Stability and realized heritability of resistance to imidacloprid in the field- collected brown planthopper Nilaparvata lugens (Hemiptera: Delphacidae) from Lampung, Indonesia

Stability and realized heritability of resistance to imidacloprid in the field- collected brown planthopper Nilaparvata lugens (Hemiptera: Delphacidae) from Lampung, Indonesia | 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 Research Article Stability and realized heritability of resistance to imidacloprid in the field- collected brown planthopper Nilaparvata lugens (Hemiptera: Delphacidae) from Lampung, Indonesia Danarsi Diptaningsari, Aziz Purwantoro, Arman Wijonarko, Y. Andi Trisyono This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-7709102/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 23 Feb, 2026 Read the published version in International Journal of Tropical Insect Science → Version 1 posted 13 You are reading this latest preprint version Abstract Availability of rice throughout the year in the Indonesian field makes the brown planthopper, ( Nilaparvata lugens ) (Stål) an important pest. Imidacloprid is an insecticide active ingredient that is commonly used for managing N. lugens populations. The data on resistance stability and realized heritability would be effective to derive a viable resistance management system against N. lugens . The resistance ratio (RR) increased after selection for five generations, from 52.36 to 180.46-fold in N. lugens collected from rice fields in the District of Central Lampung, Lampung Province, Indonesia. In order to determine the stability of resistance, five additional generations (totaling ten generations) were reared without insecticides exposure. Stability experiments revealed that resistance to imidacloprid was unstable and there was a loss of resistance from 180.46 to 39.46-fold over ten generation without any selections (DR value = -0.07). On the contrary, the resulting resistance level after continues selections with imidacloprid during the subsequent three generations might be up the resistance level from 180.46 to 235.93-fold. The realized heritability values ( h 2 ) of imidacloprid resistance were low at 0.1465 and 0.1348 in five and eight generations that were selected, respectively. This unstable resistance could be mitigated by temporarily removing selection pressure or by rotating to insecticides with different mode of actions. Having a low heritability of resistance, the sustainable wide-range resistance management plan will be required in controlling the N. lugens population in Indonesia. imidacloprid Nilaparvata lugens realized heritability resistance stability Figures Figure 1 Introduction Rice cultivation in Indonesia faces a potentially serious challenges from the brown planthopper Nilaparvata lugens Stål (Hemiptera: Delphacidae) due to the rice availability under field conditions throughout the year. The density of N. lugens in Indonesia spreads over Java, Bali, Lampung, South Sumatra, Central Sulawesi, West Kalimantan, and South Kalimantan (BB Padi, 2020). In Indonesia, the strategies used in managing N. lugens in the rice plant rely mainly on natural enemies’ conservation, resistant variety and insecticide formulations. Imidacloprid is one of the active ingredients of insecticide that has been widely used for controlling N. lugens in Indonesia since 1992 (Ditjen PSP, 2017). Overuse of the insecticides will cause insecticide resistance to develop. Imidacloprid resistance in N. lugens populations has been documented in China and Korea with a resistance ratio of 57.0 to 2029.0-fold (Bao et al., 2016 ; Zhang et al., 2016 ) and 13.0 to 237.0-fold (Min et al., 2014 ), respectively. Nilaparvata lugens populations from Vietnam and Thailand showed resistance development to imidacloprid, with the resistance ratio of 89.0 and 102.9-fold, respectively (Matsumura & Sanada-Morimura, 2010 ; Bao et al., 2016 ). Since 2010 to 2019, imidacloprid resistances of the field populations of N. lugens had been occured in the the West Java Province (13.5 to 108.1-fold) (Surahmat et al., 2016; Iswanto et al., 2019 ) and the Central Java Province (25.9 to 29.4-fold) (Londingkene et al., 2016a ; Diptaningsari et al., 2019 ). The management of resistance to insecticide requires a systematic and comprehensive strategy. An understanding of the stability and heritability of the insecticide resistance is crucial for designing insecticide resistance management (IRM) strategies to anticipate and decelerate the development of resistance. The stability of resistance plays an important role to determine an effective pest management strategy (Roush et al., 1990; Lan et al., 2001). The resistance stability was determined by rearing the selected insects in the absence of insecticides across multiple generations. The study by Ullah and Shad ( 2017 ) revealed that resistance to chlorfenapyr and triazophos in Oxycarenus hyalinipennis (Hemiptera: Lygaeidae) was unstable, on the one hand, the resistance to emamectin benzoate and fipronil was stable. The resistance in Bemisia tabaci to acetamiprid remained in the absence of selection pressure (Basit et al., 2011 ). In the house flies ( Musca domestica ) and N. lugens , the resistance to imidacloprid was not stable and dropped throughout several months (Kavi et al., 2014 , Diptaningsari et al., 2020 ). Another parameter that needs to be estimated is the realized heritability ( h 2 ) in predicting how resistance can be developed because of the use of insecticides (Tabashnik, 1992 ). The h 2 value in imidacloprid resistance was observed in N. lugens (Wang et al., 2009 , Diptaningari et al., 2019), Spodoptera litura (Abbas et al., 2012 ) and house fly ( M. domestica ) (Khan et al., 2014 ). The research results provided in this article will be of immense benefits as far as stability and heritability of resistance to imidacloprid in field population of N. lugens in Indonesia is concerned that can be potentially used in controlling N. lugens through sustainable measures. Materials and Methods Insects In the study, two groups of N. lugens were used. The resistant population was the first one, which was taken in 2018 from the rice fields (Ciherang variety) in the Village of Notoharjo (5°08'58.2"S 105°14'13.3"E), the Sub-District of Trimurjo, the District of Central Lampung, Lampung Province, Indonesia. Nilaparvata lugens (G 0 ) were collected from ten sampling points in rice fields (±2,000 adults). The insecticides used at the collection sites were BPMC, dimehipo, imidacloprid, pymetrozine, and buprofezine. The susceptible population which was obtained in 1986 was the second population that was collected from the District of Sleman, Special Region of Yogyakarta. This population was reared under laboratory conditions, kept apart with other populations, without supplementation from field population and no insecticides exposure. These populations were maintained at ±27 °C and 70–80% RH. Rice seedlings (5-7 days) were prepared in a plastic container (30 x 24 cm) in which a number of approximately 100 pairs were then placed to oviposit. The nymphs were moved with an aspirator device into a new plastic container (containing new rice plants) after these eggs had hatched and the host rice plants began to yellow. The nymphs transferred into the new rice plants, and the plants were changed every 4 days. In 5-7 days of rice plants, the newly emerged adults were reared to the next generations. Insecticide The bioassays and selections were conducted in N. lugens using a commercial 5% imidacloprid formulation (Confidor 5 WP, produced by PT. Bayer Indonesia). Selection for resistance and toxicity bioassays For eight generations (G1–G8), selection with imidacloprid was carried out by exposing third-instar nymphs of N. lugens to treated rice seedlings. Seedlings of rice (5-7 days old) were dipped in the concentration of 100 ppm on first generation and the concentration was gradually increased at every generation on selections and it reached 600 ppm on seventh and eight generation. The seeds of rice were dipped in an insecticide solution for 15 seconds, followed by air-drying for 15 minutes at room temperature. The selected rice plants treated were subsequently kept in a plastic cup (10 x 11 cm) where water had been added to such an extent that the seedlings will remain fresh. Aspirating device was used to obtain the third instar nymphs, and 20 nymphs were added to each cup. The level of mortality was taken as 120 hours after treating in order to receive at least some survivors. The untreated rice seedlings were put in plastic cups where they were left with insects for further development. The nymphs were confirmed dead when no response was observed after gentle probing with a soft brush. The selections made in the laboratory was done in the same way as already explained and imidacloprid concentration varied in all generations (Table 1). Bioassays were performed on the third instars nymphs of the resistant and the susceptible populations. Preliminary bioassay was done to establish the concentration levels that attained 5-99% deaths with leaf dipping method. Seedlings of rice (5-7 days old) were immersed at insecticide solution and air-dried under room condition (15 minutes). The treated rice seedlings were then placed in a plastic cup (10 x 11 cm), containing sufficient water to make the seedlings appear fresh. The concentrations applied in the determination of resistance were based on the outcome of the preliminary bioassays whereby the concentrations ranged between 0-640 ppm (resistant population) and 0-80 ppm (susceptible population). Each of the bioassays involved a total of 495-660 nymphs, and 60-80 nymphs were used for EACH concentration. Mortality observations were conducted daily for up to 168 hours. The LC 50 values were determined using Probit analysis (Finney, 1971) with POLO-Plus. The Abbotts formula (Abbott, 1925) was applied to correct mortality data. The resistance ratio (RR) value is a comparison between the LC 50 of the resistant population and the LC 50 of the susceptible population. If the 95% confidence limits (CLs) did not overlap, the LC 50 values were considered not significantly different. Stability of resistance Imidacloprid resistance in N. lugens was stated stable based on the LC 50 results of the resistant population developed after five generations of selection (G1 to G5), and some of the insects in the population continued to be maintained for the next 10 generations (G6 to G15) without any exposure to any insecticide. The values of LC 50 were calculated per generation. The resistant change was measured as a difference in the degree of resistance to measure the DR (Decrease in Resistance) or the estimated response on selection (Falconer and Mackay, 1996) which can further be applied to obtain the average response per generation. The DR-value was estimated following Tabashnik (1994) as follows: DR = [log (final LC 50 ) - log (initial LC 50 )] / n Where final LC 50 is the value of LC 50 of generation n reared without selection (G 5 ) and the initial LC 50 is the value of LC 50 of generation n reared with selection (G 1 ) and n is the number of generations that is reared without selection (10 generations). An estimate was done using the