Optimization of Co60 gamma radiation dose for applying sterile insect technique and inherited sterility on Tuta absoluta (Meyrick) in Iran | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Article Optimization of Co 60 gamma radiation dose for applying sterile insect technique and inherited sterility on Tuta absoluta (Meyrick) in Iran Shabnam Ashouri This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-6690207/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 13 Dec, 2025 Read the published version in Scientific Reports → Version 1 posted 16 You are reading this latest preprint version Abstract The objective of this research is to stablish standardized sterile and sub-sterile doses of Co 60 gamma radiation on Tuta absoluta (Meyrick) to facilitate the implementation of sterile insect technique for management of this invasive pest of tomato and solanaceous plants. Insects were reared on an artificial diet and pupae were irradiated at 50, 100, 150, 250 and 300 Gy. The fecundity and fertility of untreated females paired with irradiated males, and treated females mated with untreated or treated males, significantly diminished with increasing gamma doses. When pupae were exposed to irradiation at 250 and 300 Gy, completely sterile females and males were obtained, respectively. The adult’s emergence and longevity, were not significantly influenced by irradiation. To perform inherited sterility, male pupae were irradiated by 250 Gy. The F 1 sex ratio shifted in favor of males. The F 1 fecundity was diminished compared to the control, and the eggs produced were more sterile. Ultimately, the mating competitiveness of irradiated males by 250 Gy against normal males was assessed at different ratios. Based on the calculated competitiveness value, the release ratio was determined 1:1:1 (irradiated males: unirradiated males: unirradiated females), in which males subjected to irradiation are capable of competing with unirradiated males. Biological sciences/Zoology/Entomology Biological sciences/Biological techniques Biological parameters Co60 gamma ray competition index inherited sterility sterile insect technique tomato leaf miner Figures Figure 1 Figure 2 Figure 3 Introduction The continuous application of chemical pesticides for insect control intensifies the accumulation of pollutants in ecosystems and presents considerable challenges to human health and environmental integrity. The destruction of plant pollinators and natural enemies of pests represents additional harmful consequences associated with the application of chemical pesticides [1]. These environmental risks require the adoption of safe and effective integrated pest management strategies whitin sustainable and modern agriculture. The sterile insect technique (SIT) is regarded as a prominent biological and species-specific methods for pest control [2]. In this technique, sterile insects are released in large numbers in an infested area to reduce the fertility of the wild population of the same species. This method involves mass breeding of the target pest species, exposing them to gamma or X-rays to induce sexual sterility, and subsequently releasing them on a large scale into the target pest population. In essence, the released sterile males mate with wild females, thereby reducing their reproduction and fertility [2, 3]. The mutations induced through this technique exhibit distinct characteristics in lepidopterans. In lepidopterans, certain dominant lethal mutations can be expressed immediately prior to egg hatching, and most of the damaged chromosomes are transmitted to the F 1 generation. Lepidoptera possess several specific cytogenetic and cytological features which predominantly contribute to their elevated resistance to ionizing radiation. Consequently, very high radiation doses are required to achieve complete sterility in lepidopteran males. These features also significantly influence the mechanism underlying inherited sterility (IS). Therefore, inherited sterility is a type of SIT wherein semi-sterile males are released. A synergistic approach that combines SIT and IS may be employed to control lepidopteran pests [1, 2, 3, 4]. In recent years, the tomato leaf miner, Tuta absoluta (Meyrick) (Lepidoptera: Gelechiidae), has emerged as the most invasive and destructive pest affecting tomato crops and is considered a major threat to the cultivation of numerous solanaceous species, including eggplant, peppers and potatoes globally [5]. Tuta absoluta larvae causes damage to the entire plant and specially feed from the leaf’s mesophyll. Moreover, the feeding activity on the fruits affects the visual appeal of harvested products and leading to fruit deterioration. The larvae of this pest can inflict 80 to 100% damage to tomatoes if management methods are not effectively implemented [5, 6]. This insect exhibits considerable difficulty in control via chemical pesticides due to two primary factors; firstly, its larval developmental stage occurs inside the leaf mesophyll tissue, thereby the larvae are not easily exposed to chemical insecticides. The second challenging issue is the ability of insects to resist the various pesticides [5, 7]. Resistance to pyrethroids, abamectin, cartap and organophosphates (methamidophos) has been observed in South America. Partial resistance to pyrethroids has been reported in Europe, as well as to indoxacarb and spinosad in both Europe and South America. A significant increase in the application of chitin synthesis inhibitors has led to high levels of resistance to these compounds [6]. In Iran, the considerable resistance of this insect to organophosphate and pyrethroids has been reported [8], especially resistance to abamectin [9] and indoxacarb [10]. The release of sterile T. absoluta adults provides a basis for controlling this insect through SIT programs. Moreover, insects are unable to develop resistance to sterilization, and SIT may not have unintended impacts on non-target organism [11]. In Brazil, Argentina, Turkey, India, and China, laboratory assessments concerning the utilization of ionizing radiations (gamma and X-rays) for the management of T. absoluta have been conducted employing SIT and IS methods [12, 13, 14, 15, 16]. Arthur [12] reported that the lethal dose of Co 60 gamma radiation for T. absoluta pupae is quantified at 300 Gy and claimed 200 Gy is sterility dose of adults irradiated in the pupal stage. Arthur and Groppo [17] recorded the sterility dose for T. absoluta female and male adults as 150 and 200 Gy, respectively. The implementation of the inherited sterility method was also carried out on T. absoluta by Cagnotti et al. [13] utilizing X-ray. Yusef et al. [18] studied the effect of different doses of Co 60 gamma radiation on the sterility and inherited sterility of T. absoluta pupae. Cagnotti et al. [19] reported a significant reduction in the population of this insect in field experiments by irradiating pupae with 200 Gy of X-ray and releasing them in a ratio of 15:1 (untreated: treated males) within field cages compared to controls. Paladino et al. [20] reported that T. absoluta males irradiated with 300 Gy of X-ray produced a significantly more apyrene sperm than unirradiated males. All doses applied affected the morphology of eupyrene sperm bundles. Kuyulu and Genç [14] evaluated the effect of Co 60 gamma radiation at different doses on the developmental stages of eggs, fourth-instar larvae, and pupae of this insect. Sridhar et al. [15] subjected T. absoluta pupae aged two and five days to gamma irradiation at multiple doses. Their investigation encompassed the emergence rate of adult insects, the incidence of deformities, the duration of the larval and pupal stages, the rate of pupation, the longevity of adults, as well as their fertility. Zhou et al. [16] investigated the effect of C 137 gamma radiation doses on the F 0 and F 1 generations of T. absoluta . This method has also been evaluated in combination with other biological strategies like parasitoids intended for control of T. absoluta . Cagnotti et al. [21] reported that eggs laid by parents irradiated with X-rays (20834 Roentgen) during the pupal stage were acceptable for the oviposition by Trichogramma nerudai and T. pretiosum (Hymenoptera: Trichogrammatidae) and were suitable for the development of these parasitoids. Furthermore, Cagnotti et al. [22] stated that Tupiocoris cucurbitaceus (Spinola) (Hemiptera: Miridae), was capable of growth and reproduction on both irradiated and non-irradiated T. absoluta eggs. Also, Candas et al. [23] irradiated T. absoluta pupae with X-rays (20834 Roentgen) and subsequently employed them in the field experiments at a ratio of 15:1 alongside T. cucurbitaceus to manage this pest. Their findings illustrated that the integration of these two methods yielded a more pronounced and effective reduction in the population of this pest. Considering the population variation of T. absoluta across different global regions, alongside, the necessity to adapt sterile insect technique for controlling this invasive pest in Iran, the objective of the current research is to determine complete female sterility and male sub-sterility dose, as well as to assess the efficacy of these methods in T. absoluta control under laboratory conditions. To achieve these aims, the studies include: reproductive rate, sterility, adult emergence and longevity in the P 1 generation as well as sex ratio and inherited sterility in the F 1 generation. The competition ability of irradiated males against normal males was evaluated under laboratory conditions at various mating ratios, resulting in the determination of an optimal release ratio. Result Fecundity, fertility and sterility of the P 1 generation Irradiation of 6-days old male and female pupae of T. absoluta with different doses of gamma radiation significantly reduced fecundity and fertility of insects and increased the sterility rate (Table 1) according to Duncan's multiple range test (P<0.01) . This significant decrease in the number of eggs compared to the control was observed in all three combinations; IM×UF (F (5,23) =9.036, P<0.01 ), UM×IF (F( 5,23) =17.661, P<0.01 ), and IM×IF (F (5,23) =19.267, P<0.01 ). As a result of gamma irradiation, the percentage of hatched eggs across all three combinations; IM×UF (F (5,23) =18.565, P<0.01 ), UM×IF (F (5,23) =23.094, P<0.01 ) and IM×IF (F (5,23) =53.665, P<0.01 ) was significantly diminished in comparison to the control. The reduction in fecundity and fertility induced by gamma radiation exhibits a positive correlation with the radiation dose. These two parameters were used to calculate sterility rate, which increased with increasing gamma dose in all three combinations; IM×UF (F (4,19) =5.457, P<0.01 ), UM×IF (F (4,19) =2.296, P=0.107 ) and IM×IF (F (4,19) =2.39, P=0.097 ). However, this increase in the percentage of sterility was not significant in the UM×IF and IM×IF combinations. In the P 1 generation, differences in sensitivity to gamma radiation between males and females were evident. Completely sterile females and males were observed in the 250 and 300 Gy treatments, respectively. Additionally, when both of males and females irradiated at 150 Gy and crossed with each other, the fully sterility was recorded. The adult’s emergence and their longevity Following irradiation during the pupal stage, the emergence of adults (Figure 1) and their longevity (Figure 2) were recorded. Although the emergence of male (F (5,23) =2.075, P=0.116 ) (Figure 1A) and female (F (5,23) =2.597, P=0.062 ) (Figure 1B) adults decreased after irradiation compared to the control, this decrease was not significant according to Duncan's multiple range test (P<0.05) . Also, the percentage of unemerged pupae male (F (5,23) = 0.882, P=0.513 ) and female (F (5,23) =1.465, P=0.25 ) and the percentage of deformed adults male (F (5,23) =0.626, P=0.682 ) and female (F (5,23) =1.653, P=0.197 ) increased with increasing dose, but this increase was not significant according to Duncan's multiple range test (P<0.05) . The longevity of adult males (F (5,23) =0.173, P=0.969 ) and females (F (5,23) =0.384, P=0.0853 ) irradiated with different doses of gamma ray at the pupal stage did not show significant differences compared to the control according to Duncan's multiple range test (P<0.01) (Figure 2). Sex ratio and inherited sterility of F 1 generation The progeny of male parents irradiated with 250 Gy in pupal stage and combined with normal females were reared on artificial diet. The sex ratio of F 1 pupae in the treatment significantly shifted towards males compared to the control as assessed by independent T-test (t=-8.3497, P<0.01 ) (Figure 3). The number of male pupae per total number of pupae was calculated 55±1.91% in the control and 74±1.15% in the treatment. In order to assessment of inherited sterility, a dose of 250 Gy was utilized, given that in the P 1 experiments, females exhibited complete sterility at this dose, while a semi sterility was recorded in males. The fecundity, fertility and sterility rates of male and female F 1 progeny were recorded (Table 2). A significant decrease in the number of eggs per female was observed in both F 1 M×UF and UM×F 1 F crosses compared to the control (F (2,11) =211.356, P˂0.01 ). Furthermore, the percentage of eggs that hatched in these two crosses demonstrated a significant decline relative to the control (F (2,11)= 368.156, P˂0.01 ). Based