TY - JOUR
AU - Haff, Ronald P.
AB - Abstract Male adult navel orangeworm, Amyelois transitella (Walker), were irradiated using a laboratory scale x-ray irradiation unit to determine the required dose for complete egg sterility of mated female moths and inherited sterility of F1 and F2 generations. Adult male A. transitella were irradiated in two separate experiments at 100–300 Gy and 50–175 Gy. Mating frequency, fecundity, and fertility of normal females crossed with irradiated parental males was compared with the mating of nonirradiated moths. Mating frequency was 100% for females crossed with nonirradiated control males. At male treatment doses of ≥150 Gy the percentage of females found unmated increased, while multiple-mated females decreased. Female fecundity was not affected while fertility was affected in a dose-dependent relationship to exposure of parental males to x-ray irradiation. Embryonic development of eggs to the prehatch stage and egg eclosion did not occur at radiation doses ≥125 Gy. Emergence of F1 adults was low and occurred only for progeny of parental males exposed to doses ≤100 Gy, with no emergence at ≥125 Gy. Though fecundity appeared similar for control and irradiated F1 females, no F2 eggs hatched for the test exposures of 50–100 Gy. Based on our results, a dose of ≥125 Gy had efficacy in inducing both primary parental sterility in treated male moths and inherited sterility in F1 male and female moths. Results suggest that A. transitella might be considered a candidate for the sterile insect technique using adults irradiated at these relatively low x-ray exposure doses. Amyelois transitella, navel orangeworm, irradiation, x-ray, sterility The navel orangeworm, Amyelois transitella (Walker) (Lepidoptera: Pyralidae), is a severe pest of California tree nuts, causing economic injury by feeding upon the kernels of almonds, Prunus dulcis (Mill.), pistachios, Pistachia vera L., and walnuts, Juglans regia L. In addition, the boring activity and presence of A. transitella is associated with the infection of nut kernels with fungal spores of Aspergillus species and subsequent contamination by mycotoxins, primarily aflatoxins ( Palumbo et al. 2014 ). Aflatoxins and other mycotoxins have become an ever-increasing worldwide concern to food safety and public health as potential carcinogens and animal toxins, and are strictly regulated in their contamination tolerances at ppb levels ( Campbell et al. 2003 ). Historically control of A. transitella had been highly successful with use of organophosphate insecticides ( Higbee and Siegel 2012 ), but use of such products is under regulatory restriction and banning by the EPA (Food Quality Protection Act 1996) for food and environmental safety concerns. Alternative reduced-risk insecticides are effective against A. transitella but costly ( Higbee and Siegel 2012 ). Moreover, a difficult management problem occurs at “husk-split,” the critical ca . 3-wk period prior to harvest when the newly opened nuts are highly vulnerable to A. transitella attack and synthetic insecticides are restricted from use ( Hamby et al. 2011 ). Therefore, for effective control an integrated program has been developed using insecticides augmented with pheromone mating disruption ( Higbee and Burks 2008 ). The development of mating disruption for A. transitella has made considerable advances over the past 10 yr, but still is problematic, plagued with cost and efficacy concerns. A population suppression technology that would complement mating disruption and enhance control efficacy when integrated in area-wide programs is the sterile insect technique (SIT; Bloem et al. 2005 ; Bakri et al. 2005a , b ; Carpenter et al. 2005 ). SIT requires the mass-rearing, sterilization, and release of insects that then mate with a targeted wild population causing infertile offspring. SIT has been highly successful in suppressing wild populations of many Lepidopteran pests, including the codling moth, Cydia pomonella (L.) (Lepidoptera: Tortricidae), and the pink bollworm, Pectinophora gossypiella (Saunders) (Lepidoptera: Gelechiidae) ( Bloem et al. 2005 ). The radiation biology of a large number of Lepidopteran pests including worldwide pyralid moths has been investigated in support of potential development of SIT suppression programs ( Bakri et al. 2005a , b ). Induced sterility through gamma irradiation of moth adults has been studied in 18 Pyralidae species, ten pests that feed directly on host plants and eight pests that feed on mature commodities – stored products ( Table 1 ). In addition, induced sterility has been investigated in five other pyralid species through gamma-ray irradiation applied to the egg and larval stages: sugarcane shoot borer, Chilo infuscatellus Snellen ( Fatima et al. 2009 ) or pupal stage: sugarcane stalk borer, Chilo auricilius Dudgeon ( Kusumahadi and Hudaya 1988 ), pickleworm, Diaphania nitidalis (Stoll) ( Elsey and Brower 1984 ), southwest corn borer, Diatraea grandiosella Dyar ( Hallman 2004 ), and Mexican rice borer, Eoreuma loftini ( Hallman 2004 ). Sterilization effects of gamma irradiation on A. transitella life stages were first reported by Husseiny and Madsen (1964) . Johnson and Vail (1988 , 1989 ) studied gamma irradiation of A. transitella larval-infested nuts for phytosanitation purposes. Table 1. Ionizing irradiation doses (Gy) applied to adult pyralid moth pests reported to evoke complete sterility, substerility, inherited sterility in progeny, or diminished competitive mating in P1 adults Species and common name . Sterility dose . Substerility dose evoking F1–F2 sterility . Dose disrupting competitive mating (nondisruptive dose) . Reference . Amyelois transitella (Walker), navel orangeworm 500 (>540) Husseiny and Madsen 1964 Cactoblastis cactorum (Berg), cactus moth 500–males >200–females >200–males 200 females Carpenter et al. 2001 200–females <200 females Bloem et al. 2003 200 Tate et al. 2007 (>200) Marti and Carpenter 2009 Chilo partellus (Swinhoe), maize borer 150–males Bughio 1976 100–females 150–males (150) 100–F1 Bughio 1988 Chilo suppressalis (Walker), rice stem borer 230–males Chung and Ryu 1971 200–300 Chiang 1972 Corcyra cephalonica (Stainton) , rice moth 450–males (450) Chand and Sehgal 1979 300 Abdel-Baky and Hasaballa 1991 Crocidolomia binotalis Zeller, cabbage webworm >325 Sutrisno Apu 1983 200 Sutrisno Apu 2001 Diatraea saccharalis (F.), sugarcane borer 300 ≥20 (F2) Walker and Quintana 1968 ≥150 Sanford 1976 , 1977 500–males Sgrillo and Wiendl 1981 Ectomyelois ceratoniae Zeller, pomegranate fruit moth 500 Al-Izzi et al. 1990 ≥250 (300) Al-Izzi et al. 1993 <500 Dhouibi and Abderahmane 2001 Eldana saccharina Walker, African sugarcane borer >350 250 Walton et al. 2011 >200 Mudavanhu et al. 2011 250–males Mudavanhu et al. 2014 200–females Ephestia calidella (Guenee), carob moth or dried fruit moth 400–males Boshra and Mikhaiel 2006 350–females Ephestia cautella (Walker), almond moth–fig moth 450–males Calderón and Gonen 1971 400–females 400 350 350 Gonen and Calderon 1971 1,000–males, Cogburn et al. 1973 300–females 300–males 200–males Amoako-Atta et al. 1978 600–males, Brower 1979b 500–females 350 Brower 1979c 200 Brower 1980 350 200 Al-Taweel et al. 1989 Al-Taweel et al. 1990 Ephestia eleutella (Hubner), tobacco moth >150 Brower 1982 450–males, Brower and Tilton 1985 300–females Ephestia kuehniella Zeller , Mediterranean flour moth 150 Riemann 1973 175–200 Marec et al. 1999 300 Ayvaz and Tuncbilek 2006 ≥200 Ayvaz et al. 2007 Galleria mellonella (L.), greater wax moth 220–males, Walker et al. 1975 132–females >300–males, Flint and Merkle 1983 >100–females 350 Jafari et al. 2010 Omphisa anastomosalis (Guenee), sweetpotato vine borer ≥150 150 Follett 2006 Ostrinia furnacalis (Guenee), Asian corn borer 200–300 Li et al. 1988 200 Kang et al. 1993 ≥200 Wang et al. 2001 Ostrinia nubilalis (Hubner), European corn borer 250–female 250–male Zhang and Lou 1980 300–females 150–300 Rosca and Barbulescu 1989 150 (>150) Rosca and Barbulescu 1990 Plodia interpunctella (Hubner), Indian meal moth 1,000 450 Cogburn et al. 1966 >450–males Ashrafi et al. 1972 670–males, 750–1,000 Brower 1975 450–females 500 250–350 Brower 1976a (>500) Brower 1976b 500–males 350 >500–750 Ahmed et al. 1976 150 (F2) Brower 1979a 150–200 -females Brower 1981 336–388 Hallman and Phillips 2008 500 Aye et al. 2008 Species and common name . Sterility dose . Substerility dose evoking F1–F2 sterility . Dose disrupting competitive mating (nondisruptive dose) . Reference . Amyelois transitella (Walker), navel orangeworm 500 (>540) Husseiny and Madsen 1964 Cactoblastis cactorum (Berg), cactus moth 500–males >200–females >200–males 200 females Carpenter et al. 2001 200–females <200 females Bloem et al. 2003 200 Tate et al. 2007 (>200) Marti and Carpenter 2009 Chilo partellus (Swinhoe), maize borer 150–males Bughio 1976 100–females 150–males (150) 100–F1 Bughio 1988 Chilo suppressalis (Walker), rice stem borer 230–males Chung and Ryu 1971 200–300 Chiang 1972 Corcyra cephalonica (Stainton) , rice moth 450–males (450) Chand and Sehgal 1979 300 Abdel-Baky and Hasaballa 1991 Crocidolomia binotalis Zeller, cabbage webworm >325 Sutrisno Apu 1983 200 Sutrisno Apu 2001 Diatraea saccharalis (F.), sugarcane borer 300 ≥20 (F2) Walker and Quintana 1968 ≥150 Sanford 1976 , 1977 500–males Sgrillo and Wiendl 1981 Ectomyelois ceratoniae Zeller, pomegranate fruit moth 500 Al-Izzi et al. 1990 ≥250 (300) Al-Izzi et al. 1993 <500 Dhouibi and Abderahmane 2001 Eldana saccharina Walker, African sugarcane borer >350 250 Walton et