Effects of rearing and induction temperature on the temporal dynamics of heat shock protein 70 expression in a butterfly

The temporal dynamics of heat shock protein 70 (HSP70) expression in response to longer‐term acclimation and rapid hardening in the butterfly Lycaena tityrus is investigated. After a 1‐h exposure to 1 °C or 37 °C, HSP70 is quickly up‐regulated within 1 h and down‐regulated within 2 h. The fast dynamic of HSP70 expression is in contrast to the patterns found in organisms inhabiting more stable thermal environments, and is interpreted as an adaptation to the large and rapid temperature variation experienced by flying ectotherms. HSP70 expression is higher in males than in females, as well as in animals reared at 27 °C than at 20 °C, although it is very similar across the high and low induction temperatures. Animals reared at the higher temperature, however, respond less strongly to high‐temperature stress.     

Reproductive allometry does not explain the temperature-size rule in water striders (Aquarius remigis)

Most ectotherms follow the temperature-size rule meaning that individuals growing up under cool conditions are larger as adults than those growing up in warm conditions. This pattern is difficult to explain because growth is usually slower in the cold meaning it takes longer to reach a larger adult size. One potential explanation for this pattern is that the typical increase in fecundity with body size is steeper in cool environments than it is in warm environments. As such, the relative gain in fecundity for being larger in the cold would compensate for the extra time it takes to grow to that size. We present the first empirical test of this model using the water strider Aquarius remigis. Individuals were reared at either 20° or 25°C with subsampling at each instar to estimate growth trajectories. At adult eclosion, half were switched to the alternate temperature and all females isolated, mated, and reproductive output measured for 3 weeks. We found that A. remigis does follow the temperature-size rule but that fecundity was highly plastic with respect to laying temperature such that the slightly greater fecundity of those reared at 20°C was due to their larger size. Overall, those laying at 25°C were more fecund and showed a positive relationship between body size and fecundity. Those laying at 20°C did not show a significant relationship between size and fecundity. As such, the reproductive allometry shows a pattern reverse to what would be needed for a larger size in the cold to be adaptive in this species. Although A. remigis follows the temperature-size rule over this temperature range, this is likely due to a constraint on growth and development rather than being adaptive plasticity.     

Biology ofCryptolaemus montrouzieri Mulsant [Coccinellidae: Coleoptera] in relation with temperature

The effect of temperature on the developmental duration ofCryptolaemus montrouzieri Mulsant was quantified by deriving a regression equation for each developmental stage as well as the total life cycle. While the duration of life stages was shorter during summer and longer during winter, the optimum constant temperature for maximal development was found to be 30°C. The adult longevity was extended when reared at 20°C than at 30°C and ambient temperature. The longevity of adults was longer when maintained on grape mealybugMaconellicoccus hirsutus (Green) than on honey and when maintained at 20°C. The fecundity of the predator was higher at 30°C than at 20°C. Eventhough the adults could survive at 10°C, the productive capacity was impaired.      

Effects of temperature on the development of overwintering immature stages of the near‐threatened butterfly Leptalina unicolor (Bremer & Grey) (Lepidoptera: Hesperiidae)

Overwintering larvae of multivoltine and univoltine populations of Leptalina unicolor were reared under various constant and fluctuating temperatures superimposed on a photoperiod of either 12 h of light and 12 h of darkness (LD 12:12) or LD 15:9. Diapause of the larvae terminated in midwinter (by early February). All the larvae of both populations pupated after two molts without feeding and the head capsule width of the final instar larvae was smaller than that of the penultimate instar ones. The photoperiod did not significantly affect larval development, but long‐day conditions (LD 15:9) hastened pupal development. The thermoperiod had a significant effect on the development of the multivoltine population. When multivoltine population larvae were kept under a low fluctuating temperature regime (cryophase/thermophase = 14/20°C), the period until adult eclosion was shorter than that under a constant temperature of 17°C. On the contrary, when larvae were kept under a high fluctuating temperature regime (24/30°C), the period until adult eclosion was longer than that under a constant temperature of 27°C. However, the univoltine population did not show such a reaction to the fluctuating temperature. The durations of final instar larva and pupa of the multivoltine population were shorter than those of the univoltine population. The developmental zeros of penultimate and final instar larvae and pupae of the univoltine population were lower than those of the multivoltine population. The head capsule width of penultimate instar larvae and the forewing length of adults of the univoltine population were larger than those of the multivoltine population for both sexes.     

