Model and Scenario Variations in Predicted Number of Generations of Spodoptera litura Fab. on Peanut during Future Climate Change Scenario

The present study features the estimation of number of generations of tobacco caterpillar, Spodoptera litura. Fab. on peanut crop at six locations in India using MarkSim, which provides General Circulation Model (GCM) of future data on daily maximum (T.max), minimum (T.min) air temperatures from six models viz., BCCR-BCM2.0, CNRM-CM3, CSIRO-Mk3.5, ECHams5, INCM-CM3.0 and MIROC3.2 along with an ensemble of the six from three emission scenarios (A2, A1B and B1). This data was used to predict the future pest scenarios following the growing degree days approach in four different climate periods viz., Baseline-1975, Near future (NF) -2020, Distant future (DF)-2050 and Very Distant future (VDF)—2080. It is predicted that more generations would occur during the three future climate periods with significant variation among scenarios and models. Among the seven models, 1–2 additional generations were predicted during DF and VDF due to higher future temperatures in CNRM-CM3, ECHams5 & CSIRO-Mk3.5 models. The temperature projections of these models indicated that the generation time would decrease by 18–22% over baseline. Analysis of variance (ANOVA) was used to partition the variation in the predicted number of generations and generation time of S. litura on peanut during crop season. Geographical location explained 34% of the total variation in number of generations, followed by time period (26%), model (1.74%) and scenario (0.74%). The remaining 14% of the variation was explained by interactions. Increased number of generations and reduction of generation time across the six peanut growing locations of India suggest that the incidence of S. litura may increase due to projected increase in temperatures in future climate change periods.     

Herbivore- and Elicitor- Induced Resistance in Groundnut to Asian armyworm,Spodoptera litura (Fab.) (Lepidoptera: Noctuidae)

Induced defense was studied in three groundnut genotypes ICGV 86699 (resistant), NCAc 343 (resistant) and TMV 2 (susceptible) in response to Spodoptera litura infestation and jasmonic acid (JA) application. The activity of the oxidative enzymes [peroxidase (POD) and polyphenol oxidase (PPO)] and the amounts other host plant defense components [total phenols, hydrogen peroxide (H2O2), malondialdehyde (MDA), and protein content] were recorded at 24, 48, 72 and 96 h in JA pretreated (one day before) plants and infested with S. litura, and JA application and simultaneous infestation with S. litura to understand the defense response of groundnut genotypes against S. litura damage. Data on plant damage, larval survival and larval weights were also recorded. There was a rapid increase in the activities of POD and PPO and in the quantities of total phenols, H2O2, MDA and protein content in the JA pretreated + S. litura infested plants. All the three genotypes showed quick response to JA application and S. litura infestation by increasing the defensive compounds. Among all the genotypes, higher induction was recorded in ICGV 86699 in most of the parameters. Reduced plant damage, low larval survival and larval weights were observed in JA pretreated plants. It suggests that pretreatment with elicitors, such as JA could provide more opportunity for plant defense against herbivores.     

Thermal tolerance of Cyrtobagous salviniae: a biocontrol agent in a changing world

The weevil Cyrtobagous salviniae Calder & Sands 1985 (Family: Curculionidae) is a highly effective biocontrol agent for the invasive water fern Salvinia molesta D.S. Mitchell (Family: Salviniaceae). The life histories of both organisms are affected by temperature, making the potential impacts of climate change on efficacy of control an important area in which comprehensive understanding is required. Here we use warming tolerance (WT) and low temperature tolerance (LTT) as measures of C. salviniae’s sensitivity to climate warming, calculated across South Africa using critical thermal limits, lethal temperature limits and mean maximum (Tmax) and minimum (Tmin) environmental temperatures under present climatic conditions and two future periods (2040s and 2080s). From the present climate to the 2080s the WTs of C. salviniae decrease and LTTs increase indicating C. salviniae may face greater constraints on survival as Tmax nears the upper thermal limits, but increased population persistence over cool months as Tmin increases.     

Genetics of resistance to Cry1C toxin from Bacillus thuringiensis in Spodoptera litura (Fab.)