reciprocal of DR or DR-1 values to provide the figure of number of generations that are required in order to decrease LC 50 by 10-fold. To evaluate the change of resistance, the imidacloprid selection was done with the part of the fifth generation of a resistant population and the LC 50 of the next three generations (G 6 to G 8 ) was calculated. Estimation of realized heritability Realized heritability ( h 2 ) of insect resistance was assessed following the method outlined by Tabashnik, 1992 using pre insecticide selection LC 50 values and post insecticide selection LC 50 values per generation of N. lugens . The formula was used to calculate the value of the realized heritability ( h 2 ): h 2 = R / S R represents the response to insecticide selection, while S is the selection differential. An estimate of response to selection (R) was estimated following Falconer and Mackay (1996) as follows: R = [log (final LC 50 ) - log (initial LC 50 )] / n Where the final LC 50 represented the LC 50 of the offspring after n generations of selection, whereas the initial LC 50 was the LC 50 of the parental generation preceding the selections, and n was the number of the selected generations with imidacloprid. The formula used in calculating differential selection (S) (Hartl, 1988) was: S = i dp Where i refers to selection Intensity, and dp is phenotypic standard deviation. The intensity of selection ( i ) was approximated on the p (mean percent survival post-selection). The equation to use in calculating the intensity of selection ( i ) (Tabashnik, 1992) was: i = 1.583 – 0.0193336p + 0.0000428p 2 + 3.65194 / p The phenotypic standard deviation ( dp ) was calculated using the formula described by Tabashnik (1992): dp = [1/2 (initial slope + final slope)] - 1 In which the initial slope denoted the slope of the probit regression lines of the parental generation prior to selection and final slope was the slope of the probit regression lines of the offspring after n generations of selection with imidacloprid. Results Selection for resistance and toxicity bioassays The laboratory selections (Table 1) increased the resistance ratio from 52.36-fold at G 1 to 180.46-fold at G 5 and 235.93-fold at G 8 , compared to the susceptible population (Table 2). Stability of resistance With selection, resistance ratio was increasing in five generations. The resistance ratio decreased rather fast during the first fifth generations post-removal of selection (G 6 -G 10 ). Sixth generation of non-selected (G 11 ) exhibited a lower resistance ratio of 53.30-fold which was almost similar to the actual first resistance rate before selection (52.36-fold). The resistance ratio remained largely unchanged but varied a bit in the seventh (G 12 ), eighth (G 13 ), ninth (G 14 ) and tenth generations (G 15 ) without selection. These findings showed that the resistance of imidacloprid in N. lugens of the Lampung population was not stable as the value of the DR was -0.07 (Table 3). The number of the estimated generations needed to decrease the LC 50 value 10-folds was 14.29 generations. The stabilities of some insects were measured after surviving the selection with imidacloprid over 3 generations (G 6 - G 8 ). Using subsequent selections, the resistance ratio of G 6 to G 8 rose by 206.25-fold (G 6 ) to 235.93-fold (G 8 ) in comparison with the susceptible population (Table 2 and Fig 1). Estimation of realized heritability Following five consecutive generations under imidacloprid selection, the LC 50 values of the field-collected N. lugens from Lampung rose incrementally between successive generation with a G 1 value of 29.32 ppm rising to G 5 at 101.06 ppm and the slope also rose between the generations with a G 1 slope of 1.36 to G 5 slope at 1.64. Following the eighth generation of selection, the values of LC 50 were higher (132.12 ppm, G 8 ), and the slope was higher too (1.69). These values led to the calculated value of the realized heritability ( h2 ) to be 0.1465 and 0.1348 in five and eight selected generations, respectively. Ten generations were required to elevated 10-fold LC 50 value of imidacloprid even after selection of five generations and eight generations (reciprocal of R value). (Table 4). Discussion The imidacloprid resistance can be viewed as a significant problem that has led to the case of N. lugens in the recent outbreak (Gao et al., 2014, Zhang et al., 2017). The imidacloprid has been registered since the year 1992, and N. lugens has been controlled in Indonesia using this chemical (Ditjen PSP, 2017). The over application and the high use of imidacloprid is also experienced in other states (Wang et al., 2007). Resistance to the imidacloprid emerged fast within five generations of the selection in this research. The study of Wang et al. (2009) reported a 200-fold to 1298-fold change of resistance of the N. lugens to imidacloprid by selection through 23 generations. According to Iswanto et al. (2019), in the N. lugens from Karawang province, West Java Province, Indonesia, an imidacloprid resistance increase by 13.5 (fold) to 33.9-fold was observed after five generations of selection. A new selection of N. lugens after three generations (G 6 - G 8 ) in the study has raised the ratio of resistance. Such an existing resistance to imidacloprid was attributed to the accelerated detoxification rate based on the upsurge of cytochrome P450 monooxygenase enzyme activity (Zhang et al., 2017, Wen et al., 2009, Puinean et al., 2010) as well as the mutation of the target site (Wen et al., 2009, Liu et al., 2005). The imidacloprid resistance of the N. lugens populations collected in South and East Asia was linked to the overexpression of two P450 monooxygenase genes, CYP6AY1 and CYP6ER1 (Bao et al., 2016, Garrood et al., 2015). In addition, the new research of resistance mechanism to the insecticides is related to epigenetic features of metabolic resistance, such as the histone modification and the DNA methylation (Bass et al., 2014, Oppold & Müller., 2017). The high resistance ratio had fallen dramatically during the first five generations when the selection was removed, which showed that the resistance to imidacloprid was not stable in N. lugens. It could be connected with the presence of the fitness cost in the population (Londingkene et al., 2016b). Variable influence of alleles due to the levels of dominance, the initial frequency of genes, and the relative fitness of different genes can be some of the factors that may contribute towards lessening the degree of resistance in the pest population (Afzal et al., 2015). Once the selection was changed, the ratio of resistance varied between G 11 and G 15 , but showed no considerable drop. This finding was an indication that homozygous resistance alleles existed prior to reduction of the selection pressure (Roush et al., 1990). The formulation of imidacloprid is mainly used through spraying that can lead to its existence in the irrigation system. In conjunction with the systemic and long-term residual activity, this insecticide might cause long-term selection pressure which might result in N. lugens resistant (Liu et al., 2003). In an experiment where soil was used conducted in a laboratory by Sarkar et al. (2001), dissipation of imidacloprid was reported to vary between 28.7 and 47.8 days. They will result in persistent pressure of resistance among the insects although the spraying was halted. The estimated realized heritability ( h 2 ) by this research were 0.1465 and 0.1348 by five and eight selected generations respectively, which indicates the N. lugens has weak capacity to acquire resistance towards imidacloprid (Falconer and Mackay, 1996). The resistance against imidacloprid has also been found comparable with N. lugens in China ( h 2 = 0.1141) as well as in the other insects, such as S. litura ( h 2 = 0.15) and M. Domestica ( h 2 = 0.09) (Abbas et al., 2012; Khan et al., 2014). One of the parameters is an estimation of the realized heritability in predicting persistence of insecticide effectiveness in relation to population-level pest management, but the laboratory experiment cannot elaborate on the parameter (Tabashnik and McGaughey, 1994; Ullah et al., 2016;). Environmental factors will impact on phenotypic variations, e.g. insecticide rotation, migration, natural enemies, and the rest of the environmental variants. The data on resistance stability, and realized heritability would be helpful to make a recommendation about the appropriate resistance management strategy in N. lugens. Nilaparvata lugens were found to be unstable in terms of their resistance to imidacloprid whereby they experienced a drop in the level of resistance upon recovery of the selection pressure. One of the strategies to counter the imidacloprid outbreak among N. lugens would be to use insecticides with dissimilar mode of actions and lacking in cross resistance positive-tests (Ullah and Shad, 2017). Also, supporting the use of Integrated Pest Management (IPM) in rice can help to minimize the risk of development and occurrence of N. lugens outbreaks in Indonesia. Declarations Acknowledgement The author wishes to thank Sriyanto Harjanto for his support in rearing the susceptible Nilaparvata lugens population. Author contributions D. D. and Y. A. T. contributed to the conceptualization of the study, data collection, data analysis, and wrote the original draft. A. P. and A. W. contributed to the conceptualization of the study, data analysis, statistical evaluation, and manuscript editing. All authors reviewed and approved the final version of the manuscript. Funding The study was partially funded by the Indonesian Agency for Agricultural Research and Development, the Ministry of Agriculture. Declaration of Competing Interest The authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper. References Abbas, N., Shad, S.A., Razaq, M., 2012. Fitness cost, cross resistance and realized heritability of resistance to imidacloprid in Spodoptera litura (Lepidoptera: Noctuidae). J. Pestic. Biochem. Physiol. 103, 181-188. Abbott, W.S., 1925. A method of computing the effectiveness of an insecticide. J. Econ. Entomol. 18, 256-267. Afzal, M.B., Abbas, N., Shad, S.A., 2015. Inheritance, realized heritability and biochemical mechanism of acetamiprid resistance in the cotton mealybug, Phenacoccus solenopsis Tinsley (Homoptera: Pseudococcidae). J. Pestic. Biochem. Physiol. 122, 44-49. Bao, H., Gao, H., Zhang, Y., Fan, D., Fang, J., Liu, Z., 2016. 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Dynamics of imidacloprid resistance and cross‐resistance in the brown planthopper, Nilaparvata lugens . Entomologia experimentalis et applicata. 131, 20-29. Wen, Y., Liu. Z., Bao, H., Han, Z., 2009. Imidacloprid resistance and its mechanisms in field populations of brown planthopper, Nilaparvata lugens Stål in China. J. Pestic. Biochem. Physiol. 94, 36-42. Zhang, J., Zhang, Y., Wang, Y., Yang, Y., Cang, X., Liu, Z., 2016. Expression induction of P450 genes by imidacloprid in Nilaparvata lugens : A genome-scale analysis. J. Pestic. Biochem. Physiol. 