Juvenile hormone-like and anti-gonadotropic activities of Fluoromevalonate against the Egyptian cotton leaf worm Spodoptera littoralis Boisd. (Lepidoptera: Noctuidae)

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Juvenile hormone-like and anti-gonadotropic activities of Fluoromevalonate against the Egyptian cotton leaf worm Spodoptera littoralis Boisd. (Lepidoptera: Noctuidae)

Ahmad Basiouny, Karem Ghoneim
Int. J. Agron. Agri. Res.11( 4), 6-30, October 2017.
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The cotton leaf worm, Spodoptera littoralis is destructive pest of cotton and various crops all over the year in Egypt. The present study was carried out aiming to assess the effects of the anti-JH compound, Fluoromevalonate (FMev), on survival, growth, development, metamorphosis and reproductive potential of this pest. Four doses: 100, 50, 25 and 5 µg/larva had been topically applied (once) onto newly moulted 5th (penultimate) instar larvae and newly moulted 6th (last) instar larvae. FMev exhibited a weak toxicity against larvae, pupae and adults. LD50 values were estimated in 42.03 and 629.20 µg/larva, after treatment of 5th and 6th larval instars, respectively. FMev inhibited the larval growth when applied onto 5th instar larvae but promoted it after treatment of 6th instar larvae. The larval was remarkably shortened, but the pupal duration was slightly or remarkably prolonged, depending on the treated larval instar. FMev failed to exhibit anti-JH activity but a JH-like activity against this insect. The pupation rate was regressed. Treatment of 6th instar larvae with FMev resulted in the production of morphologically abnormal pupae, at the higher three doses. FMev exhibited a blocking effect on the adult emergence. FMev exhibited potent anti-gonadotropic activity against S. littoralis, since complete sterilization was recorded.


Abdullah MAR. 2009. Identification of the biological active compounds of two natural extracts for the control of the red palm weevil, Rhynchophorus ferrugineus (Olivier) (Coleoptera: Curculionidae). Egyptian Academic Journal of Biological Sciences 2(2), 35-44.

Adham FK, Rashad EM, Shoukry IF, Nasr EE. 2009. Host plants shifting affects the biology and biochemistry of Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae). Journal of Biological Sciences 2(1), 63-71.

Al-Dali AG, Ghoneim KS, Bakr RF, Bream AS, Tanani MA. 2008. Egg productivity of Schistocerca gregaria (Orthoptera: Acrididae) as affected by the non-steroidal ecdysone agonist Tebufenozide (RH- 5992). The Journal of Egyptian Academic Society for Environmental Development 9(10), 27-38.

Aliabadi FP, Sahragard A, Ghadamyari M. 2016. Lethal and sublethal effects of a chitin synthesis inhibitor, lufenuron, against Glyphodes pyloalis Walker (Lepidoptera: Pyralidae). Journal of Crop Protection 5(2), 203‐214. http://dx.doi.org/10.18869/modares.jcp.5.2.203

Al-Mekhlafi F, Mashaly AM, Abdel Mageed, A, Wadaan MA, Al-Mallah NM. 2012. Overlap effects of cyromazine concentration, treatment method and rearing temperature on the Southern cowpea weevil Callosobruchus maculatus F. reared on Mung Bean.  Pakistan Journal of Zoology 44, 285-290.

Al-Sharook Z, Balan K, Jiang Y, Rembold H. 1991. Insect growth inhibitors from two tropical Meliaceae: Effects of crude seed extracts on mosquito larvae. Journal of Applied Entomology 111, 425-430. http://dx.doi.org/10.1111/j.14390418.1991.tb00344.x

Amiri A, Bandani AR, Ravan S. 2010. Effect of an anti-juvenile hormone agent (Precocene I) on sunn pest, Eurygaster integriceps (Hemiptera: Scutelleridae) development and reproduction. African Journal of Biotechnology 9(36), 5859-5868. http://dx.doi.org/10.5897/AJB09.1751

Amsalem E, Malka O, Grozinger C, Hefetz A. 2014. Exploring the role of juvenile hormone and vitellogenin in reproduction and social behavior in bumble bees. BMC Evolution and Biology 14, Art. 45, 13pp. http://dx.doi.org/10.1186/1471-2148-14-45

Asano S, Kuwano E, Eto M. 1984. Anti-juvenile hormone activity of 1-citronellyl-5-phenylimidazole in the 3rd instar silkworm, Bombyx mori L. (Lepidoptera: Bombycidae). Applied Entomology and Zoology 19(2), 212-220. http://doi.org/10.1303/aez.19.212

Attathom T. 2002. Biotechnology for insect pest control.  Proc. Sat. Forum: “Sustainable Agricultural System in Asia”. Nagoya, Japan, 73-84 p.

Aydin MH, Gurkan MO. 2006. The efficacy of spinosad on different strains of S. littoralis (Boisduval) (Lepidoptera: Noctuidae). Turkish Journal of Biology 30, 5-9.

Bakr RFA, Guneidy NAM, El-Bermawy SM. (2005), Toxicological and biological activities of three IGRs against fourth larval instar of the cotton leafworm Spodoptera littoralis (Boisd.). Journal of Egyptian Academy of Environmental Development 6(4), 103-132.

Bakr RFA, El-barky NM, Abd Elaziz MF, Awad MH, Abd El-Halim HME. 2010. Effect of Chitin synthesis inhibitors (flufenoxuron) on some biological and biochemical aspects of the cotton leafworm Spodoptera littoralis Bosid. (Lepidoptera: Noctuidae). Egyptian Academic Journal of Biological Sciences 2(2), 43-56.

