Selection for developmental time in bean weevil (Acanthoscelides obtectus): correlated responses for other life history traits and genetic architecture of line differentiation

In this article we investigate the direct and correlated responses to selection for developmental time in order to discern differences between lines in several preadult and adult life history traits of Acanthoscelides obtectus (Coleoptera, Bruchidae). Selection for fast development was about five times as effective as selection for slow development, as judged by realized heritabilities. The correlated responses on the following life‐history traits were studied: egg size, hatching success, embryonic developmental time, egg‐to‐adult viability, body weight, first day of egg laying, total fecundity, and longevity. Analyses of the terminal generation of selection showed that all life history traits examined, except for hatching success, were affected by selection. The findings suggest that body weight, total fecundity, and longevity traded off to preadult developmental time. Unlike the adult traits, none of the preadult traits showed negative correlations with developmental time. We also present data concerning the underlying genetic basis that produces changes in preadult developmental time, body weight, and egg‐to‐adult viability in the lines selected for fast and slow preadult developmental time. Additive‐dominance genetic architecture for both preadult developmental time and body weight was found. In addition, it appears that the responses to selection for preadult developmental time involved between 10 and 28 loci, which were correlated with at least one to four genes for body weight. Epistasis makes a significant contribution to genetic divergence between fast and slow selected lines only with respect to preadult viability. The observed levels of dominance and epistasis underscore the important role of nonadditive genetic effects to the adaptive diversifications of bean weevil populations.     

EXPERIMENTAL EVOLUTION OF ACCELERATED DEVELOPMENT IN DROSOPHILA. 1. DEVELOPMENTAL SPEED AND LARVAL SURVIVAL

Developmental time is a trait of great relevance to fitness in all organisms. In holometabolous species that occupy ephemeral habitat, like Drosophila melanogaster, the impact of developmental time upon fitness is further exaggerated. We explored the trade-offs surrounding developmental time by selecting 10 independent populations from two distantly related selection treatments (CB_1-5 and CO_1-5) for faster development. After 125 generations, the resulting accelerated populations (ACB_1-5 and ACO_1-5) displayed net selection responses for development time of -33.4 hours (or 15%) for ACB and -38.6 hours (or 17%) for ACO. Since most of the change in egg-to-adult developmental time was accounted for by changes in larval duration, the “accelerated” larvae were estimated to develop 25-30% faster than their control/ancestor populations. The responses of ACB and ACO lines were remarkably parallel, despite being founded from populations evolved independently for more than 300 generations. On average, these “A” populations developed from egg to adult in less than eight days and produced fertile eggs less than 24 hours after emerging. Accelerated populations showed no change in larval feeding rate, but a reduction in pupation height, the latter being a trait relating to larval energetic expenditure in wandering prior to pupation. This experiment demonstrates the existence of a negative evolutionary correlation between preadult developmental time and viability, as accelerated populations experienced a severe cost in preadult survivorship. In the final assay generation, viability of accelerated treatments had declined by more than 10%, on average. A diallel cross demonstrated that the loss of viability in the ACO lines was not due to inbreeding depression. These results suggest the existence of a rapid development syndrome, in which the fitness benefits of fast development are balanced by fitness costs resulting from reduced preadult survivorship, marginal larval storage of metabolites, and reduced adult size.     

Life-history parameters of western flower thrips on susceptible and resistant cucumber genotypes

The life-history parameters reproduction rate, developmental time and age specific survival of the western flower thrips,Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidae], were determined on susceptible and resistant cucumber (Cucumis sativus L.) genotypes. Both newly emerged andF. occidentalis females of mixed ages showed a substantial reduction (36 to 50%) of the reproduction rate on all resistant genotypes, in particular after the second day. On the resistant genotypes 9127 and 9140,F. occidentalis had a prolonged developmental period. This was primarily due to a prolongation of the second larval stage. On all resistant genotypes,F. occidentalis suffered from high (82 to 97%) preadult mortality, predominantly at the second larval stage. It is conclude that the resistant genotypes do not cause an immediate intoxication of adult nor preadult thrips stages.     