132, 59-64. Zhang, X., Liao, X., Mao, K., Yang, P., Li, D., Alia, E., Wan, H., 2017. The role of detoxifying enzymes in field-evolved resistance to nitenpyram in the brown planthopper Nilaparvata lugens in China. J. Crop Prot. 94, 106-114. Tables Table 1. History of Nilaparvata lugens selection s with imidacloprid in the laboratory Generation (G) Concentration (ppm) No. tested insects (n) Mortality (%) No. insects alive G 1 100 186 71.51 53 G 2 200 300 65.33 104 G 3 300 300 62.33 113 G 4 400 360 65.28 125 G 5 500 540 55.19 242 G 6 500 200 47.00 106 G 7 600 400 56.75 173 G 8 600 300 36.67 190 The population was originally collected from the District of Central Lampung, Lampung Province, Indonesia in 2018. Third instars were selected by exposing them into the treated rice seedlings. Table 2. Resistance level of Nilaparvata lugens collected from Lampung Province, Indonesia, after several laboratory selections with imidacloprid Population n Slope (±SE) LC 50 (95% CI) (ppm) χ 2 (df) RR a Susceptible 552 1.22 (±0.20) 0.56 (0.24 – 0.95) 5.71 (6) 1.00 Lampung b G 1 (S) 660 1.36 (±0.10) 29.32 (23.71 – 36.26) 1.53 (7) 52.36 G 2 (S) 495 1.20 (±0.12) 40.82 (28.47 – 56.27) 2.03 (7) 72.89 G 3 (S) 495 1.24 (±0.10) 55.71 (40.01 – 79.62) 9.27 (7) 99.48 G 4 (S) 495 1.23 (±0.16) 71.73 (47.06 – 101.73) 1.93 (7) 128.09 G 5 (S) 660 1.64 (±0.22) 101.06 (73.68 – 129.82) 6.86 (7) 180.46 G 6 (S) 660 1.49 (±0.20) 115.50 (84.30 -150.89) 4.35 (7) 206.25 G 6 (Us) 660 1.56 (±0.16) 86.40 (66.82 – 109.16) 1.27 (7) 154.29 G 7 (S) 660 1.58 (±0.21) 121.19 (89.13 – 156.81) 3.35 (7) 216.40 G 7 (Us) 660 1.15 (±0.08) 63.15 (49.68 – 81.56) 6.22 (7) 112.77 G 8 (S) 660 1.69 (±0.21) 132.12 (101.46 – 167.56) 2.94 (7) 235.93 G 8 (Us) 660 1.29 (±0.09) 53.08 (42.59 – 66.78) 6.68 (7) 94.79 G 9 (Us) 660 1.21 (±0.09) 44.83 (35.68 – 56.80) 6.86 (7) 80.05 G 10 (Us) 660 1.31 (±0.09) 35.52 (28.61 – 44.29) 7.55 (7) 63.43 G 11 (Us) 660 1.36 (±0.09) 29.85 (24.16 – 36.94) 5.33 (7) 53.30 G 12 (Us) 660 1.20 (±0.08) 20.68 (16.37 – 26.01) 5.96 (7) 36.93 G 13 (Us) 660 1.43 (±0.11) 20.32 (15.65 – 25.78) 6.95 (7) 36.29 G 14 (Us) 660 1.39 (±0.12) 22.65 (16.87 – 29.37) 4.79 (7) 40.45 G 15 (Us) 660 1.66 (±0.16) 22.10 (16.58 – 28.16) 3.56 (7) 39.46 a RR (Resistance ratio), is a comparison between the LC 50 values of the Lampung population with the LC 50 value of the laboratory susceptible population. b S = Selected; Us = Unselected. Tabl e 3 . Decrease in resistance of Nilaparvata lugens from Lampung Province, Indonesia, to imida c loprid when the selections were discontinued Value Number of generations without selection 10 Initial LC 50 a (95% CI) (ppm) 101.06 (73.68 – 129.82) Final LC 50 b (95% CI) (ppm) 22.10 (16.57 – 28.16) Initial Slope a (±SE) 1.64 (±0.22) Final Slope b (±SE) 1.66 (±0.16) Initial RR a 180.46 Final RR b 39.46 DR c -0.07 GR d 14.29 a The initial LC 50 value, slope and RR (resistance ratio) were determined at G 1 . b The final LC 50 value, slope and RR (resistance ratio) were determined at G 15 . c Decrease in resistance. d The estimated number of generations needed to decrease 10-fold of the LC 50 value (DR -1 ). Table 4 . Estimation of realized heritability ( h 2 ) of imidacloprid resistance in Nilaparvata lugens from Lampung Province, Indonesia Value Selected generations 5 8 Initial LC 50 (95% CL) (ppm) a 29.32 (23.71 – 32.26) 29.32 (23.71 – 32.26) Final LC 50 (95% CL) (ppm) b 101.06 (73.68 – 129.82) 132.12 (101.46 – 167.56) Response to selection ( R ) 0.1075 0.0817 Intensity of selection ( I ) 1.1009 0.9247 Initial slope (±SE) 1.36 (±0.10) 1.36 (±0.10) Final slope (±SE) 1.64 (±0.22) 1.69 (±0.21) Phenotypic deviation ( dp ) 0.6667 0.6557 Selection differential ( S ) 0.7339 0.6063 Realized heritability ( h 2 ) 0.1465 0.1348 a The initial LC 50 value was determined at G 1. b The final LC 50 values were determined at G 5 and G 8 . Additional Declarations No competing interests reported. Supplementary Files HighlightsandGraphicalabstract.docx Cite Share Download PDF Status: Published Journal Publication published 23 Feb, 2026 Read the published version in International Journal of Tropical Insect Science → Version 1 posted Editorial decision: Revision requested 21 Jan, 2026 Reviewers agreed at journal 21 Jan, 2026 Reviews received at journal 21 Jan, 2026 Reviews received at journal 20 Jan, 2026 Reviews received at journal 19 Jan, 2026 Reviewers agreed at journal 17 Jan, 2026 Reviewers agreed at journal 15 Jan, 2026 Reviewers agreed at journal 15 Jan, 2026 Reviewers agreed at journal 15 Jan, 2026 Reviewers invited by journal 15 Jan, 2026 Editor assigned by journal 26 Sep, 2025 Submission checks completed at journal 26 Sep, 2025 First submitted to journal 25 Sep, 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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Andi Trisyono","email":"","orcid":"","institution":"Universitas Gadjah Mada","correspondingAuthor":false,"prefix":"","firstName":"Y.","middleName":"Andi","lastName":"Trisyono","suffix":""}],"badges":[],"createdAt":"2025-09-25 05:53:13","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-7709102/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-7709102/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s42690-026-01804-8","type":"published","date":"2026-02-23T15:58:21+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":100756449,"identity":"627e3dc0-6646-4fa7-9db7-a81049c22421","added_by":"auto","created_at":"2026-01-21 06:35:15","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":98413,"visible":true,"origin":"","legend":"","description":"","filename":"StabilitySubmitIJTISDanarsiDiptaningsariNEW.docx","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/e2e50b9479e1807c66be970c.docx"},{"id":100756382,"identity":"f64b13d5-f8c7-4db2-a817-8dbca1adf910","added_by":"auto","created_at":"2026-01-21 06:34:36","extension":"json","order_by":1,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":6106,"visible":true,"origin":"","legend":"","description":"","filename":"92db758f96b847afb418b149089204d8.json","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/902b9202ca9e34b626edd462.json"},{"id":100756310,"identity":"930c71ce-5bf4-4c01-8edb-e99229f7d7c0","added_by":"auto","created_at":"2026-01-21 06:34:03","extension":"docx","order_by":2,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":378657,"visible":true,"origin":"","legend":"","description":"","filename":"HighlightsandGraphicalabstract.docx","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/943e4333b9102850817085dd.docx"},{"id":100756402,"identity":"dbc8a6ef-0e07-4308-9c6e-436f30c6075a","added_by":"auto","created_at":"2026-01-21 06:34:53","extension":"xml","order_by":3,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":110023,"visible":true,"origin":"","legend":"","description":"","filename":"92db758f96b847afb418b149089204d81enriched.xml","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/e45989c701bcef9a55f08310.xml"},{"id":100756377,"identity":"33a84208-1e26-427c-8421-4e8cb777dcfe","added_by":"auto","created_at":"2026-01-21 06:34:29","extension":"xml","order_by":5,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":110252,"visible":true,"origin":"","legend":"","description":"","filename":"92db758f96b847afb418b149089204d81structuring.xml","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/51635301c23f21d0e9ee5b4c.xml"},{"id":100756399,"identity":"24f50bcb-5f0d-46fb-9bdd-a3dcee78d43a","added_by":"auto","created_at":"2026-01-21 06:34:45","extension":"html","order_by":6,"title":"","display":"","copyAsset":false,"role":"acdc-reference","size":117925,"visible":true,"origin":"","legend":"","description":"","filename":"earlyproof.html","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/9de6c125968293b90a3e926c.html"},{"id":100756414,"identity":"1603ed46-64f6-4056-9016-e10238242fa5","added_by":"auto","created_at":"2026-01-21 06:35:09","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":57326,"visible":true,"origin":"","legend":"\u003cp\u003eThe resistance stability to imidacloprid in \u003cem\u003eNilaparvata lugens \u003c/em\u003efrom Lampung Province, Indonesia. Selections were carried out every generation from the first to the eight generation (Selected). At the fifth generation, a part of this population was no longer selected for the next 10 generations (Unselected). The error bars represent the 95% confidence intervals of the resistance ratios. The LC\u003csub\u003e50\u003c/sub\u003e values for the susceptible population and the Lampung population (G\u003csub\u003e1\u003c/sub\u003e) were 0.56 (0.24-0.95) and 29.32 (23.71-36.26) ppm, respectively.\u003c/p\u003e","description":"","filename":"1.png","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/1a31f94f5b5df74d8209d04d.png"},{"id":103765686,"identity":"5c84aa41-8145-495f-9f3e-6f0e7ba97d7f","added_by":"auto","created_at":"2026-03-02 16:07:35","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1029386,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/b2ea0e78-dfec-43c1-9539-68a4514b1824.pdf"},{"id":100756401,"identity":"828e5a46-f3be-4361-8ad4-cef75d7fdd37","added_by":"auto","created_at":"2026-01-21 06:34:49","extension":"docx","order_by":0,"title":"","display":"","copyAsset":false,"role":"supplement","size":378657,"visible":true,"origin":"","legend":"","description":"","filename":"HighlightsandGraphicalabstract.docx","url":"https://assets-eu.researchsquare.com/files/rs-7709102/v1/683154e95e89b5c669faed8a.docx"}],"financialInterests":"No competing interests reported.","formattedTitle":"Stability and realized heritability of resistance to imidacloprid in the field- collected brown planthopper Nilaparvata lugens (Hemiptera: Delphacidae) from Lampung, Indonesia","fulltext":[{"header":"Introduction","content":"\u003cp\u003eRice cultivation in Indonesia faces a potentially serious challenges from the brown planthopper \u003cem\u003eNilaparvata lugens\u003c/em\u003e St\u0026aring;l (Hemiptera: Delphacidae) due to the rice availability under field conditions throughout the year. The density of \u003cem\u003eN. lugens\u003c/em\u003e in Indonesia spreads over Java, Bali, Lampung, South Sumatra, Central Sulawesi, West Kalimantan, and South Kalimantan (BB Padi, 2020). In Indonesia, the strategies used in managing \u003cem\u003eN. lugens\u003c/em\u003e in the rice plant rely mainly on natural enemies\u0026rsquo; conservation, resistant variety and insecticide formulations. Imidacloprid is one of the active ingredients of insecticide that has been widely used for controlling \u003cem\u003eN. lugens\u003c/em\u003e in Indonesia since 1992 (Ditjen PSP, 2017).