Balamani E, Nair VSK. 1989. Effects of anti-juvenile hormone agents and a juvenile hormone analog on neck-ligated post-feeding last instar larvae of Spodoptera mauritia Boisd. (Lepidoptera: Noctuidae). International Journal of Tropical Insect Science 10(5), 583-589. https://doi.org/10.1017/S1742758400021706

Banerjee S, Kalena GP, Banerji A, Singh AP. 2008. New synthetic precocenoids as potential insect control agents. Journal of Environmental Biology 29(6), 951- 957.

Barnby MA, Klocke JA. 1990. Effects of azadirachtin on levels of ecdysteroids and prothoracicotropic hormone-like activity in Heliothis virescens (Fabr) larvae. Journal of Insect Physiology 36, 125-131. https://doi.org/10.1016/0022-1910(90)90183-G

Bede JC, Teal PE, Goodman WG, Tobe SS. 2001. Biosynthetic pathway of insect juvenile hormone III in cell suspension cultures of the sedge Cyperus iria. Plant Physiology 127(2), 584–593. http://dx.doi.org/10.1104/pp.010264.

Benz G, Ren S-X. 1986. Some effects of the anti-juvenile hormone fluoromevalonate in three lepidopterous species. Bulletin de la société entomologique Suisse 59, 23-30. http://dx.doi.org/10.5169/seals-402199

Bowers WS. 1982. Toxicology of the precocenes. In: “Insecticide Mode of Action” (Coats, J.R. ed.)” pp. 403-427, New York, Academic Press. eBook ISBN: 9780323140515

Bowers WS, Aldrich JR. 1980. In vivo inactivation of denervated corpora allata by precocene II in the bug Oncopeltus fasciatus. Experientia 36, 362-364.

Bowers WS, Aregullin M. 1987. Discovery and identification of an antijuvenile hormone from Chrysanthemum conorarium. The Memórias do Instituto Oswaldo Cruz 82(III), 51-54.

Bownes M. 2004. The regulation of yolk protein gene expression in Drosophila and other flies in higher Diptera. In: “Reproductive Biology of Invertebrates” (Raikhel AS, Sappington TW, Enfield NH, eds.), Science Publishers, USA, (12), 95.

Burt ME, Kuhr RJ, Bowers WS. 1978. Metabolism of precocene II in the cabbage looper and European corn-borer. Pesticide Biochemistry and Physiology 9, 300–303. https://doi.org/10.1016/0048-3575(78)90010-X

Casida JE, Quistad GB. 1998. Golden age of insecticide research: past, present, or future? Annual Review of Entomology 43, 1-16.

Cavalcante VMS, Regis LN. 1992. The reproductive capacity of Panstrongylus  megistus  (Hemiptera:  Reduviidae)  males  treated  with precocenes. The Memórias do Instituto Oswaldo Cruz 87(4), 467-472. http://dx.doi.org/10.1590/S007402761992000400002.

Cedric G. 2005. Insects and humans. Anon: (eds.). Entomology, Third Edition. Springer, Dordrecht, 725-782 p.

Chapman RF. 1998. The insects: structure and function. 4th ed. Cambridge: Cambridge University Press, 116-118.

Chowański S, Kudlewska M, Marciniak P, Rosiński G. 2014. Synthetic Insecticides- is there an Alternative? Polish Journal of Environmental Studies 23( 2), 291-302.

Connat JL. 1988. Effects of different anti-juvenile hormone agents on the fecundity of the female cattle tick Boophilus microplus.  Pesticide Biochemistry and Physiology 30, 28-34. http://dx.doi.org/10.1016/0048-3575(88)90056-9

Connat JL, Nepa MC. 1990. Effects of different anti-juvenile hormone agents on the fecundity of the female tick Ornithodoros moubata. Pesticide Biochemistry and Physiology 37(3), 266-274. https://doi.org/10.1016/0048-3575(90)90133-M

Costa LG, Giordano G, Guizzetti M, Vitalone A. 2008. Neurotoxicity of pesticides: a brief review. Frontiers of BioSciences 13, 1240-1249. http://dx.doi.org/10.2741/2758

Damalas CA, Eleftherohorinos IG. 2011. Pesticide exposure, safety issues, and risk assessment indicators. International Journal of Environmental Research and Public Health 8(5), 1402-1419. http://dx.doi.org/10.3390/ijerph8051402

Darvas B, Kuwano E, Eto, M, Tag al-din MH, Timar T. 1990. Effects of some anti-juvenile hormone agents (Precocenes-2, J-2710, KK-110) on post embryonic development of Neobellieria bullata. Agricultural and Biological Chemistry 54(11), 3045-3047.

Davey KG, Gordon DRB. 1996. Fenoxycarb and thyroid hormones have JH-like effects on the follicle cells of Locasta migratoria in vitro. Archives of Insect Biochemistry and Physiology 32, 613- 626. http://dx.doi.org/10.1002/(SICI)1520-6327

Davies TGE, Field LM, Usherwood PNR, Williamson MS. 2007. DDT, pyrethrins and insect sodium channels. IUBMB Life 59, 151-162. http://dx.doi.org/10.1080/15216540701352042

Dempster C. 1957. The population dynamic of moraccan locust Dociostarus marcocanus in Cyprus. Anti Locust Bulletin 27 p.

Denlinger DL, Yocum GD, Rinehart JP. 2012. Hormonal control of diapause. In: “Insect Endocrinology” (Lawrence IG, ed.). pp: 430-463. Academic Press, San Diego, CA, USA.