Thermal responses in the citrus fruit fly,Dacus tsuneonis: evidence for a pupal diapause

Thermal responses controlling pupariation and adult eclosion in a citrus fruit fly,Dacus tsuneonis (Miyake), were studied to understand the winter biology of this species. When mature larvae were exposed to various temperature conditions, the highest percentage of pupariation was obtained at 15 °C, although the variance at this temperature was greater than at 20 °C or 25 °C. Pupariation occurred most rapidly at 20 °C and an alternating temperature with a mean of 15 °C. At constant 15 °C, pupae failed to emerge as adults. Pupae were characterized by a reduced respiration rate, which is typical of a diapausing pupa. When insects were stored at different temperatures for 45 days after pupariation, and then transferred to 25 °C, adult eclosion occurred earlier when the initial temperature was 10 °C than when it was 5 °C or 15 °C. Adult eclosion occurred most synchronously and pupal mortality was lowest when insects were stored at 15 °C for 90 days before incubation at 25 °C. These results strongly suggest thatD. tsuneonis enters a pupal diapause.     

Estimating reaction norms for predictive population parameters,age specific mortality,and mean longevity in temperature‐dependent cohorts of Culex quinquefasciatus Say (Diptera: Culicidae)

Culex quinquefasciatus plays a major role in the transmission of important parasites and viruses throughout the world. Because temperature is an important limiting factor on growth and longevity of all mosquito species, estimating the reaction norms provides very important basic information for understanding both plasticity and individual variations of the population. In the present study, Cx. quinquefasciatus were maintained at five different constant temperatures (15°, 20°, 23°, 27°, and 30°C) for two subsequent generations. Reproductive population parameters in blood‐fed mated females and longevities of virgin and blood‐fed mated adults reared at different temperatures were compared for the two generations. Longevity increased as temperature decreased within a range of 15° to 30°C for the unmated adults, and 15° to 27°C for the mated and blood‐fed adults. Generation times were as long as 124.07 and 106.76 days for two subsequent generations reared at 15°C, and the highest intrinsic rate of increase (r_m) values were estimated at 0.22 and 0.18, respectively, from the cohorts reared at 27°C. For survival rates, reproductive rates (R₀), and r_m values, 30°C was found to be a critical temperature for this species. These cohorts produced the smallest amount of eggs (R₀= 5.06), r_m values decreasing across generations (from 0.11 to 0.06), and the survival rates from egg to adult were found to be insufficient (16.1 and 10.8%). Additionally, the rate of exponential increase with age and age specific mortalities (b) were calculated for the virgin cohorts. Age specific mortality rates increased as temperature decreased. The increase in mortality rates started to accelerate at 27°C and was more pronounced at 30°C, for both females and males. We estimated the coefficients of variation for the b values in which females have smaller coefficients than those of the males at all temperatures.     

Effects of photoperiod and temperature on the rhythm and free‐running of adult eclosion in the Indian meal moth Plodia interpunctella

The rhythm of adult eclosion in the Indian meal moth Plodia interpunctella Hübner (Lepidoptera: Pyralidae) is investigated under various photoperiods and temperatures aiming to determine the nature of the temperature compensation and the free‐running period. Insects that are committed to a nondiapause larval development show diel rhythms of adult eclosion at 30, 25 and 20 °C. At 30 °C, the eclosion peak (i.e. the mean time of eclosion) occurs approximately 20 h after lights off under an LD 4 : 20 h photocycle, and at approximately 15 h under an LD 20 : 4 h photocycle. At 25 °C, the peak of eclosion occurs approximately 19 h after lights off under an LD 2 : 20 h photocycle and at approximately 16 h under an LD 20 : 4 h photocycle. At 20 °C, the eclosion peak is significantly advanced under long days of >12 h (i.e. approximately 20 h after lights off under an LD 4 : 20 h photocycle and approximately 9 h under an LD 20 : 4 h photocycle), indicating an effect of both lights‐off and lights‐on signals on the timing of the adult eclosion. To determine the involvement of a self‐sustained oscillator, the rhythm of adult eclosion is examined under darkness at different temperatures (30 to 21 °C). The mean free‐running periods are 22.4, 22.8, 22.0 and 22.5 h at 30, 24, 23 and 22 °C, respectively, indicating that the eclosion rhythm is temperature‐compensated. However, this rhythm does not free‐run under constant darkness at 21 °C. Because a clear diel rhythm is observed under 24‐h photocycles at 20 °C, the oscillator might be damped out within 24 h at the lower temperature.     