\u003c/p\u003e \u003cp\u003eOveruse of the insecticides will cause insecticide resistance to develop. Imidacloprid resistance in \u003cem\u003eN. lugens\u003c/em\u003e populations has been documented in China and Korea with a resistance ratio of 57.0 to 2029.0-fold (Bao et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e; Zhang et al., \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and 13.0 to 237.0-fold (Min et al., \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2014\u003c/span\u003e), respectively. \u003cem\u003eNilaparvata lugens\u003c/em\u003e populations from Vietnam and Thailand showed resistance development to imidacloprid, with the resistance ratio of 89.0 and 102.9-fold, respectively (Matsumura \u0026amp; Sanada-Morimura, \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2010\u003c/span\u003e; Bao et al., \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Since 2010 to 2019, imidacloprid resistances of the field populations of \u003cem\u003eN. lugens\u003c/em\u003e had been occured in the the West Java Province (13.5 to 108.1-fold) (Surahmat et al., 2016; Iswanto et al., \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and the Central Java Province (25.9 to 29.4-fold) (Londingkene et al., \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2016a\u003c/span\u003e; Diptaningsari et al., \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe management of resistance to insecticide requires a systematic and comprehensive strategy. An understanding of the stability and heritability of the insecticide resistance is crucial for designing insecticide resistance management (IRM) strategies to anticipate and decelerate the development of resistance. The stability of resistance plays an important role to determine an effective pest management strategy (Roush et al., 1990; Lan et al., 2001). The resistance stability was determined by rearing the selected insects in the absence of insecticides across multiple generations. The study by Ullah and Shad (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) revealed that resistance to chlorfenapyr and triazophos in \u003cem\u003eOxycarenus hyalinipennis\u003c/em\u003e (Hemiptera: Lygaeidae) was unstable, on the one hand, the resistance to emamectin benzoate and fipronil was stable. The resistance in \u003cem\u003eBemisia tabaci\u003c/em\u003e to acetamiprid remained in the absence of selection pressure (Basit et al., \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). In the house flies (\u003cem\u003eMusca domestica\u003c/em\u003e) and \u003cem\u003eN. lugens\u003c/em\u003e, the resistance to imidacloprid was not stable and dropped throughout several months (Kavi et al., \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2014\u003c/span\u003e, Diptaningsari et al., \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAnother parameter that needs to be estimated is the realized heritability (\u003cem\u003eh\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e) in predicting how resistance can be developed because of the use of insecticides (Tabashnik, \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e1992\u003c/span\u003e). The \u003cem\u003eh\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e value in imidacloprid resistance was observed in \u003cem\u003eN. lugens\u003c/em\u003e (Wang et al., \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2009\u003c/span\u003e, Diptaningari et al., 2019), \u003cem\u003eSpodoptera litura\u003c/em\u003e (Abbas et al., \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2012\u003c/span\u003e) and house fly (\u003cem\u003eM. domestica\u003c/em\u003e) (Khan et al., \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). The research results provided in this article will be of immense benefits as far as stability and heritability of resistance to imidacloprid in field population of \u003cem\u003eN. lugens\u003c/em\u003e in Indonesia is concerned that can be potentially used in controlling \u003cem\u003eN. lugens\u003c/em\u003e through sustainable measures.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cp\u003e\u003cstrong\u003eInsects\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eIn the study, two groups of \u003cem\u003eN. lugens\u003c/em\u003e were used. The resistant population was the first one, which was taken in 2018 from the rice fields (Ciherang variety) in the Village of Notoharjo (5\u0026deg;08\u0026apos;58.2\u0026quot;S 105\u0026deg;14\u0026apos;13.3\u0026quot;E), the Sub-District of Trimurjo, the District of Central Lampung, Lampung Province, Indonesia. \u003cem\u003e\u0026nbsp;Nilaparvata lugens\u003c/em\u003e (G\u003csub\u003e0\u003c/sub\u003e) were collected from ten sampling points in rice fields (\u0026plusmn;2,000 adults). The insecticides used at the collection sites were BPMC, dimehipo, imidacloprid, pymetrozine, and buprofezine. The susceptible population which was obtained in 1986 was the second population that was collected from the District of Sleman, Special Region of Yogyakarta. This population was reared under laboratory conditions, kept apart with other populations, without supplementation from field population and no insecticides exposure. These populations were maintained at \u0026plusmn;27 \u0026deg;C and 70\u0026ndash;80% RH. Rice seedlings (5-7 days) were prepared in a plastic container (30 x 24 cm) in which a number of approximately 100 pairs were then placed to oviposit. The nymphs were moved with an aspirator device into a new plastic container (containing new rice plants) after these eggs had hatched and the host rice plants began to yellow. The nymphs transferred into the new rice plants, and the plants were changed every 4 days. In 5-7 days of rice plants, the newly emerged adults were reared to the next generations.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eInsecticide\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe bioassays and selections were conducted in \u003cem\u003eN. lugens\u003c/em\u003e using a commercial 5% imidacloprid formulation (Confidor 5 WP, produced by PT. Bayer Indonesia).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eSelection for resistance and toxicity bioassays\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFor eight generations (G1\u0026ndash;G8), selection with imidacloprid was carried out by exposing third-instar nymphs of \u003cem\u003eN. lugens\u003c/em\u003e to treated rice seedlings. Seedlings of rice (5-7 days old) were dipped in the concentration of 100 ppm on first generation and the concentration was gradually increased at every generation on selections and it reached 600 ppm on seventh and eight generation. The seeds of rice were dipped in an insecticide solution for 15 seconds, followed by air-drying for 15 minutes at room temperature. The selected rice plants treated were subsequently kept in a plastic cup (10 x 11 cm) where water had been added to such an extent that the seedlings will remain fresh. Aspirating device was used to obtain the third instar nymphs, and 20 nymphs were added to each cup. The level of mortality was taken as 120 hours after treating in order to receive at least some survivors. The untreated rice seedlings were put in plastic cups where they were left with insects for further development. The nymphs were confirmed dead when no response was observed after gentle probing with a soft brush. The selections made in the laboratory was done in the same way as already explained and imidacloprid concentration varied in all generations (Table 1).\u0026nbsp;\u003c/p\u003e\n\u003cp\u003eBioassays were performed on the third instars nymphs of the resistant and the susceptible populations. Preliminary bioassay was done to establish the concentration levels that attained 5-99% deaths with leaf dipping method. \u0026nbsp;Seedlings of rice (5-7 days old) were immersed at insecticide solution and air-dried under room condition (15 minutes). The treated rice seedlings were then placed in a plastic cup (10 x 11 cm), containing sufficient water to make the seedlings appear fresh. The concentrations applied in the determination of resistance were based on the outcome of the preliminary bioassays whereby the concentrations ranged between 0-640 ppm (resistant population) and 0-80 ppm (susceptible population). Each of the bioassays involved a total of 495-660 nymphs, and 60-80 nymphs were used for EACH concentration. Mortality observations were conducted daily for up to 168 hours.\u003c/p\u003e\n\u003cp\u003eThe LC\u003csub\u003e50\u0026nbsp;\u003c/sub\u003evalues were determined using Probit analysis (Finney, 1971) with POLO-Plus. The Abbotts formula (Abbott, 1925) was applied to correct mortality data. The resistance ratio (RR) value is a comparison between the LC\u003csub\u003e50\u003c/sub\u003e of the resistant population and the LC\u003csub\u003e50\u003c/sub\u003e of the susceptible population. If the 95% confidence limits (CLs) did not overlap, the LC\u003csub\u003e50\u003c/sub\u003e values were considered not significantly different.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStability of resistance\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eImidacloprid resistance in \u003cem\u003eN. lugens\u003c/em\u003e was stated stable based on the LC\u003csub\u003e50\u003c/sub\u003e results of the resistant population developed after five generations of selection (G1 to G5), and some of the insects in the population continued to be maintained for the next 10 generations (G6 to G15) without any exposure to any insecticide. The values of LC\u003csub\u003e50\u003c/sub\u003e were calculated per generation. The resistant change was measured as a difference in the degree of resistance to measure the DR (Decrease in Resistance) or the estimated response on selection (Falconer and Mackay, 1996) which can further be applied to obtain the average response per generation. The DR-value was estimated following Tabashnik (1994) as follows:\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eDR = [log (final LC\u003csub\u003e50\u003c/sub\u003e) - log (initial LC\u003csub\u003e50\u003c/sub\u003e)] / n\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWhere\u003cem\u003e\u0026nbsp;final LC\u003csub\u003e50\u003c/sub\u003e\u003c/em\u003e is the value of \u003cem\u003eLC\u003csub\u003e50\u003c/sub\u003e\u003c/em\u003e of generation \u003cem\u003en\u003c/em\u003e reared without selection (G\u003csub\u003e5\u003c/sub\u003e) and the \u003cem\u003einitial\u003c/em\u003e \u003cem\u003eLC\u003csub\u003e50\u003c/sub\u003e\u003c/em\u003e is the value of \u003cem\u003eLC\u003csub\u003e50\u003c/sub\u003e\u003c/em\u003e of generation \u003cem\u003en\u003c/em\u003e reared with selection (G\u003csub\u003e1\u003c/sub\u003e) and \u003cem\u003en\u003c/em\u003e is the number of generations that is reared without selection (10 generations). An estimate was done using the reciprocal of DR or DR-1 values to provide the figure of number of generations that are required in order to decrease LC\u003csub\u003e50\u003c/sub\u003e by 10-fold. To evaluate the change of resistance, the imidacloprid selection was done with the part of the fifth generation of a resistant population and the LC\u003csub\u003e50\u003c/sub\u003e of the next three generations (G\u003csub\u003e6\u0026nbsp;\u003c/sub\u003eto G\u003csub\u003e8\u003c/sub\u003e) was calculated.