The present study was undertaken to determine the genetics of Cry1C resistance in Spodoptera litura. Selection of S. litura (Fab.) with Cry1C was done for eight generations to develop resistance. Reciprocal crosses between resistant and susceptible populations were made to understand the population genetics of Cry1C resistance in S. litura. Generation wise selection with Cry1C was evaluated for resistance development in S. litura. The LC₅₀ of Cry1C was 0.14 µg/cm² for the first selected generation and it increased to 23.98 µg/cm² after eight selected generations, which is a 285.47-fold increase in resistance compared with the susceptible strain. The estimated realized heritability (h²) after eight generations of selection with Cry1C insecticidal protein was 0.44. The number of generations required for the tenfold increase in LC₅₀ (1/R) was estimated to be 3.33. Response to Cry1C selection in S. litura was 0.30, the estimated selection differential was 0.69 and the pheonotypic standard deviation (dP) was 0.24. Reciprocal crosses between Cry1C resistant and susceptible strain of S. litura showed autosomal resistance.     

Transgenics in groundnut (Arachis hypogaea L.) expressing cry1AcF gene for resistance to Spodoptera litura (F.)

Large number of primary transgenic events were generated in groundnut by an Agrobacterium mediated, in planta transformation method to assess the efficacy of cry1AcF against the Spodoptera litura. The amplification of required size fragment of 750 bp with npt II primers and 901 bp with cry1AcF gene primers confirmed the integration of the gene. The expression of the cry gene was ascertained by ELISA in T₂ generation, and the maximum concentration of cry protein in transgenic plants reached approximately 0.82 μg/g FW. Further, Southern blot analysis of ten T₂ transgenic plants proved that transgene had been integrated in the genome of all the plants and Northern analysis of the same plants demonstrated the active expression of cry1AcF gene. The highest mean % larval mortalities 80.0 and 85.0 with an average mean % larval mortalities 16.25 (n = 369) and 26.0 (n = 80) were recorded in T₁ and T₂ generations, respectively. Segregation analysis of the selected lines in the T₃ generation demonstrated homozygous nature. This clearly proved that though there is considerable improvement in average mean % larval mortality in T₂ generation, the cry1AcF gene was effective against S. litura only to some extent.     

Lethal and sub‐lethal effects of spinosad on the life‐history traits of army worm,Spodoptera litura (Fabricius) (Lepidoptera: Noctuidae), and its fitness cost of resistance

Spodoptera litura is one of the most destructive polyphagous insect pests, with more than 120 host‐plant species. In our present study, a field‐collected population of S. litura when selected with spinosad for 11 consecutive generations resulted in the development of 3921‐fold resistance to spinosad as compared to the susceptible strain. The spinosad‐resistant strain of S. litura had a relatively high fitness cost (0.17) as compared to the susceptible strain. Furthermore, the lethal and sub‐lethal effects of different concentrations of spinosad were checked on the susceptible strain at different levels; i.e., LC₄₀, LC₃₀, LC₂₀ and LC₁₀, which revealed that the impact of spinosad on the life‐history traits of S. litura increased with the increase in concentration of spinosad. A significant impact of spinosad was recorded on the larval duration, pre‐pupal weight, pupal duration, pupal weight, reproductive potential and adult emergence. The outcomes of the current research clearly indicate that fitness cost of spinosad and its sub‐lethal effects have a significant impact on population dynamics of S. litura, for which it can be incorporated in integrated pest management.     

Genome-wide patterns of copy number variations in Spodoptera litura

Spodoptera litura is a polyphagous pest and can feed on more than 100 species of plants, causing great damage to agricultural production. The SNP results showed that there were gene exchanges between different regions. To explore the variations of larger segments in S. litura genome, we used genome resequencing samples from 14 regions of China, India, and Japan to study the copy number variations (CNVs). We identified 3976 CNV events and 1581 unique copy number variation regions (CNVRs) occupying the 108.5 Mb genome of S. litura. A total of 5527 genes that overlapped with CNVRs were detected. Selection signal analysis identified 19 shared CNVRs and 105 group-specific CNVRs, whose related genes were involved in various biological processes in S. litura. We constructed the first CNVs map in S. litura genome, and our findings will be valuable for understanding the genomic variations and population differences of S. litura.     

