology 13, 785–794.CrossRefGoogle Scholar
KULCSAR, P., PRESTWICH, G. G. & SONENSHINE, D. E. ( 1989). Detection binding proteins for juvenile hormone-like substances in ticks by photoaffinity labelling. In Host Regulated Developmental Mechanisms in Vector Arthropods ( ed. Borovsky, D. & Spielman, A.), pp. 18–23. Vero Beach, University of Florida Press, IFAS.
LAFONT, R., DAUPHIN-VILLEMANT, C., WARREN, J. & REES, H. H. ( 2004). Ecdysteroid chemistry and biochemistry. In Comprehensive Insect Science – Vol. 3 Endocrinology ( ed. Gilbert, L. I., Iatrou, K. & Gill, S.), pp. 125–195. Amsterdam, Elsevier.
LEAHY, M. G. & BOOTH, K. S. ( 1980). Precocene induction of tick sterility and ecdysis failure. Journal of Medical Entomology 17, 18–21.CrossRefGoogle Scholar
LEID, M., KASTNER, P. & CHAMBON, P. ( 1992). Multiplicity generates diversity in the retinoic acid signalling pathway. Trends in Biochemical Sciences 17, 427–433.CrossRefGoogle Scholar
LINDSAY, P. J. & KAUFMAN, W. R. ( 1988). Action of some steroids on salivary gland degeneration in the ixodid tick. A. americanum. Journal of Insect Physiology 34, 351–359.CrossRefGoogle Scholar
LOEB, M. J., BRANDT, E. P., WOODS, C. W. & BORKOVEC, A. B. ( 1987). An ecdysiotropic factor from brains of Heliothis virescens induces testes to produce immunodetectable ecdysteroid in vitro. Journal of Experimental Zoology 243, 275–282.CrossRefGoogle Scholar
LOMAS, L. O. & REES, H. H. ( 1998). Endocrine regulation of development and reproduction in acarines. In Recent Advances in Arthropod Endocrinology ( ed. Coast, G. M. & Webster, S. G.), pp. 91–124. Cambridge, Cambridge University Press.
LOMAS, L. O., TURNER, P. C. & REES, H. H. ( 1997). A novel neuropeptide–endocrine interaction controlling ecdysteroid production in ixodid ticks. Proceedings of the Royal Society of London B 264, 589–596.CrossRefGoogle Scholar
LUNKE, M. D. & KAUFMAN, W. R. ( 1993). Hormonal control of ovarian development in the tick Amblyomma hebraeum Koch (Acari: Ixodidae). Invertebrate Reproduction and Development 23, 25–38.CrossRefGoogle Scholar
MAGEE, R. M., JONES, L. D. & REES, H. H. ( 1996). Ecdysteroids in relation to adult development and reproduction in female Rhipicephalus appendiculatus (Acari: Ixodidae). Archives of Insect Biochemistry and Physiology 31, 197–206.3.0.CO;2-X>CrossRefGoogle Scholar
MANGELSDORF, D. J. & EVANS, R. M. ( 1995). The RXR heterodimers and orphan receptors. Cell 83, 841–850.CrossRefGoogle Scholar
MANGO, C., ODHIAMBO, T. R. & GALUN, R. ( 1976). Ecdysone and the super tick. Nature 260, 318–319.CrossRefGoogle Scholar
MAO, H. & KAUFMAN, W. R. ( 1998). DNA binding properties of the ecdysteroid receptor in the salivary gland of the female ixodid tick, Amblyomma hebraeum. Insect Biochemistry and Molecular Biology 28, 947–957.CrossRefGoogle Scholar
MAO, H. & KAUFMAN, W. R. ( 1999). Profile of the ecdysteroid hormone and its receptor in the salivary gland of the adult female tick, Amblyomma hebraeum. Insect Biochemistry and Molecular Biology 29, 33–42.CrossRefGoogle Scholar
MAO, H., McBLAIN, W. A. & KAUFMAN, W. R. ( 1995). Some properties of the ecdysteroid receptor in the salivary gland of the ixodid tick, Amblyomma hebraeum. General and Comparative Endocrinology 99, 340–348.CrossRefGoogle Scholar