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEstimation of realized heritability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eRealized heritability (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e) of insect resistance was assessed following the method outlined by Tabashnik, 1992 using pre insecticide selection LC\u003csub\u003e50\u003c/sub\u003e values and post insecticide selection LC\u003csub\u003e50\u003c/sub\u003e values per generation of \u003cem\u003eN. lugens\u003c/em\u003e. The formula was used to calculate the value of the realized heritability (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e):\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e = R / S\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eR\u003c/em\u003e represents the response to insecticide selection, while \u003cem\u003eS\u003c/em\u003e is the selection differential. An estimate of response to selection (R) was estimated following Falconer and Mackay (1996) as follows:\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eR = [log (final LC\u003csub\u003e50\u003c/sub\u003e) - log (initial LC\u003csub\u003e50\u003c/sub\u003e)] / n\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWhere the \u003cem\u003efinal LC\u003csub\u003e50\u003c/sub\u003e\u003c/em\u003e represented the LC\u003csub\u003e50\u003c/sub\u003e of the offspring after \u003cem\u003en\u003c/em\u003e generations of selection, whereas the \u003cem\u003einitial LC\u003csub\u003e50\u003c/sub\u003e\u003c/em\u003e was the LC\u003csub\u003e50\u003c/sub\u003e of the parental generation preceding the selections, and \u003cem\u003en\u003c/em\u003e was the number of the selected generations with imidacloprid. The formula used in calculating differential selection (S) (Hartl, 1988) was:\u003c/p\u003e\n\u003cp\u003e\u003cem\u003eS = i dp\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eWhere \u003cem\u003ei\u003c/em\u003e refers to selection Intensity, and \u003cem\u003edp\u003c/em\u003e is phenotypic standard deviation. The intensity of selection (\u003cem\u003ei\u003c/em\u003e) was approximated on the \u003cem\u003ep\u003c/em\u003e (mean percent survival post-selection). The equation to use in calculating the intensity of selection (\u003cem\u003ei\u003c/em\u003e) (Tabashnik, 1992) was:\u003c/p\u003e\n\u003cp\u003e\u003cem\u003ei = 1.583 \u0026ndash; 0.0193336p + 0.0000428p\u003csup\u003e2\u003c/sup\u003e + 3.65194 / p\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eThe phenotypic standard deviation (\u003cem\u003edp\u003c/em\u003e) was calculated using the formula described by Tabashnik (1992):\u003c/p\u003e\n\u003cp\u003e\u003cem\u003edp = [1/2 (initial slope + final slope)] \u003csup\u003e- 1\u003c/sup\u003e\u003c/em\u003e\u003c/p\u003e\n\u003cp\u003eIn which the \u003cem\u003einitial slope\u003c/em\u003e denoted the slope of the probit regression lines of the parental generation prior to selection and \u003cem\u003efinal slope\u003c/em\u003e was the slope of the probit regression lines of the offspring after \u003cem\u003en\u003c/em\u003e generations of selection with imidacloprid.\u003c/p\u003e"},{"header":"Results","content":"\u003cp\u003e\u003cstrong\u003eSelection for resistance and toxicity bioassays\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe laboratory selections (Table 1) increased the resistance ratio from 52.36-fold at G\u003csub\u003e1\u003c/sub\u003e to 180.46-fold at G\u003csub\u003e5\u0026nbsp;\u003c/sub\u003eand 235.93-fold at G\u003csub\u003e8\u003c/sub\u003e, compared to the susceptible population (Table 2).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eStability of resistance\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eWith selection, resistance ratio was increasing in five generations. The resistance ratio decreased rather fast during the first fifth generations post-removal of selection (G\u003csub\u003e6\u003c/sub\u003e-G\u003csub\u003e10\u003c/sub\u003e). Sixth generation of non-selected (G\u003csub\u003e11\u003c/sub\u003e) exhibited a lower resistance ratio of 53.30-fold which was almost similar to the actual first resistance rate before selection (52.36-fold). The resistance ratio remained largely unchanged but varied a bit in the seventh (G\u003csub\u003e12\u003c/sub\u003e), eighth (G\u003csub\u003e13\u003c/sub\u003e), ninth (G\u003csub\u003e14\u003c/sub\u003e) and tenth generations (G\u003csub\u003e15\u003c/sub\u003e) without selection. These findings showed that the resistance of imidacloprid in \u003cem\u003eN. lugens\u003c/em\u003e of the Lampung population was not stable as the value of the DR was -0.07 (Table 3). The number of the estimated generations needed to decrease the LC\u003csub\u003e50\u0026nbsp;\u003c/sub\u003evalue 10-folds was 14.29 generations. The stabilities of some insects were measured after surviving the selection with imidacloprid over 3 generations (G\u003csub\u003e6\u003c/sub\u003e- G\u003csub\u003e8\u003c/sub\u003e). Using subsequent selections, the resistance ratio of G\u003csub\u003e6\u003c/sub\u003e to G\u003csub\u003e8\u003c/sub\u003e rose by 206.25-fold (G\u003csub\u003e6\u003c/sub\u003e) to 235.93-fold (G\u003csub\u003e8\u003c/sub\u003e) in comparison with the susceptible population (Table 2 and Fig 1).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEstimation of realized heritability\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eFollowing five consecutive generations under imidacloprid selection, the LC\u003csub\u003e50\u0026nbsp;\u003c/sub\u003evalues of the field-collected \u003cem\u003eN. lugens\u003c/em\u003e from Lampung rose incrementally between successive generation with a G\u003csub\u003e1\u0026nbsp;\u003c/sub\u003evalue of 29.32 ppm rising to G\u003csub\u003e5\u0026nbsp;\u003c/sub\u003eat 101.06 ppm and the slope also rose between the generations with a G\u003csub\u003e1\u0026nbsp;\u003c/sub\u003eslope of 1.36 to G\u003csub\u003e5\u0026nbsp;\u003c/sub\u003eslope at 1.64. Following the eighth generation of selection, the values of\u0026nbsp;LC\u003csub\u003e50\u0026nbsp;\u003c/sub\u003ewere higher (132.12 ppm,\u0026nbsp;G\u003csub\u003e8\u003c/sub\u003e), and the slope was higher too (1.69). These values led to the calculated value of the realized heritability (\u003cem\u003eh2\u003c/em\u003e) to be 0.1465 and 0.1348 in five and eight selected generations, respectively. Ten generations were required to elevated 10-fold\u0026nbsp;LC\u003csub\u003e50\u0026nbsp;\u003c/sub\u003evalue of imidacloprid even after selection of five generations and eight generations (reciprocal of R value). (Table 4).\u003c/p\u003e"},{"header":"Discussion","content":"\u003cp\u003eThe imidacloprid resistance can be viewed as a significant problem that has led to the case of \u003cem\u003eN. lugens\u003c/em\u003e in the recent outbreak (Gao et al., 2014, Zhang et al., 2017). The imidacloprid has been registered since the year 1992, and \u003cem\u003eN. lugens\u003c/em\u003e has been controlled in Indonesia using this chemical (Ditjen PSP, 2017). The over application and the high use of imidacloprid is also experienced in other states (Wang et al., 2007). Resistance to the imidacloprid emerged fast within five generations of the selection in this research. The study of Wang et al. (2009) reported a 200-fold to 1298-fold change of resistance of the \u003cem\u003eN. lugens\u003c/em\u003e to imidacloprid by selection through 23 generations. According to Iswanto et al. (2019), in the \u003cem\u003eN. lugens\u003c/em\u003e from Karawang province, West Java Province, Indonesia, an imidacloprid resistance increase by 13.5 (fold) to 33.9-fold was observed after five generations of selection. A new selection of \u003cem\u003eN. lugens\u003c/em\u003e after three generations (G\u003csub\u003e6\u003c/sub\u003e- G\u003csub\u003e8\u003c/sub\u003e) in the study has raised the ratio of resistance.\u003c/p\u003e\n\u003cp\u003eSuch an existing resistance to imidacloprid was attributed to the accelerated detoxification rate based on the upsurge of cytochrome P450 monooxygenase enzyme activity (Zhang et al., 2017, Wen et al., 2009, Puinean et al., 2010) as well as the mutation of the target site (Wen et al., 2009, Liu et al., 2005). The imidacloprid resistance of the \u003cem\u003eN. lugens\u003c/em\u003e populations collected in South and East Asia was linked to the overexpression of two P450 monooxygenase genes, CYP6AY1 and CYP6ER1 (Bao et al., 2016, Garrood et al., 2015). In addition, the new research of resistance mechanism to the insecticides is related to epigenetic features of metabolic resistance, such as the histone modification and the DNA methylation (Bass et al., 2014, Oppold \u0026amp; M\u0026uuml;ller., 2017).\u003c/p\u003e\n\u003cp\u003eThe high resistance ratio had fallen dramatically during the first five generations when the selection was removed, which showed that the resistance to imidacloprid was not stable in \u003cem\u003eN. lugens.\u0026nbsp;\u003c/em\u003eIt could be connected with the presence of the fitness cost in the population (Londingkene et al., 2016b). Variable influence of alleles due to the levels of dominance, the initial frequency of genes, and the relative fitness of different genes can be some of the factors that may contribute towards lessening the degree of resistance in the pest population (Afzal et al., 2015).\u003c/p\u003e\n\u003cp\u003eOnce the selection was changed, the ratio of resistance varied between G\u003csub\u003e11\u0026nbsp;\u003c/sub\u003eand G\u003csub\u003e15\u003c/sub\u003e, but showed no considerable drop. This finding was an indication that homozygous resistance alleles existed prior to reduction of the selection pressure (Roush et al., 1990). The formulation of imidacloprid is mainly used through spraying that can lead to its existence in the irrigation system. In conjunction with the systemic and long-term residual activity, this insecticide might cause long-term selection pressure which might result in \u003cem\u003eN. lugens\u003c/em\u003e resistant (Liu et al., 2003). In an experiment where soil was used conducted in a laboratory by Sarkar et al. (2001), dissipation of imidacloprid was reported to vary between 28.7 and 47.8 days. They will result in persistent pressure of resistance among the insects although the spraying was halted.\u003c/p\u003e\n\u003cp\u003eThe estimated realized heritability (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e) by this research were 0.1465 and 0.1348 by five and eight selected generations respectively, which indicates the \u003cem\u003eN. lugens\u003c/em\u003e has weak capacity to acquire resistance towards imidacloprid (Falconer and Mackay, 1996). The resistance against imidacloprid has also been found comparable with \u003cem\u003eN. lugens\u003c/em\u003e in China (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e = 0.1141) as well as in the other insects, such as \u003cem\u003eS. litura\u003c/em\u003e (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e= 0.15) and \u003cem\u003eM. Domestica\u003c/em\u003e (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e = 0.09) (Abbas et al., 2012; Khan et al., 2014). One of the parameters is an estimation of the realized heritability in predicting persistence of insecticide effectiveness in relation to population-level pest management, but the laboratory experiment cannot elaborate on the parameter (Tabashnik and McGaughey, 1994; Ullah et al., 2016;). Environmental factors will impact on phenotypic variations, e.g. insecticide rotation, migration, natural enemies, and the rest of the environmental variants.\u003c/p\u003e\n\u003cp\u003eThe data on resistance stability, and realized heritability would be helpful to make a recommendation about the appropriate resistance management strategy in \u003cem\u003eN. lugens. Nilaparvata lugens\u003c/em\u003e were found to be unstable in terms of their resistance to imidacloprid whereby they experienced a drop in the level of resistance upon recovery of the selection pressure. One of the strategies to counter the imidacloprid outbreak among \u003cem\u003eN. lugens\u003c/em\u003e would be to use insecticides with dissimilar mode of actions and lacking in cross resistance positive-tests (Ullah and Shad, 2017). Also, supporting the use of Integrated Pest Management (IPM) in rice can help to minimize the risk of development and occurrence of \u003cem\u003eN. lugens\u003c/em\u003e outbreaks in Indonesia.