Toxicity of Insecticides and Metabolism of Deltamethrin in Helicoverpa assulta Guenéee and Spodoptera litura Fabricius

This experiment was carried out to evaluate the effects of synergists on toxicity of deltamethrin and to investigate the pharmacokinetics of deltamethrin in Helicoverpa assulta Guenéee and Spodoptera litura Fabricius. Susceptibilities of H. assulta and Haman-collected S. litura to 7 insecticides revealed different responses between the 2 species. The different susceptibility of the 2 species to deltamethrin was correlated with the differences in the penetration and metabolism. Larvae of H. assulta pretreated with synergists (piperonyl butoxide (PB), S-benzyl-O, O-diisopropyl phosphorothioate (IBP), and diethyl maleate (DM) were more sensitive to deltamethrin, with the highest synergism ratio (SR = 6.8) being observed with PB. Significant differences in synergism among synergists were not observed. For S. litura, the highest SR(=8.8) was observed with PB. At all time periods, the penetration of ¹⁴C-deltamethrin was relatively faster to S. litura than to H. assulta. Time (hrs) required for 20% (t₂₀) of the applied doses to be excreted from treated S. litura was 1.5-fold faster than that of H. assulta. Higher synergism and longer retention in larval body of S. litura by the addition of PB suggest that the mixed function oxidase be a major factor in detoxification of deltamethrin. The combined use of PB with deltamethrin, and possibly other pyrethroids, might indicate a potential for controlling field populations resistant to deltamethrin and other pyrethroids.     

Significant strength gains observed in rugby players after specific resistance exercise protocols based on individual salivary testosterone responses

Our previous work has demonstrated that professional athletes show protocol-dependent variability in salivary testosterone (T) responses to resistance exercise (RE). The current study examines the consistency and functional outcomes of prescribing a RE regimen based on T response. We hypothesized that prescribing an individual-specific RE protocol based on T response would enhance weight training gains. Sixteen amateur rugby players [(mean +/- SD) age: 20 +/- 2 years; height: 181.5 +/- 8.2 cm; weight: 94.2 +/- 11.1 kg] were characterized by their maximal (Tmax) and minimal (Tmin) T response to four RE protocols: four sets of 10 repetitions (reps) at 70% of one repetition maximum (1RM) with 2 minutes' rest between sets (4 x10-70%); three sets of five reps at 85% 1RM with 3 minutes' rest (3 x 5-85%); five sets of 15 reps at 55% of 1RM with 1 minute's rest (5 x 15-55%); and three sets of 5 reps at 40% 1RM with 3 minutes' rest (3 x 5-40%). Eight athletes then performed a 3-week training block performing only their Tmax protocol. The remaining eight only performed Tmin. After 3 weeks, the athletes were retested on the RE protocols and then crossed over and performed the alternate 3-week training block. All 16 athletes showed significant increases in estimated bench and leg press 1RM strength and bodyweight while performing Tmax. When Tmin was performed, 75% of athletes showed either no change or a significant decline in 1RM performance. Consistent protocol-responses over the experimental period were seen for both the Tmax and Tmin protocols in 12 of 16 athletes. Thus, a relationship between an individual's biologically available T response to RE and enhanced functional gains is reported.     

The toxicity and biological effect of Pedalium murex L. extracts on the tobacco cutworm,Spodoptera litura (Fabr.) larvae

Petroleum ether, acetone, water and aqueous extract of Pedalium murex L. (Pedalaceae) leaves, root and fruit were tested against fourth instar larvae of the tobacco cutworm, Spodoptera litura (Fabr.) under laboratory conditions using leaf dip method. Larval mortality; larval, pupal and adult periods; pupal weight, pupation and adult emergence; larval and pupal deformities of S. litura were recorded. All the tested solvent extracts of P. murex were effective against S. litura life stages by causing mortality in a dose dependent manner. However, the efficacy was more significant with respect to acetone (leaf and root) and petroleum ether (root) at higher concentrations (0.8%) which leads to 100% larval mortality. The water and aqueous extracts of root caused 86.6 and 88.0% larval mortality respectively at 4% concentration. Among the plant parts tested, root had more importance followed by leaf and fruit. Plant extracts extended the larval, pupal and adult periods; reduced the pupal weight, pupal and adult emergence and caused larval–pupal deformities. Total life time was highly prolonged in acetone extracts of P. murex root. Though all the extracts were found to have insecticide activity, acetone and petroleum ether extracts of P. murex root can be used as an effective alternative to modern synthetic insecticides. Bioactive principles from these extracts can be isolated, identified and integrated in S. litura management.     