on these two parameters, the percentage of sterility was calculated. In the F 1 M×UF cross, 100% F 1 sterility was calculated, while the UM×F 1 F cross yielded a sterility rate of 98%. This finding suggests that the impact of irradiation is more pronounced in F 1 males compared to F 1 females, indicating a greater inheritance of sterility in F 1 males than in F 1 females. Male mating competitiveness In the application of SIT for controlling pest insects, it is imperative that irradiated males possess the capacity to compete with wild males in order to successfully mate with wild females. Considering he outcomes from both P 1 and F 1 generations, 250 Gy was determined as the optimal dose for sterility. Therefore, the competitive ability of male moths obtained from pupae irradiated with 250 Gy gamma was assessed against unirradiated males. Through the calculation of the competitiveness value (CV) utilizing the Fried’s formula [24], males subjected to treatment competed perfectly with untreated males in the ratios of 1:1:1, 2:1:1, 3:1:1 and 4:1:1 (IM: UM: UF). The CV values in all ratios are greater than 1. (Table 3). The number of eggs laid by females copulating with both treated and untreated males across various ratios exhibited a significant reduction in comparison to the control (F (5,17) =3.516, P˂0.05 ). Furthermore, the proportion of eggs that hatched at different ratios was significantly diminished compared to the control (F (5,17) =6.324, P˂0.01 ). Based on the chi-square test (X 2 ), the observed egg hatching percentage was significantly ( P˂0.05 ) lower than the expected values at all ratios. Therefore, the 1:1:1 ratio emerged as a suitable proportion of treated males to untreated males for mating with normal females that were released under experimental conditions. Discussion The mass rearing of insects on artificial diet is a basic prerequisite for conducting evaluations related to the sterile insect technique (SIT) and F 1 inherited sterility (IS) [3]. In the extant literature concerning the application of the SIT for control of T. absoluta , specific artificial diet has not been introduced for mass rearing of this insect, and tomato plant was used for rearing. Therefore, it can be acknowledged that the present study is the first case of investigating the SIT method on T. absoluta that artificial diet was employed for mass rearing of this insect. The selection of an appropriate sterile and sub-sterile dose for the implementation of the SIT and IS methods represents is a very sensitive task. This dose must be selected in a manner such that it does not impose adverse effects on the insects, fertile females are not released into the ecosystem, and is calibrated to sustain the mating competitiveness of the male. In addition, this dose should not affect the biological parameters like adult emergence and longevity. The objective of inherited sterility is to identify a dose that is fully sterilizing for females and semi-sterilizing for males [25]. In previous studies concerning the application of SIT and IS techniques in the management of this insect, different radiation sources have been utilized, including X-ray, C 137 and Co 60 gamma rays. Arthur [12] irradiated T. absoluta pupae with Co 60 gamma ray and documented sterilizing doses of 150 and 200 Gy for females and males, respectively. At these doses, the oviposition did not occur, while a dose of 100 Gy yielded a 40% hatching rate of eggs. In the current investigation, regarding the IM×UF combination, similarly an egg hatching rate of 42.85% was noted at a dose of 100 Gy. However, at doses of 150, 200, 250, and 300 Gy, the hatching rates were recorded as 38.14, 28.27, 27.69 and 0%, respectively. Therefore, the sterility dose of males was determined as 300 Gy. Cagnotti et al. [13] claimed that 200 Gy of X-rays causes complete sterility on T. absoluta female pupae. Nevertheless, in the UM×IF combination of present study, at doses of 100, 150, 200, 250, and 300 Gy, the hatching rates were observed 35.3, 36.7, 12.31, 0 and 0% and the dose of 250 Gy was recognized as achieving complete sterility for females. Additionally, they asserted that at a dose of 300 Gy for the irradiation of male pupae, normal females that copulated with these males were unable to produce an adequate number of eggs (about 8 eggs), suggesting that the mating ability of males is compromised at a dose of 300 Gy. It confirms the findings of the current study, in which male and females irradiated separately at 300 Gy produced a limited quantity of eggs (~ 14 and 3 eggs, respectively), and even in the combination contained irradiated males and females, no eggs were observed. Kuyulu and Genç [14] stated that when males of T. absoluta were treated at doses of 100, 200 and 400 Gy Co 60 gamma ray, the number of eggs was 77, 82 and 49 and the percentage of egg survival was 78, 29 and 0%, respectively. When the females were treated at 100 and 200 Gy, 82 and 27 eggs with 59 and 26% survival rate were recorded, respectively. No eggs were observed at doses of 300 and 400 Gy, indicating the failure of mating in females at these higher doses. The sensitivity to radiation in females compared to males was also seen in current research. Sridhar et al. [15] recorded 43.8% egg hatching for T. absoluta at 300 Gy gamma ray, which is in conflict with the results of the present study. In our results, complete sterility occurred at 300 Gy. They stated that differences in insect sterilizing doses could be related to the age of the pupa, laboratory conditions, suitability of insect growth, and etc. Therefore, the difference in sensitivity to radiation can be related to small differences in the total DNA content and the developmental stage of the insects at the time of irradiation. On the other hand, the type of radiation is also important, and X-rays have different energy levels than gamma rays, so these two rays can affect chromatin differently [26]. Yamada et al. [27] explained the reasons behind the differences in dosage requirements among certain SIT researchers, who necessitate significantly greater doses than others to attain complete sterilization in the same insect species using same radian sources. They claimed dose-rate-dependent effects in inducing sterilization in insects are very important. Difference in required dose for complete sterility has been observed in Lepidoptera and mosquitos. This highlights the urgent need for SIT programs to perform routine and periodic quality control in dosimetry because the source of irradiators decays over time. However, additional investigations are necessary to enhance the comprehension of dose-rate-dependent effects in insect populations [27]. Inherited sterility occurs when insects are exposed to low radiation doses, causing their next generation to be more sterile than the parents. It is effective for pest control due to improved insect quality and compatibility with other control methods [2]. Cagnotti et al. [13] claimed that the quantity of second-generation (F 1 ) larvae of T. absoluta also exhibited a decline in correlation with increased radiation exposure. They indicated that when semi-sterile males were paired with fertile wild females, the detrimental consequences of radiation exposure were inherited and expressed for next generations, predominantly in the F 1 generation. In other words, egg laying was reduced in the F 1 generation and their progeny demonstrated a significantly higher sterility rate than irradiated parental moths. These findings corroborate the outcomes observed in the current investigation. They found complete sterility of females at 200 Gy X-ray radiation and reported this dose could be utilized to induced inherited sterility in this insect. Yusef et al. [18] reported that T. absoluta females were completely sterile at a dose of 100 Gy Co 60 gamma ray and considered a dose of 130 Gy to be suitable for inherited sterility in which males were partially sterile. Sridhar et al. [15] reported 65% hatching at a dose of 150 Gy and stated that the complete sterility did not occur in two consecutive generations at this dose (26 and 42% egg hatching). However, they considered the dose of 150 Gy to be suitable for inherited sterility. Zhou et al. [16] reported 300 Gy gamma ray of C 137 as relative sterilizing dose for males and complete sterilizing dose for females and they selected this dose for inherited sterility. They showed that the number of eggs in the combinations; F 1 M×UF and UM×F 1 F was reduced by 84.16 and 85.02% compared to the control, and 0 and 1.85% hatching rate were recorded, respectively. This suggests that males are more likely to inherit sterility than females. These results are similar with results of the current study, in which irradiated males at a dose of 250 Gy caused 27.69% of eggs to hatch in the P 1 generation and females were completely sterile at this dose, therefore this dose was selected for IS assessment. In the next generation, adult insects obtained from surviving eggs were crossed as F 1 M×UF and UM×F 1 F, and complete sterility was occurred (100 and 98.57%, respectively). One notable advantage of inherited sterility in lepidopterans is the alteration of the F 1 sex ratio favoring male offspring, attributed to their WZ/ZZ (male/female) chromosomal sex determination system. This shift is the result of recessive lethal mutations induced in the Z sex chromosomes of the treated parents, which are responsible for the death of F 1 females [2]. Zhou et al. [16] investigated the effect of C 137 gamma ray (100 to 400 Gy) on the sex ratio of T. absoluta F 1 generation. They claimed that with increasing radiation dose of parent males, the sex ratio of their offspring shifted more towards males. Yusef et al. [18] also showed that radiation treatment of Co 60 gamma ray causes a greater male bias in F 1 and F 2 generations of T. absoluta compared to the control, which is similar to the results of the present study. Nevertheless, Cagnotti et al. [13] reported that different doses of X-rays on male pupae exhibited no significant impact on the sex ratio of their F 1 generation and reported an overall male ratio of 66%. The observed variability in the outcomes of these researches can be attributed to the utilization of different radiation sources (X-rays versus gamma rays). Since longer exposure time in the higher doses may induce adverse effects on biological parameters and cause physiological defects on insects that decrease the competition ability of sterile moths, it is essential to ensure that the used dose rate for radiation does not have any negative effects [2, 26]. Arthur [12] elucidated that upon exposure of T. absoluta pupae to Co 60 gamma radiation doses of 100 and 200 Gy, the adult emergence was quantified at 60 and 52%, respectively. These findings align with the outcomes observed in the current study. However, they reported 300 Gy completely inhibited the emergence of adults, but the current study recorded an emergence rate of 43 to 46% at the 300 Gy treatments. Cagnotti et al. [13] stated that with increasing X-ray dose, the rate of adult emergence decreased, females that were irradiated with 250 Gy during the pupal stage did not fully emerge, and the number of males that emerged was very low. Additionally, they reported external abnormalities such as deformed wings and bent legs at doses higher than 350 Gy. Nevertheless, these researchers did not document any malformations at radiation doses ranging from 0 to 300 Gy, which is contradictory to the findings of our study. In this investigation, the percentage of malformed insects was recorded between 15 and 29% at doses between 0 and 300 Gy. Yusef et al. [18] also stated that the emergence of adults decreases with increasing dose and they did not consider doses higher than 200 Gy Co 60 gamma ray suitable for mating because they observed deformation of wings and bent legs at these doses. Kuyulu and Genç [14] showed that the adult’s emergence was 67, 42, 17 and 8% at doses of 100, 200, 300 and 400 Gy Co 60 gamma ray, respectively, and the emerged adults were deformed at 400 Gy. In contrast, in the present study, 43 to 57% emerged adults were observed at doses between 100 and 300 Gy. Sridhar et al. [15] reported the adult’s emergence was 31% at 300 Gy. Zhou et al. [16] stated that C 137 gamma radiation did not have a negative effect on the emergence of adult males and females, but males irradiated at 400 Gy were four times more deformed than controls. In our results, the rate of adult’s emergence and deformation did not increase significantly with increasing dose. Cagnotti et al. [13] showed that the longevity of T. absoluta males and females’ adults under X-rays treatment is not affected and recorded their average lifespan as 17 and 12 days respectively. Sridhar et al. [15] reported the lifespan of adults as 3.17 days at 300 Gy treatment of five-days old male pupae and 7.65 day in control. Zhou et al. [16] stated that males irradiated with 400 Gy C 137 gamma rays had a 16-day shorter lifespan than control, but at doses of 100 and 200 Gy, there was no significant difference. In the present study, the average longevity of adults was recorded as about 13 to 14 days and showed no significant difference under the influence of radiation compared to the control. These differing results depend on the various types of radiation source and the radiosensitivity of different geographic