\u003c/p\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgement\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe author wishes to thank Sriyanto Harjanto for his support in rearing the susceptible \u003cem\u003eNilaparvata lugens\u003c/em\u003e population.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eD. D. and Y. A. T. contributed to the conceptualization of the study, data collection, data analysis, and wrote the original draft. A. P. and A. W. contributed to the conceptualization of the study, data analysis, statistical evaluation, and manuscript editing. All authors reviewed and approved the final version of the manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe study was partially funded by the Indonesian Agency for Agricultural Research and Development, the Ministry of Agriculture.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eDeclaration of Competing Interest\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing financial interests or personal relationships that could have appeared to influence the work reported in this paper.\u003cstrong\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAbbas, N., Shad, S.A., Razaq, M., 2012. Fitness cost, cross resistance and realized heritability of resistance to imidacloprid in \u003cem\u003eSpodoptera litura\u003c/em\u003e (Lepidoptera: Noctuidae). 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Outbreaks of the brown planthopper \u003cem\u003eNilaparvata lugens\u003c/em\u003e (St\u0026aring;l) in the Yangtze River Delta: immigration or local reproduction?. Plos One. 9, 1-12.\u003c/li\u003e\n\u003cli\u003eGarrood, W.T., Zimmer, C.T., Gorman, K.J., Nauen, R., Bass, C.B., Davies, T.G.E., 2015. Field-evolved resistance to imidacloprid and ethiprole in populations of brown planthopper \u003cem\u003eNilaparvata lugens\u003c/em\u003e collected from across South and East Asia. Pest. Manag. Sci. 72, 140-149.\u003c/li\u003e\n\u003cli\u003eHartl, D.L., 1988. A Primer of Population Genetics. 2nd ed. Sinauer Associates, Inc., Sunderland, MA.\u003c/li\u003e\n\u003cli\u003eIswanto, E.H., Dadang, Winasa, I.W., Rahmini., 2019. Pengaruh insektisida terhadap kemampuan adaptasi wereng batang cokelat pada varietas padi. Penelitian Pertanian Tanaman Pangan. 3, 125-133.\u003c/li\u003e\n\u003cli\u003eKavi, L.A.K., Kaufman, P.E., Scott, J.G., 2014. Genetics and mechanisms of imidacloprid resistance in house flies. J. Pestic. Biochem. Physiol. 109, 64-69.\u003c/li\u003e\n\u003cli\u003eKhan, H., Abbas, N., Shad, S.A., Afzal, M.B., 2014. Genetics and realized heritability of resistance to imidacloprid in a poultry population of house fly, \u003cem\u003eMusca domestica\u003c/em\u003e L. (Diptera: Muscidae) from Pakistan. J. Pestic. Biochem. Physiol. 114, 38-43.\u003c/li\u003e\n\u003cli\u003eLan, Y.Q., Zhao, S.X., 2001. The stability of resistance to three pyrethroids in \u003cem\u003eSpodoptera exigua\u003c/em\u003e H\u0026uuml;bner. Chinese J. Pestic. Sci. 6, 77\u0026ndash;80.\u003c/li\u003e\n\u003cli\u003eLiu, Z.W., Han, Z.J., Wang, Y.C., Zhang, L.C., Zhang, H.W., Liu, C.J., 2003. Selection for imidacloprid resistance in \u003cem\u003eNilaparvata lugens\u003c/em\u003e (St\u0026aring;l): Cross-resistance patterns and possible mechanisms. Pest Manag. Sci. 59, 1355\u0026ndash;1359.\u003c/li\u003e\n\u003cli\u003eLiu, Z., Williamson, M.S., Lansdell, S.J., Denholm, I., Han, Z., Millar, N.S., 2005. A nicotinic acetylcholine receptor mutation conferring target-site resistance to imidacloprid in \u003cem\u003eNilaparvata lugens\u003c/em\u003e (brown planthopper). PNAS. 102, 8420-8425. \u003c/li\u003e\n\u003cli\u003eLondingkene, J.A., Trisyono, Y.A., Witjaksono., Martono, E., 2016a. Resistance to Imidacloprid and Effect of Three Synergist on the Resistance Level of Brown Planthopper, in: Nuringtyas, T.R., Roto, R., Widyaparaga, A., Mahardika, M., Advances of Science and Technology for Society: Proceedings of the 1\u003csup\u003est\u003c/sup\u003e International Conference on Science and Technology 2015 (ICST-2015). AIP Conference Proceedings, Yogyakarta, pp. 1-5.\u003c/li\u003e\n\u003cli\u003eLondingkene, J.A., Trisyono, Y.A., Witjaksono., Martono, E., 2016b. Relative fitness and feeding capacity of imidacloprid resistant \u003cem\u003eNilaparvata lugens\u003c/em\u003e. J. Perlindungan Tanaman Indonesia. 1, 43-39.\u003c/li\u003e\n\u003cli\u003eMatsumura, M., Sanada-Morimura, S., 2010. Recent status of insecticide resistance in Asian rice Planthoppers. JARQ. 44, 225-230.\u003c/li\u003e\n\u003cli\u003eMin, S., Lee, S.W., Choi, B.R., Lee, S.H., Kwon, D.H., 2014. Insecticide resistance monitoring and correlation analysis to select appropriate insecticides against \u003cem\u003eNilaparvata lugens\u003c/em\u003e (St\u0026aring;l), a migratory pest in Korea. J. Asia-Pac. Entomol. 17, 711-716.\u003c/li\u003e\n\u003cli\u003eOppold, A.M., M\u0026uuml;ller, R., 2017. Epigenetics: A Hidden Target of Insecticides, ed. by Verlinden H, Advances in Insect Physiology, Elsevier, London.\u003c/li\u003e\n\u003cli\u003ePuinean, A.M., Denholm, I., Millar, N.S., Nauen, R., Williamson, M.S., 2010. Characterization of imidacloprid resistance mechanisms in the brown planthopper, \u003cem\u003eNilaparvata lugens\u003c/em\u003e Stal. (Hemiptera: Delphacidae). J. Pestic. Biochem. Physiol. 97, 129-132.\u003c/li\u003e\n\u003cli\u003eRoush, R.T., Hoy, M.A., 1990. Laboratory, glasshouse and field studies of artificially selected carboxyl resistance in \u003cem\u003eMetaseuilus occidentalis\u003c/em\u003e. J. Econ. Entomol. 74, 142\u0026ndash;147.\u003c/li\u003e\n\u003cli\u003eSarkar, M.A., Roy, S., Kole, R.K., Chowdhury, A., 2001. Persistence and metabolism of imidacloprid in different soils of West Bengal. Pest Manag. Sci. 57, 598-602.\u003c/li\u003e\n\u003cli\u003eSurahmat, E.C., Dadang., Prijono, D., 2016. Kerentanan wereng batang cokelat (\u003cem\u003eNilaparvata lugens\u003c/em\u003e) dari enam lokasi di pulau Jawa terhadap tiga jenis insektisida. JHPT Tropika 16, 71-81.\u003c/li\u003e\n\u003cli\u003eTabashnik, B.E., 1992. Resistance risk assessment: realized heritability of resistance to \u003cem\u003eBacillus thuringiensis\u003c/em\u003e in diamondback moth (Lepidoptera: Plutellidae), tobacco budworm (Lepidoptera: Noctuidae), and Colorado potato beetle (Coleoptera: Chrysomelidae). J. Econ. Entomol. 85, 1551-1559.\u003c/li\u003e\n\u003cli\u003eTabashnik, B.E., 1994. Evolution of resistance to \u003cem\u003eBacillus thuringiensis\u003c/em\u003e. Ann. Rev. Entomol. 39, 47-79.\u003c/li\u003e\n\u003cli\u003eTabashnik, B.E., McGaughey, W.H., 1994. Resistance risk assessment for single and multiple insecticide: responses for Indianmeal Moth (Lepidoptera: Pyrallidae) to \u003cem\u003eBacillus thuringiensis\u003c/em\u003e. J. Econ. Entomol. 87, 834-841.\u003c/li\u003e\n\u003cli\u003eUllah, S., Shah, R.M., Shad, S.A., 2016. Genetics, realized heritability and possible mechanism of chlorfenapyr resistance in \u003cem\u003eOxycarenus hyalinipennis\u003c/em\u003e (Lygaeidae: Hemiptera). J. Pestic. Biochem. Physiol. 133, 91-96.\u003c/li\u003e\n\u003cli\u003eUllah, S., Shad, S.A., 2017. Toxicity of insecticides, cross-resistance and stability of chlorfenapyr resistance in different strains of \u003cem\u003eOxycarenus hyalinipennis\u003c/em\u003e Costa (Hemiptera: Lygaeidae). J. Crop Prot. 99, 132-136.\u003c/li\u003e\n\u003cli\u003eWang, Y.H., Wang, M.H., 2007. Factors affecting the outbreak and management tactics of brown planthopper, \u003cem\u003eNilaparvata lugens\u003c/em\u003e (St\u0026aring;l) in China in recent years. Pestic. Sci. Adm. 29, 49\u0026ndash;54.\u003c/li\u003e\n\u003cli\u003eWang, Y.H., Wu, S.G., Zhu, Y.C., Chen, J., Liu, F.Y., Zhao, X.P., Wang, Q., Li, Z., Bo, X.P. and Shen, J.L., 2009. Dynamics of imidacloprid resistance and cross‐resistance in the brown planthopper, \u003cem\u003eNilaparvata lugens\u003c/em\u003e. Entomologia experimentalis et applicata. 131, 20-29. \u003c/li\u003e\n\u003cli\u003eWen, Y., Liu. Z., Bao, H., Han, Z., 2009. Imidacloprid resistance and its mechanisms in field populations of brown planthopper, \u003cem\u003eNilaparvata lugens\u003c/em\u003e St\u0026aring;l in China. J. Pestic. Biochem. Physiol. 94, 36-42.\u003c/li\u003e\n\u003cli\u003eZhang, J., Zhang, Y., Wang, Y., Yang, Y., Cang, X., Liu, Z., 2016. Expression induction of P450 genes by imidacloprid in \u003cem\u003eNilaparvata lugens\u003c/em\u003e: A genome-scale analysis. J. Pestic. Biochem. Physiol. 132, 59-64.\u003c/li\u003e\n\u003cli\u003eZhang, X., Liao, X., Mao, K., Yang, P., Li, D., Alia, E., Wan, H., 2017. The role of detoxifying enzymes in field-evolved resistance to nitenpyram in the brown planthopper \u003cem\u003eNilaparvata lugens\u003c/em\u003e in China. J. Crop Prot. 94, 106-114.\u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Tables","content":"\u003cp\u003e\u003cstrong\u003eTable 1. History of \u003cem\u003eNilaparvata lugens\u003c/em\u003e selection\u003c/strong\u003e\u003cstrong\u003es\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003ewith imidacloprid in the laboratory\u003c/strong\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"603\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003cp\u003eGeneration (G)\u003c/p\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003eConcentration\u003c/p\u003e\n \u003cp\u003e(ppm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003eNo. tested insects\u003c/p\u003e\n \u003cp\u003e(n)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003eMortality\u0026nbsp;\u003c/p\u003e\n \u003cp\u003e(%)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 123px;\"\u003e\n \u003cp\u003eNo.\u0026nbsp;\u003c/p\u003e\n \u003cp\u003einsects alive\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e1\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e100\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e186\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e71.51\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e53\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e2\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e65.33\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e104\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e3\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e62.33\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e113\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e4\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e400\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e360\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e65.28\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e125\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e5\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e500\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e540\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e55.19\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e242\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e6\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e500\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e200\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e47.00\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e106\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e7\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e600\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e400\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e56.75\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e173\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 111px;\"\u003e\n \u003cp\u003eG\u003csub\u003e8\u003c/sub\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e600\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e300\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e36.67\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 123px;\"\u003e\n \u003cp\u003e190\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003eThe population was originally collected from the District of Central Lampung, Lampung Province, Indonesia in 2018. Third instars were selected by exposing them into the treated rice seedlings.