Electrophoretic Pattern of Larval Esterases in Field and Laboratory-selected Strains of the Tobacco Cutworm,Spodoptera litura (Fabricius)

This research was performed to find out and characterize the specific esterase isozymes related to OP and pyrethroid resistance in the tobacco cutworm Spodoptera litura (Fabricius). Two laboratory strains (DSR₅ and CSR₄) of S. litura, selected with deltamethrin and chlorpyrifos-methyl, had higher larval esterase activities than Hamancollected (HC) strain. Ten esterase isozymes were separated in the wild HC strain on 6.5% nondenaturing polyacrylamide gel electrophoresis (pH 8.3), but only 5 of them were detected in the two strains selected with insecticides. The isozymes were designated from E1 to E10 according to their mobility to cathode. E4 was stained more strongly in both laboratory-selected strains than in HC strain. The frequency of E2 in DSR₅ strain was higher than in HC strain. E2 and E4 were proved to be arylesterase and carboxyesterase, respectively, according to enzyme inhibition tests. Thus decreased number and increased intensity of esterase bands in DSR₅ and CSR₄ strain may be associated with insecticide resistance, resulting from selections successively with insecticides for several generations.     

Toxic Effect of Destruxin A on Abnormal Wing Disc-Like (SLAWD) in Spodoptera litura Fabricius (Lepidoptera: Noctuidae)

Background

Destruxin A (DA) is a microbial insecticide with potent bioactivity against Spodoptera litura larvae. A previous proteomic analysis of S. litura (SL-1) cells exposed to DA showed the abnormal expression of wing disc-like protein of S. litura (SLAWD). To further understand the effect of DA on SLAWD expression, a functional study was carried out.

Principal Findings

The SLAWD gene (SLAWD) was cloned and an open reading frame of 537 bp encoding a polypeptide of 178 amino acids was detected. Real-time fluorescence quantitative PCR (qRT-PCR) suggested that SLAWD is expressed in all developmental stages of S. litura, but expression was highest during the pupal and adult stages. RNAi knockdown of SLAWD expression in 6th-stage larvae was achieved by the microinjection of a specific double-stranded RNA (dsRNA). The results showed a significant decrease in SLAWD mRNA expression levels between the prepupal and adult stages. Interestingly, SLAWD expression was similarly down-regulated by treating 6th-stage larvae with DA. Growth- and development-related statistics confirmed the observed abnormalities in S. litura development after either RNAi or DA treatment.

Conclusions

SLAWD appears to have a biosynthetic function in the pupal and adult stages of S. litura. The toxic effect of DA on S. litura development is due the negative effect of DA on SLAWD gene expression.     

Biochemical mechanism of chlorantraniliprole resistance in Spodoptera litura (Fab) (Lepidoptera: Noctuidae)

Spodoptera litura, a polyphagus insect pest of economic importance, having the ability to develop resistance to various classes of insecticides was selected for the study. Leaf dip bioassay studies were done after ten generations of selection pressure reported the development of resistance up to 80 folds by S. litura against chlorantraniliprole compared to lab susceptible strain. Bioassay studies conducted using enzyme inhibitors such as triphenyl phosphate (TPP) followed by diethyl-maleate (DEM) and piperonyl butoxide (PBO) showed good amount of synergism with chlorantraniliprole and improved efficacy against resistant strain. Findings of bioassay studies were supported by in-vitro enzyme inhibition assays. Esterase activity in gut homogenate of resistant strain was significantly inhibited by TPP suggesting esterase mediated biochemical resistance development in S. litura against chlorantraniliprole.     