populations in different regions [26]. When SIT and IS are applied for insect control in the field and greenhouse, the mating competition ability of the released sterile males has great importance in the method’s success [26]. Zhou et al. [16] irradiated T. absoluta pupae with optimal dose of 300 Gy of C 137 gamma ray and released emerged adults in different treatments. They recorded egg hatching rate of 96% at the 0:1:1 (IM: UM: UF) ratio, 38% for the 1:0:1 ratio, and 66% for the 1:1:1 ratio in the competitiveness assessment. Subsequently they calculated a competitiveness value (CV) of 1.07, signifying that irradiated male maintain competitive with wild males during mating at 1:1:1 ratio. In the current research, when pupae irradiated with 250 Gy of Co 60 gamma ray and emerged adults released in different ratios, egg hatching rate was recorded 78, 22 and 37% at the 0:1:1, 1:0:1, and 1:1:1 ratios, respectively. Also, the competitiveness value was calculated 2.6, 3.92, 4.57 and 5.32 in 1:1:1, 2:1:1, 3:1:1 and 4:1:1 ratios, respectively. In all ratios, the competition index was higher than 1 and increased by increasing mating ratio, reflecting that these sterile males are more competitive than those of wild males for mating with wild females. Consequently, the initial ratio of 1:1:1 is recommended for release in controlled laboratory conditions. Conclusion In conclusion, a dose of 250 Gy Co 60 gamma ray can induce complete sterility of P 1 females and partial sterility of P 1 males, as well as complete sterility in the male progeny (F 1 ) of T. absoluta . A release ratio of 1:1:1 (IM: UM: UF) was sufficient for treated males to compete with normal males. Moreover, the irradiation of pupae at 250 Gy had no adverse effects on adult emergence and longevity. In view of these results, the use of partial sterility of P 1 male moths alongside their inherited sterility in F 1 progeny achieved through gamma-irradiation of 250 Gy Co 60 is recommended for the control of T. absoluta population in Iran. Materials and Methods Insect rearing Tomato leaves infested with T. absoluta were collected from our research greenhouse, and the larvae and pupae were maintained until the adult’s emergence. The emerged adults were transferred to egg-laying containers equipped with a cotton wick saturated with a 15% sugar solution. Crepe paper served as the substrate for egg deposition. The laid eggs were collected daily and transferred to the larval rearing containers (8 cm in diameter and 6 cm in height) which have a hole for ventilation and covered with a non-porous cloth. For rearing of larvae, the semi-artificial diet introduced by Greene et al. [28] was employed with some modifications derived from the diet proposed by Mihsfeldt and Parra [29]. This specific artificial diet was applied in a thin layer along the inner walls of the containers. The larval rearing containers were cleaned daily and new pieces of diet was added. The pupae formed between diet tissue were removed daily and kept in glass Petri dishes (100 mm × 15 mm) containing wet filter paper. Insects were reared at a temperature of 25 ± 1°C, a relative humidity of 40 ± 5%, and a photoperiod of 16:8 (L:D) hours. The effect of gamma radiation on P 1 and F 1 generations The pupae were collected daily from rearing containers and sexed under stereomicroscope based on the morphology of the last abdominal segment [30]. The six-days old pupae (48 to 72 hours prior to emergence) were subjected to gamma radiation of 100, 150, 200, 250, and 300 Gy. For irradiation, an Issledovatle PX30 gamma cell with a CO 60 source with a total activity of 200 Curies was used at a dose rate of 118 Gy/hr (~ 0.033 Gy/sec). This process was performed in four replicates. Subsequently, the percentage of emerged adults was calculated. The percentage of non-emerged pupae and deformed adults with non-flying wings were also recorded. Three emerged males and females were paired in egg-laying containers at a ratio of 1:1 across four replicates, in the following four combinations: Unirradiated Female (UF) × Unirradiated Male (UM) Unirradiated Female (UF) × Irradiated Male (IM) Irradiated Female (IF) × Unirradiated Male (UM) Irradiated Female (IF) × Irradiated Male (IM) To determine the appropriate sterility dose for males and females, the number of eggs laid and the percentage of eggs hatching were recorded and compared with the control. The sterility index was determined in accordance with Eq. (1) [31]; $$\:\%\:Sterility\:=100-(\frac{a\times\:b}{A\times\:B}\times\:100)$$ (1) Where: a: number of eggs in treatment, b: % hatching eggs in treatment, A: number of eggs in control, and B: % hatching eggs in control. The males and female adults’ longevity was recorded until the death of the last moth. To investigate inherited sterility, the male pupae were subjected to further irradiation at a dose of 250 Gy, based on the obtained results. Emerged male adults were mated with normal females, and control mating were concurrently prepared with normal pairs. Their offspring was reared on the artificial diet. The sex ratio of F 1 pupae in the control and treatment was recorded. F 1 adult males and females obtained from this IM×UF combination, were re-mated with pairs derived from the control combination UM×UF. Three pairs of F 1 adult males and females were crossed as F 1 M×UF, UM×F 1 F, and UM×UF as control in four replicates. The number of eggs, the percentage of egg hatching, and sterility rate were calculated. The effect of gamma radiation on male mating competitiveness The mating competitiveness ability of irradiated males compared to normal males was calculated under controlled laboratory conditions. According to the obtained results, 250 Gy was applied on male pupae. The irradiated and unirradiated emerged male adults (IM and UM) were mated with unirradiated females (UF) at different mating ratios of 1:1:1, 2:1:1, 3:1:1 and 4:1:1 (IM: UM: UM). The ratios of 0:1:1 and 1:0:1 (IM: UM: UM) were considered as controls. The number 1 in all ratios is three individuals. The number of eggs and egg hatch were counted daily. The observed egg hatch percentage and the expected egg hatch percentage at each ratio were compared using the chi-square (X 2 ) test. The expected egg-hatch percentage (Ee) were calculated with Eq. (2) [24]; $$\:Ee=\frac{N\left(Ha\right)+S\left(Hs\right)}{N+S}$$ (2) Where: Ee: Expected egg-hatch percentage, Ha: Hatched eggs percentage in 0:1:1 (IM: UM: UM) combination, Hs: Hatched eggs percentage in 1:0:1 (IM: UM: UM) combination, N: The number of normal males, S: The number of sterile males. The competitiveness values (CV) of the irradiated males based on the observed egg hatched (Eo) were evaluated according to Eq. (3) [24]; $$\:\:CV=\frac{Ha-Eo}{Eo-Hs}÷\frac{S}{N}$$ (3) This value when equal to 1, indicates a comparable level of competition between irradiated and unirradiated males. Values close to zero indicate superior competitiveness of unirradiated males [24]. Statistical analysis The biological and reproductive parameters were evaluated using one-way analysis of variance. A completely randomized design incorporating four replications was implemented. Mean values were compared using Duncan's multiple range test. The independent T-test was employed to compare means of F 1 sex ratio. Data were analyzed using SPSS software version 22 (IBM ©). Data Availability All data supporting the findings of this study are available within the paper. The datasets generated during the current study are available from the corresponding author on reasonable request. Declarations Acknowledgements I would like to thank Mr. Siyavash Farhadi and Dr. Saeid Moori for their technical help in the insects rearing and Nuclear Science and Technology Research Institute, Atomic Energy Organization of Iran (AEOI). Competing interests There is no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. Funding declaration No specific funding has been provided for this research. Author Contribution S. A. is responsible for all matters related to conducting experiments and writing the article. References Simmons, G. S., Bloem, K. A., Bloem, S., Carpenter, J. E. & Suckling, D. M. Impact of moth suppression/eradication programmes using the sterile insect technique or inherited sterility. In Sterile insect technique. Principles and practice in area-wide integrated pest management 1007-1050 (Dordrecht: Springer Netherlands, 2021). Marec, F. & Vreysen, M. J. Advances and challenges of using the sterile insect technique for the management of pest lepidoptera. Insects 10(11), 371 (2019). Dyck, V. A., Hendrichs, J. & Robinson, A. S. Sterile insect technique: principles and practice in area-wide integrated pest management p. 1216 (Taylor & Francis. 2021). Marec, F., Bloem, S. & Carpenter, J. E. Inherited sterility in insects. In Sterile insect technique. Principles and practice in area-wide integrated pest management 163-200 (Dordrecht: Springer Netherlands, 2021). Kumari, D. A., Anitha, G., Hirur, M. E., Suresh, V. & Nayak, M. Tomato leaf miner, Tuta absoluta : A review of its biology and management. J. Econ. Entomol. 45, 1050-1058 (2021). Biondi, A., Guedes, R. N. C., Wan, F. H. & Desneux, N. Ecology, worldwide spread, and management of the invasive South American tomato pinworm, Tuta absoluta : past, present, and future. Annu. Rev. Entomol. 63, 239-258 (2018). Aynalem, B. Tomato leafminer [( Tuta absoluta Meyrick) (Lepidoptera: Gelechiidae)] and its current ecofriendly management strategies: A review. J. Agric. Biotechnol. Sustainable Dev. 10(2), 11-24 (2018). Zibaee, I., Mahmood, K., Esmaeily, M., Bandani, A. R. & Kristensen, M. Organophosphate and pyrethroid resistances in the tomato leaf miner Tuta absoluta (Lepidoptera: Gelechiidae) from Iran. J. Appl. Entomol. 142(1-2), 181-191 (2018). Azizi, M. & Khajehali, J. Evaluation of resistance to abamectin in the populations of Tuta absoluta (Lepidoptera: gelechiidae), collected from Isfahan province, Iran. J. Agric. Sci. Tech. 24(2), 379-391 (2022). Aboutalebian-Soureshjani, A., Rafiee-Dastjerdi, H., Naseri, B., Hassanpour, M. & Khajehali, J. Indoxacarb resistance in Iranian populations of Tuta absoluta (Lepidoptera: Gelechiidae): Cross-resistance, biochemical and molecular mechanisms. Pestic. Biochem. Physiol . 196, 105633 (2023). Mawcha, K. T., et al. An overview of sustainable management strategies for Tuta absoluta . Int. J. Pest Manage. 1-24 (2025). Arthur, V. Use of gamma radiation to control three Lepidopteran pests in Brazil. Irradiation as a phytosanitary treatment of food and agricultural commodities. In: Proceedings of the final research coordination meeting organized by the joint FAO/IAEA Division of Nuclear Techniques in Food and Agriculture, 2002, Vienna, Austria, 45-50 (2004). Cagnotti, C. L., et al. Effects of X-rays on Tuta absoluta for use in inherited sterility programs. J. Pest Sci. 85, 413-421 (2012). Kuyulu, A. & Genç, H. Effects of gamma radiation on tomato leafminer, Tuta absoluta (Meyrick)(Lepidoptera: Gelechiidae). Int. J. Aerosp. Mech. Eng. 10(7), 451-455 (2016). Sridhar, V., Reddy, P. R., Vidyashree, L. K., Chandana, P. S. & Hadapad, B. A. Standardisation of gamma irradiation dose for Sterile Insect Technique to manage South American tomato moth [ Phthorimaea (Tuta) absoluta (Meyrick)]. J. Hortic. Sci. 18(2) 453-459 (2023). Zhou, S., et al. Screening the optimal dose of gamma radiation for Tuta absoluta sterility: paving the way for sterile insect technique programs. Entomol. Gen. 44(2) , 415-422 (2024). Arthur, V. & Groppo, G. A. Influence of gamma radiation on adults of Tuta absoluta (Meyrick) (Lep.: Gelechiidae). Boletín de Sanidad Vegetal, Plagas 33(1), 43-44 (2007). Yusef, M. V., et al. Determination of the range of gamma radiation dose to generate inherited sterility in the 'Tuta absoluta ' tomato moth (Meyrick, 1917)(Lepidoptera: Gelechiidae). In XLI Annual meeting of the Argentine Association of Nuclear Technology (AATN 2014) , no. INIS-AR-C-1707, pp. 15 (2014). Cagnotti, C. L., et al. Inherited sterility in Tuta absoluta (Lepidoptera: Gelechiidae): Pest population suppression and potential for combined use with a generalist predator. Fla. Entomol. 99 (sp1), 87-94 (2016). Paladino, L. Z. C., et al. The effect of X-rays on cytological traits of Tuta absoluta (Lepidoptera: Gelechiidae). Fla. Entomol. 99(sp1), 43-53 (2016). Cagnotti, C. L., et al. Acceptability and suitability of Tuta absoluta eggs from irradiated parents to parasitism by Trichogramma nerudai and Trichogramma pretiosum (Hymenoptera: Trichogrammatidae). Agric. Forest Entomol. 18(3), 198-205 (2016). Cagnotti, C. L., et al. Life history study of the mirid Tupiocoris cucurbitaceus feeding on Tuta absoluta eggs: implications for biological control and its combination with inherited sterility. Biocontrol 66, 207-216 (2021). Candas, L., Cagnotti, C. L. & López, S. N. Integration of inherited sterility and inoculative releases of a miridae predator for the control of the tomato leaf miner, Tuta absoluta . BioControl 69(1), 29-37 (2024). Fried, M. Determination of sterile insect competitiveness. J. Econ. Entomol. 6, 869-872 (1971). Bloem, S., Carpenter, J. E. & Hofmeyr, J. H. Radiation biology and inherited Sterility in false codling moth (Lepidoptera: Tortricidae). J. Econ. Entomol. 