al. 2011 >200 Mudavanhu et al. 2011 250–males Mudavanhu et al. 2014 200–females Ephestia calidella (Guenee), carob moth or dried fruit moth 400–males Boshra and Mikhaiel 2006 350–females Ephestia cautella (Walker), almond moth–fig moth 450–males Calderón and Gonen 1971 400–females 400 350 350 Gonen and Calderon 1971 1,000–males, Cogburn et al. 1973 300–females 300–males 200–males Amoako-Atta et al. 1978 600–males, Brower 1979b 500–females 350 Brower 1979c 200 Brower 1980 350 200 Al-Taweel et al. 1989 Al-Taweel et al. 1990 Ephestia eleutella (Hubner), tobacco moth >150 Brower 1982 450–males, Brower and Tilton 1985 300–females Ephestia kuehniella Zeller , Mediterranean flour moth 150 Riemann 1973 175–200 Marec et al. 1999 300 Ayvaz and Tuncbilek 2006 ≥200 Ayvaz et al. 2007 Galleria mellonella (L.), greater wax moth 220–males, Walker et al. 1975 132–females >300–males, Flint and Merkle 1983 >100–females 350 Jafari et al. 2010 Omphisa anastomosalis (Guenee), sweetpotato vine borer ≥150 150 Follett 2006 Ostrinia furnacalis (Guenee), Asian corn borer 200–300 Li et al. 1988 200 Kang et al. 1993 ≥200 Wang et al. 2001 Ostrinia nubilalis (Hubner), European corn borer 250–female 250–male Zhang and Lou 1980 300–females 150–300 Rosca and Barbulescu 1989 150 (>150) Rosca and Barbulescu 1990 Plodia interpunctella (Hubner), Indian meal moth 1,000 450 Cogburn et al. 1966 >450–males Ashrafi et al. 1972 670–males, 750–1,000 Brower 1975 450–females 500 250–350 Brower 1976a (>500) Brower 1976b 500–males 350 >500–750 Ahmed et al. 1976 150 (F2) Brower 1979a 150–200 -females Brower 1981 336–388 Hallman and Phillips 2008 500 Aye et al. 2008 Open in new tab Table 1. Ionizing irradiation doses (Gy) applied to adult pyralid moth pests reported to evoke complete sterility, substerility, inherited sterility in progeny, or diminished competitive mating in P1 adults Species and common name . Sterility dose . Substerility dose evoking F1–F2 sterility . Dose disrupting competitive mating (nondisruptive dose) . Reference . Amyelois transitella (Walker), navel orangeworm 500 (>540) Husseiny and Madsen 1964 Cactoblastis cactorum (Berg), cactus moth 500–males >200–females >200–males 200 females Carpenter et al. 2001 200–females <200 females Bloem et al. 2003 200 Tate et al. 2007 (>200) Marti and Carpenter 2009 Chilo partellus (Swinhoe), maize borer 150–males Bughio 1976 100–females 150–males (150) 100–F1 Bughio 1988 Chilo suppressalis (Walker), rice stem borer 230–males Chung and Ryu 1971 200–300 Chiang 1972 Corcyra cephalonica (Stainton) , rice moth 450–males (450) Chand and Sehgal 1979 300 Abdel-Baky and Hasaballa 1991 Crocidolomia binotalis Zeller, cabbage webworm >325 Sutrisno Apu 1983 200 Sutrisno Apu 2001 Diatraea saccharalis (F.), sugarcane borer 300 ≥20 (F2) Walker and Quintana 1968 ≥150 Sanford 1976 , 1977 500–males Sgrillo and Wiendl 1981 Ectomyelois ceratoniae Zeller, pomegranate fruit moth 500 Al-Izzi et al. 1990 ≥250 (300) Al-Izzi et al. 1993 <500 Dhouibi and Abderahmane 2001 Eldana saccharina Walker, African sugarcane borer >350 250 Walton et al. 2011 >200 Mudavanhu et al. 2011 250–males Mudavanhu et al. 2014 200–females Ephestia calidella (Guenee), carob moth or dried fruit moth 400–males Boshra and Mikhaiel 2006 350–females Ephestia cautella (Walker), almond moth–fig moth 450–males Calderón and Gonen 1971 400–females 400 350 350 Gonen and Calderon 1971 1,000–males, Cogburn et al. 1973 300–females 300–males 200–males Amoako-Atta et al. 1978 600–males, Brower 1979b 500–females 350 Brower 1979c 200 Brower 1980 350 200 Al-Taweel et al. 1989 Al-Taweel et al. 1990 Ephestia eleutella (Hubner), tobacco moth >150 Brower 1982 450–males, Brower and Tilton 1985 300–females Ephestia kuehniella Zeller , Mediterranean flour moth 150 Riemann 1973 175–200 Marec et al. 1999 300 Ayvaz and Tuncbilek 2006 ≥200 Ayvaz et al. 2007 Galleria mellonella (L.), greater wax moth 220–males, Walker et al. 1975 132–females >300–males, Flint and Merkle 1983 >100–females 350 Jafari et al. 2010 Omphisa anastomosalis (Guenee), sweetpotato vine borer ≥150 150 Follett 2006 Ostrinia furnacalis (Guenee), Asian corn borer 200–300 Li et al. 1988 200 Kang et al. 1993 ≥200 Wang et al. 2001 Ostrinia nubilalis (Hubner), European corn borer 250–female 250–male Zhang and Lou 1980 300–females 150–300 Rosca and Barbulescu 1989 150 (>150) Rosca and Barbulescu 1990 Plodia interpunctella (Hubner), Indian meal moth 1,000 450 Cogburn et al. 1966 >450–males Ashrafi et al. 1972 670–males, 750–1,000 Brower 1975 450–females 500 250–350 Brower 1976a (>500) Brower 1976b 500–males 350 >500–750 Ahmed et al. 1976 150 (F2) Brower 1979a 150–200 -females Brower 1981 336–388 Hallman and Phillips 2008 500 Aye et al. 2008 Species and common name . Sterility dose . Substerility dose evoking F1–F2 sterility . Dose disrupting competitive mating (nondisruptive dose) . Reference . Amyelois transitella (Walker), navel orangeworm 500 (>540) Husseiny and Madsen 1964 Cactoblastis cactorum (Berg), cactus moth 500–males >200–females >200–males 200 females Carpenter et al. 2001 200–females <200 females Bloem et al. 2003 200 Tate et al. 2007 (>200) Marti and Carpenter 2009 Chilo partellus (Swinhoe), maize borer 150–males Bughio 1976 100–females 150–males (150) 100–F1 Bughio 1988 Chilo suppressalis (Walker), rice stem borer 230–males Chung and Ryu 1971 200–300 Chiang 1972 Corcyra cephalonica (Stainton) , rice moth 450–males (450) Chand and Sehgal 1979 300 Abdel-Baky and Hasaballa 1991 Crocidolomia binotalis Zeller, cabbage webworm >325 Sutrisno Apu 1983 200 Sutrisno Apu 2001 Diatraea saccharalis (F.), sugarcane borer 300 ≥20 (F2) Walker and Quintana 1968 ≥150 Sanford 1976 , 1977 500–males Sgrillo and Wiendl 1981 Ectomyelois ceratoniae Zeller, pomegranate fruit moth 500 Al-Izzi et al. 1990 ≥250 (300) Al-Izzi et al. 1993 <500 Dhouibi and Abderahmane 2001 Eldana saccharina Walker, African sugarcane borer >350 250 Walton et al. 2011 >200 Mudavanhu et al. 2011 250–males Mudavanhu et al. 2014 200–females Ephestia calidella (Guenee), carob moth or dried fruit moth 400–males Boshra and Mikhaiel 2006 350–females Ephestia cautella (Walker), almond moth–fig moth 450–males Calderón and Gonen 1971 400–females 400 350 350 Gonen and Calderon 1971 1,000–males, Cogburn et al. 1973 300–females 300–males 200–males Amoako-Atta et al. 1978 600–males, Brower 1979b 500–females 350 Brower 1979c 200 Brower 1980 350 200 Al-Taweel et al. 1989 Al-Taweel et al. 1990 Ephestia eleutella (Hubner), tobacco moth >150 Brower 1982 450–males, Brower and Tilton 1985 300–females Ephestia kuehniella Zeller , Mediterranean flour moth 150 Riemann 1973 175–200 Marec et al. 1999 300 Ayvaz and Tuncbilek 2006 ≥200 Ayvaz et al. 2007 Galleria mellonella (L.), greater wax moth 220–males, Walker et al. 1975 132–females >300–males, Flint and Merkle 1983 >100–females 350 Jafari et al. 2010 Omphisa anastomosalis (Guenee), sweetpotato vine borer ≥150 150 Follett 2006 Ostrinia furnacalis (Guenee), Asian corn borer 200–300 Li et al. 1988 200 Kang et al. 1993 ≥200 Wang et al. 2001 Ostrinia nubilalis (Hubner), European corn borer 250–female 250–male Zhang and Lou 1980 300–females 150–300 Rosca and Barbulescu 1989 150 (>150) Rosca and Barbulescu 1990 Plodia interpunctella (Hubner), Indian meal moth 1,000 450 Cogburn et al. 1966 >450–males Ashrafi et al. 1972 670–males, 750–1,000 Brower 1975 450–females 500 250–350 Brower 1976a (>500) Brower 1976b 500–males 350 >500–750 Ahmed et al. 1976 150 (F2) Brower 1979a 150–200 -females Brower 1981 336–388 Hallman and Phillips 2008 500 Aye et al. 2008 Open in new tab Use of x-rays as an alternative irradiation technology to gamma-rays has been advocated on an increasing scale to avoid the expenditures required in safely protecting gamma facilities and securing their ionizing isotope sources ( Suckling 2003 , Mastrangelo et al. 2010 , Mehta and Parker 2011 ). X-ray technology presents a highly secure and less vulnerable method of irradiation than gamma sources, thus increasing the potential for the safe expansion of SIT programs worldwide. Recent studies investigating use of x-ray irradiation to sterilize Lepidoptera have included a pyralid pest, the sweetpotato vine borer, O. anastomosalis ( Follett 2006 ), and two tortricid pests the Mexican leafroller, Amorbia emigratella Busck ( Follett 2008 ), and the light brown apple moth, Epiphyas postvittans (Walker) ( Follett and Snook 2012) , and a gelechiid, Tuta absoluta (Meyrick) ( Cagnotti et al. 2012 ). The scope of this study was to demonstrate the suitability and sensitivity of adult A. transitella for sterilization by x-ray irradiation and thereby contribute knowledge critical to the assessment of the feasibility of a SIT program for population suppression of A. transitella. In the initial gamma radiation study on A. transitella , Husseiny and Madsen (1964) found that adult A. transitella were more radiotolerant than larvae and that male moths were more radiotolerant and challenging for sterilization than females, as has been reported for many other pyralids and insects ( Bakri et al. 2005a , b ; Carpenter et al. 2005 ; Table 1 ). Therefore, this study investigated the efficacy of a broad range of doses of x-ray irradiation for sterilizing parental males. These irradiated males were crossed with normal nonirradiated females and their resultant egg fecundity, fertility, and eclosion of F1 progeny were compared to the productivity of the mating of normal moths. Also, inherited sterility was assessed through emergence of F1 adults and their outcrossing with fertile counterparts followed again by assessment