Di Ilio V, Cristofaro M, Marchini D, Nobili P, Dallai R. 1999. Effects of a neem compound on the fecundity and longevity of Ceratitis capitata (Diptera: Tephritidae). Journal of Economic Entomology 92, 76-82. https://doi.org/10.1093/jee/92.1.76

Djeghader NEH, Aïssaoui L, Amira K, Boudjelida H. 2014. Impact of a chitin synthesis inhibitor, Novaluron, on the development and the reproductive performance of mosquito Culex pipiens. World Applied Science Journal 29(7), 954-960. http://dx.doi.org/10.5829/idosi.wasj.2014.29.07.82190

Edwards JP, Bergot BJ, Staal GB. 1983. Effects of three compounds with anti-juvenile hormone activity and a juvenile hormone analogue on endogenous juvenile hormone levels in the tobacco horn worm, Manduca sexta. J. Insect. Physiol., 29(1), 83-89. https://doi.org/10.1016/0022-1910(83)90109-9

Edwards JP, Cerf DC, Staal GB. 1985. Inhibition of oötheca production in Periplaneta americana (L.) with the anti juvenile hormone Fluoromevalonate. Journal of Insect Physiology 31(9), 723-728. https://doi.org/10.1016/0022-1910(85)90053-8

Eid MA, Salem MS, El-Ibrashy MT, Abdel-Hamid M. 1982. Effects of precocene II, Cycloheximide and C16-JH on the growth rate and adult morphometrics of Schistocerca gregaria Forsk. Bulletin of the Entomological Society of Egypt, Econnomic Series 13, 95-105.

El-Ibrahsy MT. 1971. The effect of allatectomy upon reproduction and development of the cotton 1eaf-worm Spodoptera littoralis. Journal of Insect Physiology 17, 1783-1790.

El-Ibrashy MT. 1982. Juvenile hormone mimics in retrospect and antagonists in prospect.  Zeitsclirift für angewandte Entomologie 94, 217-236. http://dx.doi.org/10.1111/j.14390418.1982.tb02569.x

El-Ibrashy MT, Aref NB. 1985. Effects of certain juvenoids on growth and morphogenesis in Spodoptera littoralis Boisduval. Journal of Plant Protection in the Tropics, 2(2), 105-116.

El-Khawas MAM, Abd El-Gawad HAS. 2002. The efficiency of two plant extracts (Fenugreek and Lupine) and commercial biofungicide (Biofly) on the cotton leafworm, Spodoptera littoralis (Boisd) (Lepidoptera: Noctuidae) larvae as a new approach of control. Journal of Egyptian German Society of Zoology 37, 39-57. http://dx.doi.org/10.11648/j.cbe.20170202.15

Farag AI, Varjas L. 1983. Precocious metamorphosis and moulting deficiencies induced by an anti-JH compound FMev in the fall webworm, Hyphantria cunea. Entomologia Experimentalis et  Applicata 34, 65-70. http://dx.doi.org/10.1111/j.1570-7458.1983.tb03291.x

Farazmand H, Chaika SYu. 2008. Effects of Precocene-I and II on the development of Colorado potato beetle, Leptinotarsa decemlineata (Col.: Chrysomelidae). Applied Entomology and Phytopathology 76(1), 14 p.

Fescemyer HW, Masler EP, Davis RNE, Kelly TJ. 1992. Vitellogenin synthesis in female larvae of the gypsy moth Lymantria dispar (L.): suppression by juvenile hormone. Comparative Biochemistry and Physiology 103B(3), 533-54. https://doi.org/10.1016/0305-0491(92)90366-Y

Finney DJ. 1971. Probit analysis. 3rd ed. Cambridge, England: Cambridge University Press, 318 p.

Flatt T, Heyland A, Rus F, Porpiglia E, Sherlock C, Yamamoto R, Garbuzov A, Palli SR, Tatar M, Silverman N. 2008. Hormonal regulation of the humoral innate immune response in Drosophila melanogaster. Journal of Experimental Biology 211, 2712-2724. http://dx.doi.org/10.1242/jeb.014878

Gäde G, Goldsworthy GJ. 2003. Insect peptide hormones: a selective review of their physiology and potential application for pest control. Pest Management Science 59(10), 1063–1075. http://dx.doi.org/10.1002/ps.755

Gaur R, Kumar K. 2009. Effect of precocene on morphogenesis in housefly, Musca domestica Linn. Entomon 34, 107-110.

Gaur R, Kumar K. 2010. Insect growth-regulating effects of Withania somnifera in a polyphagous pest, Spodoptera litura. Phytoparasitica 38(3), 237–241. http://dx.doi.org/10.1007/s12600-010-0092-x

Ghoneim KS. 1985. Physiological studies on endocrine and reproductive systems of the cotton leafworm Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae). Ph.D. Thesis, Fac. of Sci., Al-Azhar Univ., Cairo, Egypt.

Ghoneim KS, Mohamed HA, Bream AS. 2000. Efficacy of the neem seed extract, Neemazal, on growth and development of the Egyptian cotton leafworrn, Spodoptera littoralis Boisd. (Lepidoptera: Noctuidae). Journal of Egyptian German Society of Zoology 33, 161-179.

Ghoneim K, Tanani M, Hamadah Kh, Basiouny A, Waheeb H. 2014. Inhibited reproductive capacity of Egyptian cotton leaf worm Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae) by the chitin synthesis inhibitor Novaluron. Egyptian Academic Journal of Biological Sciences 7(2), 105-118.