MARZOUK, A. S., MOHAMED, F. S. A. & KHALIL, G. M. ( 1985). Neurohemal-endocrine organs in the camel tick, Hyalomma dromedarii (Acari: Ixodoidea: Ixodidae). Journal of Medical Entomology 22, 385–391.CrossRefGoogle Scholar
McSWAIN, J. L., TUCKER, J. S., ESSENBERG, R. C. & SAUER, J. R. ( 1989). Brain factor induced formation of inositol phosphates in tick salivary glands. Insect Biochemistry 19, 343–349.CrossRefGoogle Scholar
NÄSSEL, D. R. ( 1996). Neuropeptides, amines, and amino acids in an elementary insect ganglion: functional and chemical anatomy of the unfused abdominal ganglion. Progress in Neurobiology 48, 325–420.CrossRefGoogle Scholar
NEESE, P. A., SONENSHINE, D. E., KALLAPUR, V. L., APPERSON, C. S. & ROE, R. M. ( 2000). Absence of insect juvenile hormones in the American dog tick, Dermacentor variabilis (Say) (Acari: Ixodidae), and in Ornithodoros parkeri Cooley (Acari: Argasidae). Journal of Insect Physiology 46, 477–490.CrossRefGoogle Scholar
OBENCHAIN, F. D. & MANGO, C. K. A. ( 1980). Effects of exogenous ecdysteroids and juvenile hormones on female reproductive development in Ornithodoros p. porcinus. American Zoologist 20, Abstract No. 1192.Google Scholar
OBENCHAIN, F. D. & OLIVER, J. H. Jr. ( 1975). Neurosecretory system of the American dog tick, Dermacentor variabilis (Acari: Ixodidae). II. Distribution of secretory cell types, axonal pathways and putative nerohemal-neuroendocrine associations: comparative histological and anatomical implications. Journal of Morphology 145, 269–294.Google Scholar
OGIHARA, K. ( 2003). Ecdysteroid hormone titer and expression of ecdysone receptor mRNA as related to vitellogenesis in the soft tick, Ornothodoros moubata (Acari: Argasidae). Master of Agricultural Science Thesis, University of Tsukuba, Japan.
OLIVER, J. H. Jr. ( 1986 a). Relationship among feeding, gametogenesis, mating and syngamy in ticks. In Host Regulated Development Mechanisms in Vector Arthropods ( ed. Borovsky, D. & Spielman, A.), pp. 93–99. Vero Beach, University of Florida Press, IFAS.
OLIVER, J. H. Jr. ( 1986 b). Induction of oogenesis and oviposition in ticks. In Morphology, Physiology and Behavioral Biology of Ticks ( ed. Sauer, J. R. & Hair, J. A.), pp. 233–247. Chichester, Ellis Horwood.
OLIVER, J. H. Jr. & DOTSON, E. M. ( 1993). Hormonal control of molting and reproduction in ticks. American Zoologist 33, 384–396.CrossRefGoogle Scholar
OLIVER, J. H. Jr., ZHU, X. X., VOGEL, G. N. & DOTSON, E. M. ( 1992). Role of synganglion in oogenesis of the tick Ornithodoros parkeri (Acari: Argasidae). Journal of Parasitology 78, 93–98.CrossRefGoogle Scholar
PALMER, M. J., HARMON, M. A. & LAUDET, V. ( 1999). Characterization of EcR and RXR homologues in the Ixodid tick, Amblyomma americanum (L.). American Zoologist 39, 747–757.CrossRefGoogle Scholar
PALMER, M. J., WARREN, J. T., JIN, X., GUO, X. & GILBERT, L. I. ( 2002). Developmental profiles of ecdysteroids, ecdysteroid receptor mRNAs and DNA binding properties of ecdysteroid receptors in the Ixodid tick. Amblyomma americanum (L.). Insect Biochemistry and Molecular Biology 32, 465–476.CrossRefGoogle Scholar