of resultant egg fecundity, fertility, and eclosion of F2 progeny. Materials and Methods Insect Rearing A laboratory colony of A. transitella was established in the winter of 2010 from both larvae and adults emerging from field-infested walnuts of various varieties picked from the ground in orchards of USDA-ARS National Clonal Germplasm Repository (Davis, CA) located at the Wolfskill Experimental Station, Winters, CA. Emergent adults were placed in 1-gal glass mating-jars, provided cotton-wicked vials containing a 4% honey water solution and twigs of almond foliage, and allowed to mate. Jar openings were covered by paper towel sheets having a dimpled surface to encourage oviposition. Sheets of eggs for colony rearing were then disinfected for microbes by soaking for 15 min in a 10% formaldehyde solution followed by 15 min continuous distilled water rinsing. Colony rearing was conducted in 1-gal jars with fine-mesh copper screen lids. Treated egg sheets were placed on a simple, established red-flake organic wheat bran-based diet (with glycerin, honey, brewers’ yeast, and Vanderzants vitamin stock; USDA-ARS-SJVASC, Parlier, CA) upon which hatched larvae were reared until adult emergence. Rearing room was maintained at 28–32°C and a photoperiod of 16:8 (L:D) h. To provide virgin moths for the irradiation experiments, rearing jars were emptied in the morning of moths, then in the afternoon as new moths emerged they were singularly transferred to individual clear plastic 30-ml portion-cups with lids (4 cm opening by 5 cm height) and their gender determined under a dissecting microscope. X-Ray Irradiation Treatment Irradiation treatments were performed in a custom laboratory x-ray cabinet (CXR-105 x-ray tubes, Comet Technologies USA Inc., Stamford, CT). Four x-ray tubes were powered with 100 kV at 10mA and situated around a rotating carousel target area at a distance of 11 cm from each source to the center of the target cup. The applied dose in the sample cups was measured with a radiation measurement system (Accu-Dose MNL/2086, Radcal Corporation, Monrovia, CA) using a high dose rate ion chamber (10X6-0.18, Radcal Corporation) at an emission ranging from 50 to 60 Gy/min with treatment doses based on exposure time. Two sets of treatments were administered in separate experiments. Experimental Design Adult sterility tests were conducted on unmated 1-d-old moths. Cohorts of three males were placed in replicate individual plastic portion-cups. Cups of males were either irradiated or designated as nonirradiated controls, while all females were untreated. Two dose range experiments on male moths were conducted in a randomized design: 1) a five-dose test of exposure to 100–300 Gy in 50 Gy intervals ( n  = 5) and 2) a following six-dose test of exposure to 50–175 Gy in 25 Gy intervals ( n  = 10). Similarly for the nonirradiated controls, cups of males were placed in the irradiation chamber but no x-rays were applied. Once treated the three males within an exposure-cup were transferred to a mating–oviposition jar containing three normal nonirradiated virgin females newly emerged from the colony (<4 h old). The mating–oviposition jars were small, 116-ml canning jelly-jars (6 cm opening by 5.5 cm tall; Kerr, Hearthmark LLC, Daleville, IN) with paper towel lids secured by metal lid-bands. Moths remained together to mate for 5–7 d to allow expression of their fecundity, then the oviposited-upon paper towels were placed in plastic petri dishes (6 cm diam.). Dead moths were removed from mating jars and placed in 4-ml vials with a 50% ethanol–water solution. To determine mating status or success, the bursa copulatrix of female moths were dissected and absence or number of spermatophores determined ( Husseiny and Madsen 1964 ). Petri dishes of eggs laid for the replicated treatments were observed for a period of 6–21 d posttreatment to determine fecundity (number of eggs laid) and fertility (number of eggs hatched). Numbers of unfertilized (yellow), fertilized (red), and total eggs laid, prehatch developed larvae (black-heads), and hatched F1 larvae were recorded. The second experiment was extended in length and scope beyond F1 larval hatching to observe the heritability of irradiation effects on the growth–development of the F1 larvae and emergence of F1 adults, followed by the recording of the F1 adults’ fecundity, fertility, and hatch of progeny of the F2 generation. For this experiment using the irradiation dose range of 50–175 Gy, neonate F1 larvae were removed from the egg-paper petri dishes shortly (< ½ d) after hatch and placed in plastic portion-cups with wheat bran diet. For the irradiation treatments, all neonates that hatched were placed in diet cups, with cups having between one and 20 larvae. For the nonirradiated controls, a sampling of newly hatched neonates from the 10 replicate petri dishes were placed in diet cups with between 7 and >14 individuals each. Larvae were followed through pupation and emergence of F1 adults. Upon emergence individual F1 adults were placed singularly in mating portion-cups containing a small 2 - by 2-cm piece of paper towel. Then newly emerged (<4 h) and unmated moths were removed from the colony and placed in the mating-cups as an opposite sex pairing with a treated F1 adult. As described similarly above, after a period of 5 d, adults were removed, placed in aqueous 50% ethanol, and then the females’ mating status was determined through dissection of their bursa copulatrix. The F2 eggs deposited in the mating cups were counted and observed for hatching over a period of 21 d. Statistical Analyses Data trends were interpreted by performing regression analyses using linear, quadratic, and cubic formula to determine best curve fit (R 2 , F value, and P value) and proportion of variance of mating and egg development that is predictable from irradiation dose levels. Because a substantial low hatch rate and mortality was observed in the control group, additionally before analysis, the data were supplemented and corrected using Abbott’s formula ( Abbott 1925 ). In concurrent analyses, a square-root transformation was used to normalize count data while percentage data were arcsine transformed prior to analysis of variance (SigmaStat, 2010). One-way ANOVA was used to determine significant differences among mean values for dose-treatment groups and controls (SigmaStat, 2010). A P -value of 0.05 was used to establish significance in all tests using the Tukey’s method to detect significant pair-wise comparisons. Data are presented in the tables as recorded and not transformed or corrected. Results Significant and predictive effects on mating, fecundity, fertility, and inherited sterility were observed for the dose exposure levels (≥50 Gy) of x-ray irradiation applied to parental male moths. Irradiation dose-dependent effects were determined for degree and frequency of mating, fecundity, and fertility, but varied greatly in their degree of predictability of the derived regression equations from poor to high, with R 2 values ranging from <0.1 to ≤0.9. Dose thresholds were determined that induced complete sterility in parental and F1 generations without detrimental effects on degree of mating and competitiveness with untreated moths. 100–300 Gy Dose Experiment The total percentage of females mated showed a high degree of irradiation dose-dependent effects (R 2  = 0.82; Table 2 ). Evidence of mating was observed for all females regardless for those crossed with the nonirradiated males or with males treated with doses of 100–150 Gy ( Table 2 ). Above 200 Gy doses treatments the percentage of mated females progressively decreased for both total matings and females that mated twice, while the occurrence of unmated females and those that mated once progressively increased. Table 2. Mean percentage (±SE) mating by nonirradiated female A. transitella crossed with nonirradiated or irradiated males at five x-ray dose levels Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Total . 0 0c a 46.7 (20.0)bc 46.7 (17.0)ab 100a 100 0c 40.0 (12.5)bc 60.0 (12.5)a 100a 150 0c 80.0 (13.3)a 20.0 (13.3)bc 100a 200 20.0 (13.3)bc 73.3 (12.5)ab 6.7 (6.7)c 80.0 (13.3)ab 250 33.3 (10.5)b 46.7 (13.3)bc 20.0 (8.2)bc 66.7 (10.5)b 300 93.3 (6.7)a 6.7 (6.7)c 0c 6.7 (6.7)c Regression y = −20.95 + (0.272 × Dose) R 2  = 0.543, F  = 33.27, P  < 0.001 y = 45.23 − (0.388 × Dose) + (0.0062 × Dose 2 ) − (0.000018 × Dose 3 ) R 2  = 0.385, F  = 5.42, P  = 0.028 y = 48.43 + (0.351 × Dose) − (0.0045 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.364, F  = 4.95, P  = 0.035 y = 100.30 − (0.151 × Dose) + (0.0023 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.819, F  = 39.24, P  < 0.001 ANOVA F5,24  = 23.89, P  < 0.001 F5,24  = 3.71, P  = 0.012 F5,24  = 4.42, P  = 0.005 F5,24  = 23.89, P  < 0.001 Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Total . 0 0c a 46.7 (20.0)bc 46.7 (17.0)ab 100a 100 0c 40.0 (12.5)bc 60.0 (12.5)a 100a 150 0c 80.0 (13.3)a 20.0 (13.3)bc 100a 200 20.0 (13.3)bc 73.3 (12.5)ab 6.7 (6.7)c 80.0 (13.3)ab 250 33.3 (10.5)b 46.7 (13.3)bc 20.0 (8.2)bc 66.7 (10.5)b 300 93.3 (6.7)a 6.7 (6.7)c 0c 6.7 (6.7)c Regression y = −20.95 + (0.272 × Dose) R 2  = 0.543, F  = 33.27, P  < 0.001 y = 45.23 − (0.388 × Dose) + (0.0062 × Dose 2 ) − (0.000018 × Dose 3 ) R 2  = 0.385, F  = 5.42, P  = 0.028 y = 48.43 + (0.351 × Dose) − (0.0045 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.364, F  = 4.95, P  = 0.035 y = 100.30 − (0.151 × Dose) + (0.0023 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.819, F  = 39.24, P  < 0.001 ANOVA F5,24  = 23.89, P  < 0.001 F5,24  = 3.71, P  = 0.012 F5,24  = 4.42, P  = 0.005 F5,24  = 23.89, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Table 2. Mean percentage (±SE) mating by nonirradiated female A. transitella crossed with nonirradiated or irradiated males at five x-ray dose levels Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Total . 