Ghoneim K, Tanani M, Hamadah Kh, Basiouny A, Waheeb H. 2015. Bioefficacy of Novaluron, a chitin synthesis inhibitor, on survival and development of Spodoptera littoralis (Boisd.)(Lepidoptera: Noctuidae). Journal of Advances in Zoology 1(1), 24-36.

Ghoneim K, Hassan HA, Tanani MA, Bakr NA. 2017a. Toxic and disruptive effects of Novaluron, a chitin synthesis inhibitor, on development of the pink bollworm Pectinophora gossypiella (Saunders)(Lepidoptera: Gelechiidae). International Journal of Entomology Research 2(2), 36-47.

Ghoneim K, Hamadah Kh, Mansour AN, Abo Elsoud AA. 2017b. Toxicity and disruptive impacts of Novaluron, a chitin synthesis inhibitor, on development and metamorphosis of the olive leaf moth Palpita unionalis (Hübner) (Lepidoptera: Pyralidae). International Journal of Trend in Research and Development 4(3), 184-193. Available Online @www.ijtsrd.com

Gilbert LI, Granger NA, Roe RM. 2000. The juvenile hormones: historical facts and speculations on future research directions. Insect Biochemistry and Molecular Biology 30, 617–644. http://dx.doi.org/10.1016/S0965-1748(00)00034-5

Hagedorn HH. 1985. The role of ecdysteroids in reproduction. In: “Comprehensive Insect Physiology, Biochemistry and Pharmacology”(Kerkut GA, Gilbert LI, eds.). Pergamon, Oxford, 8, 205–262.

Hamadah Kh, Ghoneim K, Mansour AN, Abo Elsoud AA. 2017. Deranged adult performance and reproductive potential of the olive leaf moth Palpita unionalis (Hübner)(Lepidoptera: Pyralidae) by the non-steroidal ecdysone agonist, Methoxyfenozide. International Journal of Information Research and Review 04(06), 4228-4240.

Hassan HA, Ghoneim K, Tanani MA, Bakr NA. 2017. Impairing effectiveness of the chitin synthesis inhibitor, Novaluron, on adult performance and reproductive potential of the pink bollworm Pectinophora gossypiella (Saunders) (Lepidoptera: Gelechiidae). Journal of Entomology and Zoology Studies 5(2), 581-592.

Haunerland NH, Bowers WS. 1985. Comparative studies on pharmacokinetics and metabolism of the anti-juvenile hormone precocene II. Archives of Insect Biochemistry and Physiology 2, 55-63. http://dx.doi.org/10.1002/arch.940020106

Hosny MM, Topper CP, Moawad GG, El-Saadany GB. 1986. Economic damage thresholds of Spodoptera littoralis (Boisd.) (Lepidoptera: Noctuidae) on cotton in Egypt. Crop Protection 5, 100-104. https://doi.org/10.1016/0261-2194(86)90088-8

Indrasith L, Sasaki ST, Yaginuma T, Yamashita O. 1988. The occurrence of premature form of egg-specific protein in vitellogenic follicles of Bombyx mori. Journal of Comparative Physiology 158, 1-7.

Ishaaya I, Yablonski S, Horowitz AR. 1995. Comparative toxicity of two ecdysteroid agonists, RH-2485 and RH-5992, on susceptible and pyrethroid-resistant strains of the Egyptian cotton leafworm, Spodoptera littoralis. Phytoparasitica, 23, 139–145.

Islam MS. 1995. Endocrine manipulation in crowd-reared desert locust Schistocerca gregaria (Forskal). II. Effects of Precocene on mortality and reproductive parameters. Bangladesh Journal of Zoology 23, 199-208.

Josephrajkumar A, Subrahmanyam B, Srinivasan S. 1999. Plumbagin and azadirachtin deplete haemolymph ecdysteroid levels and alter the activity profiles of two lysosomal enzymes in the fat body of Helicoverpa armigera (Lepidoptera: Noctuidae). European Journal of Entomology 96: 347-353.

Kadono-Okuda K, Kuwano E, Eto M, Yamashita O. 1987. Inhibitory action of an imidazole compound on ecdysone synthesis in prothoracic glands of the silkworm, Bombyx mori. Growth Differentiation 29, 527-533. http://dx.doi.org/10.1111/j.1440-169X.1987.00527.x

Khafagi WE, Hegazi EM. 1999. Latent effects of precocenes (I and II) and juvenile hormone I on Spodoptera littoralis (Boisd.) larvae. Archives of Phytopathology and Plant Protection 32, 337–350. http://dx.doi.org/10.1080/03235409909383302

Khafagi WE, Hegazi EM. 2001. Effects of juvenile hormones I & II and precocenes I & II on the immune response of Spodoptera littoralis larvae to supernumerary larvae of the solitary parasitoid, Microplitis rufiventris Kok. Journal of Insect Physiology 47, 1249–1259.

Khafagi WE, Hegazi EM. 2004. Effects of juvenile hormone I and precocene I & II on Microplitis rufiventris when administered via its host, Spodoptera littoralis. BioControl 49, 517–536. Article in BioControl 49(5), 517-536 · http://dx.doi.org/10.1023/B:BICO.0000036435.76314.ad

Khan IA, Kumar K. 2000. Precocene II acts as insect growth regulator in polyphagous pest, Pericallia ricini F. (Lepidoptera: Arctiidae). The Proceedings of the National Academy of Sciences, India 70(B), 279-285.

Khan IA, Kumar K. 2005. Effect of precocene II on imaginal differentiation in flesh fly, Sarcophaga ruficornis F. (Diptera: Sarcophagidae). Entomon 30, 187-191.