96, 1724-1731 (2003). Bakri, A., Mehta, K. & Lance, D. R. Sterilizing insects with ionizing radiation. In Sterile insect technique. Principles and practice in area-wide integrated pest management 355-398 (Dordrecht: Springer Netherlands, 2021). Yamada, H., et al. Radiation dose-rate is a neglected critical parameter in dose–response of insects. Sci. Rep. 12, 6242 (2022). Greene, G. L., Leppla, N. C. & Dickerson, W. A. Velvet bean caterpillar: a rearing procedure and artificial medium. J. Econ. Entomol. 69, 487-488 (1976). Mihsfeldt, L. H. & Parra, J. R. P. Biologia de Tuta absoluta (Meyrick, 1917) em dieta artificial. Sci. Agric. 56(4), 769-776 (1999). Genç, H. The tomato leafminer, Tuta absoluta (Meyrick)(Lepidoptera: Gelechiidae): pupal key characters for sexing individuals. Turk. J. Zool. 40(5), 801-805 (2016). Toppazada, A., Abdallah, S. & Eldefrawi, M. E. Chemosterilization of larvae and adults of the Egyptian cotton leaf worm, Prodenia litura by Apholate, Metepa and Tepa. J. Econ. Entomol. 59, 1125-1128 (1966). Tables Table 1. Fecundity, fertility and sterility (Mean ± SE) of Tuta absoluta parental generation that emerged from pupae irradiated a with different gamma ray doses (Gy). U: unirradiated, I: irradiated, M: male and F: female. Means indicated by different letters in each separate columns are significantly different according to Duncan's multiple range test (P<0.01). Combination Dose (Gy) Egg number ± SE Egg hatch (%) ± SE Sterility (%) ± SE IM×UF 0 98.25±12.60 a 80.17±4.45 a 100 47.5±11.93 b 42.85±4.21 b 73.34±7.31 c 150 34.25±12.68 bc 38.13±12.18 b 87.62±3.45 bc 200 37.50±11.85 bc 28.26±3.95 b 86.03±4.43 bc 250 17.75±3.40 bc 27.69± 4.47 b 93.33±2.16 ab 300 13.75±2.17 c 0.00±0.00 c 100±0.00 a UM×IF 0 98.25±12.60 a 80.17±4.45 a 100 25.00±9.65 b 35.30±2.44 b 87.93±5.45 a 150 18.75±6.99 b 36.70±12.84 b 88.57±5.89 a 200 19.50±10.27 b 12.31±7.19 c 94.29±3.30 a 250 6.25±5.26 b 0.00±0.00 c 100±0.00 a 300 2.50±2.50 b 0.00±0.00 c 100±0.00 a IM×IF 0 98.25±12.60 a 80.17±4.45 a 100 25.25±14.69 b 14.42±9.76 b 91.11±5.45 a 150 8.5±5.68 b 0.00±0.00 c 100±0.00 a 200 9.25±5.45 b 0.00±0.00 c 100±0.00 a 250 1.75±1.43 b 0.00±0.00 c 100±0.00 a 300 0.00±0.00 b 0.00±0.00 c 100±0.00 a Table 2 . F 1 fecundity, fertility and sterility percentage of Tuta absoluta resulting from irradiation of P 1 male pupae with 250 Gy gamma ray. U: unirradiated, M: male and F: female. Means indicated by different letters in each separate row are significantly different according to Duncan's multiple range test (P<0.01) . Combination Egg number ± SE Egg hatch (%) ± SE Sterility (%) ± SE UM×UF 9 5.5±3.66 76.32 ±3 .31 a F 1 M×UF 1 8.5±3.12 b 0.00± 0.00 b 100.00±0.00 UM×F 1 F 1 6.25±2.39 b 5.44±1.95 b 98.57±0.78 Table 3 . Number of eggs, observed and expected egg hatching percentages, and competitiveness value of Tuta absoluta resulting from different release ratios of irradiated males by 250 Gy gamma ray alongside unirradiated males and females. U: unirradiated, I: irradiated, M: male and F: female. Means indicated by different letters in each column are significantly different according to Duncan's multiple range test (P<0.01) . Ratio (IM: UM: UF) Egg number± SE Observed egg hatch % Expected egg hatch % X 2 df=3 P ˂ 0.05 CV 0:1:1 102.00±17.01 a 78.16±5.62 a 1:1:1 23.67±9.94 b 37.17±16.15 b 50.17 43.14 * 2.60 2:1:1 21.00±2.31 b 28.53±7.12 b 40.85 17.40 * 3.92 3:1:1 59.00±12.85 ab 26.00±8.24 b 36.18 17.76 * 4.57 4:1:1 38.33±2.848 b 24.71±3.40 b 33.38 10.60 * 5.32 1:0:1 45.33±30.90 b 22.19±2.3 b X 2 values marked with asterisk (*) indicate a significant difference between the observed and expected percentage of egg hatching (P˂0.05) . CV: Competitiveness value [24]. The number 1 in all ratios is equivalent to three insects Additional Declarations No competing interests reported. Cite Share Download PDF Status: Published Journal Publication published 13 Dec, 2025 Read the published version in Scientific Reports → Version 1 posted Editorial decision: Revision requested 10 Oct, 2025 Reviews received at journal 07 Oct, 2025 Reviewers agreed at journal 06 Oct, 2025 Reviewers agreed at journal 05 Oct, 2025 Reviews received at journal 02 Oct, 2025 Reviewers agreed at journal 30 Sep, 2025 Reviewers agreed at journal 29 Sep, 2025 Reviewers agreed at journal 21 Jul, 2025 Reviews received at journal 29 Jun, 2025 Reviewers agreed at journal 17 Jun, 2025 Reviewers agreed at journal 15 Jun, 2025 Reviewers invited by journal 15 Jun, 2025 Editor assigned by journal 15 Jun, 2025 Editor invited by journal 05 Jun, 2025 Submission checks completed at journal 22 May, 2025 First submitted to journal 22 May, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. 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Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6690207","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Article","associatedPublications":[],"authors":[{"id":472644700,"identity":"2c25a93c-0fdd-4fa5-979b-8eb30883d1bc","order_by":0,"name":"Shabnam Ashouri","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAA4UlEQVRIiWNgGAWjYDACCQYGZhDNxt4AJA0gggeI08JzgFQtDBIJRLpLfnbz488FFYfz+SQfP3zwoYBBzrx/AePhCjxaDO4cM5OeceawZZt0mrHhDAMGY5kbDxgOnsGnRSLBjJm37bABm3SCmTSPAUPiDIkDDAcb8DlsRvrnz2Atkse//yZKC8ONHANpsBYJHjNmsBb+BvxaDG7klEnznEk3YOPJKZacYSBhLCHB2EDIYZs/81RYG8i3H9/44cMfGzkJ/sOHP+J1GBoARpNEIikawID/AKk6RsEoGAWjYJgDAFtfR6frNIZvAAAAAElFTkSuQmCC","orcid":"","institution":"Nuclear Science and Technology Research Institute","correspondingAuthor":true,"prefix":"","firstName":"Shabnam","middleName":"","lastName":"Ashouri","suffix":""}],"badges":[],"createdAt":"2025-05-18 07:08:11","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6690207/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6690207/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1038/s41598-025-32383-2","type":"published","date":"2025-12-13T15:59:43+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":84982445,"identity":"f1d50cd2-8c00-4c08-be2b-4425740ffa41","added_by":"auto","created_at":"2025-06-19 13:42:44","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":77042,"visible":true,"origin":"","legend":"\u003cp\u003ePercentage of adult’s emergence, unemerged pupae and deformed male (A) and female (B) adults of \u003cem\u003eTuta absoluta\u003c/em\u003e originating from pupae subjected to different gamma ray doses (Gy).\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-6690207/v1/9a8b5d59ba1b6c02c1cf6de5.png"},{"id":84982977,"identity":"cc03b60c-aa6e-4543-ba9a-1849537742f4","added_by":"auto","created_at":"2025-06-19 13:50:44","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":23318,"visible":true,"origin":"","legend":"\u003cp\u003eLongevity of \u003cem\u003eTuta absoluta\u003c/em\u003e male and female adults originating from pupae subjected to different gamma ray doses (Gy).\u003c/p\u003e","description":"","filename":"2.png","url":"https://assets-eu.researchsquare.com/files/rs-6690207/v1/04526acc5d78eff705f3235e.png"},{"id":84982446,"identity":"05cd3834-2fb6-4ba4-92a5-c60d4661cbe9","added_by":"auto","created_at":"2025-06-19 13:42:44","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":16122,"visible":true,"origin":"","legend":"\u003cp\u003eF\u003csub\u003e1\u003c/sub\u003e progeny sex ratio (male pupae to total pupae) of \u003cem\u003eTuta absoluta\u003c/em\u003e resulting from the irradiation of male P\u003csub\u003e1\u003c/sub\u003e pupae with 250 Gy gamma ray and emerged males was crossed with normal females, in comparison with control. Two columns display significant differences as determined by independent T-test \u003cem\u003e(P\u0026lt;0.01)\u003c/em\u003e.\u003c/p\u003e","description":"","filename":"3.png","url":"https://assets-eu.researchsquare.com/files/rs-6690207/v1/7e7f961163a7d30451f01248.png"},{"id":98244072,"identity":"4e6075f5-a661-4e7f-9720-d6f329efdfb7","added_by":"auto","created_at":"2025-12-15 16:12:48","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1008922,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6690207/v1/cb9064df-7f84-4ab0-89d0-7a8e56f1c277.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"\u003cp\u003eOptimization of Co\u003csup\u003e60\u003c/sup\u003e gamma radiation dose for applying sterile insect technique and inherited sterility on \u003cem\u003eTuta absoluta\u003c/em\u003e (Meyrick) in Iran\u003c/p\u003e","fulltext":[{"header":"Introduction","content":"\u003cp\u003eThe continuous application of chemical pesticides for insect control intensifies the accumulation of pollutants in ecosystems and presents considerable challenges to human health and environmental integrity. The destruction of plant pollinators and natural enemies of pests represents additional harmful consequences associated with the application of chemical pesticides [1]. These environmental risks require the adoption of safe and effective integrated pest management strategies whitin sustainable and modern agriculture. The sterile insect technique (SIT) is regarded as a prominent biological and species-specific methods for pest control [2]. In this technique, sterile insects are released in large numbers in an infested area to reduce the fertility of the wild population of the same species. This method involves mass breeding of the target pest species, exposing them to gamma or X-rays to induce sexual sterility, and subsequently releasing them on a large scale into the target pest population. In essence, the released sterile males mate with wild females, thereby reducing their reproduction and fertility [2, 3]. The mutations induced through this technique exhibit distinct characteristics in lepidopterans. In lepidopterans, certain dominant lethal mutations can be expressed immediately prior to egg hatching, and most of the damaged chromosomes are transmitted to the F\u003csub\u003e1\u003c/sub\u003e generation. Lepidoptera possess several specific cytogenetic and cytological features which predominantly contribute to their elevated resistance to ionizing radiation. Consequently, very high radiation doses are required to achieve complete sterility in lepidopteran males. These features also significantly influence the mechanism underlying inherited sterility (IS). Therefore, inherited sterility is a type of SIT wherein semi-sterile males are released. A synergistic approach that combines SIT and IS may be employed to control lepidopteran pests [1, 2, 3, 4].\u003c/p\u003e \u003cp\u003eIn recent years, the tomato leaf miner, \u003cem\u003eTuta absoluta\u003c/em\u003e (Meyrick) (Lepidoptera: Gelechiidae), has emerged as the most invasive and destructive pest affecting tomato crops and is considered a major threat to the cultivation of numerous solanaceous species, including eggplant, peppers and potatoes globally [5]. \u003cem\u003eTuta absoluta\u003c/em\u003e larvae causes damage to the entire plant and specially feed from the leaf\u0026rsquo;s mesophyll. Moreover, the feeding activity on the fruits affects the visual appeal of harvested products and leading to fruit deterioration. The larvae of this pest can inflict 80 to 100% damage to tomatoes if management methods are not effectively implemented [5, 6]. This insect exhibits considerable difficulty in control via chemical pesticides due to two primary factors; firstly, its larval developmental stage occurs inside the leaf mesophyll tissue, thereby the larvae are not easily exposed to chemical insecticides. The second challenging issue is the ability of insects to resist the various pesticides [5, 7]. Resistance to pyrethroids, abamectin, cartap and organophosphates (methamidophos) has been observed in South America. Partial resistance to pyrethroids has been reported in Europe, as well as to indoxacarb and spinosad in both Europe and South America. A significant increase in the application of chitin synthesis inhibitors has led to high levels of resistance to these compounds [6]. In Iran, the considerable resistance of this insect to organophosphate and pyrethroids has been reported [8], especially resistance to abamectin [9] and indoxacarb [10]. The release of sterile \u003cem\u003eT. absoluta\u003c/em\u003e adults provides a basis for controlling this insect through SIT programs. Moreover, insects are unable to develop resistance to sterilization, and SIT may not have unintended impacts on non-target organism [11]. In Brazil, Argentina, Turkey, India, and China, laboratory assessments concerning the utilization of ionizing radiations (gamma and X-rays) for the management of \u003cem\u003eT. absoluta\u003c/em\u003e have been conducted employing SIT and IS methods [12, 13, 14, 15, 16].