LABORATORY EVOLUTION OF LIFE-HISTORY TRAITS IN THE BEAN WEEVIL (ACANTHOSCELIDES OBTECTUS): THE EFFECTS OF SELECTION ON DEVELOPMENTAL TIME IN POPULATIONS WITH DIFFERENT PREVIOUS HISTORY

In this study we examined the direct and correlated responses for fast and slow preadult development time in three laboratory populations of the bean weevil (Acanthoscelides obtectus). The first population (“base,” B) has experienced laboratory conditions for more than 10 years; the second (“young,” Y) and the third (“old,” O) populations were selected for early and late reproduction, respectively, before the onset of the present experiments. All three populations are successfully selected for both fast and slow preadult development. The realized heritabilities are very similar in all populations, suggesting a similar level of the additive genetic variance for preadult development. We studied the correlated responses on the following life-history traits: egg-to-adult viability, wet body weight, early fecundity, late fecundity, total realized female fecundity, and adult longevity. All life-history traits examined here, except for the egg-to-adult viability, are affected by selection for preadult development in at least in one of the studied populations. In all three populations, beetles selected for slow preadult development are heavier and live longer than those from the fast-selected lines. The findings with respect to adult longevity are unexpected, because the control Y and O populations, selected for short- and long-lived beetles, respectively, do not show significant differences in preadult development. Thus, our results indicate that some kind of asymmetrical correlated responses occur for preadult development and adult longevity each time that direct selection has been imposed on one or the other of these two traits. In contrast to studies with Drosophila, it appears that for insect species that are aphagous as adults, selection for preadult development entails selection for alleles that also change the adult longevity, but that age-specific selection (applied in the Y and O populations) mostly affects the alleles that have no significant influence on the preadult development. Implications of these findings on the developmental and evolutionary theories of aging are also discussed.     

Effect of host plants on developmental time and life table parameters of <Emphasis Type="Italic">Amphitetranychus viennensis</Emphasis> (Acari: Tetranychidae)

The effect of host plant species including black cherry (Prunus serotina cv. Irani), cherry (Prunus avium cv. siahe Mashhad) and apple (Malus domestica cv. shafi Abadi) was studied on biological parameters of Amphitetranychus viennensis (Zacher) in the laboratory at 25 ± 1°C, 70 ± 10% RH and 16L: 8D photoperiod. Duration of each life stage, longevity, reproduction rate, the intrinsic rate of natural increase (r _m ), net reproductive rate (R ₀ ), mean generation time (T), doubling time (DT), and finite rate of increase (λ) of the hawthorn spider mite on the three host plants were calculated. Differences in fertility life table parameters of the spider mite among host plants were analyzed using pseudo-values, which were produced by jackknife re-sampling. The results indicated that black cherry might be the most suitable plant for hawthorn spider mite due to the shorter developmental period (10.6 days), longer adult longevity (25.5 days), higher reproduction (65.6 eggs), and intrinsic rate of natural increase (0.194 females/female/day). Cherry was the least suitable host plant. To determine the effect of host shifts, the mite was transferred from black cherry onto cherry and apple. In the first generation after shifting to apple, the developmental period, reproduction and life table parameters were negatively influenced. However, population growth parameters in the first generation on cherry were actually better than after three generations on this new host. This underscores the relevance of the mites’ recent breeding history for life table studies.     

Testing competing measures of profitability for mobile resources

Optimal diet theory often fails to predict a forager’s diet choice when prey are mobile. Because they escape or defend themselves, mobile prey are likely to increase the forager’s handling time, thereby decreasing its fitness gain rate. Many animals have been shown to select their prey so as to maximize either their fitness gain or their fitness gain rate. However, no study has yet compared directly these two measures of profitability by generating testable predictions about the choice of the forager. Under laboratory conditions, we compared these two measures of profitability, using the aphid parasitoid Aphidius colemani and its host, Myzus persicae. Fitness gain was calculated for parasitoids developing in each host instar by measuring life-history traits such as developmental time, sex ratio and fecundity. Fitness gain rate was estimated by dividing fitness gain by handling time, the time required to subdue the host. Fourth instar aphids provided the best fitness gain to parasitoids, whereas second instar aphids were the most profitable in terms of fitness gain rate. Host choice tests showed that A. colemani females preferred second instar hosts, suggesting that their decision maximizes fitness gain rate over fitness gain. Our results indicate that fitness gain rate is a reliable predictor of animal’s choice for foragers exploiting resources that impose additional time cost due to their mobility. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Effect of African nightshade species (Solanaceae) on developmental time and life table parameters of <Emphasis Type="Italic">Tetranychus evansi</Emphasis> (Acari: Tetranychidae)