Khan M, Hossain MA, Islam MS. 2007. Effects of neem leaf dust and a commercial formulation of a neem compound on the longevity, fecundity and ovarian development of the melon fly, Bactocera cucurbitae (Coquillett) and the oriental fruit fly, Bactrocera dorsalis (Hendel) (Diptera: Tephritidae). Pakistan Journal of Biological Sciences 10, 3656-3661. http://dx.doi.org/10.3923/pjbs.2007.3656.3661

Kiuchi M, Kimura K, Akai H. 1985. Induction of trimolters from a tetramolter strain of Bombyx mori by anti-juvenoid treatment. Journal of Sericultural Science of Japan 54(1), 77-81. http://dx.doi.org/10.11416/kontyushigen1930.54.77

Kumar K, Khan IA. 2004. Effect of precocene on development of ovarian follicles in flesh fly, Sarcophaga ruficornis F. Indian Journal of Experimental Biology 42(1), 74-80.

Kuwano E, Eto M. 1983. Terpenoid imidazoles: New anti-juvenile hormones and insecticidal compounds. Proceedings of Tenth International Congress of Plant Protection, Brighton 1, 278.

Kuwano E, Takeya T, Eto M. 1983. Terpenoid imidazoles: new antijuvenile hormones. Agricultural and Biological Chemistry 47, 921-923. http://dx.doi.org/10.1080/00021369.1983.10865749

Kuwano E, Tanaka Y, Kikuchi M, Eto M. 1988. Effects of anti-juvenile hormones and related compounds on development in the larvae of Bombyx mori. Journal of the Faculty of Agriculture, Kyushu University 33(1,2), 17-28.

Lange AB, Phillips DR, Loughton BG. 1983. The effects of precocene II on early adult development in male Locusta. Journal of Insect Physiology 29(1), 73-81. https://doi.org/10.1016/0022-1910(83)90108-7

Lehmann PH, Lyytinen A, Piiroinen S, Lindström L. 2015. Is a change in juvenile hormone sensitivity involved in range expansion in an invasive beetle? Frontiers in Zoology 12, 20. http://dx.doi.org/10.1186/s12983-015-0113-1

Li YM, Kai ZP, Huang J, Tobe SS. 2017. Lepidopteran HMG-CoA reductase is a potential selective target for pest control.  PeerJournal 5, e2881. http://dx.doi.org/10.7717/peerj.2881

Linton YM, Nisbet AJ, Mordue (Luntz) AJ. 1997. The effect of azadirachtin on the testes of the desert locust Schistocerca gregaria (Forskal). Journal of Insect Physiology 43, 1077-1084.  http://dx.doi.org/10.1016/S0022-1910(97)00060-7

Liu XC, Liu ZL. 2014. Evaluation of larvicidal activity of the essential oil of Ageratum conyzoides L. aerial parts and its major constituents against Aedes albopictus. Journal of Entomology and Zoology Studies; 2(4), 345-350.

Lu XN, Liu XC, Liu QZ, Liu ZL. 2014. Isolation of insecticidal constituents from the essential oil of Ageratum houstonianum Mill. against Liposcelis bostrychophila. Journal of Chemistry Article ID http://dx.doi.org/10.1155/2014/6456.87

Magd El-Din S, El-Gengaihi SE. 2000. Joint action of some botanical extracts against the Egyptian cotton leafworm Spodoptera littoralis Bosid645687, 6 p.. (Lepidoptera: Noctuidae). Egyptian Journal of Biological Pest Control 10(1), 51-56.

Mahmoudvand M, Abbasipour H, Sheikhi Garjan A, Bandani AR. 2012. Decrease in pupation and adult emergence of Plutella xylostella (L.) treated with hexaflumuron. Chilean Journal of Agricultural Research 72(2), 206-211. http://dx.doi.org/10.4067/S071858392012000200007.

Mathai S, Nair VSK. 1983. Effects of precocene II on last instar larvae of Spodoptera mauritia (Lepidoptera: Noctuidae). Current Science 52, 376-377.

Mathai S, Nair VSK. 1984. Morphogenetic effects of precocene II on the larval stages of Spodoptera mauritia Boisd. (Lepidoptera: Noctuidae). In: Proceedings of Fifth All India Symposium on Developmental Biology (Agarwal SK, Goel SC, eds.). 59-66 P.

Mayer RT, Witt W, Kitschka GE, Chen AC. 1988. Evidence that chitin synthesis inhibitors affect cell membrane transport. In: “Endocrinological Frontiers in Physiological Insect Ecology”. (Sehnal F, Zabza A, Denlinger DL, eds.). Wroclow Tech. Univ. Press Wroclaw.

Menn JJ. 1985. Prospects of exploitation of insect antijuvenile hormones for selective insect control. In: “Approaches to New Leads for Insecticides” (von Keyserlingk HC, Jäger A, von Szczepanski C. eds). ©Springer-Verlag Berlin Heidelberg.

Miles M, Lysandrou M. 2002. Evidence for negative cross resistance to insecticides in field collected Spodoptera littoralis (Boisd.) from Lebanon in laboratory bioassays. Mededelingen (Rijksuniversiteit te Gent. Fakulteit van de Landbouwkundige en Toegepaste Biologische Wetenschappen) 67, 665.