\u003c/p\u003e \u003cp\u003eArthur [12] reported that the lethal dose of Co\u003csup\u003e60\u003c/sup\u003e gamma radiation for \u003cem\u003eT. absoluta\u003c/em\u003e pupae is quantified at 300 Gy and claimed 200 Gy is sterility dose of adults irradiated in the pupal stage. Arthur and Groppo [17] recorded the sterility dose for \u003cem\u003eT. absoluta\u003c/em\u003e female and male adults as 150 and 200 Gy, respectively. The implementation of the inherited sterility method was also carried out on \u003cem\u003eT. absoluta\u003c/em\u003e by Cagnotti et al. [13] utilizing X-ray. Yusef et al. [18] studied the effect of different doses of Co\u003csup\u003e60\u003c/sup\u003e gamma radiation on the sterility and inherited sterility of \u003cem\u003eT. absoluta\u003c/em\u003e pupae. Cagnotti et al. [19] reported a significant reduction in the population of this insect in field experiments by irradiating pupae with 200 Gy of X-ray and releasing them in a ratio of 15:1 (untreated: treated males) within field cages compared to controls. Paladino et al. [20] reported that \u003cem\u003eT. absoluta\u003c/em\u003e males irradiated with 300 Gy of X-ray produced a significantly more apyrene sperm than unirradiated males. All doses applied affected the morphology of eupyrene sperm bundles. Kuyulu and Gen\u0026ccedil; [14] evaluated the effect of Co\u003csup\u003e60\u003c/sup\u003e gamma radiation at different doses on the developmental stages of eggs, fourth-instar larvae, and pupae of this insect. Sridhar et al. [15] subjected \u003cem\u003eT. absoluta\u003c/em\u003e pupae aged two and five days to gamma irradiation at multiple doses. Their investigation encompassed the emergence rate of adult insects, the incidence of deformities, the duration of the larval and pupal stages, the rate of pupation, the longevity of adults, as well as their fertility. Zhou et al. [16] investigated the effect of C\u003csup\u003e137\u003c/sup\u003e gamma radiation doses on the F\u003csub\u003e0\u003c/sub\u003e and F\u003csub\u003e1\u003c/sub\u003e generations of \u003cem\u003eT. absoluta\u003c/em\u003e.\u003c/p\u003e \u003cp\u003eThis method has also been evaluated in combination with other biological strategies like parasitoids intended for control of \u003cem\u003eT. absoluta\u003c/em\u003e. Cagnotti et al. [21] reported that eggs laid by parents irradiated with X-rays (20834 Roentgen) during the pupal stage were acceptable for the oviposition by \u003cem\u003eTrichogramma nerudai\u003c/em\u003e and \u003cem\u003eT. pretiosum\u003c/em\u003e (Hymenoptera: Trichogrammatidae) and were suitable for the development of these parasitoids. Furthermore, Cagnotti et al. [22] stated that \u003cem\u003eTupiocoris cucurbitaceus\u003c/em\u003e (Spinola) (Hemiptera: Miridae), was capable of growth and reproduction on both irradiated and non-irradiated \u003cem\u003eT. absoluta\u003c/em\u003e eggs. Also, Candas et al. [23] irradiated \u003cem\u003eT. absoluta\u003c/em\u003e pupae with X-rays (20834 Roentgen) and subsequently employed them in the field experiments at a ratio of 15:1 alongside \u003cem\u003eT. cucurbitaceus\u003c/em\u003e to manage this pest. Their findings illustrated that the integration of these two methods yielded a more pronounced and effective reduction in the population of this pest.\u003c/p\u003e \u003cp\u003eConsidering the population variation of \u003cem\u003eT. absoluta\u003c/em\u003e across different global regions, alongside, the necessity to adapt sterile insect technique for controlling this invasive pest in Iran, the objective of the current research is to determine complete female sterility and male sub-sterility dose, as well as to assess the efficacy of these methods in \u003cem\u003eT. absoluta\u003c/em\u003e control under laboratory conditions. To achieve these aims, the studies include: reproductive rate, sterility, adult emergence and longevity in the P\u003csub\u003e1\u003c/sub\u003e generation as well as sex ratio and inherited sterility in the F\u003csub\u003e1\u003c/sub\u003e generation. The competition ability of irradiated males against normal males was evaluated under laboratory conditions at various mating ratios, resulting in the determination of an optimal release ratio.\u003c/p\u003e"},{"header":"Result","content":"\u003cp\u003e\u003cstrong\u003eFecundity, fertility and sterility of the P\u003csub\u003e1\u003c/sub\u003e generation\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIrradiation of 6-days old male and female pupae of \u003cem\u003eT. absoluta\u003c/em\u003e with different doses of gamma radiation significantly reduced fecundity and fertility of insects and increased the sterility rate (Table 1) according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.01)\u003c/em\u003e. This significant decrease in the number of eggs compared to the control was observed in all three combinations; IM\u0026times;UF (F\u003csub\u003e(5,23)\u003c/sub\u003e=9.036, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e), UM\u0026times;IF (F(\u003csub\u003e5,23)\u003c/sub\u003e=17.661, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e), and IM\u0026times;IF (F\u003csub\u003e(5,23)\u003c/sub\u003e=19.267, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e).\u003c/p\u003e\n\u003cp\u003eAs a result of gamma irradiation, the percentage of hatched eggs across all three combinations; IM\u0026times;UF (F\u003csub\u003e(5,23)\u003c/sub\u003e=18.565, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e), UM\u0026times;IF (F\u003csub\u003e(5,23)\u003c/sub\u003e=23.094, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e) and IM\u0026times;IF (F\u003csub\u003e(5,23)\u003c/sub\u003e=53.665, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e) was significantly diminished in comparison to the control.\u003c/p\u003e\n\u003cp\u003eThe reduction in fecundity and fertility induced by gamma radiation exhibits a positive correlation with the radiation dose. These two parameters were used to calculate sterility rate, which increased with increasing gamma dose in all three combinations; IM\u0026times;UF (F\u003csub\u003e(4,19)\u003c/sub\u003e=5.457, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e), UM\u0026times;IF (F\u003csub\u003e(4,19)\u003c/sub\u003e=2.296, \u003cem\u003eP=0.107\u003c/em\u003e) and IM\u0026times;IF (F\u003csub\u003e(4,19)\u003c/sub\u003e=2.39, \u003cem\u003eP=0.097\u003c/em\u003e). However, this increase in the percentage of sterility was not significant in the UM\u0026times;IF and IM\u0026times;IF combinations. In the P\u003csub\u003e1\u003c/sub\u003e generation, differences in sensitivity to gamma radiation between males and females were evident. Completely sterile females and males were observed in the 250 and 300 Gy treatments, respectively. Additionally, when both of males and females irradiated at 150 Gy and crossed with each other, the fully sterility was recorded.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eThe adult\u0026rsquo;s emergence and their longevity\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFollowing irradiation during the pupal stage, the emergence of adults (Figure 1) and their longevity (Figure 2) were recorded. Although the emergence of male (F\u003csub\u003e(5,23)\u003c/sub\u003e=2.075, \u003cem\u003eP=0.116\u003c/em\u003e) (Figure 1A) and female (F\u003csub\u003e(5,23)\u003c/sub\u003e=2.597, \u003cem\u003eP=0.062\u003c/em\u003e) (Figure 1B) adults decreased after irradiation compared to the control, this decrease was not significant according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.05)\u003c/em\u003e. Also, the percentage of unemerged pupae male (F\u003csub\u003e(5,23)\u003c/sub\u003e= 0.882, \u003cem\u003eP=0.513\u003c/em\u003e) and female (F\u003csub\u003e(5,23)\u003c/sub\u003e=1.465, \u003cem\u003eP=0.25\u003c/em\u003e) and the percentage of deformed adults male (F\u003csub\u003e(5,23)\u003c/sub\u003e=0.626, \u003cem\u003eP=0.682\u003c/em\u003e) and female (F\u003csub\u003e(5,23)\u003c/sub\u003e=1.653, \u003cem\u003eP=0.197\u003c/em\u003e) increased with increasing dose, but this increase was not significant according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.05)\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eThe longevity of adult males (F\u003csub\u003e(5,23)\u003c/sub\u003e=0.173, \u003cem\u003eP=0.969\u003c/em\u003e) and females (F\u003csub\u003e(5,23)\u003c/sub\u003e=0.384, \u003cem\u003eP=0.0853\u003c/em\u003e) irradiated with different doses of gamma ray at the pupal stage did not show significant differences compared to the control according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.01)\u003c/em\u003e (Figure 2).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSex ratio and inherited sterility of F\u003csub\u003e1\u003c/sub\u003e generation\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe progeny of male parents irradiated with 250 Gy in pupal stage and combined with normal females were reared on artificial diet. The sex ratio of F\u003csub\u003e1\u003c/sub\u003e pupae in the treatment significantly shifted towards males compared to the control as assessed by independent T-test (t=-8.3497, \u003cem\u003eP\u0026lt;0.01\u003c/em\u003e) (Figure 3). The number of male pupae per total number of pupae was calculated 55\u0026plusmn;1.91% in the control and 74\u0026plusmn;1.15% in the treatment. \u003c/p\u003e\n\u003cp\u003eIn order to assessment of inherited sterility, a dose of 250 Gy was utilized, given that in the P\u003csub\u003e1\u003c/sub\u003e experiments, females exhibited complete sterility at this dose, while a semi sterility was recorded in males. The fecundity, fertility and sterility rates of male and female F\u003csub\u003e1\u003c/sub\u003e progeny were recorded (Table 2). A significant decrease in the number of eggs per female was observed in both F\u003csub\u003e1\u003c/sub\u003eM\u0026times;UF and UM\u0026times;F\u003csub\u003e1\u003c/sub\u003eF crosses compared to the control (F\u003csub\u003e(2,11)\u003c/sub\u003e=211.356, \u003cem\u003eP˂0.01\u003c/em\u003e). Furthermore, the percentage of eggs that hatched in these two crosses demonstrated a significant decline relative to the control (F\u003csub\u003e(2,11)=\u003c/sub\u003e368.156, \u003cem\u003eP˂0.01\u003c/em\u003e). Based on these two parameters, the percentage of sterility was calculated. In the F\u003csub\u003e1\u003c/sub\u003eM\u0026times;UF cross, 100% F\u003csub\u003e1\u003c/sub\u003e sterility was calculated, while the UM\u0026times;F\u003csub\u003e1\u003c/sub\u003eF cross yielded a sterility rate of 98%. This finding suggests that the impact of irradiation is more pronounced in F\u003csub\u003e1\u003c/sub\u003e males compared to F\u003csub\u003e1\u003c/sub\u003e females, indicating a greater inheritance of sterility in F\u003csub\u003e1\u003c/sub\u003e males than in F\u003csub\u003e1\u003c/sub\u003e females.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eMale mating competitiveness\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn the application of SIT for controlling pest insects, it is imperative that irradiated males possess the capacity to compete with wild males in order to successfully mate with wild females. Considering he outcomes from both P\u003csub\u003e1\u003c/sub\u003e and F\u003csub\u003e1\u003c/sub\u003e generations, 250 Gy was determined as the optimal dose for sterility. Therefore, the competitive ability of male moths obtained from pupae irradiated with 250 Gy gamma was assessed against unirradiated males. Through the calculation of the competitiveness value (CV) utilizing the Fried\u0026rsquo;s formula [24], males subjected to treatment competed perfectly with untreated males in the ratios of 1:1:1, 2:1:1, 3:1:1 and 4:1:1 (IM: UM: UF). The CV values in all ratios are greater than 1. (Table 3). The number of eggs laid by females copulating with both treated and untreated males across various ratios exhibited a significant reduction in comparison to the control (F\u003csub\u003e(5,17)\u003c/sub\u003e=3.516, \u003cem\u003eP˂0.05\u003c/em\u003e). Furthermore, the proportion of eggs that hatched at different ratios was significantly diminished compared to the control (F\u003csub\u003e(5,17)\u003c/sub\u003e=6.324, \u003cem\u003eP˂0.01\u003c/em\u003e). Based on the chi-square test (X\u003csup\u003e2\u003c/sup\u003e), the observed egg hatching percentage was significantly (\u003cem\u003eP˂0.05\u003c/em\u003e) lower than the expected values at all ratios. Therefore, the 1:1:1 ratio emerged as a suitable proportion of treated males to untreated males for mating with normal females that were released under experimental conditions. \u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe mass rearing of insects on artificial diet is a basic prerequisite for conducting evaluations related to the sterile insect technique (SIT) and F\u003csub\u003e1\u003c/sub\u003e inherited sterility (IS) [3]. In the extant literature concerning the application of the SIT for control of \u003cem\u003eT. absoluta\u003c/em\u003e, specific artificial diet has not been introduced for mass rearing of this insect, and tomato plant was used for rearing. Therefore, it can be acknowledged that the present study is the first case of investigating the SIT method on \u003cem\u003eT. absoluta\u003c/em\u003e that artificial diet was employed for mass rearing of this insect.