0 0c a 46.7 (20.0)bc 46.7 (17.0)ab 100a 100 0c 40.0 (12.5)bc 60.0 (12.5)a 100a 150 0c 80.0 (13.3)a 20.0 (13.3)bc 100a 200 20.0 (13.3)bc 73.3 (12.5)ab 6.7 (6.7)c 80.0 (13.3)ab 250 33.3 (10.5)b 46.7 (13.3)bc 20.0 (8.2)bc 66.7 (10.5)b 300 93.3 (6.7)a 6.7 (6.7)c 0c 6.7 (6.7)c Regression y = −20.95 + (0.272 × Dose) R 2  = 0.543, F  = 33.27, P  < 0.001 y = 45.23 − (0.388 × Dose) + (0.0062 × Dose 2 ) − (0.000018 × Dose 3 ) R 2  = 0.385, F  = 5.42, P  = 0.028 y = 48.43 + (0.351 × Dose) − (0.0045 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.364, F  = 4.95, P  = 0.035 y = 100.30 − (0.151 × Dose) + (0.0023 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.819, F  = 39.24, P  < 0.001 ANOVA F5,24  = 23.89, P  < 0.001 F5,24  = 3.71, P  = 0.012 F5,24  = 4.42, P  = 0.005 F5,24  = 23.89, P  < 0.001 Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Total . 0 0c a 46.7 (20.0)bc 46.7 (17.0)ab 100a 100 0c 40.0 (12.5)bc 60.0 (12.5)a 100a 150 0c 80.0 (13.3)a 20.0 (13.3)bc 100a 200 20.0 (13.3)bc 73.3 (12.5)ab 6.7 (6.7)c 80.0 (13.3)ab 250 33.3 (10.5)b 46.7 (13.3)bc 20.0 (8.2)bc 66.7 (10.5)b 300 93.3 (6.7)a 6.7 (6.7)c 0c 6.7 (6.7)c Regression y = −20.95 + (0.272 × Dose) R 2  = 0.543, F  = 33.27, P  < 0.001 y = 45.23 − (0.388 × Dose) + (0.0062 × Dose 2 ) − (0.000018 × Dose 3 ) R 2  = 0.385, F  = 5.42, P  = 0.028 y = 48.43 + (0.351 × Dose) − (0.0045 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.364, F  = 4.95, P  = 0.035 y = 100.30 − (0.151 × Dose) + (0.0023 × Dose 2 ) + (0.0000095 × Dose 3 ) R 2  = 0.819, F  = 39.24, P  < 0.001 ANOVA F5,24  = 23.89, P  < 0.001 F5,24  = 3.71, P  = 0.012 F5,24  = 4.42, P  = 0.005 F5,24  = 23.89, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Female fecundity in total eggs laid was not significantly affected in the first experimental series using 100–300 Gy doses (R 2  = 0.19; Table 3 ). Male exposure to dose levels of ≥100 Gy resulted in significantly (R 2  = 0.77) fewer eggs showing visible embryonic development (VED; e.g., developing to and beyond the “red” color stage) compared to controls. In contrast, numbers of eggs laid did not differ significantly for those eggs showing no visible embryonic development (NVED; R 2  = 0.02). Significant increases in the percentage of NVED eggs laid and complementary decreases in percentage of VED eggs laid occurred with all irradiation doses ≥100 Gy compared to the nonirradiated controls (R 2  = 0.91). Fertility of females in both numbers of larvae hatched and percentage of total eggs hatched showed a high degree of irradiation dose dependency (R 2  = 0.76 and 0.89, respectively). The frequency of hatching was significantly reduced at the 100 Gy irradiation of males compared to controls, with only nine neonates hatching from 15 females mated with irradiated males compared with 918 neonates produced from the crossing of females with nonirradiated males ( Table 3 ). Moreover, no neonates hatched with the ≥150 Gy doses and no eggs developed to the prehatch stage with ≥200 Gy doses. Table 3. Mean (±SE) reproduction of groups of three female A. transitella crossed with nonirradiated or x-ray irradiated males at five dose levels, [overall total], n  = 5 Male Irradiation Dose (Gy) . Number of eggs . % eggs . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . 0 80.6 (14.3) 198.6 (45.2)a a 279.2 (34.3) ND 183.6 (47.1) a 32.4 (8.1)b 67.6 (8.1)a ND 61.9 (9.5)a [993] [1396] [918] 100 217.2 (47.7) 20.4 (10.5)b 237.6 (54.7) 4.8 (2.9) 1.8 (1.6) b 93.6 (3.2)a 6.4 (3.2)b 1.5 (0.9) 2.1 (1.2)b [102] [1188] [24] [9] 150 174.8 (39.0) 14.0 (11.8)b 188.8 (49.7) 0.4 (0.4) 0b 95.5 (3.0)a 4.6 (3.0)b 0.1 (0.1) 0b [70] [944] [2] 200 205.3 (37.0) 0 b 205.3 (37.0) 0 0b 100a 0 b 0 0b [1026] 250 155.6 (37.6) 0.2 (0.2)b 155.8 (37.7) 0 0b 99.9 (0.1)a 0.1 (0.1)b 0 0b [1] [779] 300 146.2 (57.1) 0.4 (0.2)b 146.6 (57.2) 0 0b 99.7 (0.2)a 0.3 (0.2)b 0 0b [2] [733] Regression y = 139.06 + (0.145 ×Dose) R 2  = 0.0236, F  = 0.677, P  = 0.418 y = 197.97 − (2.829 × Dose) + (0.0133 × Dose 2 ) − 0.0000202 × Dose 3 ) R 2  = 0.767, F  = 28.50, P  < 0.001 y = 276.91 − (0.448 × Dose) R 2  = 0.185, F  = 6.365, P  = 0.018 y = 31.040 − (0.422 × Dose) + (0.00186 × Dose 2 ) − (0.0000027 × Dose 3 ) R 2  = 0.346, F  = 3.704, P  = 0.068 y = 182.72 − (3.088 × Dose) + (0.0160 × Dose 2 ) + (0.0000259 × Dose 3 ) R 2  = 0.755, F  = 5.42, P  < 0.001 y = 32.76 + (0.974 × Dose) − (0.00461 × Dose b ) + (0.0000071 × Dose 3 ) R 2  = 0.908, F  = 85.52, P  < 0.001 y = 67.23 − (0.973 × Dose) + (0.0046 × Dose 2 ) − (0.0000070 × Dose 3 ) R 2  = 0.908, F  = 85.51, P  < 0.001 y = 9.725 − (0.133 × Dose) + (0.00059 × Dose 2 ) − (0.00000085 × Dose 3 ) R 2  = 0.359, F  = 3.92, P  = 0.061 y = 61.683 − (1.010 × Dose) + (0.00513 × Dose 2 ) + (0.0000082 × Dose 3 ) R 2  = 0.893, F  = 72.129, P  < 0.001 ANOVA F5, 24  = 1.43, P  = 0.249 F5, 24  = 16.18, P  < 0.001 F5, 24  = 1.19, P  = 0.342 F4, 20  = 2.68, P  = 0.061 F5, 24  = 15.10, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F4, 20  = 2.84, P  = 0.051 F5, 24  = 41.46, P  < 0.001 Male Irradiation Dose (Gy) . Number of eggs . % eggs . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . 0 80.6 (14.3) 198.6 (45.2)a a 279.2 (34.3) ND 183.6 (47.1) a 32.4 (8.1)b 67.6 (8.1)a ND 61.9 (9.5)a [993] [1396] [918] 100 217.2 (47.7) 20.4 (10.5)b 237.6 (54.7) 4.8 (2.9) 1.8 (1.6) b 93.6 (3.2)a 6.4 (3.2)b 1.5 (0.9) 2.1 (1.2)b [102] [1188] [24] [9] 150 174.8 (39.0) 14.0 (11.8)b 188.8 (49.7) 0.4 (0.4) 0b 95.5 (3.0)a 4.6 (3.0)b 0.1 (0.1) 0b [70] [944] [2] 200 205.3 (37.0) 0 b 205.3 (37.0) 0 0b 100a 0 b 0 0b [1026] 250 155.6 (37.6) 0.2 (0.2)b 155.8 (37.7) 0 0b 99.9 (0.1)a 0.1 (0.1)b 0 0b [1] [779] 300 146.2 (57.1) 0.4 (0.2)b 146.6 (57.2) 0 0b 99.7 (0.2)a 0.3 (0.2)b 0 0b [2] [733] Regression y = 139.06 + (0.145 ×Dose) R 2  = 0.0236, F  = 0.677, P  = 0.418 y = 197.97 − (2.829 × Dose) + (0.0133 × Dose 2 ) − 0.0000202 × Dose 3 ) R 2  = 0.767, F  = 28.50, P  < 0.001 y = 276.91 − (0.448 × Dose) R 2  = 0.185, F  = 6.365, P  = 0.018 y = 31.040 − (0.422 × Dose) + (0.00186 × Dose 2 ) − (0.0000027 × Dose 3 ) R 2  = 0.346, F  = 3.704, P  = 0.068 y = 182.72 − (3.088 × Dose) + (0.0160 × Dose 2 ) + (0.0000259 × Dose 3 ) R 2  = 0.755, F  = 5.42, P  < 0.001 y = 32.76 + (0.974 × Dose) − (0.00461 × Dose b ) + (0.0000071 × Dose 3 ) R 2  = 0.908, F  = 85.52, P  < 0.001 y = 67.23 − (0.973 × Dose) + (0.0046 × Dose 2 ) − (0.0000070 × Dose 3 ) R 2  = 0.908, F  = 85.51, P  < 0.001 y = 9.725 − (0.133 × Dose) + (0.00059 × Dose 2 ) − (0.00000085 × Dose 3 ) R 2  = 0.359, F  = 3.92, P  = 0.061 y = 61.683 − (1.010 × Dose) + (0.00513 × Dose 2 ) + (0.0000082 × Dose 3 ) R 2  = 0.893, F  = 72.129, P  < 0.001 ANOVA F5, 24  = 1.43, P  = 0.249 F5, 24  = 16.18, P  < 0.001 F5, 24  = 1.19, P  = 0.342 F4, 20  = 2.68, P  = 0.061 F5, 24  = 15.10, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F4, 20  = 2.84, P  = 0.051 F5, 24  = 41.46, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Table 3. Mean (±SE) reproduction of groups of three female A. transitella crossed with nonirradiated or x-ray irradiated males at five dose levels, [overall total], n  = 5 Male Irradiation Dose (Gy) . Number of eggs . % eggs . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . 0 80.6 (14.3) 198.6 (45.2)a a 279.2 (34.3) ND 183.6 (47.1) a 32.4 (8.1)b 67.6 (8.1)a ND 61.9 (9.5)a [993] [1396] [918] 100 217.2 (47.7) 20.4 (10.5)b 237.6 (54.7) 4.8 (2.9) 1.8 (1.6) b 93.6 (3.2)a 6.4 (3.2)b 1.5 (0.9) 2.1 (1.2)b [102] [1188] [24] [9] 150 174.8 (39.0) 14.0 (11.8)b 188.8 (49.7) 0.4 (0.4) 0b 95.5 (3.0)a 4.6 (3.0)b 0.1 (0.1) 0b [70] [944] [2] 200 205.3 (37.0) 0 b 205.3 (37.0) 0 0b 100a 0 b 0 0b [1026] 250 155.6 (37.6) 0.2 (0.2)b 155.8 (37.7) 0 0b 99.9 (0.1)a 0.1 (0.1)b 0 0b [1] [779] 300 146.2 (57.1) 0.4 (0.2)b 146.6 (57.2) 0 0b 99.7 (0.2)a 0.3 (0.2)b 0 0b [2] [733] Regression y = 139.06 + (0.145 ×Dose) R 2  = 0.0236, F  = 0.677, P  = 0.418 y = 197.97 − (2.829 × Dose) + (0.0133 × Dose 2 ) − 0.0000202 × Dose 3 ) R 2  = 0.767, F  = 28.50, P  < 0.001 y = 276.91 − (0.448 × Dose) R 2  = 0.185, F  = 6.365, P  = 0.018 y = 31.040 − (0.422 × Dose) + (0.00186 × Dose 2 ) − (0.0000027 × Dose 3 ) R 2  = 0.346, F  = 3.704, P  = 0.068 y = 182.72 − (3.088 × Dose) + (0.0160 × Dose 2 ) + (0.0000259 × Dose 3 ) R 2  = 0.755, F  = 5.42, P  < 0.001 y = 32.76 + (0.974 × Dose) − (0.00461 × Dose b ) + (0.0000071 × Dose 3 ) R 2  = 0.908, F  = 85.52, P  < 0.001 y = 67.23 − (0.973 × Dose) + (0.0046 × Dose 2 ) − (0.0000070 × Dose 3 ) R 2  = 0.908, F  = 85.51, P  < 0.001 y = 9.725 − (0.133 × Dose) + (0.00059 × Dose 2 ) − (0.00000085 × Dose 3 ) R 2  = 0.359, F  = 3.92, P  = 0.061 y = 61.683 − (1.010 × Dose) + (0.00513 × Dose 2 ) + (0.0000082 × Dose 3 ) R 2  = 0.893, F  = 72.129, P  < 0.001 ANOVA F5, 24  = 1.43, P  = 0.249 F5, 24  = 16.18, P  < 0.001 F5, 24  = 1.19, P  = 0.342 F4, 20  = 2.68, P  = 0.061 F5, 24  = 15.10, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F4, 20  = 2.84, P  = 0.051 F5, 24  = 41.46, P  < 0.001 Male Irradiation Dose (Gy) . Number of eggs . % eggs . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . 0 80.6 (14.3) 198.6 (45.2)a a 279.2 (34.3) ND 183.6 (47.1) a 32.4 (8.1)b 67.6 (8.1)a ND 61.9 (9.5)a [993] [1396] [918] 100 217.2 (47.7) 20.4 (10.5)b 237.6 (54.7) 4.8 (2.9) 1.8 (1.6) b 93.6 (3.2)a 6.4 (3.2)b 1.5 (0.9) 2.1 (1.2)b [102] [1188] [24] [9] 150 174.8 (39.0) 14.0 (11.8)b 188.8 (49.7) 0.4 (0.4) 0b 95.5 (3.0)a 4.6 (3.0)b 0.1 (0.1) 0b [70] [944] [2] 200 205.3 (37.0) 0 b 205.3 (37.0) 0 0b 100a 0 b 0 0b [1026] 250 155.6 (37.6) 0.2 (0.2)b 155.8 (37.7) 0 0b 99.9 (0.1)a 0.1 (0.1)b 0 0b [1] [779] 300 146.2 (57.1) 0.4 (0.2)b 146.6 (57.2) 0 0b 99.7 (0.2)a 0.3 (0.2)b 0 0b [2] [733] Regression y = 139.06 + (0.145 ×Dose) R 2  = 0.0236, F  = 0.677, P  = 0.418 y = 197.97 − (2.829 × Dose) + (0.0133 × Dose 2 ) − 0.0000202 × Dose 3 ) R 2  = 0.767, F  = 28.50, P  < 0.001 y = 276.91 − (0.448 × Dose) R 2  = 0.185, F  = 6.365, P  = 0.018 y = 31.040 − (0.422 × Dose) + (0.00186 × Dose 2 ) − (0.0000027 × Dose 3 ) R 2  = 0.346, F  = 3.704, P  = 0.068 y = 182.72 − (3.088 × Dose) + (0.0160 × Dose 2 ) + (0.0000259 × Dose 3 ) R 2  = 0.755, F  = 5.42, P  < 0.001 y = 32.76 + (0.974 × Dose) − (0.00461 × Dose b ) + (0.0000071 × Dose 3 ) R 2  = 0.908, F  = 85.52, P  < 0.001 y = 67.23 − (0.973 × Dose) + (0.0046 × Dose 2 ) − (0.0000070 × Dose 3 ) R 2  = 0.908, F  = 85.51, P  < 0.001 y = 9.725 − (0.133 × Dose) + (0.00059 × Dose 2 ) − (0.00000085 × Dose 3 ) R 2  = 0.359, F  = 3.92, P  = 0.061 y = 61.683 − (1.010 × Dose) + (0.00513 × Dose 2 ) + (0.0000082 × Dose 3 ) R 2  = 0.893, F  = 72.129, P  < 0.001 ANOVA F5, 24  = 1.43, P  = 0.249 F5, 24  = 16.18, P  < 0.001 F5, 24  = 1.19, P  = 0.342 F4, 20  = 2.68, P  = 0.061 F5, 24  = 15.10, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F5, 24  = 50.67, P  < 0.001 F4, 20  = 2.84, P  = 0.051 F5, 24  = 41.46, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab 50–175 Gy Dose Experiment The goal of the second experiment, a series of exposures from 50–175 Gy, was to better resolve dose–sterility activity at irradiation doses below and above the 100 Gy activity level of the first test. The observed mating status data for the nonirradiated and the 50–175 Gy irradiation treatments of males was not significantly predicted by either quadratic or cubic regression equations (R 2 ranging from 0.05 to 0.33). All females become mated for the nonirradiated controls and similarly 97–100% of the females mated when crossed with males treated at doses of 50–125 Gy ( Table 4 ). However, at male treatment doses of ≥150 Gy the percentage of females found unmated increased ( Table 4 ). No differences were found in percentage of matings that were single or double in frequency between females paired with nonirradiated control males and females paired with males exposed to any of the irradiated doses. No significant differences in frequency of multiple mating were found for females crossed with nonirradiated males and those crossed with males irradiated between 50 and 125 Gy ( Table 4 ). At exposure doses of >150 Gy the frequency of multiple mating significantly decreased from the levels for females mated with nonirradiated males and those males irradiated at doses of 50–125 Gy. Table 4. Mean percentage ( ± SE) mating by nonirradiated female A. transitella crossed with nonirradiated or irradiated males at five x-ray dose levels Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Multiple . Total . 0 0 b a 33.3 (8.6) 50.0 (9.0) 66.7 (8.6)abc 100a 50 0 b 20.0 (5.4) 40.0 (8.3) 80.0 (5.4)ab 100a 75 3.3 (3.3)b 16.7 (9.0) 50.0 (10.2) 80.0 (8.9)ab 96.7 (3.3)a 100 3.3 (3.3)b 26.7 (6.7) 33.3 (9.9) 70.0 (7.8)abc 96.7 (3.3)a 125 0 b 13.3 (5.4) 23.3 (7.1) 86.7 (5.4)a 100a 150 33.3 (7.0)a 30.0 (7.8) 30.0 (7.8) 36.7 (7.8)c 66.7 (7.0)b 175 23.3 (7.1)a 30.0 (7.8) 40.0 (8.3) 6.7 (6.7) 0 b Regression y = 0.487 − (0.110 × Dose) + (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 y = 32.658 − (0.311 × Dose) + (0.00174 × Dose 2 ) R 2  = 0.052, F  = 1.85, P  = 0.178 y = 48.91 + (0.277 × Dose) – (0.00767 × Dose 2 ) + (0.000033 × Dose 3 ) R 2  = 0.078, F  = 1.86, P  = 0.177 y = 66.855 + (0.421 × Dose) − (0.00328 × Dose 2 ) R 2  = 0.211, F  = 8.94, P  = 0.004 y = 99.513 + (0.110 × Dose) − (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 ANOVA F6,63  = 10.55, P  < 0.001 F6,63  = 1.08, P  = 0.387 F6,63  = 1.31, P  = 0.268 F6,63  = 5.25, P  < 0.001 F6,63  = 10.55, P  < 0.001 Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Multiple . Total . 0 0 b a 33.3 (8.6) 50.0 (9.0) 66.7 (8.6)abc 100a 50 0 b 20.0 (5.4) 40.0 (8.3) 80.0 (5.4)ab 100a 75 3.3 (3.3)b 16.7 (9.0) 50.0 (10.2) 80.0 (8.9)ab 96.7 (3.3)a 100 3.3 (3.3)b 26.7 (6.7) 33.3 (9.9) 70.0 (7.8)abc 96.7 (3.3)a 125 0 b 13.3 (5.4) 23.3 (7.1) 86.7 (5.4)a 100a 150 33.3 (7.0)a 30.0 (7.8) 30.0 (7.8) 36.7 (7.8)c 66.7 (7.0)b 175 23.3 (7.1)a 30.0 (7.8) 40.0 (8.3) 6.7 (6.7) 0 b Regression y = 0.487 − (0.110 × Dose) + (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 y = 32.658 − (0.311 × Dose) + (0.00174 × Dose 2 ) R 2  = 0.052, F  = 1.85, P  = 0.178 y = 48.91 + (0.277 × Dose) – (0.00767 × Dose 2 ) + (0.000033 × Dose 3 ) R 2  = 0.078, F  = 1.86, P  = 0.177 y = 66.855 + (0.421 × Dose) − (0.00328 × Dose 2 ) R 2  = 0.211, F  = 8.94, P  = 0.004 y = 99.513 + (0.110 × Dose) − (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 ANOVA F6,63  = 10.55, P  < 0.001 F6,63  = 1.08, P  = 0.387 F6,63  = 1.31, P  = 0.268 F6,63  = 5.25, P  < 0.001 F6,63  = 10.55, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Table 4. Mean percentage ( ± SE) mating by nonirradiated female A. transitella crossed with nonirradiated or irradiated males at five x-ray dose levels Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Multiple . Total . 0 0 b a 33.3 (8.6) 50.0 (9.0) 66.7 (8.6)abc 100a 50 0 b 20.0 (5.4) 40.0 (8.3) 80.0 (5.4)ab 100a 75 3.3 (3.3)b 16.7 (9.0) 50.0 (10.2) 80.0 (8.9)ab 96.7 (3.3)a 100 3.3 (3.3)b 26.7 (6.7) 33.3 (9.9) 70.0 (7.8)abc 96.7 (3.3)a 125 0 b 13.3 (5.4) 23.3 (7.1) 86.7 (5.4)a 100a 150 33.3 (7.0)a 30.0 (7.8) 30.0 (7.8) 36.7 (7.8)c 66.7 (7.0)b 175 23.3 (7.1)a 30.0 (7.8) 40.0 (8.3) 6.7 (6.7) 0 b Regression y = 0.487 − (0.110 × Dose) + (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 y = 32.658 − (0.311 × Dose) + (0.00174 × Dose 2 ) R 2  = 0.052, F  = 1.85, P  = 0.178 y = 48.91 + (0.277 × Dose) – (0.00767 × Dose 2 ) + (0.000033 × Dose 3 ) R 2  = 0.078, F  = 1.86, P  = 0.177 y = 66.855 + (0.421 × Dose) − (0.00328 × Dose 2 ) R 2  = 0.211, F  = 8.94, P  = 0.004 y = 99.513 + (0.110 × Dose) − (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 ANOVA F6,63  = 10.55, P  < 0.001 F6,63  = 1.08, P  = 0.387 F6,63  = 1.31, P  = 0.268 F6,63  = 5.25, P  < 0.001 F6,63  = 10.55, P  < 0.001 Male irradiation dose (Gy) . Number of matings post irradiation . 0 . 1 . 2 . Multiple . Total . 0 0 b a 33.3 (8.6) 50.0 (9.0) 66.7 (8.6)abc 100a 50 0 b 20.0 (5.4) 40.0 (8.3) 80.0 (5.4)ab 100a 75 3.3 (3.3)b 16.7 (9.0) 50.0 (10.2) 80.0 (8.9)ab 96.7 (3.3)a 100 3.3 (3.3)b 26.7 (6.7) 33.3 (9.9) 70.0 (7.8)abc 96.7 (3.3)a 125 0 b 13.3 (5.4) 23.3 (7.1) 86.7 (5.4)a 100a 150 33.3 (7.0)a 30.0 (7.8) 30.0 (7.8) 36.7 (7.8)c 66.7 (7.0)b 175 23.3 (7.1)a 30.0 (7.8) 40.0 (8.3) 6.7 (6.7) 0 b Regression y = 0.487 − (0.110 × Dose) + (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 y = 32.658 − (0.311 × Dose) + (0.00174 × Dose 2 ) R 2  = 0.052, F  = 1.85, P  = 0.178 y = 48.91 + (0.277 × Dose) – (0.00767 × Dose 2 ) + (0.000033 × Dose 3 ) R 2  = 0.078, F  = 1.86, P  = 0.177 y = 66.855 + (0.421 × Dose) − (0.00328 × Dose 2 ) R 2  = 0.211, F  = 8.94, P  = 0.004 y = 99.513 + (0.110 × Dose) − (0.00154 × Dose 2 ) R 2  = 0.334, F  = 16.81, P  < 0.001 ANOVA F6,63  = 10.55, P  < 0.001 F6,63  = 1.08, P  = 0.387 F6,63  = 1.31, P  = 0.268 F6,63  = 5.25, P  < 0.001 F6,63  = 10.55, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Female fecundity, egg embryonic development, and fertility were significantly affected in a dose-dependent manner of male exposure (R 2 ranging from 0.50 to 0.82). Fecundity and number of total eggs laid decreased relative to controls for male treatments ≥125 Gy (R 2  = 0.10 and 0.39, respectively). As dose increased above 50 Gy the number of VED eggs laid significantly decreased compared to nonirradiated controls ( Table 5 ). Moreover, significantly lower numbers of VED eggs were laid at the 150–175 Gy exposure range than the 50–75 Gy level (R 2  = 0.51). Also numbers of embryos that successfully developed to the prehatch-larval stage significantly decreased at doses of ≥100 Gy (R 2  = 0.50; Table 5 ). At the ≥100 Gy exposure the percentage of eggs laid that were NVED increased over controls (R 2  = 0.59), while the percentage of VED eggs laid and percentage of eggs reaching the prehatch stage significantly decreased (R 2  = 0.59 and 0.58, respectively). Most importantly the fertility of females, in both numbers of larvae hatching and percentage hatching of eggs, was significantly reduced for all dose levels ≥50 Gy irradiation of males (R 2  = 0.72 and 0.82, respectively), with only 11 neonates hatching (0.3% of total eggs laid) from 30 females mated with irradiated males at the 100 Gy dose. Complete sterility was observed with no neonates hatched when irradiation doses ≥125 Gy were administered to parental males. Furthermore, no eggs developed to the prehatch stage at the 175 Gy dose. Also, a significant prolonging of the embryonic development period from laying to both prehatch egg stage and to larval hatch was observed for the lower doses from 50 to 150 Gy (R 2  = 0.75 and 0.56, respectively; Table 5 ). Table 5. Average reproduction of groups of three female A. transitella crossed with nonirradiated or x-ray irradiated males at six dose levels, means (±SE) [overall total], n  = 10 Male irradia-tion dose (Gy) . Number of eggs . % eggs . Interval (d) to . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . Prehatch . Hatch . 