Mitsuoka T, Takita M, Kanke E, Kawasaki H. 2001. Ecdysteroid titer, responsiveness of prothoracic gland to prothoracicotropic hormone (PTTH), and PTTH release of the recessive trimolter strain of Bombyx mori in extra-ecdysed larvae by JHA and 20E application. Zoological Science, Japan 18(2), 235-240. https://doi.org/10.2108/zsj.18.235

Mondal K, Parween S. 2000. Insect growth regulators and their potential in the management of stored-product insect pests. Integrated Pest Management Review 5, 255-295. http://dx.doi.org/10.1023/A:1012901832162

Moroney MJ. 1956. Facts from figures (3rd ed.). Penguin Books Ltd., Harmondsworth. Middle Sex.

Mosallanejad H, Smagghe G. 2009. Biochemical mechanisms of methoxyfenozide resistance in the cotton leafworm Spodoptera littoralis. Pest Management Science 65, 732-736. http://dx.doi.org/10.1002/ps.1753

Moya P, Castillo M, Primo-Yufera E, Couillaud F, Martinez-Manez R, Garcera MD, Miranda HA, Primo J, Martinez-Pardo R. 1997. Brevioxime: a new juvenile hormone biosynthesis inhibitor isolated from Penicillium brevicompactum. Journal of Organic Chemistry 62, 8544-8545. http://dx.doi.org/10.1021/jo970397y

Muraleedharan D, Varghese A, Abraham G, Mathews RA. 1986. Effect of precocene-II on endocrines, feeding and digestion in the semilooper caterpillar, Achoea Janata. In: “Insect Neurochemistry and Neurophysiology”, 307-314 P. Publisher: Springer. http://dx.doi.org/10.1007/978-1-4612-4832-3_39

Nair VSK, Rajalekshmi E. 1988. Effects of fluoromevalonate on penultimate and last instar larvae of Spodoptera litura Boisd. Indian Journal of Experimental Biology 27, 170-173.

Nandi PS, Chakravarty K. 2011. Juvenoids and anti-Juvenoids as third generation pesticide to control lepidopteran field crop pests. Indian Streams Research Journal 1(6), 15pp. http://dx.doi.org/10.9780/22307850

Nasiruddin M, Mordue (Luntz) AJ. 1994. The protection of barley seedlings from attack by Schistocerca gregaria using azadirachtin and related analogues. Entomologia Experimentalis et Applicata 70, 247-252. http://dx.doi.org/10.1111/j.15707458.1994.tb00753.x

Nassar MM, Hafez ST, Farrag AM, Abahussain MO. 1999.  Efficacy of BaySir-8514 and precocene II against the grey flesh fly Parasarcophaga dux (Thomson) (Diptera: Sarcophagidae). Journal of the Egyptian Society of Parasitology 29(3), 697-707.

Nave JF. d’Orchymont H, Ducep JB, Jung MJ. 1985. Mechanism of inhibition of cholesterol bisynthesis by 6-fluoromevalonate. Biochemistry Journal 227, 247-254.

Newitt RA, Hammock BD. 1986. Relationship between juvenile hormone and ecdysteroids in larval-pupal development of Trichoplusia ni (Lepidoptera:  Noctuidae). Journal of Insect Physiology 32(I0), 835-844. https://doi.org/10.1016/0022-1910(86)90098-3

Okunade AL. 2002. Review: Ageratum conyzoides L. (Asteraceae). Fitoterapia 73, 1-16.

Osman KSA. 1988.  Effect of precocene-3 on the hormonal regulation of oogenesis and larval mortality in Aiolopus thalassinus (Fabr.) (Saltatoria: Acrididae). Mitteilungen der Deutschen Gesellschaft für Allgemeine und Angewandte Entomologie 6, 464-487.

Osman EE, Rarwash I, El-Samadisi MM. 1984. Effect of the anti-moulting agent “Dimilin” on the blood picture and cuticle formation in Spodoptera littoralis (Boisd.) larvae. Bulletin of Entomological Society of Egypt (Economic Series) 14, 3-46.

Parween S, Faruki SI, Begum M. 2001. Impairment of reproduction in the red flour beetle, Tribolium castaneum (Herbst.)(Coleoptera: Tenebrionidae) due to larval feeding on triflumuron-treated diet. Journal of Applied Entomology 125, 1-4. http://dx.doi.org/10.1046/j.1439-0418.2001.00576.x

Pradeep AR, Nair VSK. 2000. Antigonadotropic effects of precocene 2: allaticidal action in females of Nilaparvata lugens (Stal). Philippine Entomologist 14(2), 175-183.

Quistad GB, Cerf DC, Schooley DA, Staal GB. 1981. Fluoromevalonate acts as an inhibitor of insect juvenile hormone biosynthesis. Nature 289, 176-177.

Raikhel AS, Brown MR, Belles X. 2005. Hormonal control of reproductive processes. In: “Comprehensive Molecular Insect Science” (Gilbert LI, Iatrou SS, Gill SS. eds.). 3, 433–492. Oxford, UK: Elsevier.

Raslan SAA. 2002. Preliminary report on initial and residual mortality of the natural product, spinosad for controlling cotton leaf worm egg masses. In: 2002 Cotton Season At Sharkia Governorate, Egypt. 2nd International Conference, Plant Protection Research Institute, Cairo, Egypt, 21-24 December, 2002. 1, 635-637.

Relyea RA. 2009. A cocktail of contaminants: how mixtures of pesticides at low concentrations affect aquatic communities. Oecologia 159, 363-376.

Richard AG. 1957. Cumulative effects of optimum and suboptimum temperatures on insect development. In: “Influence of Temperature on Biological Systems” (Johnson, FH. ed.). Ronald Press Comp., New York, 145-162.