POUND, J. M. & OLIVER, J. H. Jr. ( 1979). Juvenile hormone: evidence of its role in the reproduction of ticks. Science 206, 355–357.CrossRefGoogle Scholar
POUND, J. M. & OLIVER, J. H. Jr. ( 1982). Synganglial and neurosecretory morphology of female Ornithodoros parkeri (Cooley) (Acari: Argasidae). Journal of Morphology 173, 159–177.CrossRefGoogle Scholar
POUND, J. M., OLIVER, J. H. Jr. & ANDREWS, R. H. ( 1984). Induction of apolysis and cuticle formation in female Ornithodoros parkeri (Acari: Argasidae) by hemocoelic injections of β-ecdysone. Journal of Medical Entomology 21, 612–614.CrossRefGoogle Scholar
REES, H. H. & ISAAC, R. E. ( 1984). Biosynthesis of ovarian ecdysteroid phosphates and their metabolic fate during embryogenesis in Schistocerca gregaria. In Biosynthesis, Metabolism and Mode of Action of Invertebrate Hormones. ( ed. Hoffmann, J. & Porchet, M.), pp. 181–195. Berlin, Springer Verlag.CrossRef
RIDDIFORD, L. M. ( 1996). Juvenile hormone: the status of its “status quo” action. Archives of Insect Biochemistry and Physiology 32, 271–286.3.0.CO;2-W>CrossRefGoogle Scholar
RIDDIFORD, L. M., CHERBAS, P. & TRUMAN, J. W. ( 2001). Ecdysone receptors and their biological actions. Vitamins and Hormones – Advances in Research and Applications 60, 1–73.Google Scholar
ROBINSON, P. D., MORGAN, E. D., WILSON, Y. D. & LAFONT, R. ( 1987). The metabolism of ingested and injected [3H]ecdysone by final instar larvae of Heliothis armigera. Physiological Entomology 12, 321–330.CrossRefGoogle Scholar
ROSELL, R. & COONS, L. B. ( 1991). Purification and partial characterization of vitellin from the eggs of the hard tick Dermacentor variabilis. Insect Biochemistry 21, 871–885.CrossRefGoogle Scholar
ROSELL, R. & COONS, L. B. ( 1992). The role of the fat body, midgut and ovary in vitellogenin production and vitellogenesis in the female tick Dermacentor variabilis. International Journal for Parasitology 22, 341–349.CrossRefGoogle Scholar
ROSELL-DAVIS, R. & COONS, L. B. ( 1989). Relationship between feeding, mating, vitellogenin production and vitellogenesis in tick Dermacentor variabilis. Experimental and Applied Acarology 7, 95–105.CrossRefGoogle Scholar
SANKHON, N., LOCKEY, T., ROSELL, R. C., ROTHSCHILD, M. & COONS, L. ( 1999). Effect of methoprene and 20-hydroxyecdysone on vitellogenin production in cultured fat bodies and backless explants from unfed female Dermacentor variabilis. Journal of Insect Physiology 45, 755–761.CrossRefGoogle Scholar
SANNASI, A. & SUBRAMONIAM, T. ( 1972). Hormonal rupture of larval diapause in the tick Rhipicephalus sanguineus (Lat.). Experientia 28, 666–667.CrossRefGoogle Scholar
SAUER, J. R., ESSENBERG, R. C. & BOWMAN, A. S. ( 2000). Salivary glands in ixodid ticks: control and mechanism of secretion. Journal of Insect Physiology 46, 1069–1078.CrossRefGoogle Scholar
SAUER, J. R., McSWAIN, J. L., BOWMAN, A. S. & ESSENBERG, R. C. ( 1995). Tick salivary gland physiology. Annual Review of Entomology 40, 245–267.CrossRefGoogle Scholar
SCHRIEFER, M. E. ( 1991). Vitellogenesis in Hyalomma dromedarii (Acari: Ixodidae): a model for analysis of endocrine regulation in ixodid ticks. PhD Dissertation, Old Dominion University & East Virginia Medical School, Norfolk.