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003c/strong\u003e\u003cstrong\u003eTable 2. Resistance level of \u003cem\u003eNilaparvata lugens\u003c/em\u003e collected from Lampung Province, Indonesia, after several laboratory selections with imidacloprid\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\" width=\"600\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003e\u003cstrong\u003ePopulation\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e\u003cstrong\u003en\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eSlope (\u0026plusmn;SE)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eLC\u003csub\u003e50\u003c/sub\u003e (95% CI) (ppm)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026chi;\u003csup\u003e2\u003c/sup\u003e (df)\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eRR\u003csup\u003ea\u003c/sup\u003e\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eSusceptible\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e552\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.22 (\u0026plusmn;0.20)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;0.56 (0.24 \u0026ndash; 0.95)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e5.71 (6)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e1.00\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eLampung\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\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: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e1\u003c/sub\u003e (S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.36 (\u0026plusmn;0.10)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp;29.32 (23.71 \u0026ndash; 36.26)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1.53 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e52.36\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e2\u0026nbsp;\u003c/sub\u003e(S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e495\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.20 (\u0026plusmn;0.12)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 40.82 (28.47 \u0026ndash; 56.27)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e2.03 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e72.89\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e3\u0026nbsp;\u003c/sub\u003e(S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e495\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.24 (\u0026plusmn;0.10)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 55.71 (40.01 \u0026ndash; 79.62)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e9.27 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e99.48\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e4\u0026nbsp;\u003c/sub\u003e(S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e495\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.23 (\u0026plusmn;0.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 71.73 (47.06 \u0026ndash; 101.73)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1.93 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e128.09\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e5\u0026nbsp;\u003c/sub\u003e(S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.64 (\u0026plusmn;0.22)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e101.06 (73.68 \u0026ndash; 129.82)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e6.86 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e180.46\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e6\u003c/sub\u003e (S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.49 (\u0026plusmn;0.20)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e115.50 (84.30 -150.89)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e4.35 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e206.25\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e6\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.56 (\u0026plusmn;0.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 86.40 (66.82 \u0026ndash; 109.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e1.27 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e154.29\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e7\u003c/sub\u003e (S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.58 (\u0026plusmn;0.21)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e121.19 (89.13 \u0026ndash; 156.81)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e3.35 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e216.40\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e7\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.15 (\u0026plusmn;0.08)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 63.15 (49.68 \u0026ndash; 81.56)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e6.22 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e112.77\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e8\u003c/sub\u003e (S)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.69 (\u0026plusmn;0.21)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e132.12 (101.46 \u0026ndash; 167.56)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e2.94 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e235.93\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e8\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.29 (\u0026plusmn;0.09)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 53.08 (42.59 \u0026ndash; 66.78)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e6.68 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e94.79\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e9\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.21 (\u0026plusmn;0.09)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 44.83 (35.68 \u0026ndash; 56.80)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e6.86 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e80.05\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e10\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.31 (\u0026plusmn;0.09)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 35.52 (28.61 \u0026ndash; 44.29)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e7.55 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e63.43\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e11\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.36 (\u0026plusmn;0.09)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 29.85 (24.16 \u0026ndash; 36.94)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e5.33 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e53.30\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e12\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.20 (\u0026plusmn;0.08)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 20.68 (16.37 \u0026ndash; 26.01)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e5.96 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e36.93\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e13\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.43 (\u0026plusmn;0.11)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 20.32 (15.65 \u0026ndash; 25.78)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e6.95 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e36.29\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e14\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.39 (\u0026plusmn;0.12)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 22.65 (16.87 \u0026ndash; 29.37)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e4.79 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e40.45\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 113px;\"\u003e\n \u003cp\u003eG\u003csub\u003e15\u0026nbsp;\u003c/sub\u003e(Us)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 47px;\"\u003e\n \u003cp\u003e660\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 104px;\"\u003e\n \u003cp\u003e1.66 (\u0026plusmn;0.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 179px;\"\u003e\n \u003cp\u003e\u0026nbsp; 22.10 (16.58 \u0026ndash; 28.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 84px;\"\u003e\n \u003cp\u003e3.56 (7)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 72px;\"\u003e\n \u003cp\u003e39.46\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003csup\u003ea\u003c/sup\u003eRR (Resistance ratio), is a comparison between the LC\u003csub\u003e50\u003c/sub\u003e\u003csub\u003e\u0026nbsp;\u003c/sub\u003evalues of the Lampung population with\u0026nbsp;the\u0026nbsp;LC\u003csub\u003e50\u003c/sub\u003e\u003csub\u003e\u0026nbsp;\u003c/sub\u003evalue of the laboratory susceptible population.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003csup\u003eb\u003c/sup\u003eS = Selected; Us = Unselected.\u003c/p\u003e\u003cp\u003e\u003cstrong\u003eTabl\u003c/strong\u003e\u003cstrong\u003ee\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e3\u003c/strong\u003e\u003cstrong\u003e.\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eDecrease in resistance of\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003eNilaparvata lugens\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003e\u003cem\u003e\u0026nbsp;\u003c/em\u003e\u003c/strong\u003e\u003cstrong\u003efrom Lampung Province, Indonesia, to\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;imida\u003c/strong\u003e\u003cstrong\u003ec\u003c/strong\u003e\u003cstrong\u003eloprid\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003ewhen the selections were discontinued\u003c/strong\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 301px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd style=\"width: 300px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eValue\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eNumber of generations without selection\u0026nbsp;\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp; 10\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eInitial LC\u003csub\u003e50\u003c/sub\u003e\u003csup\u003ea\u003c/sup\u003e (95% CI) (ppm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e101.06 (73.68 \u0026ndash; 129.82)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eFinal LC\u003csub\u003e50\u003c/sub\u003e\u003csup\u003eb\u003c/sup\u003e (95% CI) (ppm)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp;22.10 (16.57 \u0026ndash; 28.