0 241.9 (30.9) 263.6 (43.9)a a 505.5 (50.5)a 218.7 (23.6) a 205.6 (22.4)a 48.9 (5.9)a 51.1 (5.9)a 43.3 (3.3)a 40.7 (3.1)a 9.3 (0.3)a 10.1 (0.3)a [2,419] [2,636] [5,055] [2,187] [2,056] 50 259.1 (33.1) 143.8 (35.9) bc 402.9 (59.9)ab 107.1 (29.4) ab 33.2 (9.8)b 68.8 (6.1)ab 31.2 (6.1)b 22.4 (6.0)ab 6.9 (1.4)b 10.9 (0.5)ab 12.1 (0.6)b [2,591] [1,438] [4,029] [1,071] [332] 75 242.7 (14.7) 157.4 (34.5)ab 400.1 (30.0)ab 113.7 (26.5) ab 9.1 (4.2)b 64.1 (5.9)ab 35.9 (5.9)ab 26.0 (4.6)a 2.0 (0.8)c 11.9 (0.4)bc 14.0 (0.5)bc [2,427] [1,574] [4,001] [1,137] [91] 100 291.3 (35.3) 98.4 (23.1)bcd 389.7 (49.1)abc 49.2 (13.4) bc 1.1 (0.6)b 77.0 (5.0)bc 23.0 (5.0)bc 11.5 (2.8)bc 0.3 (0.2)c 13.2 (0.4)c 15.3 (0.6)c [2,913] [984] [3,897] [492] [11] 125 270.0 (21.8) 31.1 (11.5)cd 301.1(28.3)bcd 4.6 (2.4) c 0b 91.4 (3.0)cd 8.6 (3.0)cd 1.3 (0.7)c 0c 16.3 (0.2)d – [2,700] [311] [3,011] [46] 150 205.1 (29.1) 11.7 (7.9)d 216.8 (34.3)cd 1.7 (1.1) c 0b 96.8 (2.2)d 3.2 (2.2)d 0.5 (0.3)c 0c 16.5 (0.2)d – [2,051] [117] [2,168] [17] 175 184.9 (29.3) 0.4 (0.2)d 185.3 (29.3)d 0 c 0b 99.8 (0.1)d 0.2 (0.1)d 0c 0c – – [1,849] [4] [1,853] Regression y = 234.07 + (1.075 × Dose) − (0.0077 × Dose 2 ) R 2  = 0.0955, F  = 3.537, P  = 0.064 y = 249.501 − (1.541 × Dose) R 2  = 0.514, F  = 71.85, P  < 0.001 y = 519.47 − (1.829 × Dose) R 2  = 0.387, F  = 42.87, P  < 0.001 y = 159.26 − (1.011 × Dose) R 2  = 0.502, F  = 68.51, P  < 0.001 y = 114.91 − (1.874 × Dose) + (0.00717 × Dose 2 ) R 2  = 0.715, F  = 84.03, P  < 0.001 y = 48.75 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 51.25 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 32.82 − (0.204 × Dose) R 2  = 0.581, F  = 94.39, P  < 0.001 y = 22.11 − (0.355 × Dose) + (0.00134 × Dose 2 ) R 2  = 0.820, F  = 152.79, P  < 0.001 y = 9.30 + (0.0191 × Dose) + (0.000224 × Dose 2 ) R 2  = 0.752, F  = 62.31, P  < 0.001 y = 10.06 + (0.0167 × Dose) + (0.000567 × Dose 2 ) R 2  = 0.559, F  = 23.41, P  < 0.001 ANOVA F6,63  = 1.67, P =  0.145 F6,63  = 12.36, P  < 0.001 F6,63  = 7.41, P  < 0.001 F6,63  = 12.86, P  < 0.001 F6,63  = 28.08, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 20.54, P  < 0.001 F6,63  = 51.99, P  < 0.001 F5,38  = 26.11, P  < 0.001 F3,32  = 17.92, P  < 0.001 Male irradia-tion dose (Gy) . Number of eggs . % eggs . Interval (d) to . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . Prehatch . Hatch . 0 241.9 (30.9) 263.6 (43.9)a a 505.5 (50.5)a 218.7 (23.6) a 205.6 (22.4)a 48.9 (5.9)a 51.1 (5.9)a 43.3 (3.3)a 40.7 (3.1)a 9.3 (0.3)a 10.1 (0.3)a [2,419] [2,636] [5,055] [2,187] [2,056] 50 259.1 (33.1) 143.8 (35.9) bc 402.9 (59.9)ab 107.1 (29.4) ab 33.2 (9.8)b 68.8 (6.1)ab 31.2 (6.1)b 22.4 (6.0)ab 6.9 (1.4)b 10.9 (0.5)ab 12.1 (0.6)b [2,591] [1,438] [4,029] [1,071] [332] 75 242.7 (14.7) 157.4 (34.5)ab 400.1 (30.0)ab 113.7 (26.5) ab 9.1 (4.2)b 64.1 (5.9)ab 35.9 (5.9)ab 26.0 (4.6)a 2.0 (0.8)c 11.9 (0.4)bc 14.0 (0.5)bc [2,427] [1,574] [4,001] [1,137] [91] 100 291.3 (35.3) 98.4 (23.1)bcd 389.7 (49.1)abc 49.2 (13.4) bc 1.1 (0.6)b 77.0 (5.0)bc 23.0 (5.0)bc 11.5 (2.8)bc 0.3 (0.2)c 13.2 (0.4)c 15.3 (0.6)c [2,913] [984] [3,897] [492] [11] 125 270.0 (21.8) 31.1 (11.5)cd 301.1(28.3)bcd 4.6 (2.4) c 0b 91.4 (3.0)cd 8.6 (3.0)cd 1.3 (0.7)c 0c 16.3 (0.2)d – [2,700] [311] [3,011] [46] 150 205.1 (29.1) 11.7 (7.9)d 216.8 (34.3)cd 1.7 (1.1) c 0b 96.8 (2.2)d 3.2 (2.2)d 0.5 (0.3)c 0c 16.5 (0.2)d – [2,051] [117] [2,168] [17] 175 184.9 (29.3) 0.4 (0.2)d 185.3 (29.3)d 0 c 0b 99.8 (0.1)d 0.2 (0.1)d 0c 0c – – [1,849] [4] [1,853] Regression y = 234.07 + (1.075 × Dose) − (0.0077 × Dose 2 ) R 2  = 0.0955, F  = 3.537, P  = 0.064 y = 249.501 − (1.541 × Dose) R 2  = 0.514, F  = 71.85, P  < 0.001 y = 519.47 − (1.829 × Dose) R 2  = 0.387, F  = 42.87, P  < 0.001 y = 159.26 − (1.011 × Dose) R 2  = 0.502, F  = 68.51, P  < 0.001 y = 114.91 − (1.874 × Dose) + (0.00717 × Dose 2 ) R 2  = 0.715, F  = 84.03, P  < 0.001 y = 48.75 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 51.25 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 32.82 − (0.204 × Dose) R 2  = 0.581, F  = 94.39, P  < 0.001 y = 22.11 − (0.355 × Dose) + (0.00134 × Dose 2 ) R 2  = 0.820, F  = 152.79, P  < 0.001 y = 9.30 + (0.0191 × Dose) + (0.000224 × Dose 2 ) R 2  = 0.752, F  = 62.31, P  < 0.001 y = 10.06 + (0.0167 × Dose) + (0.000567 × Dose 2 ) R 2  = 0.559, F  = 23.41, P  < 0.001 ANOVA F6,63  = 1.67, P =  0.145 F6,63  = 12.36, P  < 0.001 F6,63  = 7.41, P  < 0.001 F6,63  = 12.86, P  < 0.001 F6,63  = 28.08, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 20.54, P  < 0.001 F6,63  = 51.99, P  < 0.001 F5,38  = 26.11, P  < 0.001 F3,32  = 17.92, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Table 5. Average reproduction of groups of three female A. transitella crossed with nonirradiated or x-ray irradiated males at six dose levels, means (±SE) [overall total], n  = 10 Male irradia-tion dose (Gy) . Number of eggs . % eggs . Interval (d) to . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . Prehatch . Hatch . 0 241.9 (30.9) 263.6 (43.9)a a 505.5 (50.5)a 218.7 (23.6) a 205.6 (22.4)a 48.9 (5.9)a 51.1 (5.9)a 43.3 (3.3)a 40.7 (3.1)a 9.3 (0.3)a 10.1 (0.3)a [2,419] [2,636] [5,055] [2,187] [2,056] 50 259.1 (33.1) 143.8 (35.9) bc 402.9 (59.9)ab 107.1 (29.4) ab 33.2 (9.8)b 68.8 (6.1)ab 31.2 (6.1)b 22.4 (6.0)ab 6.9 (1.4)b 10.9 (0.5)ab 12.1 (0.6)b [2,591] [1,438] [4,029] [1,071] [332] 75 242.7 (14.7) 157.4 (34.5)ab 400.1 (30.0)ab 113.7 (26.5) ab 9.1 (4.2)b 64.1 (5.9)ab 35.9 (5.9)ab 26.0 (4.6)a 2.0 (0.8)c 11.9 (0.4)bc 14.0 (0.5)bc [2,427] [1,574] [4,001] [1,137] [91] 100 291.3 (35.3) 98.4 (23.1)bcd 389.7 (49.1)abc 49.2 (13.4) bc 1.1 (0.6)b 77.0 (5.0)bc 23.0 (5.0)bc 11.5 (2.8)bc 0.3 (0.2)c 13.2 (0.4)c 15.3 (0.6)c [2,913] [984] [3,897] [492] [11] 125 270.0 (21.8) 31.1 (11.5)cd 301.1(28.3)bcd 4.6 (2.4) c 0b 91.4 (3.0)cd 8.6 (3.0)cd 1.3 (0.7)c 0c 16.3 (0.2)d – [2,700] [311] [3,011] [46] 150 205.1 (29.1) 11.7 (7.9)d 216.8 (34.3)cd 1.7 (1.1) c 0b 96.8 (2.2)d 3.2 (2.2)d 0.5 (0.3)c 0c 16.5 (0.2)d – [2,051] [117] [2,168] [17] 175 184.9 (29.3) 0.4 (0.2)d 185.3 (29.3)d 0 c 0b 99.8 (0.1)d 0.2 (0.1)d 0c 0c – – [1,849] [4] [1,853] Regression y = 234.07 + (1.075 × Dose) − (0.0077 × Dose 2 ) R 2  = 0.0955, F  = 3.537, P  = 0.064 y = 249.501 − (1.541 × Dose) R 2  = 0.514, F  = 71.85, P  < 0.001 y = 519.47 − (1.829 × Dose) R 2  = 0.387, F  = 42.87, P  < 0.001 y = 159.26 − (1.011 × Dose) R 2  = 0.502, F  = 68.51, P  < 0.001 y = 114.91 − (1.874 × Dose) + (0.00717 × Dose 2 ) R 2  = 0.715, F  = 84.03, P  < 0.001 y = 48.75 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 51.25 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 32.82 − (0.204 × Dose) R 2  = 0.581, F  = 94.39, P  < 0.001 y = 22.11 − (0.355 × Dose) + (0.00134 × Dose 2 ) R 2  = 0.820, F  = 152.79, P  < 0.001 y = 9.30 + (0.0191 × Dose) + (0.000224 × Dose 2 ) R 2  = 0.752, F  = 62.31, P  < 0.001 y = 10.06 + (0.0167 × Dose) + (0.000567 × Dose 2 ) R 2  = 0.559, F  = 23.41, P  < 0.001 ANOVA F6,63  = 1.67, P =  0.145 F6,63  = 12.36, P  < 0.001 F6,63  = 7.41, P  < 0.001 F6,63  = 12.86, P  < 0.001 F6,63  = 28.08, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 20.54, P  < 0.001 F6,63  = 51.99, P  < 0.001 F5,38  = 26.11, P  < 0.001 F3,32  = 17.92, P  < 0.001 Male irradia-tion dose (Gy) . Number of eggs . % eggs . Interval (d) to . No visible embryonic development . Visible embryonic development . Total . Prehatch . Hatched . No visible embryonic development . Visible embryonic development . Prehatch . Hatched . Prehatch . Hatch . 0 241.9 (30.9) 263.6 (43.9)a a 505.5 (50.5)a 218.7 (23.6) a 205.6 (22.4)a 48.9 (5.9)a 51.1 (5.9)a 43.3 (3.3)a 40.7 (3.1)a 9.3 (0.3)a 10.1 (0.3)a [2,419] [2,636] [5,055] [2,187] [2,056] 50 259.1 (33.1) 143.8 (35.9) bc 402.9 (59.9)ab 107.1 (29.4) ab 33.2 (9.8)b 68.8 (6.1)ab 31.2 (6.1)b 22.4 (6.0)ab 6.9 (1.4)b 10.9 (0.5)ab 12.1 (0.6)b [2,591] [1,438] [4,029] [1,071] [332] 75 242.7 (14.7) 157.4 (34.5)ab 400.1 (30.0)ab 113.7 (26.5) ab 9.1 (4.2)b 64.1 (5.9)ab 35.9 (5.9)ab 26.0 (4.6)a 2.0 (0.8)c 11.9 (0.4)bc 14.0 (0.5)bc [2,427] [1,574] [4,001] [1,137] [91] 100 291.3 (35.3) 98.4 (23.1)bcd 389.7 (49.1)abc 49.2 (13.4) bc 1.1 (0.6)b 77.0 (5.0)bc 23.0 (5.0)bc 11.5 (2.8)bc 0.3 (0.2)c 13.2 (0.4)c 15.3 (0.6)c [2,913] [984] [3,897] [492] [11] 125 270.0 (21.8) 31.1 (11.5)cd 301.1(28.3)bcd 4.6 (2.4) c 0b 91.4 (3.0)cd 8.6 (3.0)cd 1.3 (0.7)c 0c 16.3 (0.2)d – [2,700] [311] [3,011] [46] 150 205.1 (29.1) 11.7 (7.9)d 216.8 (34.3)cd 1.7 (1.1) c 0b 96.8 (2.2)d 3.2 (2.2)d 0.5 (0.3)c 0c 16.5 (0.2)d – [2,051] [117] [2,168] [17] 175 184.9 (29.3) 0.4 (0.2)d 185.3 (29.3)d 0 c 0b 99.8 (0.1)d 0.2 (0.1)d 0c 0c – – [1,849] [4] [1,853] Regression y = 234.07 + (1.075 × Dose) − (0.0077 × Dose 2 ) R 2  = 0.0955, F  = 3.537, P  = 0.064 y = 249.501 − (1.541 × Dose) R 2  = 0.514, F  = 71.85, P  < 0.001 y = 519.47 − (1.829 × Dose) R 2  = 0.387, F  = 42.87, P  < 0.001 y = 159.26 − (1.011 × Dose) R 2  = 0.502, F  = 68.51, P  < 0.001 y = 114.91 − (1.874 × Dose) + (0.00717 × Dose 2 ) R 2  = 0.715, F  = 84.03, P  < 0.001 y = 48.75 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 51.25 + (0.305 × Dose) R 2  = 0.588, F  = 97.13, P  < 0.001 y = 32.82 − (0.204 × Dose) R 2  = 0.581, F  = 94.39, P  < 0.001 y = 22.11 − (0.355 × Dose) + (0.00134 × Dose 2 ) R 2  = 0.820, F  = 152.79, P  < 0.001 y = 9.30 + (0.0191 × Dose) + (0.000224 × Dose 2 ) R 2  = 0.752, F  = 62.31, P  < 0.001 y = 10.06 + (0.0167 × Dose) + (0.000567 × Dose 2 ) R 2  = 0.559, F  = 23.41, P  < 0.001 ANOVA F6,63  = 1.67, P =  0.145 F6,63  = 12.36, P  < 0.001 F6,63  = 7.41, P  < 0.001 F6,63  = 12.86, P  < 0.001 F6,63  = 28.08, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 17.27, P  < 0.001 F6,63  = 