Riddiford LM. 1994. Cellular and molecular action of juvenile hormone. I. General considerations and premetamorphic actions. Advances in Insect Physiology 24, 213-243. https://doi.org/10.1016/S0065-2806(08)60084-3

Riddiford LM. 2008. Juvenile hormone action: A 2007 perspective. Journal of Insect Physiology 54, 895–901. https://doi.org/10.1016/j.jinsphys.2008.01.014

Ringo J, Talyn B, Brannan M. 2005. Effects of precocene and low protein diet on reproductive behavior in Drosophila melanogaster (Diptera: Drosophilidae). Annals of Entomological Society of America 98, 601-607. https://doi.org/10.1603/00138746(2005)098[0601:EOPALP]2.0.CO;2

Roberto CF, Sosa ME, Favier LS, Penna F, Guerreiro E, Giordano OS, Tonn CE. 1998. Growth inhibitory activities of benzofuron and chromene derivatives towards Tenebrio molitor. Journal of Natural Products 61, 1209-1211. http://dx.doi.org/10.1021/np9800248

Rust MK, Hemsarth WLH. 2017. Intrinsic activity of IGRs against larval cat fleas. Journal of Medical Entomology 54(2), 418-421. https://doi.org/10.1093/jme/tjw201

Salem H, Smagghe G, Degheele D. 1997. Effects of tebufenozide on oocyte growth in Plodia interpunctella. Medical Faculty, Landbouww, Gent University 62(1), 9-13.

Sallam MH. 1999. Effect of Diflubenzuron on embryonic development of the acridid, Heteracris littoralis. Journal of Egyptian German Society of Zoology 30(E), 17-26.

Sammour EA, Kandit MA, Abdel-Aziz NF. 2008. The reproductive potential and fat of chlorfluazuron and lufenuron against cotton leafworm, Spodoptera littoralis (Boisd). American-Eurasian Journal of Agricultural and Environmental Sciences 4(1), 62-67.

Sánchez CM, Martín JMP, Jin J-S, Dávalos A, Zhang W, de la Peña G, Martínez-Botas J, Rodríguez-Acebes S, Suárez Y, Hazen MJ, Gómez-Coronado D, Busto R, Cheng Y-C, Lasunción MA. 2015. Disruption of the mevalonate pathway induces dNTP depletion and DNA damage. Biochimica et Biophysica Acta 1851, 1240-1253. https://doi.org/10.1016/j.bbalip.2015.06.001

Sariaslani FS, McGee LR, Ovenall DW. 1987. Microbial transformation of precocene II: Oxidative reactions by Streptomyces griseus.  Applied and Environmental Microbiology 53(8), 1780-1784.

Saxena RC, Sharma M, Kumar ML, Bansal SK, Shrivastava D. 1994. Developmental effects of 6, 7-dimethoxy-2,2-dimethyl chromene on the preimaginal stages of Anopheles stephensi.  Proceedings of Academy of Environmental Biology 3, 181-184.

Schwedes CC, Carney GE. 2012. Ecdysone signaling in adult Drosophila melanogaster. Journal of Insect Physiology 58, 293–302. https://doi.org/10.1016/j.jinsphys.2012.01.013

Shonouda ML, Osman SL. 2000. New botanical derivatives, used in medicinal reparations, showing bioactive action on insect pests.1-Toxicological effect on the development of Spodoptera littoralis (Boisduval) (Lepidoptera: Noctuidae). Journal of Egyptian German Society of Zoology 31, 227-234.

Shuto A, Kuwano E, Eto M. 1988. Synthesis of two optical isomers of the insect anti-juvenile hormone agent fluoromevalonolactone (FMev) and their biological activities. Agricultural and Biological Chemistry 52(4), 915-919. http://dx.doi.org/10.1080/00021369.1988.10868759

Singh S, Kumar K. 2011. Sensitivity of last larval stadium of Chrysomya megacephala (Fabricius) to anti-allatin ageratochromene Precocene II. Indian journal of Experimental Biology 49, 600-608.

Smagghe G, Degheele D. 1994. The  significance  of  pharmacokinetics  and  metabolism  to  the  biological  activity  of  RH-5992 (tebufenozide)  in  Spodoptera  exemptaSpodoptera  exigua and Leptinotarsa decemlineata. Pesticide Biochemistry and Physiology 49, 224-234. https://doi.org/10.1006/pest.1994.1050

Smagghe G, Carton B, Wesemael W, Ishaaya I, Tirry L. 1999. Ecdysone agonists-mechanism of action and application on Spodoptera species. Pesticide Science 55, 343-389. http://dx.doi.org/10.1002/(SICI)10969063(199903)55:3<386::AID-PS922>3.0.CO;2-5

Soller M, Bownes M, Kubli E. 1997. Mating and sex peptide stimulate the accumulation of yolk in oocytes of Drosophila melanogaster. European Journal of Biochemistry 243, 732–738.

Soltani N, Mazouni A. 1992. Diflubenzuron and oogenesis in the codling moth, Cydia pomonella. Pesticide Science 34, 257-261. http://dx.doi.org/10.1002/ps.2780340311

Sparks TC, Roe RM, Buehler A, Hammock BD. 1987. The evaluation of anti-juvenile hormones using last stadium larvae of the cabbage looper Trichplusia ni (Hubner). Insect Biochemistry 17(7), 1011-1016. https://doi.org/10.1016/0020-1790(87)90112-0

Srivastava S, Kumar K. 1996. Effect of precocenes on metamorphosis of flesh fly, Sarcophaga ruficornis F. Indian Journal of Experimental Biology 34, 547-553.