Ants (Hymenoptera: Formicidae) as potential egg predators of lepidopteran pests of sweet potato, <Emphasis Type="Italic">Ipomoea batatas</Emphasis> (Solanales: Convolvulaceae), in Japan

The sweet potato Ipomoea batatas (L.) Lam., which has extrafloral nectaries, was visited by nine indigenous ant species in south Kyushu, Japan. These ant species were evaluated for their ability to remove the eggs of two lepidopteran species—Aedia leucomelas (Linnaeus) and Spodoptera litura (Fabricius)—through ant-exclusion experiments. The results showed that Tetramorium bicarinatum (Nylander), one of the dominant ant species, is an efficient predator of lepidopteran eggs. This ant species removed A. leucomelas eggs from ant-visited plants in six trials conducted from mid-July to mid-October, with high removal rates (90–100%) occurring from mid-September onward. On ant-excluded plants, only 0–18% of exposed eggs were removed by ants that accidentally invaded the plants. Tetramorium bicarinatum also removed egg masses and the hatchlings of S. litura. The mean percent area of egg-laden leaves eaten by S. litura larvae was lower on ant-visited plants (0.9%) than on ant-excluded plants (57.3%). In contrast, Formica japonica Motschulsky, another dominant ant species, was not observed removing the eggs or larvae of the lepidopteran species. These results show that certain indigenous ant species are efficient as biocontrol agents. Thus, individual ant species need to be investigated for their ability to control insect pests in various crops.     

Herbivore-induced resistance in different groundnut germplasm lines to Asian armyworm, Spodoptera litura (Fab.) (Lepidoptera: Noctuidae)

Three groundnut germplasm lines, ICGV86699 (resistant), NCAC 343 (resistant) and TMV 2 (susceptible), were examined for Spodoptera litura (Fab.) resistance. Biochemical parameters such as oxidative enzyme activities, peroxidase (POD) and polyphenol oxidase (PPO), other defensive components such as total phenols, hydrogen peroxide (H₂O₂), malondialdehyde (MDA) and protein contents were evaluated in these germplasm lines after 24, 48, 72 and 96 h following S. litura infestation to characterize the mechanism of resistance. Enzyme activities and total phenols, H₂O₂, MDA and protein contents were increased following infestation; however, significance varied at different time intervals and among germplasm lines depending upon the induced level of resistance. The three germplasm lines differed in resistance mechanisms to S. litura and the resistance may be partly due to higher enzyme activities, and other components studied. Among the three germplasms tested, ICGV86699 showed greater elevation in POD and PPO activities and in phenolic and H₂O₂ contents at different time intervals as compared to NCAC 343 and TMV 2.     

Characterization of ovarian follicular dynamics in dromedary camels (Camelus dromedarius)

Ovarian follicular dynamics was monitored by transrectal ultrasonography, for a period of 60 to 90 days, and its correlation with plasma estradiol-17β (E2) and progesterone (P4) were studied in seventeen, multiparous, non-lactating, 12 to 20-year-old dromedary camels. The average number of follicles recruited (12.77 ± 0.93) in each wave between animals varied (P < 0.001). The number of follicles recruited during different follicular waves was highly repeatable (0.95) within individual animals. The growth and mature phase periods of the dominant follicle (DF) were 6.10 ± 0.15 and 10.20 ± 0.47 days, respectively with a linear growth rate of 1.17 ± 0.02 mm/day between Day 0 and 10 of the follicular wave. There was an inverse relationship between the diameter of the largest DF and number of follicles (r = −0.95, P < 0.001). The DF development did not regularly alternate between the ovaries and the incidence of codominance was 45%. The mean maximum diameter of DF during its mature phase was 27.30 ± 0.78 mm and oversized follicle was 38.43 ± 1.41 mm. In 73.3% waves, the DF continued its growth for a period of 10.64 ± 1.53 days even after losing its dominance and developed into oversized follicle. The duration of the regression phase of DF and oversized follicle were 24.71 ± 3.79 and 18.50 ± 2.23 days. The mean duration of a complete follicular wave was 47.11 ± 2.94 days with an interwave interval (IWI) of 16.36 ± 0.37 days. The IWI within an individual was repeatable (0.88) and between the animals was variable (P < 0.001). Plasma E2 concentration profiles showed a wave like pattern. The peak plasma E2 concentrations were attained approximately 12 days after beginning of the growth phase, when the largest DF grew to a diameter of 18.7 mm. Plasma concentration of P4 was below 1.0 ng/mL in 85% of waves and above 1.0 ng/mL in 15% of the waves for a period of 3 to 6 days in the absence of spontaneous ovulation. It is concluded that ovarian follicular development and plasma E2 concentrations occurs in a wave like pattern in dromedary camels and the IWI and follicle numbers recruited per wave are variable between the animals and repeatable within an individual animal.     