\u003c/p\u003e \u003cp\u003eThe selection of an appropriate sterile and sub-sterile dose for the implementation of the SIT and IS methods represents is a very sensitive task. This dose must be selected in a manner such that it does not impose adverse effects on the insects, fertile females are not released into the ecosystem, and is calibrated to sustain the mating competitiveness of the male. In addition, this dose should not affect the biological parameters like adult emergence and longevity. The objective of inherited sterility is to identify a dose that is fully sterilizing for females and semi-sterilizing for males [25]. In previous studies concerning the application of SIT and IS techniques in the management of this insect, different radiation sources have been utilized, including X-ray, C\u003csup\u003e137\u003c/sup\u003eand Co\u003csup\u003e60\u003c/sup\u003e gamma rays. Arthur [12] irradiated \u003cem\u003eT. absoluta\u003c/em\u003e pupae with Co\u003csup\u003e60\u003c/sup\u003e gamma ray and documented sterilizing doses of 150 and 200 Gy for females and males, respectively. At these doses, the oviposition did not occur, while a dose of 100 Gy yielded a 40% hatching rate of eggs. In the current investigation, regarding the IM\u0026times;UF combination, similarly an egg hatching rate of 42.85% was noted at a dose of 100 Gy. However, at doses of 150, 200, 250, and 300 Gy, the hatching rates were recorded as 38.14, 28.27, 27.69 and 0%, respectively. Therefore, the sterility dose of males was determined as 300 Gy. Cagnotti et al. [13] claimed that 200 Gy of X-rays causes complete sterility on \u003cem\u003eT. absoluta\u003c/em\u003e female pupae. Nevertheless, in the UM\u0026times;IF combination of present study, at doses of 100, 150, 200, 250, and 300 Gy, the hatching rates were observed 35.3, 36.7, 12.31, 0 and 0% and the dose of 250 Gy was recognized as achieving complete sterility for females. Additionally, they asserted that at a dose of 300 Gy for the irradiation of male pupae, normal females that copulated with these males were unable to produce an adequate number of eggs (about 8 eggs), suggesting that the mating ability of males is compromised at a dose of 300 Gy. It confirms the findings of the current study, in which male and females irradiated separately at 300 Gy produced a limited quantity of eggs (~\u0026thinsp;14 and 3 eggs, respectively), and even in the combination contained irradiated males and females, no eggs were observed. Kuyulu and Gen\u0026ccedil; [14] stated that when males of \u003cem\u003eT. absoluta\u003c/em\u003e were treated at doses of 100, 200 and 400 Gy Co\u003csup\u003e60\u003c/sup\u003e gamma ray, the number of eggs was 77, 82 and 49 and the percentage of egg survival was 78, 29 and 0%, respectively. When the females were treated at 100 and 200 Gy, 82 and 27 eggs with 59 and 26% survival rate were recorded, respectively. No eggs were observed at doses of 300 and 400 Gy, indicating the failure of mating in females at these higher doses. The sensitivity to radiation in females compared to males was also seen in current research. Sridhar et al. [15] recorded 43.8% egg hatching for \u003cem\u003eT. absoluta\u003c/em\u003e at 300 Gy gamma ray, which is in conflict with the results of the present study. In our results, complete sterility occurred at 300 Gy. They stated that differences in insect sterilizing doses could be related to the age of the pupa, laboratory conditions, suitability of insect growth, and etc. Therefore, the difference in sensitivity to radiation can be related to small differences in the total DNA content and the developmental stage of the insects at the time of irradiation. On the other hand, the type of radiation is also important, and X-rays have different energy levels than gamma rays, so these two rays can affect chromatin differently [26]. Yamada et al. [27] explained the reasons behind the differences in dosage requirements among certain SIT researchers, who necessitate significantly greater doses than others to attain complete sterilization in the same insect species using same radian sources. They claimed dose-rate-dependent effects in inducing sterilization in insects are very important. Difference in required dose for complete sterility has been observed in Lepidoptera and mosquitos. This highlights the urgent need for SIT programs to perform routine and periodic quality control in dosimetry because the source of irradiators decays over time. However, additional investigations are necessary to enhance the comprehension of dose-rate-dependent effects in insect populations [27].\u003c/p\u003e \u003cp\u003eInherited sterility occurs when insects are exposed to low radiation doses, causing their next generation to be more sterile than the parents. It is effective for pest control due to improved insect quality and compatibility with other control methods [2]. Cagnotti et al. [13] claimed that the quantity of second-generation (F\u003csub\u003e1\u003c/sub\u003e) larvae of \u003cem\u003eT. absoluta\u003c/em\u003e also exhibited a decline in correlation with increased radiation exposure. They indicated that when semi-sterile males were paired with fertile wild females, the detrimental consequences of radiation exposure were inherited and expressed for next generations, predominantly in the F\u003csub\u003e1\u003c/sub\u003e generation. In other words, egg laying was reduced in the F\u003csub\u003e1\u003c/sub\u003e generation and their progeny demonstrated a significantly higher sterility rate than irradiated parental moths. These findings corroborate the outcomes observed in the current investigation. They found complete sterility of females at 200 Gy X-ray radiation and reported this dose could be utilized to induced inherited sterility in this insect. Yusef et al. [18] reported that \u003cem\u003eT. absoluta\u003c/em\u003e females were completely sterile at a dose of 100 Gy Co\u003csup\u003e60\u003c/sup\u003egamma ray and considered a dose of 130 Gy to be suitable for inherited sterility in which males were partially sterile. Sridhar et al. [15] reported 65% hatching at a dose of 150 Gy and stated that the complete sterility did not occur in two consecutive generations at this dose (26 and 42% egg hatching). However, they considered the dose of 150 Gy to be suitable for inherited sterility. Zhou et al. [16] reported 300 Gy gamma ray of C\u003csup\u003e137\u003c/sup\u003e as relative sterilizing dose for males and complete sterilizing dose for females and they selected this dose for inherited sterility. They showed that the number of eggs in the combinations; F\u003csub\u003e1\u003c/sub\u003eM\u0026times;UF and UM\u0026times;F\u003csub\u003e1\u003c/sub\u003eF was reduced by 84.16 and 85.02% compared to the control, and 0 and 1.85% hatching rate were recorded, respectively. This suggests that males are more likely to inherit sterility than females. These results are similar with results of the current study, in which irradiated males at a dose of 250 Gy caused 27.69% of eggs to hatch in the P\u003csub\u003e1\u003c/sub\u003e generation and females were completely sterile at this dose, therefore this dose was selected for IS assessment. In the next generation, adult insects obtained from surviving eggs were crossed as F\u003csub\u003e1\u003c/sub\u003eM\u0026times;UF and UM\u0026times;F\u003csub\u003e1\u003c/sub\u003eF, and complete sterility was occurred (100 and 98.57%, respectively).\u003c/p\u003e \u003cp\u003eOne notable advantage of inherited sterility in lepidopterans is the alteration of the F\u003csub\u003e1\u003c/sub\u003e sex ratio favoring male offspring, attributed to their WZ/ZZ (male/female) chromosomal sex determination system. This shift is the result of recessive lethal mutations induced in the Z sex chromosomes of the treated parents, which are responsible for the death of F\u003csub\u003e1\u003c/sub\u003e females [2]. Zhou et al. [16] investigated the effect of C\u003csup\u003e137\u003c/sup\u003e gamma ray (100 to 400 Gy) on the sex ratio of \u003cem\u003eT. absoluta\u003c/em\u003e F\u003csub\u003e1\u003c/sub\u003e generation. They claimed that with increasing radiation dose of parent males, the sex ratio of their offspring shifted more towards males. Yusef et al. [18] also showed that radiation treatment of Co\u003csup\u003e60\u003c/sup\u003e gamma ray causes a greater male bias in F\u003csub\u003e1\u003c/sub\u003e and F\u003csub\u003e2\u003c/sub\u003e generations of \u003cem\u003eT. absoluta\u003c/em\u003e compared to the control, which is similar to the results of the present study. Nevertheless, Cagnotti et al. [13] reported that different doses of X-rays on male pupae exhibited no significant impact on the sex ratio of their F\u003csub\u003e1\u003c/sub\u003e generation and reported an overall male ratio of 66%. The observed variability in the outcomes of these researches can be attributed to the utilization of different radiation sources (X-rays versus gamma rays).\u003c/p\u003e \u003cp\u003eSince longer exposure time in the higher doses may induce adverse effects on biological parameters and cause physiological defects on insects that decrease the competition ability of sterile moths, it is essential to ensure that the used dose rate for radiation does not have any negative effects [2, 26]. Arthur [12] elucidated that upon exposure of \u003cem\u003eT. absoluta\u003c/em\u003e pupae to Co\u003csup\u003e60\u003c/sup\u003e gamma radiation doses of 100 and 200 Gy, the adult emergence was quantified at 60 and 52%, respectively. These findings align with the outcomes observed in the current study. However, they reported 300 Gy completely inhibited the emergence of adults, but the current study recorded an emergence rate of 43 to 46% at the 300 Gy treatments. Cagnotti et al. [13] stated that with increasing X-ray dose, the rate of adult emergence decreased, females that were irradiated with 250 Gy during the pupal stage did not fully emerge, and the number of males that emerged was very low. Additionally, they reported external abnormalities such as deformed wings and bent legs at doses higher than 350 Gy. Nevertheless, these researchers did not document any malformations at radiation doses ranging from 0 to 300 Gy, which is contradictory to the findings of our study. In this investigation, the percentage of malformed insects was recorded between 15 and 29% at doses between 0 and 300 Gy. Yusef et al. [18] also stated that the emergence of adults decreases with increasing dose and they did not consider doses higher than 200 Gy Co\u003csup\u003e60\u003c/sup\u003e gamma ray suitable for mating because they observed deformation of wings and bent legs at these doses. Kuyulu and Gen\u0026ccedil; [14] showed that the adult\u0026rsquo;s emergence was 67, 42, 17 and 8% at doses of 100, 200, 300 and 400 Gy Co\u003csup\u003e60\u003c/sup\u003e gamma ray, respectively, and the emerged adults were deformed at 400 Gy. In contrast, in the present study, 43 to 57% emerged adults were observed at doses between 100 and 300 Gy. Sridhar et al. [15] reported the adult\u0026rsquo;s emergence was 31% at 300 Gy. Zhou et al. [16] stated that C\u003csup\u003e137\u003c/sup\u003e gamma radiation did not have a negative effect on the emergence of adult males and females, but males irradiated at 400 Gy were four times more deformed than controls. In our results, the rate of adult\u0026rsquo;s emergence and deformation did not increase significantly with increasing dose.\u003c/p\u003e \u003cp\u003eCagnotti et al. [13] showed that the longevity of \u003cem\u003eT. absoluta\u003c/em\u003e males and females\u0026rsquo; adults under X-rays treatment is not affected and recorded their average lifespan as 17 and 12 days respectively. Sridhar et al. [15] reported the lifespan of adults as 3.17 days at 300 Gy treatment of five-days old male pupae and 7.65 day in control. Zhou et al. [16] stated that males irradiated with 400 Gy C\u003csup\u003e137\u003c/sup\u003e gamma rays had a 16-day shorter lifespan than control, but at doses of 100 and 200 Gy, there was no significant difference. In the present study, the average longevity of adults was recorded as about 13 to 14 days and showed no significant difference under the influence of radiation compared to the control. These differing results depend on the various types of radiation source and the radiosensitivity of different geographic populations in different regions [26].\u003c/p\u003e \u003cp\u003eWhen SIT and IS are applied for insect control in the field and greenhouse, the mating competition ability of the released sterile males has great importance in the method\u0026rsquo;s success [26]. Zhou et al. [16] irradiated \u003cem\u003eT. absoluta\u003c/em\u003e pupae with optimal dose of 300 Gy of C\u003csup\u003e137\u003c/sup\u003e gamma ray and released emerged adults in different treatments. They recorded egg hatching rate of 96% at the 0:1:1 (IM: UM: UF) ratio, 38% for the 1:0:1 ratio, and 66% for the 1:1:1 ratio in the competitiveness assessment. Subsequently they calculated a competitiveness value (CV) of 1.07, signifying that irradiated male maintain competitive with wild males during mating at 1:1:1 ratio. In the current research, when pupae irradiated with 250 Gy of Co\u003csup\u003e60\u003c/sup\u003e gamma ray and emerged adults released in different ratios, egg hatching rate was recorded 78, 22 and 37% at the 0:1:1, 1:0:1, and 1:1:1 ratios, respectively. Also, the competitiveness value was calculated 2.6, 3.92, 4.57 and 5.32 in 1:1:1, 2:1:1, 3:1:1 and 4:1:1 ratios, respectively. In all ratios, the competition index was higher than 1 and increased by increasing mating ratio, reflecting that these sterile males are more competitive than those of wild males for mating with wild females. Consequently, the initial ratio of 1:1:1 is recommended for release in controlled laboratory conditions.\u003c/p\u003e"},{"header":"Conclusion","content":"\u003cp\u003eIn conclusion, a dose of 250 Gy Co\u003csup\u003e60\u003c/sup\u003e gamma ray can induce complete sterility of P\u003csub\u003e1\u003c/sub\u003e females and partial sterility of P\u003csub\u003e1\u003c/sub\u003e males, as well as complete sterility in the male progeny (F\u003csub\u003e1\u003c/sub\u003e) of \u003cem\u003eT. absoluta\u003c/em\u003e. A release ratio of 1:1:1 (IM: UM: UF) was sufficient for treated males to compete with normal males. Moreover, the irradiation of pupae at 250 Gy had no adverse effects on adult emergence and longevity. In view of these results, the use of partial sterility of P\u003csub\u003e1\u003c/sub\u003e male moths alongside their inherited sterility in F\u003csub\u003e1\u003c/sub\u003e progeny achieved through gamma-irradiation of 250 Gy Co\u003csup\u003e60\u003c/sup\u003e is recommended for the control of \u003cem\u003eT. absoluta\u003c/em\u003e population in Iran.