SHANBAKY, N. M. & KHALIL, G. M. ( 1975). The sub-genus Persicargus (Ixodoidea: Argasidae: Argas). 22. The effect of feeding on hormonal control of egg development in Argas (Persicargas) arboreus. Experimental Parasitology 37, 361–366.Google Scholar
SHANBAKY, N. M., MANSOUR, M. M., MAIN, A. J., EL-SAID, A. & HELMY, N. ( 1990). Hormonal control of vitellogenesis in Argas (Argas) hermanni (Acari: Argasidae). Journal of Medical Entomology 27, 968–974.CrossRefGoogle Scholar
SHEPHERD, J., OLIVER, J. H. Jr. & HALL, J. D. ( 1982). A polypeptide from male accessory glands which triggers maturation of tick spermatozoa. International Journal of Invertebrate Reproduction 5, 129–137.CrossRefGoogle Scholar
SLINGER, A. J. & ISAAC, R. E. ( 1988). Ecdysteroid titers during embryogenesis of the cockroach, Periplaneta americana. Journal of Insect Physiology 34, 1119–1125.CrossRefGoogle Scholar
SMITH, W. A., VARGHESE, A. H., HEALY, M. S. & LOU, K. J. ( 1996). Cyclic AMP is a prerequisite messenger in the action of big PTTH in the prothoracic glands of pupal Manduca sexta. Insect Biochemistry and Molecular Biology 26, 161–170.CrossRefGoogle Scholar
SOLLER, M., BOWNES, M. & KUBLI, E. ( 1999). Control of oocyte maturation in sexually mature Drosophila females. Developmental Biology 208, 337–351.CrossRefGoogle Scholar
SOLOMON, K. R., MANGO, C. K. A. & OBENCHAIN, F. D. ( 1982). Endocrine mechanisms in ticks: effects of insect hormones and their mimics on development and reproduction. In Physiology of Ticks ( ed. Obenchain, F. D. & Galun, R.), pp. 399–438. Oxford, Pergamon.CrossRef
SONENSHINE, D. E. ( 1986). Tick pheromones: an overview. In Morphology, Physiology, and Behavioural Biology of Ticks ( ed. Sauer, J. R. & Hair, J. A.), pp. 342–360. Chichester, Ellis Horwood.
SONENSHINE, D. E. ( 1991). Biology of Ticks, Vol. 1. New York, Oxford University Press.
SONENSHINE, D. E., ROE, R. M., VENKATESH, K., APPERSON, C., WINDER, B., SCHRIEFER, M. E. & BAEHR, J. C. ( 1989). Biochemical evidence of the occurrence of a juvenoid in ixodid ticks. In Host Regulated Developmental Mechanisms In Vector Arthropods ( ed. Borovsky, D. & Spielman, A.), pp. 9–17. Vero Beach, University of Florida Press, IFAS.
STAUFFER, A. & CONNAT, J.-L. ( 1990). Anterioposterior gradient during nymphal-adult moulting cycle of the tropical bont tick, Amblyomma variegatum (Acarina: Ixodidae). Correlations between ecdysteroid titers and integument structure. Roux's Archives of Developmental Biology 198, 309–321.Google Scholar
STREY, O. F., TEEL, P. D. & LONGNECKER, M. T. ( 2001). Effects of pyriproxyfen on off-host water-balance and survival of adult lone star ticks (Acari: Ixodidae). Journal of Medical Entomology 38, 589–595.CrossRefGoogle Scholar
TAYLOR, D., CHINZEI, Y., ITO, K., HIGUCHI, N. & ANDO, K. ( 1991 a). Stimulation of vitellogenesis by pyrethroids in mated and virgin female adults, and fourth instar females of Ornithodoros moubata. Journal of Medical Entomology 28, 322–329.Google Scholar
TAYLOR, D., CHINZEI, Y., MIURA, K. & ANDO, K. ( 1991 b). Vitellogenin synthesis, processing and hormonal regulation in the tick, Ornithodoros parkeri (Acari: Argasidae). Insect Biochemistry 21, 723–733.Google Scholar
TAYLOR, D., NAKAJIMA, Y. & CHINZEI, Y. ( 2000). Ecdysteroids and vitellogenesis in the soft tick, Ornithodoros moubata (Acari: Argasidae). In Proceedings of the 3rd International Conference on “Ticks and Tick-borne Pathogens: Into the 21st Century” ( ed. Kazimírová, M., Labuda, M. & Nuttall, P. A.), pp. 223–227. Bratislava, Slovak Republic, Institute of Zoology, Slovak Academy of Sciences.