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eInitial Slope\u003csup\u003ea\u003c/sup\u003e (\u0026plusmn;SE)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;1.64 (\u0026plusmn;0.22)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eFinal Slope\u003csup\u003eb\u003c/sup\u003e (\u0026plusmn;SE)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 1.66 (\u0026plusmn;0.16)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eInitial RR\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e180.46\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eFinal RR\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp; 39.46\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eDR\u003csup\u003ec\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp; -0.07\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd style=\"width: 301px;\"\u003e\n \u003cp\u003eGR\u003csup\u003ed\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 300px;\"\u003e\n \u003cp\u003e\u0026nbsp;14.29\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003csup\u003ea\u003c/sup\u003e\u003csup\u003e\u0026nbsp;\u003c/sup\u003eThe initial LC\u003csub\u003e50\u003c/sub\u003e\u003csub\u003e\u0026nbsp;\u003c/sub\u003evalue, slope and\u0026nbsp;RR (resistance ratio)\u0026nbsp;were determined at G\u003csub\u003e1\u003c/sub\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003csup\u003eb\u003c/sup\u003e\u003csup\u003e\u0026nbsp;\u003c/sup\u003eThe final LC\u003csub\u003e50\u003c/sub\u003e\u003csub\u003e\u0026nbsp;\u003c/sub\u003evalue, slope\u0026nbsp;and\u0026nbsp;RR (resistance ratio)\u0026nbsp;were determined at G\u003csub\u003e15\u003c/sub\u003e.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ec\u003c/sup\u003e\u003csup\u003e\u0026nbsp;\u003c/sup\u003eDecrease in resistance.\u003c/p\u003e\n\u003cp\u003e\u003csup\u003ed\u003c/sup\u003e\u003csup\u003e\u0026nbsp;\u003c/sup\u003eThe estimated number of generations needed to decrease 10-fold of the LC\u003csub\u003e50\u003c/sub\u003e value (DR\u003csup\u003e-1\u003c/sup\u003e).\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003e\u003c/strong\u003e\u003cstrong\u003eTable\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003e4\u003c/strong\u003e\u003cstrong\u003e. Estimation of realized heritability (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e) of imidacloprid resistance in \u003cem\u003eNilaparvata lugens\u003c/em\u003e from\u0026nbsp;\u003c/strong\u003e\u003cstrong\u003eLampung\u003c/strong\u003e\u003cstrong\u003e\u0026nbsp;Province, Indonesia\u003c/strong\u003e\u003c/p\u003e\n\u003ctable border=\"1\" cellspacing=\"0\" cellpadding=\"0\"\u003e\n \u003ctbody\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003e\u003cstrong\u003e\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd colspan=\"2\" style=\"width: 399px;\"\u003e\n \u003cp\u003e\u003cstrong\u003eValue\u003c/strong\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eSelected generations\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 5\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 8\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eInitial LC\u003csub\u003e50\u003c/sub\u003e (95% CL) (ppm)\u003csup\u003ea\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; 29.32 (23.71 \u0026ndash; 32.26)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp;29.32 (23.71 \u0026ndash; 32.26)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eFinal LC\u003csub\u003e50\u003c/sub\u003e (95% CL) (ppm)\u003csup\u003eb\u003c/sup\u003e\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e101.06 (73.68 \u0026ndash; 129.82)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e132.12 (101.46 \u0026ndash; 167.56)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eResponse to selection (\u003cem\u003eR\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.1075\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; \u0026nbsp; \u0026nbsp;0.0817\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eIntensity of selection (\u003cem\u003eI\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 1.1009\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.9247\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eInitial slope (\u0026plusmn;SE)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 1.36 (\u0026plusmn;0.10)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 1.36 (\u0026plusmn;0.10)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eFinal slope (\u0026plusmn;SE)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 1.64 (\u0026plusmn;0.22)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 1.69 (\u0026plusmn;0.21)\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003ePhenotypic deviation (\u003cem\u003edp\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.6667\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.6557\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eSelection differential (\u003cem\u003eS\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.7339\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.6063\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003ctr\u003e\n \u003ctd valign=\"top\" style=\"width: 217px;\"\u003e\n \u003cp\u003eRealized heritability (\u003cem\u003eh\u003csup\u003e2\u003c/sup\u003e\u003c/em\u003e)\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 198px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.1465\u003c/p\u003e\n \u003c/td\u003e\n \u003ctd valign=\"top\" style=\"width: 201px;\"\u003e\n \u003cp\u003e\u0026nbsp; \u0026nbsp; 0.1348\u003c/p\u003e\n \u003c/td\u003e\n \u003c/tr\u003e\n \u003c/tbody\u003e\n\u003c/table\u003e\n\u003cp\u003e\u003csup\u003ea\u003c/sup\u003e The initial LC\u003csub\u003e50\u003c/sub\u003e\u003csub\u003e\u0026nbsp;\u003c/sub\u003evalue was determined at\u0026nbsp;G\u003csub\u003e1.\u003c/sub\u003e\u003c/p\u003e\n\u003cp\u003e\u003csup\u003eb\u003c/sup\u003e The final LC\u003csub\u003e50\u003c/sub\u003e values were determined at G\u003csub\u003e5\u003c/sub\u003e and G\u003csub\u003e8\u003c/sub\u003e.\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":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"imidacloprid, Nilaparvata lugens, realized heritability, resistance, stability","lastPublishedDoi":"10.21203/rs.3.rs-7709102/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-7709102/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eAvailability of rice throughout the year in the Indonesian field makes the brown planthopper, (\u003cem\u003eNilaparvata lugens\u003c/em\u003e) (St\u0026aring;l) an important pest. Imidacloprid is an insecticide active ingredient that is commonly used for managing \u003cem\u003eN. lugens\u003c/em\u003e populations. The data on resistance stability and realized heritability would be effective to derive a viable resistance management system against \u003cem\u003eN. lugens\u003c/em\u003e. The resistance ratio (RR) increased after selection for five generations, from 52.36 to 180.46-fold in \u003cem\u003eN. lugens\u003c/em\u003e collected from rice fields in the District of Central Lampung, Lampung Province, Indonesia. In order to determine the stability of resistance, five additional generations (totaling ten generations) were reared without insecticides exposure. Stability experiments revealed that resistance to imidacloprid was unstable and there was a loss of resistance from 180.46 to 39.46-fold over ten generation without any selections (DR value = -0.07). On the contrary, the resulting resistance level after continues selections with imidacloprid during the subsequent three generations might be up the resistance level from 180.46 to 235.93-fold. The realized heritability values (\u003cem\u003eh\u003c/em\u003e\u003csup\u003e\u003cem\u003e2\u003c/em\u003e\u003c/sup\u003e) of imidacloprid resistance were low at 0.1465 and 0.1348 in five and eight generations that were selected, respectively. This unstable resistance could be mitigated by temporarily removing selection pressure or by rotating to insecticides with different mode of actions. Having a low heritability of resistance, the sustainable wide-range resistance management plan will be required in controlling the \u003cem\u003eN. lugens\u003c/em\u003e population in Indonesia.\u003c/p\u003e","manuscriptTitle":"Stability and realized heritability of resistance to imidacloprid in the field- collected brown planthopper Nilaparvata lugens (Hemiptera: Delphacidae) from Lampung, Indonesia","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2026-01-21 06:27:14","doi":"10.21203/rs.3.rs-7709102/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Revision requested","date":"2026-01-21T09:12:04+00:00","index":"","fulltext":""},{"type":"reviewerAgreed","content":"189528927557744656771000135525593540884","date":"2026-01-21T08:51:26+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-21T05:58:37+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-20T23:30:47+00:00","index":"hide","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2026-01-19T09:29:01+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"170757642907061044349043410756263323657","date":"2026-01-17T06:42:18+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"148661411975140425922979924958035521974","date":"2026-01-16T04:37:20+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"305963563094204458390608758067702279802","date":"2026-01-16T00:30:49+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"217953260185120274913790612433414126346","date":"2026-01-15T17:32:58+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2026-01-15T17:04:00+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-09-26T09:15:05+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2025-09-26T09:13:33+00:00","index":"","fulltext":""},{"type":"submitted","content":"International Journal of Tropical Insect Science","date":"2025-09-25T05:38:24+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"international-journal-of-tropical-insect-science","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jtis","sideBox":"Learn more about [International Journal of Tropical Insect Science](http://link.springer.com/journal/42690)","snPcode":"42690","submissionUrl":"https://www.editorialmanager.com/jtis/default2.aspx","title":"International Journal of Tropical Insect Science","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"71ba8273-c0bd-443c-90ac-38dd2bf52041","owner":[],"postedDate":"January 21st, 2026","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2026-03-02T16:04:17+00:00","versionOfRecord":{"articleIdentity":"rs-7709102","link":"https://doi.org/10.1007/s42690-026-01804-8","journal":{"identity":"international-journal-of-tropical-insect-science","isVorOnly":false,"title":"International Journal of Tropical Insect Science"},"publishedOn":"2026-02-23 15:58:21","publishedOnDateReadable":"February 23rd, 2026"},"versionCreatedAt":"2026-01-21 06:27:14","video":"","vorDoi":"10.1007/s42690-026-01804-8","vorDoiUrl":"https://doi.org/10.1007/s42690-026-01804-8","workflowStages":[]},"version":"v1","identity":"rs-7709102","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-7709102","identity":"rs-7709102","version":["v1"]},"buildId":"XKTyCvWXoU3ODBz1xrDgd","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}