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec10\" class=\"Section2\"\u003e \u003ch2\u003eInsect rearing\u003c/h2\u003e \u003cp\u003eTomato leaves infested with \u003cem\u003eT. absoluta\u003c/em\u003e were collected from our research greenhouse, and the larvae and pupae were maintained until the adult\u0026rsquo;s emergence. The emerged adults were transferred to egg-laying containers equipped with a cotton wick saturated with a 15% sugar solution. Crepe paper served as the substrate for egg deposition. The laid eggs were collected daily and transferred to the larval rearing containers (8 cm in diameter and 6 cm in height) which have a hole for ventilation and covered with a non-porous cloth. For rearing of larvae, the semi-artificial diet introduced by Greene et al. [28] was employed with some modifications derived from the diet proposed by Mihsfeldt and Parra [29]. This specific artificial diet was applied in a thin layer along the inner walls of the containers. The larval rearing containers were cleaned daily and new pieces of diet was added. The pupae formed between diet tissue were removed daily and kept in glass Petri dishes (100 mm \u0026times; 15 mm) containing wet filter paper. Insects were reared at a temperature of 25\u0026thinsp;\u0026plusmn;\u0026thinsp;1\u0026deg;C, a relative humidity of 40\u0026thinsp;\u0026plusmn;\u0026thinsp;5%, and a photoperiod of 16:8 (L:D) hours.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eThe effect of gamma radiation on P\u003csub\u003e1\u003c/sub\u003e and F\u003csub\u003e1\u003c/sub\u003e generations\u003c/h2\u003e \u003cp\u003eThe pupae were collected daily from rearing containers and sexed under stereomicroscope based on the morphology of the last abdominal segment [30]. The six-days old pupae (48 to 72 hours prior to emergence) were subjected to gamma radiation of 100, 150, 200, 250, and 300 Gy. For irradiation, an Issledovatle PX30 gamma cell with a CO\u003csup\u003e60\u003c/sup\u003e source with a total activity of 200 Curies was used at a dose rate of 118 Gy/hr (~\u0026thinsp;0.033 Gy/sec). This process was performed in four replicates. Subsequently, the percentage of emerged adults was calculated. The percentage of non-emerged pupae and deformed adults with non-flying wings were also recorded. Three emerged males and females were paired in egg-laying containers at a ratio of 1:1 across four replicates, in the following four combinations:\u003c/p\u003e \u003cp\u003eUnirradiated Female (UF) \u0026times; Unirradiated Male (UM)\u003c/p\u003e \u003cp\u003eUnirradiated Female (UF) \u0026times; Irradiated Male (IM)\u003c/p\u003e \u003cp\u003eIrradiated Female (IF) \u0026times; Unirradiated Male (UM)\u003c/p\u003e \u003cp\u003eIrradiated Female (IF) \u0026times; Irradiated Male (IM)\u003c/p\u003e \u003cp\u003eTo determine the appropriate sterility dose for males and females, the number of eggs laid and the percentage of eggs hatching were recorded and compared with the control. The sterility index was determined in accordance with Eq.\u0026nbsp;(1) [31];\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:\\%\\:Sterility\\:=100-(\\frac{a\\times\\:b}{A\\times\\:B}\\times\\:100)$$\u003c/div\u003e\u003c/div\u003e(1) \u003c/p\u003e \u003cp\u003eWhere:\u003c/p\u003e \u003cp\u003ea: number of eggs in treatment,\u003c/p\u003e \u003cp\u003eb: % hatching eggs in treatment,\u003c/p\u003e \u003cp\u003eA: number of eggs in control,\u003c/p\u003e \u003cp\u003eand B: % hatching eggs in control.\u003c/p\u003e \u003cp\u003eThe males and female adults\u0026rsquo; longevity was recorded until the death of the last moth. To investigate inherited sterility, the male pupae were subjected to further irradiation at a dose of 250 Gy, based on the obtained results. Emerged male adults were mated with normal females, and control mating were concurrently prepared with normal pairs. Their offspring was reared on the artificial diet. The sex ratio of F\u003csub\u003e1\u003c/sub\u003e pupae in the control and treatment was recorded. F\u003csub\u003e1\u003c/sub\u003e adult males and females obtained from this IM\u0026times;UF combination, were re-mated with pairs derived from the control combination UM\u0026times;UF. Three pairs of F\u003csub\u003e1\u003c/sub\u003e adult males and females were crossed as F\u003csub\u003e1\u003c/sub\u003eM\u0026times;UF, UM\u0026times;F\u003csub\u003e1\u003c/sub\u003eF, and UM\u0026times;UF as control in four replicates. The number of eggs, the percentage of egg hatching, and sterility rate were calculated.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eThe effect of gamma radiation on male mating competitiveness\u003c/h2\u003e \u003cp\u003eThe mating competitiveness ability of irradiated males compared to normal males was calculated under controlled laboratory conditions. According to the obtained results, 250 Gy was applied on male pupae. The irradiated and unirradiated emerged male adults (IM and UM) were mated with unirradiated females (UF) at different mating ratios of 1:1:1, 2:1:1, 3:1:1 and 4:1:1 (IM: UM: UM). The ratios of 0:1:1 and 1:0:1 (IM: UM: UM) were considered as controls. The number 1 in all ratios is three individuals. The number of eggs and egg hatch were counted daily. The observed egg hatch percentage and the expected egg hatch percentage at each ratio were compared using the chi-square (X\u003csup\u003e2\u003c/sup\u003e) test. The expected egg-hatch percentage (Ee) were calculated with Eq.\u0026nbsp;(2) [24];\u003cdiv id=\"Equb\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equb\" name=\"EquationSource\"\u003e\n$$\\:Ee=\\frac{N\\left(Ha\\right)+S\\left(Hs\\right)}{N+S}$$\u003c/div\u003e\u003c/div\u003e(2) \u003c/p\u003e \u003cp\u003eWhere:\u003c/p\u003e \u003cp\u003eEe: Expected egg-hatch percentage,\u003c/p\u003e \u003cp\u003eHa: Hatched eggs percentage in 0:1:1 (IM: UM: UM) combination,\u003c/p\u003e \u003cp\u003eHs: Hatched eggs percentage in 1:0:1 (IM: UM: UM) combination,\u003c/p\u003e \u003cp\u003eN: The number of normal males,\u003c/p\u003e \u003cp\u003eS: The number of sterile males.\u003c/p\u003e \u003cp\u003eThe competitiveness values (CV) of the irradiated males based on the observed egg hatched (Eo) were evaluated according to Eq.\u0026nbsp;(3) [24];\u003cdiv id=\"Equc\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equc\" name=\"EquationSource\"\u003e\n$$\\:\\:CV=\\frac{Ha-Eo}{Eo-Hs}\u0026divide;\\frac{S}{N}$$\u003c/div\u003e\u003c/div\u003e(3) \u003c/p\u003e \u003cp\u003eThis value when equal to 1, indicates a comparable level of competition between irradiated and unirradiated males. Values close to zero indicate superior competitiveness of unirradiated males [24].\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eThe biological and reproductive parameters were evaluated using one-way analysis of variance. A completely randomized design incorporating four replications was implemented. Mean values were compared using Duncan's multiple range test. The independent T-test was employed to compare means of F\u003csub\u003e1\u003c/sub\u003e sex ratio. Data were analyzed using SPSS software version 22 (IBM \u0026copy;).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec14\" class=\"Section2\"\u003e \u003ch2\u003eData Availability\u003c/h2\u003e \u003cp\u003eAll data supporting the findings of this study are available within the paper. The datasets generated during the current study are available from the corresponding author on reasonable request.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAcknowledgements\u003c/h2\u003e\n\u003cp\u003eI would like to thank Mr. Siyavash Farhadi and Dr. Saeid Moori for their technical help in the insects rearing and Nuclear Science and Technology Research Institute, Atomic Energy Organization of Iran (AEOI).\u003c/p\u003e\n\u003ch2\u003eCompeting interests\u003c/h2\u003e\n\u003cp\u003eThere is no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003c/p\u003e\n\u003ch2\u003eFunding declaration\u003c/h2\u003e\n\u003cp\u003eNo specific funding has been provided for this research.\u003c/p\u003e\u003ch2\u003eAuthor Contribution\u003c/h2\u003e\u003cp\u003eS. A. is responsible for all matters related to conducting experiments and writing the article.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eSimmons, G. S., Bloem, K. A., Bloem, S., Carpenter, J. E. \u0026amp; Suckling, D. M. Impact of moth suppression/eradication programmes using the sterile insect technique or inherited sterility. In \u003cem\u003eSterile insect technique. Principles and practice in area-wide integrated pest management\u003c/em\u003e 1007-1050 (Dordrecht: Springer Netherlands, 2021).\u003c/li\u003e\n\u003cli\u003eMarec, F. \u0026amp; Vreysen, M. J. Advances and challenges of using the sterile insect technique for the management of pest lepidoptera. \u003cem\u003eInsects\u003c/em\u003e \u003cstrong\u003e10(11),\u003c/strong\u003e 371 (2019).\u003c/li\u003e\n\u003cli\u003eDyck, V. A., Hendrichs, J. \u0026amp; Robinson, A. 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Zool.\u003c/em\u003e \u003cstrong\u003e40(5),\u003c/strong\u003e 801-805 (2016).\u003c/li\u003e\n\u003cli\u003eToppazada, A., Abdallah, S. \u0026amp; Eldefrawi, M. E. Chemosterilization of larvae and adults of the Egyptian cotton leaf worm, \u003cem\u003eProdenia litura\u003c/em\u003e by Apholate, Metepa and Tepa. \u003cem\u003eJ. Econ. Entomol. \u003c/em\u003e\u003cstrong\u003e59,\u003c/strong\u003e 1125-1128 (1966).\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1.\u0026nbsp;\u003c/strong\u003eFecundity, fertility and sterility (Mean \u0026plusmn; SE) of \u003cem\u003eTuta absoluta\u003c/em\u003e parental generation that emerged from pupae irradiated a with different gamma ray doses (Gy). U: unirradiated, I: irradiated, M: male and F: female. Means indicated by different letters in each separate columns are significantly different according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.01).\u003c/em\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"624\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 90px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCombination\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eDose (Gy)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eEgg number \u0026plusmn; SE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eEgg hatch (%) \u0026plusmn; SE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSterility (%) \u0026plusmn; SE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"6\" valign=\"top\" style=\"width: 90px;\"\u003e\n \u003cp\u003eIM\u0026times;UF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e98.25\u0026plusmn;12.60 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e80.17\u0026plusmn;4.45 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e47.5\u0026plusmn;11.93 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e42.85\u0026plusmn;4.21 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e73.34\u0026plusmn;7.31 c\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e150\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e34.25\u0026plusmn;12.68 bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e38.13\u0026plusmn;12.18 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e87.62\u0026plusmn;3.45 bc\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e37.50\u0026plusmn;11.85 bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e28.26\u0026plusmn;3.95 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e86.03\u0026plusmn;4.43 bc\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e250\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e17.75\u0026plusmn;3.40 bc\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e27.69\u0026plusmn; 4.47 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e93.33\u0026plusmn;2.16 ab\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e13.75\u0026plusmn;2.17 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"6\" valign=\"top\" style=\"width: 90px;\"\u003e\n \u003cp\u003eUM\u0026times;IF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e98.25\u0026plusmn;12.60 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e80.17\u0026plusmn;4.45 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e25.00\u0026plusmn;9.65 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e35.30\u0026plusmn;2.44 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e87.93\u0026plusmn;5.45 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e150\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e18.75\u0026plusmn;6.99 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e36.70\u0026plusmn;12.84 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e88.57\u0026plusmn;5.89 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e19.50\u0026plusmn;10.27 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e12.31\u0026plusmn;7.19 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e94.29\u0026plusmn;3.30 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e250\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e6.25\u0026plusmn;5.26 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e2.50\u0026plusmn;2.50 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd rowspan=\"6\" valign=\"top\" style=\"width: 90px;\"\u003e\n \u003cp\u003eIM\u0026times;IF\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e0\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e98.25\u0026plusmn;12.60 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e80.17\u0026plusmn;4.45 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e25.25\u0026plusmn;14.69 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e14.42\u0026plusmn;9.76 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e91.11\u0026plusmn;5.45 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e150\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e8.5\u0026plusmn;5.68 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e9.25\u0026plusmn;5.45 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e250\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e1.75\u0026plusmn;1.43 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 144px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 156px;\"\u003e\n \u003cp\u003e0.00\u0026plusmn;0.00 c\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 138px;\"\u003e\n \u003cp\u003e100\u0026plusmn;0.00 a\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003cbr\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003e\u0026nbsp;\u003cstrong\u003eTable 2\u003c/strong\u003e\u003cstrong\u003e\u003cspan dir=\"RTL\"\u003e.