20.54, P  < 0.001 F6,63  = 51.99, P  < 0.001 F5,38  = 26.11, P  < 0.001 F3,32  = 17.92, P  < 0.001 a Mean values in columns followed by the same letter are not significantly different, P  > 0.05, ANOVA followed by Tukey’s method to detect significant pair-wise comparisons. Open in new tab Inherited Sterility Experiment The experiment using exposures of 50–175 Gy was extended in scope to study inherited sterility by placing a portion or all of the eclosed F1 neonates into diet cups, allowing feeding and development, then observing the pupation and emergence of F1 adults. The emerged F1 adults were placed individually in cups, provided a nonirradiated virgin counterpart, and allowed to mate, followed by the observation of their resultant egg–fecundity and hatch–fertility. No F1 neonates hatched for the irradiated doses >125 Gy applied to P1 males ( Table 5 ). For the nonirradiated, 50 Gy, and 75 Gy treatments subsamples of 212, 126, and 73 eclosed F1 neonates respectively were placed onto diet, while all 11 F1 neonates for the 100 Gy treatment were placed onto diet for rearing ( Table 6 ). A low emergence rate of F1 moths occurred, ranging from 22 to five moths for the nonirradiated to the 100 Gy treatment, respectively. The sex ratio of F1 adults was 6.3: 1 female: male for the nonirradiated moths but overall reversed to 1.5–4: 1 male: female for the irradiated treatments of 50 and 100 Gy ( Table 6 ). Both the F1 development periods from neonate to adult and the F1 adult life span periods were similar for nonirradiated and irradiated treatments ( Table 6 ). The mating frequency of untreated–control females was 29% mated multiple times and 71% mated once. Mating decreased for the progeny of 50–75 Gy irradiation for either the F1 females or colony females crossed with F1 males, with a decrease in females mated multiple times and mated once. However, 42–50% females remained unmated. Most importantly, for the 100 Gy parental dose no mating was observed for the singular female that survived long enough to attempt to mate. Table 6. Mating and reproduction of F2 adults emerged from F1 hatched larvae produced by x-ray irradiated and nonirradiated parental male moths Dose (Gy) . No. F1 neonates into diet cups a . No. F1 emergent moths (male : female) . Interval (d) F 1 larval to adult, mean (+SE) . F 1 adult life span (d), mean (+SE) . F 1 female mating status, no. females (percentage) . Not mated . Single mated . Multiple mated . 0 212 3 m : 19 f 54.1 (2.5) b 9.9 (0.9) 0 10 (71%) 4 (29%) 50 126 9 m : 6 f 64.0 (1.8) 10.5 (1.2) 5 (50%) 5 (50%) 0 75 73 4 m : 6 f 63.6 (7.7) 8.2 (0.8) 3 (42%) 2 (29%) 2 (29%) 100 11 4 m : 1 f 59.0 (8.0) 9.0 (0) 2 (100%) 0 0 125 None – – – – – – 150 None – – – – – – 175 None – – – – – – Dose (Gy) . No. F1 neonates into diet cups a . No. F1 emergent moths (male : female) . Interval (d) F 1 larval to adult, mean (+SE) . F 1 adult life span (d), mean (+SE) . F 1 female mating status, no. females (percentage) . Not mated . Single mated . Multiple mated . 0 212 3 m : 19 f 54.1 (2.5) b 9.9 (0.9) 0 10 (71%) 4 (29%) 50 126 9 m : 6 f 64.0 (1.8) 10.5 (1.2) 5 (50%) 5 (50%) 0 75 73 4 m : 6 f 63.6 (7.7) 8.2 (0.8) 3 (42%) 2 (29%) 2 (29%) 100 11 4 m : 1 f 59.0 (8.0) 9.0 (0) 2 (100%) 0 0 125 None – – – – – – 150 None – – – – – – 175 None – – – – – – a Subsamples of total hatched larvae, totals were: 1,187 for 0 Gy, 332 for 50 Gy, 91 for 75 Gy, and 11 for 100 Gy. b No significant differences for development, life span, nor mating status data. Open in new tab Table 6. Mating and reproduction of F2 adults emerged from F1 hatched larvae produced by x-ray irradiated and nonirradiated parental male moths Dose (Gy) . No. F1 neonates into diet cups a . No. F1 emergent moths (male : female) . Interval (d) F 1 larval to adult, mean (+SE) . F 1 adult life span (d), mean (+SE) . F 1 female mating status, no. females (percentage) . Not mated . Single mated . Multiple mated . 0 212 3 m : 19 f 54.1 (2.5) b 9.9 (0.9) 0 10 (71%) 4 (29%) 50 126 9 m : 6 f 64.0 (1.8) 10.5 (1.2) 5 (50%) 5 (50%) 0 75 73 4 m : 6 f 63.6 (7.7) 8.2 (0.8) 3 (42%) 2 (29%) 2 (29%) 100 11 4 m : 1 f 59.0 (8.0) 9.0 (0) 2 (100%) 0 0 125 None – – – – – – 150 None – – – – – – 175 None – – – – – – Dose (Gy) . No. F1 neonates into diet cups a . No. F1 emergent moths (male : female) . Interval (d) F 1 larval to adult, mean (+SE) . F 1 adult life span (d), mean (+SE) . F 1 female mating status, no. females (percentage) . Not mated . Single mated . Multiple mated . 0 212 3 m : 19 f 54.1 (2.5) b 9.9 (0.9) 0 10 (71%) 4 (29%) 50 126 9 m : 6 f 64.0 (1.8) 10.5 (1.2) 5 (50%) 5 (50%) 0 75 73 4 m : 6 f 63.6 (7.7) 8.2 (0.8) 3 (42%) 2 (29%) 2 (29%) 100 11 4 m : 1 f 59.0 (8.0) 9.0 (0) 2 (100%) 0 0 125 None – – – – – – 150 None – – – – – – 175 None – – – – – – a Subsamples of total hatched larvae, totals were: 1,187 for 0 Gy, 332 for 50 Gy, 91 for 75 Gy, and 11 for 100 Gy. b No significant differences for development, life span, nor mating status data. Open in new tab Fecundity of F1 females and normal females mated to F1 males was similar for nonirradiated and P1 male irradiated treatments, with numerous >200 NVED eggs laid per female for all treatments. However, differences in F1 fertility were observed, with numerous VED F2 eggs laid and developing to eclosed neonates for the nonirradiated controls, while in contrast at only the 50 Gy treatment half the number of VED F2 eggs were produced, while none occurred at the 75 and 100 Gy doses. Moreover, no F2 eggs hatched at any irradiated dose ≥50 Gy administered to P1 males. Discussion The goal of SIT is to disrupt–prevent reproduction of females of the targeted invasive or established pest population through the release of mass-reared sterile insects with a focus on sterile males. Key to success of SIT is that the released sterile males are truly adequate, equivalent, and competitive with the fertile males of the targeted population ( Calkins and Parker 2005 , Lance and McInnis 2005 , Perez-Staples et al. 2012 ). Released sterilized males must be fully competitive in their mating abilities, including finding of females, courtship, copulation, and delivery of spermataphores and sperm ( Marec et al. 1999 , Wang et al. 2001 , Bloem et al. 2003 , Calkins and Parker 2005 , Lance and McInnis 2005 , Marti and Carpenter 2009 , Mudavanhu et al. 2011 , Perez-Staples et al. 2012 ). In addition, a crucial factor is the competitiveness of steriles in their frequency or degree of multiple mating with target wilds ( Calkins and Parker 2005 , Lance and McInnis 2005 , Perez-Staples et al. 2012 ). The sperm of released sterile males must be competitive in ability to fertilize eggs of normal wild females, while disrupting reproduction ( Marec et al. 1999 , Carpenter et al. 2005 ). The sterility in mating by irradiated males with wild females can be expressed at various stages in reproductive development, including nondevelopment of fertilized eggs, noneclosion of fertilized eggs, nondevelopment–death of larvae, nonpupation, nonemergence of adults, defective adults, and inherited sterility with the germ cells—gametes of emergent F1 adults being sterile ( Calkins and Parker 2005 , Carpenter et al. 2005 , Lance and McInnis 2005 , Perez-Staples et al. 2012 ). Successful SIT requires the determination of the optimal threshold for x-ray irradiation doses that effective induce sterility without diminishing ability, drive, and frequency of mating ( Bakri et al. 2005a , 2005b , Calkins and Parker 2005 , Carpenter et al. 2005 , Lance and McInnis 2005 , Perez-Staples et al. 2012 ). Sterility–dose thresholds can be derived from the studies reported here where normal females were crossed with irradiated parental males and reproduction was compared to nonirradiated control moth mating, fecundity, and fertility. Mating occurrence and frequency of mating in P1 adults were equivalent to nonirradiated controls for irradiation doses to males of 125 Gy and lower. At doses of 150 Gy significant changes in mating frequency were observed, with an increase in frequency of unmated females and a reduction in multiple-mating and overall mating. Fecundity, in numbers of total eggs laid, was similar for control and irradiated females up through a dose of 100 Gy, above which significant decreases in total eggs laid occurred. As reported here, colony females crossed with nonirradiated control males oviposit a large portion (32–49%, Tables 3 and 5 ) of eggs that showed no embryonic development. Husseiny and Madsen (1964) reported that nonviable eggs showing no visible embryonic development averaged ca. 49% (range of 34–68%) of eggs laid by A. transitella females in mating experiments. Similarly, when periodic infusions of wild A. transitella with colony stock are routinely used the percentage of eggs showing NVED is observed to be “highly variable,” ranging from 20 to >60% as experienced at various current laboratory rearing sites in California (personal communications: S. Tebbets, USDA-ARS-SJVASC, Parlier and B. Higbee, Wonderful Orchards, Shafter). This relative high proportion of nondeveloping eggs laid by A. transitella appears to represent a common occurrence in this species, the practice of laying–eliminating infertile eggs when reared in laboratory colonies. Contributing to this occurrence of nonviable eggs could be a negative fitness factor of undetermined cause, though potentially due to nonviable sperm, lack of sperm transfer during mating, or early embryonic death. However, the level of NVED eggs was fairly constant for all test females, whether crossed with nonirradiated control males