TAYLOR, D., SONENSHINE, D. E. & PHILLIPS, J. S. ( 1991 c). Ecdysteroids as a component of the genital sex pheromone in two species of hard ticks, Dermacentor variabilis (Say) and Dermacentor andersoni Stiles (Acari: Ixodidae). Experimental and Applied Acarology 12, 275–296.Google Scholar
THUMMEL, C. S. ( 2001) Molecular mechanisms of developmental timing in C. elegans and Drosophila. Developmental Cell 1, 453–465.CrossRefGoogle Scholar
THUMMEL, C. S. ( 2002). Ecdysone-regulated puff genes 2000. Insect Biochemistry and Molecular Biology 32, 113–120.CrossRefGoogle Scholar
TOMASCHKO, K.-H. ( 1994). Ecdysteroids from Pycnogonium litorale (Arthropoda, Pantopoda) act as chemical defense against Carcinus maenas (Crustacea, Decapoda). Journal of Chemical Ecology 20, 1445–1455.CrossRefGoogle Scholar
VENKATESH, K., ROE, R. M., APPERSON, C. S., SONENSHINE, D. E., SCHRIEFER, M. E. & BOLAND, L. M. ( 1990). Metabolism of juvenile hormone during adult development of Dermacentor variabilis (Acari: Ixodidae). Journal of Medical Entomology 27, 36–42.CrossRefGoogle Scholar
WAINWRIGHT, G. & REES, H. H. ( 2001). Hormonal regulation of reproductive development in crustaceans. In Environment and Animal Development (ed. Atkinson, D. & Thorndyke, M.), pp. 71–84. Oxford, Bios.
WEISS, B. L. & KAUFMAN, W. R. ( 2001). The relationship between ‘critical weight’ and 20-hydroxyecdysone in the female ixodid tick, Amblyomma hebraeum. Journal of Insect Physiology 47, 1261–1267.CrossRefGoogle Scholar
WHITEHEAD, D. L., OSIR, E. W., OBENCHAIN, F. D. & THOMAS, L. S. ( 1986). Evidence for the presence of ecdysteroids and preliminary characterization of their carrier proteins in the eggs of the brown ear tick Rhipicephalus appendiculatus (Neumann). Insect Biochemistry 19, 112–133.CrossRefGoogle Scholar
WIGGLESWORTH, K. P., LEWIS, D. & REES, H. H. ( 1985). Ecdysteroid titre and metabolism to novel apolar derivatives in adult female Boophilus microplus (Ixodidae). Archives of Insect Biochemisty and Physiology 2, 39–54.CrossRefGoogle Scholar
WRIGHT, J. E. ( 1969). Hormonal temination of larval diapause in Dermacentor albipictus. Science 163, 390–391.CrossRefGoogle Scholar
ZHU, X. X. & OLIVER, J. H. Jr. ( 2001). Cockroach allatostatin-like immunoreactivity in the synganglion (Acari: Ixodidae). Experimental and Applied Acarology 25, 1005–1013.CrossRefGoogle Scholar
ZHU, X. X., OLIVER, J. H. Jr. & DOTSON, E. M. ( 1991 a). Immunocytochemical localization of an insulin-like substance in the synganglion of the tick, Ornithodoros parkeri (Acari: Argasidae). Experimental and Applied Acarology 13, 153–159.Google Scholar
ZHU, X. X., OLIVER, J. H. Jr. & DOTSON, E. M. ( 1991 b). Epidermis as the source of ecdysone in an argasid tick. Proceedings of the National Academy of Sciences, USA 88, 3744–3747.Google Scholar
ZHU, X. X., OLIVER, J. H. Jr., DOTSON, E. M. & REN, H. L. ( 1994). Correlation between ecdysteroids and cuticulogenesis in nymphs of the tick Ornithodoros parkeri (Acari: Argasidae). Journal of Medical Entomology 31, 479–485.CrossRefGoogle Scholar