Gene effects for specific leaf area and harvest index in three crosses of groundnut (Arachis hypogaea)

Groundnut productivity is low in the semi‐arid tropics mainly because of drought caused by low and erratic rainfall. Genotypes that have ability to use limited available water efficiently are required to enhance productivity of the crop. In groundnut, water use efficiency (WUE) is correlated with specific leaf area (SLA). The latter can be used as a surrogate trait for selecting for WUE. Partitioning of assimilates as measured by the harvest index (HI) has the greatest effect on pod yield. In order to improve SLA and in turn WUE and HI, a good knowledge of genetic systems controlling the expression of these traits is essential for the choice of an efficient breeding procedure. This study was conducted to investigate inheritance of SLA and HI in three crosses involving Chico, TMV 2 NLM, and ICGV 86031 groundnut genotypes. The study included parents, F1, F2, and backcross generations. Generation means analysis indicated that the additive effects were more important than the dominance effects in the expression of SLA and HI. In addition to additive and dominance effects, additive × additive type of epistasis, which can be fixed in groundnut (a self pollinated crop), was also significant for SLA in all the three and for HI, in two crosses. The selection for SLA and HI can be effective in early generations in some crosses and to exploit the additive × additive type of interaction, it can be done in large populations of later generations.     

The effect of dietary nickel on the immune responses of Spodoptera litura Fabricius larvae

By exposing Spodoptera litura Fabricius larvae to nickel (Ni) in artificial diets for successive three generations, we investigated the impacts of the dietary Ni on growth and immune response of the fifth and sixth instar larvae at 24 h intervals. The time of newly moulted fifth instar larvae was labelled as 0 h. After exposure to 5 mg/kg Ni for two generations, Ni exposure significantly improved larval phenoloxidase activity and encapsulation grade in fifth instar larvae when compared to controls, except for encapsulation grade at 72-120 h in the second generation. However, higher concentrations of Ni (≥10 mg/kg) only significantly reduced encapsulation grade at 72-120 h. In the third generation, insects given higher dietary levels of Ni (≥10 mg/kg) showed lower immune responses and retarded relative growth rate (RGR) compared to controls, but those exposed to lower Ni levels (≤5 mg/kg) had a significantly improved encapsulation grade at 24-72 h. Larvae at lower Ni level (≤5 mg/kg) treatments had significantly higher RGR in comparison with that in controls. There was no significant difference in food relative consumption rate (RCR) and RGR among any treatment of the fifth instar larvae in three successive generations. These results indicated that the type and extent of effects on growth and immune responses of S. litura varied with the Ni concentrations and exposure periods.     

Opposite latitudinal gradients in projected ocean acidification and bleaching impacts on coral reefs

Coral reefs and the services they provide are seriously threatened by ocean acidification and climate change impacts like coral bleaching. Here, we present updated global projections for these key threats to coral reefs based on ensembles of IPCC AR5 climate models using the new Representative Concentration Pathway (RCP) experiments. For all tropical reef locations, we project absolute and percentage changes in aragonite saturation state (Ωarag) for the period between 2006 and the onset of annual severe bleaching (thermal stress >8 degree heating weeks); a point at which it is difficult to believe reefs can persist as we know them. Severe annual bleaching is projected to start 10–15 years later at high‐latitude reefs than for reefs in low latitudes under RCP8.5. In these 10–15 years, Ωarag keeps declining and thus any benefits for high‐latitude reefs of later onset of annual bleaching may be negated by the effects of acidification. There are no long‐term refugia from the effects of both acidification and bleaching. Of all reef locations, 90% are projected to experience severe bleaching annually by 2055. Furthermore, 5% declines in calcification are projected for all reef locations by 2034 under RCP8.5, assuming a 15% decline in calcification per unit of Ωarag. Drastic emissions cuts, such as those represented by RCP6.0, result in an average year for the onset of annual severe bleaching that is ~20 years later (2062 vs. 2044). However, global emissions are tracking above the current worst‐case scenario devised by the scientific community, as has happened in previous generations of emission scenarios. The projections here for conditions on coral reefs are dire, but provide the most up‐to‐date assessment of what the changing climate and ocean acidification mean for the persistence of coral reefs.