The effect of five African nightshade (Solanum sp.) species on the biological and demographic parameters of the tomato spider mite, Tetranychus evansi Baker and Pritchard, was examined in the laboratory at 25 ± 1°C, 70–80% RH and 12L:12D photoperiod. Duration of each development stage, reproduction rate, longevity, intrinsic rate of natural increase (r _m), and doubling time (DT) of the tomato spider mite on the five nightshade species were calculated. The results indicated that S. villosum, S. scabrum, S. tarderemotum and S. americanum are more suitable for T. evansi due to a shorter developmental period, longer adult longevity, higher reproduction and intrinsic rate of natural increase ranging between 0.180 and 0.196 females/female/day compared with S. sarrachoides which cannot support T. evansi populations as the r _m (−0.063 females/female/day) and DT were negative on this host. Differences in developmental time and life table parameters among the other host plants were not significant.     

Comparative life table statistics for the ladybeetle Scymnus syriacus reared on the green citrus aphid,Aphis spiraecola,fed on two host plants

Demographic statistics for the ladybeetle Scymnus syriacus Marseul (Coleoptera: Coccinellidae) were investigated when reared on the green citrus aphid, Aphis spiraecola Patch (Homoptera: Aphididae), fed on two host plants: spirea (Spirea sp.) and Thompson orange (Citrus sinensis cv. Thompson) under laboratory conditions (25 ± 1°C, 65 ± 5% relative humidity and 16 h light : 8 h dark). Mean developmental times from egg to adult were 20.32 ± 0.18 and 22.07 ± 0.15 days for spirea and Thompson orange, respectively. The survival rate from egg to adult was higher for spirea (85%) than for Thompson orange (72.5%). The oviposition periods were 41.8 ± 2.47 and 39.71 ± 2.6 days and the total number of eggs per female were 657.6 ± 29.5 and 587.9 ± 32.6 on spirea and Thompson orange, respectively. Fertility life tables were also constructed to compare the effects of host plants. The intrinsic rate of increase (r_m) on spirea was higher than that on Thompson orange. Jackknife estimates of r_m varied from 0.149 on spirea to 0.133 on Thompson orange. The mean population generation times on these host plants were 38.16 and 40.65 days, respectively. These results indicate that spirea‐fed A. spiraecola is more suitable for S. syriacus, producing a higher survival rate and shorter developmental time.     

Sublethal effects of Beauveria bassiana on life table parameters of two–spotted spider mite,Tetranychus urticae (Acari: Tetranychidae)

Effects of the entomopathogenic fungus Beauveria bassiana were studied on life table parameters of two-spotted spider mite, Tetranychus urticae feeding on bean and cucumber under laboratory conditions. The developmental periods for all immature stages were not affected by fungal infection on each host plant but the duration of larval stage was significantly longer on bean. The female and male longevity, oviposition period and fecundity were significantly lower on fungus-treated mites but were not different between two host plants. Significant reductions were found on the intrinsic rate of increase (r _m ), the net reproductive rate (R ₀), the finite rate of increase (λ), the mean generation time (T _c ) and the population doubling time (D _t ) as a result of mycosis. Only the mean generation time (T _c ) was influenced considering the effect of host plant, which was shorter on cucumber.     

Development of a positive preference–performance relationship in an oligophagous beetle: adaptive learning?