Srivastava S, Kumar K. 1997. Precocene I and II induced metamorphosis in a noctuid moth, Spodoptera litura Fabr. The Proceedings of the National Academy of Sciences, India 67(B), 213-226.

Srivastava S, Kumar K. 1999. Juvenilizing effects of ethoxyprecocene in a lepidopteran insect. Indian Journal of Experimental Biology 37(4), 379-383. http://hdl.handle.net/123456789/19032

Staal GB. 1982. Insect control with growth regulation interfering with the endocrine system. Entomologia Experimentalis et Applicata 31, 15–23. http://dx.doi.org/10.1111/j.1570-7458.1982.tb03115.x

Staal GB. 1986. Anti-juvenile hormone agents. Annual Review of Entomology 31, 391-429. https://doi.org/10.1146/annurev.en.31.010186.002135

Subrahmanzam B, Müller T, Rembold H. 1989. Inhibition of turnover of neurosecretion by azadirachtin in Locusta migratoria. Journal of Insect Physiology 35, 493-500. https://doi.org/10.1016/0022-1910(89)90056-5

Szczepanik M, Obara R, Szumny A, Gabryś B, Halarewicz-Pacan A, Nawrot J, Wawrzeńczyk C. 2005. Synthesis and insect antifeedant activity of precocene derivatives with lactone moiety. Journal of Agriculture and Food Chemistry 53, 5905-5910. http://dx.doi.org/10.1021/jf058034j

Tarrant C, Cupp EW, Bowers WS. 1982. The effects of precocene II on reproduction and development of triatomine bugs (Reduviidae: Triatominae). American Journal of Tropical Medicine and Hygiene 31(2), 416-420. http://dx.doi.org/10.4269/ajtmh.1982.31.416

Tatar M, Kopelman A, Epstein D, Tu M-P, Yin C-M, Garofalo RS. 2001a. A mutant Drosophila insulin receptor homolog that extends life-span and impairs neuroendocrine function. Science 292, 107-110. http://dx.doi.org/10.1126/science.1057987

Tatar M, Priest N, Chien S. 2001b. Negligible senescence during reproductive diapause in D. melanogaster. The American Naturalist 158, 248–258. http://dx.doi.org/10.1086/321320

Tateishi K, Kiuchi M, Takeda S. 1993. New cuticle formation and moult inhibition by RH-5849 in the common cutworm, Spodoptera litura (Lepidoptera: Noctuidae). Applied Entomology and Zoology 28, 177-184. http://doi.org/10.1303/aez.28.177

Telfer WH. 2009. Egg formation in Lepidoptera. Journal of Insect Science 9, 50. insectscience.org/9.50.

Tiryaki D, Temur C. 2010. The fate of pesticide in the environment. Journal of Biological and Environmental Sciences 4(10), 29-32.

Toppozada A, Abd-allah S, El-Defrawi ME. 1966. Chemosterilization of larvae and adults of the Egyptian cotton leafworm, Prodenia litura by apholate, metepa and hempa. Journal of Economic Entomologt 59, 1125-1128.

Trigo JR, Campos S, Pereira AM. 1988. Presença de alcalóides pirrolizidinicos em Ageratum conyzoides L. In: “Simposio de Plantas Medicinais do Brasil”, Sao Paulo. (Resumos). p. 13.

Truman JW, Riddiford LM. 2007. The morphostatic actions of juvenile hormone. Insect Biochemical and Molecular Biology 37, 761–770. https://doi.org/10.1016/j.ibmb.2007.05.011

Van Der Gaag N. 2000. Pick your poison. New Internationalist, 323, 9-11.

Vyas AV, Mulchandani NB. 1984. New chromenes from Ageratum conyzoides. In: “Progress Report1980-1983” (Choughley ASU, Heblo MR.eds)  BARC, Bio-organic Division. p.7.

Walkowiak K, Spochacz M, Rosinski G. 2015. Peptidomimetics- A new class of bioinsecticides. Postepy Biologii Komorki 42(2), 235-254.

Wang Y, Wang M. 2007. The research of IGRs. World Pesticides 29, 8-11.

Wigglesworth VB. 1984. Insect Physiology. 8th ed., Chapman & Hall, London, 191 p.

Wilson TG. 2004. The molecular site of action of juvenile hormone and juvenile hormone insecticides during metamorphosis: how these compounds kill insects. Journal of Insect Physiology 50(2/3), 111-121. https://doi.org/10.1016/j.jinsphys.2003.12.004

Wu J. 2002. The application of IGRs in agricultural pest control. Pesticides 41, 6-8.

Yasyukevich VV, Zvantsov AB. 1999. Effects of antihormones on the susceptibility of Anopheles sacharovi mosquitoes to the malaria causative organism Plasmodium gallinaceum.  Meditsinskaya Parazitologiya i Parazitarnye Bolezni 68, 46-48. 

Yoshida T, Shiotsuki T, Kuwano E. 2000. Synthesis and precocious metamorphosis-inducing activity of 3-(1-alkenyl)pyridines. Journal of Pesticide Science 25, 253-258. http://doi.org/10.1584/jpestics.25.253

Zhou Z, Deng G, Luo S. 2003. Study and application of IGRs. Journal of Guangxi Agricultural Sciences 1, 34-36.

Zhou F, Zhu G, Zhao H, Wang Z, Xue M, Li X, Xu H, Ma X, Liu Y. 2016. Sterilization effects of adult-targeted baits containing insect growth regulators on Delia antiqua. Scientific Reports 6,  32855; 9 p. http://dx.doi.org/10.1038/srep32855.