or with males exposed to all levels of irradiation tested, from 50 to 300 Gy. In contrast, compared with controls these irradiation levels as expected caused dramatic dose-dependent decreases in numbers of viable VED eggs laid, prehatch larvae, and hatched larvae ( Tables 3 and 5 ). Fertility significantly decreased with X-ray doses of 50 Gy and greater for the percentage of both VED eggs laid and eggs that hatched. The percentage egg hatch of A. transitella larvae reported here for the control (0 Gy) crosses (62% and 41%, in Tables 3 and 5 ) is relatively low and variable between the tests, but again, this has been found to be typical and similar to the 67% hatch rate reported by Husseiny and Madsen (1964) and the variable range (30 to 80%) experienced in colony rearing of this species in California laboratories (personal communications: S. Tebbets and B. Higbee). Complete F1 sterility was reached at a dose of 125 Gy with no neonate hatching. Prior studies on A. transitella by Johnson and Vail (1988 , 1989 ) in phytosaniation studies of nut commodities reported that gamma irradiation of A. transitella and P. interpunctella larvae with doses of 149 to >600 Gy caused a lack of adult emergence. Based on our results, a dose of ≥125 Gy had efficacy in inducing both primary parental infertility and sterility in treated male moths and inherited infertility and sterility in F1 male and female moths. Thus, an x-ray dose of 125 Gy could be considered optimal for complete sterility of P1 treated male A. transitella moths. Similar efficacy in irradiating adult males has been observed in other pyralid pests, though the threshold and optimal irradiation doses were generally higher ( Table 1 ). Prior studies established doses of generally 130–400 Gy (mean of 260 Gy) applied to P1 male moths were required for complete sterility in 16 pyralid pests ( Bakri et al. 2005a , 2005b ). These prior tests over a 50-yr period utilized gamma irradiation (primarily Cobalt 60). The dose level of 125 Gy for complete sterility reported here for x-ray irradiation of P1 male A. transitella would suggest that males of this species are radiosensitive, while other pyralids are relatively more radiotolerant in their sterilizing dose from gamma exposures ( Table 1 ). One exemption appears to be the sweetpotato vine borer, O. anastomosalis , with a reported sterilizing dose of ≥150 Gy, interestingly also by means of x-ray irradiation ( Follett 2006 ). Inherited sterility in the F2 generation as expressed in the lethal effects upon F2 eggs was achieved here with exposure of P1 male A. transitella to substerilizing x-ray doses of 50–75 Gy, with no F2 egg hatch nor fertile F2 eggs laid, respectively. This result also supports the classification of A. transitella as a radiosensitive species. A substerilizing gamma dose to P1 adults has been reported for pyralids to generally be in the range of 150 to >350 Gy to evoke F1–F2 sterility in progeny, with a sterilizing dose of 200 Gy reported in 15 of the 37 studies ( Table 1 ). Mating competiveness is fundamental to SIT success; sterilized moths must mate as effectively as wild moths to attain the goal of introducing and spreading the radiation-induced lethal mutations through a target pest population ( Calkins and Parker 2005 ). We observed here that x-ray irradiation doses of 125 Gy had no adverse effect on the ability to mate nor on the frequency of mating compared with the mating behavior and efficiency of nonirradiated normal moth pairs. However, at the 150 Gy dose and above administered to parental males, we observed a decrease in mating occurrence and efficiency, resulting in an increase percentage of crossed females remaining unmated and a decreasing degree of multiple mating. The suggested complete sterility dose of 125 Gy for A. transitella males is lower than many other reports on effects of irradiation on mating by pyralid pests, where detrimental effects on mating ability and frequency occurred at relatively high dose levels of 350–>500 Gy ( Ashrafi et al. 1972 , Brower 1975 ; Table 1 ). As reported in many prior studies a number of key phenomena in mating, mating frequency, development rates, longevity, and sex ratios were affected by the irradiation of parental moths ( Bakri et al. 2005a , 2005b ; Carpenter et al. 2005 ). In this study, these increases in activity were observed in a dose-dependent manner with the exposure of parental males to the lower range of x-ray doses, from 50 to 125 Gy, while exposure to higher dose rates above 150 Gy were detrimental causing a reversal and rapid decline to zero of developmental and behavioral activities. An increase in mating frequency for crosses with sterile moths has been reported for other pyralids, e.g., E. cautella , P. interpunctella , D. saccharalis , and E. saccharina ( Gonen and Calderon 1971 ; Brower 1975 , 1976b ; Sanford 1976 ; Mudavanhu et al. 2011 ; reviewed in, Lance and McInnis 2005 ). Here we observed an increase, though not a significant increase, in overall frequency of multiple mating by A. transitella females when crossed with irradiated males exposed to doses from 50 to 125 Gy dose compared with nonirradiated controls ( Table 4 ). At exposures of 150 Gy and above the ability and propensity of A. transitella males to mate decreased as the dose of radiation is increased. Also, observed here with A. transitella was a significant increase in the embryonic development period for viable eggs to the prehatch stage and larval hatch observed for the lower doses from 50 to 150 Gy compared to development of normal progeny ( Tzanakakis and Barnes 1988 , Sanderson et al. 1989 ; Table 5 ). Similar increases in developmental periods with irradiation were reported for other pyralids e.g., D. saccharalis , D. nitidalis , C. cactorum , and E. kuehniella ( Sanford 1976 , Elsey and Brower 1984 , Carpenter et al. 2001 , Ayvaz and Tuncbilek 2006 ). In addition, the sex ratio of emergent F1 A. transitella adults shifted in our study from predominantly females (86%) for nonirradiated controls to predominantly males (80%) for irradiation doses up to 100 Gy ( Table 6 ). However, flight ability–quality of these F1 adult progeny of irradiation was not tested here and should be assessed in future tests. A shift in sex ratio of filial adults from female to male dominated evoked in a dose-dependent manner by gamma irradiation exposure has been well reported for insects and Lepidoptera, including Pyralidae, e.g., D. saccharalis and E. cautella ( Sanford 1976 , Gonen and Calderon 1971 , Brower 1980 , Al-Taweel et al. 1989 , 1990 ). In addition to its radiosensitivity, A. transitella has many life history, growth, and development attributes that would make it a good candidate for mass-rearing and the exploration of the feasibility and development of a SIT program with over-flooding releases of steriles. A. transitella is a multivoltine, nondiapausing, noncannibalistic, preharvest, and stored-product pest of tree nuts ( Wade 1961 ). In the wild A. transitella has been reported to mate predominantly a single time with multiple mating being at a low eight percent frequency ( Wade 1961 , Husseiny and Madsen 1964 , Burks et al. 2011 ). Though eggs are laid singularly they can reach high egg densities on egg traps in the field and on paper surfaces in colony rearing jars. The feeding activity of A. transitella on both natural in-shell nuts and rearing diet is gregarious, communal, and noncannibalistic, with compatibility of simultaneous feeding of multiple development stages and generations. Moreover, A. transitella larvae will continue feeding on a diet source over generations, efficiently accepting, consuming, and reprocessing the diet and residual feces. A basis of this consumption ability of A. transitella larvae is their high tolerance to microbes and mycotoxins ( Lee and Campbell 2000 , Niu et al. 2009 ). Rearing is relatively inexpensive and easy, with diet ingredients being few, natural, and whole (wheat bran, honey, brewers’ yeast, and salts), requiring simply mixing. Also, the rearing process, from egg hatch to adult emergence and oviposition, all occurs easily and continuously in the rearing jars. Thereby, the rearing of A. transitella can readily be scaled to mass-rearing and generate nearly continuous output of adults for SIT irradiation. Reported here is an initial study on irradiating adult male A. transitella to assess their possible candidacy for SIT. Additional studies on irradiation of the egg, larval, pupal, and female stages of A. transitella are needed to assess the potential of x-rays for phytosanitary and quarantine control purposes ( Johnson and Vail, 1988 , 1989 , Follett and Snook 2012 ). Though additional research is necessary, the results suggest that A. transitella might be considered a candidate for SIT using adults irradiated at these low–moderate exposure doses provided by secure and safe x-ray sources. Being able and confident to use a lower irradiation rate might allow for increased mating fitness and perhaps better SIT efficacy, especially when integrated with mating disruption suppression programs. 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TI - Effects of X-Ray Irradiation on Male Navel Orangeworm Moths (Lepidoptera: Pyralidae) on Mating, Fecundity, Fertility, and Inherited Sterility
JF - Journal of Economic Entomology
DO - 10.1093/jee/tov201
DA - 2015-10-01
UR - https://www.deepdyve.com/lp/oxford-university-press/effects-of-x-ray-irradiation-on-male-navel-orangeworm-moths-M7WyYt5i0G
SP - 2200
EP - 2212
VL - 108
IS - 5
DP - DeepDyve
ER -