The relationship between oviposition preference and larval performance is a central topic in insect–plant biology. In this study, we investigate whether the oligophagous flea beetle, Altica fragariae Nakane (Coleoptera: Chrysomelidae), exhibits a positive preference–performance relationship, and whether oviposition preference develops over time. We tested the beetles using four sympatric plant species: Duchesnea indica (Andrews) Focke (the normal host plant), Agrimonia pilosa Ledeb. (a secondary host plant), and Potentilla chinensis Ser. and Sanguisorba officinalis L. (host plants of two related Altica species) (all Rosaceae). In no‐choice experiments, both oviposition rate and offspring fitness parameters (eclosion rate, development time, and body mass) were highest on D. indica. Oviposition rate was much lower on P. chinensis than on A. pilosa, whereas offspring fitness parameters did not differ significantly between these two host plants. Offspring fitness were lowest for S. officinalis, and adult females refused to oviposit on this acceptable non‐host in a no‐choice situation. Repeated two‐choice experiments showed that the proportion of oviposition on one of the novel host plants decreased significantly over time when the alternative host plant was D. indica. In repeated two‐choice experiments using A. pilosa and P. chinensis, females mainly fed on A. pilosa but distributed their eggs equally over the two host plants, in accordance with the lack of difference in offspring fitness on those hosts. Together, these results showed that A. fragariae females develop a positive preference–performance relationship over time. We suggest that A. fragariae achieves this through adaptive learning of oviposition preference: not only does the female learn to discriminate among the host plants when there is a fitness difference for her offspring, but the female also fails to discriminate when there is no fitness difference.     

外寄生性花绒寄甲的寄生选择及其发育表现

"选择-表现"假说认为,成虫应该选择有利于子代发育的高品质寄主,但在寄主选择中,除了寄主品质外,其他因素也可能影响寄主选择决策。寄主选择研究通常以成虫为对象,而对那些初龄幼虫选择寄主的寄生性昆虫很少关注。以1龄幼虫积极搜寻寄主的寄生性花绒寄甲为模式生物,采用双选试验设计,观察了花绒寄甲初孵幼虫在不同体重青杨天牛幼虫之间、在已被寄生与健康的黄粉虫蛹之间的寄生选择性;然后采用回归设计,观察了花绒寄甲寄生若干不同体重的青杨天牛幼虫后的发育表现。研究结果表明,花绒寄甲1龄幼虫对体型较大的青杨天牛幼虫的选择偏好显著大于对体型较小的寄主幼虫的选择,选择大体型幼虫的比值比是选择小体型幼虫的4.55倍;对已被寄生的寄主黄粉虫蛹的选择偏好显著大于对健康寄主蛹的选择,选择已被寄生寄主的比值比是选择健康寄主的12.57倍。寄生青杨天牛幼虫的花绒寄甲幼虫发育历期平均为11.49 d、蛹历期为26.67 d、幼虫发育至成虫的羽化率50%,这些发育表现与寄生时青杨天牛幼虫的体重没有显著关系。但刚羽化寄甲成虫体重与寄生时寄主的体重存在显著的正直线关系:寄生时的寄主体重每增大0.01 g,羽化出的寄甲成虫体重增大近0.08%;方差分析寄甲成虫体重在不同寄主体重水平之间的差异表明,从体型较大寄主中羽化的寄甲成虫体重显著大于从体型较小寄主中羽化的成虫。研究结果说明,花绒寄甲初孵幼虫在寄主选择决策时,在寄主体型大小与被寄生状态之间可能采取折衷对策,而且对体型大小不同的寄主选择与子代发育适合度表现存在一致性,从而支持"选择-表现"假说。     

THE EVOLUTION OF DEVELOPMENT IN DROSOPHILA MELANOGASTER SELECTED FOR POSTPONED SENESCENCE

The role of development in the evolution of postponed senescence is poorly understood despite the existence of a major gerontological theory connecting developmental rate to aging. We investigate the role of developmental rate in the laboratory evolution of aging using 24 distinct populations of Drosophila melanogaster. We have found a significant difference between the larval developmental rates of our Drosophila stocks selected for early (B) and late-life (O) fertility. This larval developmental time difference of approximately 12% (O > B) has been stable for at least 5 yr, occurs under a wide variety of rearing conditions, responds to reverse selection, and is shown for two other O-like selection treatments. Emerging adults from lines with different larval developmental rates show no significant differences in weight at emergence, thorax length, or starvation resistance. Long-developing lines (O, CO, and CB) have greater survivorship from egg to pupa and from pupa to adult, with and without strong larval competition. Crosses between slower developing populations, and a variety of other lines of evidence, indicate that neither mutation accumulation nor inbreeding depression are responsible for the extended development of our late-reproduced selection treatments. These results stand in striking contrast to other recent studies. We argue that inbreeding depression and inadvertent direct selection in other laboratories' culture regimes explain their results. We demonstrate antagonistic pleiotropy between developmental rate and preadult viability. The absence of any correlation between longevity and developmental time in our stocks refutes the developmental theory of aging.     