\u003c/span\u003e\u003c/strong\u003e F\u003csub\u003e1\u0026nbsp;\u003c/sub\u003efecundity, fertility and sterility percentage of \u003cem\u003eTuta absoluta\u003c/em\u003e resulting from irradiation of P\u003csub\u003e1\u003c/sub\u003e male pupae with 250 Gy gamma ray. U: unirradiated, M: male and F: female. Means indicated by different letters in each separate row are significantly different according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.01)\u003c/em\u003e.\u003c/em\u003e\u003c/p\u003e\n\u003cdiv align=\"\"\u003e\n \u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cstrong\u003e\u003cspan dir=\"LTR\"\u003eCombination\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cstrong\u003e\u003cspan dir=\"LTR\"\u003eEgg number \u0026plusmn; SE\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 134px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cstrong\u003e\u003cspan dir=\"LTR\"\u003eEgg hatch (%) \u0026plusmn; SE\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cstrong\u003e\u003cspan dir=\"LTR\"\u003eSterility (%) \u0026plusmn; SE\u003c/span\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 89px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003eUM\u0026times;UF\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e9\u003c/span\u003e\u003cspan dir=\"LTR\"\u003e5.5\u0026plusmn;3.66\u0026nbsp;\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 134px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e76.32 \u0026plusmn;3\u003c/span\u003e\u003cspan dir=\"LTR\"\u003e.31 a\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e\u0026nbsp;\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003eF\u003csub\u003e1\u003c/sub\u003eM\u0026times;UF\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e1\u003c/span\u003e\u003cspan dir=\"LTR\"\u003e8.5\u0026plusmn;3.12 b\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 134px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e0.00\u0026plusmn; 0.00 b\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e100.00\u0026plusmn;0.00\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003eUM\u0026times;F\u003csub\u003e1\u003c/sub\u003eF\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e1\u003c/span\u003e\u003cspan dir=\"LTR\"\u003e6.25\u0026plusmn;2.39 b\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 134px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e5.44\u0026plusmn;1.95 b\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 116px;\"\u003e\n \u003cp dir=\"RTL\"\u003e\u003cspan dir=\"LTR\"\u003e98.57\u0026plusmn;0.78\u003c/span\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003e\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u0026nbsp;\u003cstrong\u003eTable 3\u003c/strong\u003e\u003cstrong\u003e.\u0026nbsp;\u003c/strong\u003eNumber of eggs, observed and expected egg hatching percentages, and competitiveness value of \u003cem\u003eTuta absoluta\u003c/em\u003e resulting from different release ratios of irradiated males by 250 Gy gamma ray alongside unirradiated males and females. U: unirradiated, I: irradiated, M: male and F: female. Means indicated by different letters in each column are significantly different according to Duncan\u0026apos;s multiple range test \u003cem\u003e(P\u0026lt;0.01)\u003c/em\u003e.\u003c/p\u003e\n\u003cdiv align=\"\"\u003e\n \u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eRatio\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e(IM: UM: UF)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eEgg number\u0026plusmn; SE\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eObserved egg hatch %\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eExpected egg hatch %\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eX\u003csup\u003e2\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u003csup\u003edf=3\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003cp\u003e\u003cstrong\u003e\u003cem\u003eP\u003c/em\u003e\u003c/strong\u003e\u003cem\u003e˂\u003cstrong\u003e0.05\u003c/strong\u003e\u003c/em\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eCV\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e0:1:1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e102.00\u0026plusmn;17.01 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e78.16\u0026plusmn;5.62 a\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e1:1:1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e23.67\u0026plusmn;9.94 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e37.17\u0026plusmn;16.15 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e50.17\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e43.14\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e2.60\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e2:1:1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e21.00\u0026plusmn;2.31 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e28.53\u0026plusmn;7.12 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e40.85\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e17.40\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e3.92\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e3:1:1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e59.00\u0026plusmn;12.85 ab\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e26.00\u0026plusmn;8.24 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e36.18\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e17.76\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e4.57\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e4:1:1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e38.33\u0026plusmn;2.848 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e24.71\u0026plusmn;3.40 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e33.38\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e10.60\u003csup\u003e*\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e5.32\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 108px;\"\u003e\n \u003cp\u003e1:0:1\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 114px;\"\u003e\n \u003cp\u003e45.33\u0026plusmn;30.90 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 96px;\"\u003e\n \u003cp\u003e22.19\u0026plusmn;2.3 b\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 89px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 55px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 44px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n \u003c/table\u003e\n\u003c/div\u003e\n\u003cp\u003eX\u003csup\u003e2\u003c/sup\u003e values marked with asterisk (*) indicate a significant difference between the observed and expected percentage of egg hatching \u003cem\u003e(P˂0.05)\u003c/em\u003e.\u003c/p\u003e\n\u003cp\u003eCV: Competitiveness value [24].\u003c/p\u003e\n\u003cp\u003eThe number 1 in all ratios is equivalent to three insects\u003c/p\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"
[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true},"keywords":"Biological parameters, Co60 gamma ray, competition index, inherited sterility, sterile insect technique, tomato leaf miner","lastPublishedDoi":"10.21203/rs.3.rs-6690207/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6690207/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThe objective of this research is to stablish standardized sterile and sub-sterile doses of Co\u003csup\u003e60\u003c/sup\u003e gamma radiation on \u003cem\u003eTuta absoluta\u003c/em\u003e (Meyrick) to facilitate the implementation of sterile insect technique for management of this invasive pest of tomato and solanaceous plants. Insects were reared on an artificial diet and pupae were irradiated at 50, 100, 150, 250 and 300 Gy. The fecundity and fertility of untreated females paired with irradiated males, and treated females mated with untreated or treated males, significantly diminished with increasing gamma doses. When pupae were exposed to irradiation at 250 and 300 Gy, completely sterile females and males were obtained, respectively. The adult\u0026rsquo;s emergence and longevity, were not significantly influenced by irradiation. To perform inherited sterility, male pupae were irradiated by 250 Gy. The F\u003csub\u003e1\u003c/sub\u003e sex ratio shifted in favor of males. The F\u003csub\u003e1\u003c/sub\u003e fecundity was diminished compared to the control, and the eggs produced were more sterile. Ultimately, the mating competitiveness of irradiated males by 250 Gy against normal males was assessed at different ratios. Based on the calculated competitiveness value, the release ratio was determined 1:1:1 (irradiated males: unirradiated males: unirradiated females), in which males subjected to irradiation are capable of competing with unirradiated males.\u003c/p\u003e","manuscriptTitle":"Optimization of Co60 gamma radiation dose for applying sterile insect technique and inherited sterility on Tuta absoluta (Meyrick) in Iran","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-06-19 13:42:40","doi":"10.21203/rs.3.rs-6690207/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2025-10-10T11:27:03+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-07T20:31:31+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"33477633572616707243303595007544951966","date":"2025-10-06T07:07:44+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"69963525117211247109751278644728607197","date":"2025-10-05T15:36:04+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-10-02T23:01:29+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"141629074960755434698261424410196948254","date":"2025-09-30T16:27:09+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"223770109040244586646583099741591432795","date":"2025-09-29T13:42:16+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"208878550447229708955841532320333793577","date":"2025-07-21T12:10:45+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2025-06-29T04:47:34+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"322069385886457610185584012186057610755","date":"2025-06-17T14:11:48+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"13464027816928161869754772580063627232","date":"2025-06-16T01:30:33+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-06-15T12:13:54+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-06-15T11:57:56+00:00","index":"","fulltext":""},{"type":"editorInvited","content":"","date":"2025-06-06T03:20:13+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-05-22T13:22:30+00:00","index":"","fulltext":""},{"type":"submitted","content":"Scientific Reports","date":"2025-05-22T13:21:24+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"
[email protected]","identity":"scientific-reports","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"scirep","sideBox":"Learn more about [Scientific Reports](http://www.nature.com/srep/)","snPcode":"","submissionUrl":"","title":"Scientific Reports","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"stoa","reportingPortfolio":"Scientific Reports","inReviewEnabled":true,"inReviewRevisionsEnabled":true}}],"origin":"","ownerIdentity":"8be0d076-f2d6-475c-92f3-a5a829a546fb","owner":[],"postedDate":"June 19th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[{"id":50301734,"name":"Biological sciences/Zoology/Entomology"},{"id":50301735,"name":"Biological sciences/Biological techniques"}],"tags":[],"updatedAt":"2025-12-15T16:05:58+00:00","versionOfRecord":{"articleIdentity":"rs-6690207","link":"https://doi.org/10.1038/s41598-025-32383-2","journal":{"identity":"scientific-reports","isVorOnly":false,"title":"Scientific Reports"},"publishedOn":"2025-12-13 15:59:43","publishedOnDateReadable":"December 13th, 2025"},"versionCreatedAt":"2025-06-19 13:42:40","video":"","vorDoi":"10.1038/s41598-025-32383-2","vorDoiUrl":"https://doi.org/10.1038/s41598-025-32383-2","workflowStages":[]},"version":"v1","identity":"rs-6690207","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6690207","identity":"rs-6690207","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}
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