Sublethal effects of indoxacarb on the diamondback moth, <Emphasis Type="Italic">Plutella xylostella</Emphasis> (L.) (Lepidoptera: Yponomeutidae)

The effects of sublethal concentrations of indoxacarb on Plutella xylostella (L.) (Lepidoptera: Yponomeutidae) were examined. In comparison with the control, sublethal concentrations led to a significant decrease in adult emergence, the percentage of pupation and pupal weight of the parent generation, fecundity, and adult longevity. They also significantly decreased the net reproductive rate (R ₀), intrinsic rate of increase (r _m), finite rate of increase (λ), and gross reproduction rate (GRR), and had significantly increased egg developmental time and doubling time (Dt) of P. xylostella. However, sublethal concentrations of indoxacarb had no significant affect on adult emergence, percentage of pupation, larva, prepupa and pupa developmental periods, hatchability, and sex ratio of offspring. The results of this study indicated that sublethal concentrations can decrease the biological parameters of P. xylostella.     

RESPONSES AND CORRELATED RESPONSES TO ARTIFICIAL SELECTION ON THORAX LENGTH IN DROSOPHILA MELANOGASTER

Two sets of four replicate lines of Drosophila melanogaster were selected for large and small thorax with controls. F, progeny of crosses between the selected lines within each size category showed (a) a reduction in preadult viability in large lines relative to control and small lines when they were cultured at medium or high density in competition with a standard mutant marked competitor stock, and (b) an increase in larval development time in large lines relative to control and small lines. Natural selection for increased body size in adults may therefore be opposed by adverse effects on larval viability. The results are discussed in terms of the developmental mechanisms probably responsible for the change in body size. The preadult survival of the large and control lines was measured at three different temperatures, and there was no evidence for a significant interaction between size and temperature. The observed evolutionary increase in body size in response to reduced temperature in Drosophila must therefore involve either different genes from those subject to selection for size at a single temperature, or a fitness component other than preadult survival. There was no significant asymmetry in response to selection, and thorax length showed heterosis in crosses between the selected lines.     

Comparison of larval performance and oviposition preference of Spodoptera frugiperda among three host plants: Potential risks to potato and tobacco crops

Fall armyworm (FAW), Spodoptera frugiperda (J.E. Smith), is a major polyphagous pest with the potential to seriously damage various crops. A better understanding of FAW's performance on different hosts may help to predict which plants will be attacked when the preferred host is absent, and facilitate the development of effective pest management practices. We compared the larval performance of FAW fed on maize with that of FAW fed on potato and tobacco, which are important crops in China, using an age-stage two-sex life table and adult female oviposition preference experiments. In cage experiments with potato, tobacco, or maize as the host, FAW reared on maize exhibited the strongest performance with shorter developmental time in the larval stage, longer longevity, and a higher reproductive rate in adults. Females oviposited on maize in preference to potato or tobacco. Compared with larvae fed on maize, those fed on potato and tobacco exhibited significantly lower survival, with only 31.61% and 8.13% developing to the adult stage, respectively. Several life table parameters, including the mean generation time (T), net reproductive rate (R₀), finite rate of increase (λ), and intrinsic rate of natural increase (r) were negatively affected in FAW fed on potato and tobacco. Our results support the preference–performance hypothesis, that is, that herbivore females maximize fitness by choosing host plants associated with strong larval performance. Although larvae and adults performed poorly on potato and tobacco, their offspring will be capable of establishing populations on them, posing a potential threat to these crops in China.     

Temperature‐dependent development of Neoseiulus barkeri (Acari: Phytoseiidae) on Tetranychus urticae (Acari: Tetranychidae) at seven constant temperatures

Abstract The effect of seven constant temperatures of 15, 20, 25, 27, 30, 35 and 37°C on developmental time of Neoseiulus barkeri Hughes were determined in laboratory conditions under 65%± 5% RH and a photoperiod of 12 : 12 (L : D) h on nymphal stages of Tetranychus urticae Koch. Total developmental time of females (from egg to adult emergence) at the above‐mentioned temperatures was 26.59, 14.43, 6.32, 5.64, 4.59, 3.98 and 4.67 days, respectively. Developmental rate of the N. barkeri increased as temperature increased from 15 to 35°C, but declined at 37°C. A linear and two nonlinear models were fitted to developmental rate of immature stages of N. barkeri to predict the developmental rate as a function of temperature, as well as to estimate the thermal constant (K) and critical temperatures (i.e., T_min, T_opt and T_max). The estimated values of the T_min and K for total developmental time using the linear model were 12.07°C and 86.20 degree‐days (DD), respectively. The T_min and T_max estimated by the Sharpe‐Schoolfield‐Ikemoto (SSI) model were 11.90°C and 37.41°C, respectively. The estimated T_opt for overall immature stage development of N. barkeri by the Lactin and SSI models were 33.89°C and 24.51°C, respectively. Based on the biological criteria of model evaluation, the linear and SSI models were found to be the best models for describing the developmental rate of overall immature stages of N. barkeri and estimating the temperature thresholds.     

Adaptation of the gypsy moth to an unsuitable host plant

The pattern of adaptation with regard to life history traits and traits thought to be important in feeding habits of caterpillars in two populations of the gypsy moth (Lymantria dispar L.; Lepidoptera: Lymantriidae) originating from the locust tree (Robinia pseudoacacia; Fabaceae) and oak (Quercus petrea; Fagaceae) forests were investigated in the laboratory. The Robinia population has experienced unsuitable locust tree leaves as an exclusive food resource for more than 40 years. Since Quercus species are the principal host plants of the gypsy moth, the specific objectives of this study have been to measure the extent of differentiation between ancestral and derived populations in several life history traits (egg-to-adult viability, duration of larval and pupal stages, and pupal weight) and nutritional indices – relative growth rate (RGR), relative consumption rate (RCR), assimilation efficiency (AD), gross growth efficiency (ECI), and net growth efficiency (ECD). Significant differences between the Quercus and Robinia populations were detected in pupal duration, RGR, RCR, and AD. The presence of a significant population × host interaction in traits such as preadult viability, duration of pupal stage, RGR, and ECI suggests that adaptation of the gypsy moth to the unsuitable host might be ongoing. Using a full-sib design, we screened for genetic variation in life history traits within both populations, and examined the genetic correlations of performance across oak and locust leaves within both populations. The genetic variances for analyzed life history traits were lower under conditions that are commonly encountered in nature. Our data show that positive cross-host genetic correlations preponderate within both populations.     

Physiological and morphological development of Anagyrus ananatis at constant temperatures

The lower developmental temperature threshold (T ₀) and the Degree Days (DD) required for the encyrtid endoparasitoid Anagyrus ananatis Gahan to develop from egg to adult on the pink pineapple mealybug (PPM), Dysmicoccus brevipes (Cockerell) (Hemiptera: Pseudococcidae), were determined. The T ₀ was estimated to be about 12.65 °C for both females and males. In contrast, females and males required about 275 and 265 DD, respectively, to complete development from egg to adult. Temperatures from 19 to 29 °C were optimal for mass rearing of A. ananatis, with the optimal temperature being around 24 °C. At this temperature, A. ananatis could complete almost two generations in the time it takes PPM to complete only one generation. Although A. ananatis is a koinobiont, the mealybug host was killed within a few (6–8) days after parasitization. The developmental stages of A. ananatis were described (e.g., appearance, size, color) and their time periods quantified when reared on PPM at 23.5 ± 0.5°C. Encyrtiform eggs were inserted through the dorsal surface of the PPM and were attached to the host via a slender stalk. This immature parasitoid remained attached to the host cuticle via the stalk until entering the prepupal stage. The host mealybug mummified during the parasitoid’s prepupal stage. First adult eclosion occurred at 24 days post-parasitization.