Factors affecting the cold hardiness of the peach-potato aphid Myzus persicae

Supercooling point studies were used to investigate the factors influencing the cold hardiness of the peach-potato aphid Myzus persicae, a freezing-susceptible insect. Overwintering adults lost cold hardiness as winter progressed, with a variable proportion showing a marked reduction in supercooling ability. Cold hardiness increased in spring so that all individuals demonstrated extensive supercooling ability typical of aphids reared in the laboratory at 20°C with a long photoperiod; these levels of cold hardiness were maintained in the field during summer and early autumn. First instar nymphs demonstrated considerable cold hardiness all year. Surface moisture caused inoculative freezing in some first instar nymphs and adults when supercooled, but the majority were unaffected. In the laboratory, adults starved for 7 days at 5°C showed distinct losses of supercooling potential equivalent to those observed in the field during mid to late winter. No loss of cold hardiness was found in first instar nymphs starved under the same conditions. The results demonstrate that the cold hardiness characteristics of M. persicae are atypical of those observed in other freezing-susceptible insects and it is suggested that continued feeding during mild winter conditions allows maintenance of cold hardiness particularly in adult aphids, and provides a possible explanation for the successful anholocyclic overwintering of M. persicae during such winters.     

Effects of acclimation on the thermal tolerance of the brown planthopper Nilaparvata lugens (Stål)

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Induction of adult diapause and of low and high reproductive states in a parasitoid wasp,Ooencyrtus nezarae, by photoperiod and temperature

Ooencyrtus nezarae Ishii (Hymenoptera: Encyrtidae) was reared on eggs ofRiptortus clavatus (Thunberg) (Heteroptera: Alydidae) at various temperatures under long-day (L16:D8) or short-day (L10:D14) conditions. There was no diapause during egg, larval or pupal stages under any set of conditions examined. However, at 15°C under short-day conditions, vitellogenesis was arrested in all adult females and they entered diapause. At 15°C under long-day conditions, or at 20°C under short-day conditions, some adult females entered diapause. Under the latter set of conditions, the adult females laid eggs but they laid fewer eggs than under long-day conditions, Even at 25°C, under short-day conditions, adult females laid fewer eggs than under long-day conditions, and this low rate of oviposition was attributed to the retarded development of ovaries. Diapause adults reared at 15°C were more resistant to low temperature than nondiapause adults reared at 25°C.     

Effects of brief exposures to low temperature on the development, longevity and fecundity of the grain aphid Sitobion auenae (Hemiptera: Aphididae)

First instar nymphs and adults of the grain aphid Sirobion auenae that had been reared at 10°C and 20°C over a number of generations, were cooled to -5°C and -10°C for 1 h and 6 h and returned to 20°C to assess the effects of brief exposures to low temperatures (cold-pulses) on their survival. rate of development, longevity and fecundity. A strong acclimation response was observed in first instar nymphs, with significantly less mortality in groups reared to 10°C compared to 20°C. Mean development time from first instar to adult was not significantly affected by low temperature exposure at the first nymphal stage. Longevity in all groups cooled as first instars was reduced by the sub-zero cold-pulses, and was also dependent on temperature and exposure time. Acclimated aphids survived longer than non-acclimated individuals. Reproductive rate, in terms of the number of nymphs born per aphid per day, was unaffected by cold stress applied at the first instar stage. Total fecundity was however reduced, being a function of the number and longevity of the survivors. Adult aphids were less cold hardy than nymphs; mortality was higher at -10°C than -5°C increasing with duration of exposure from 1 h to 6 h. Mean fecundity was reduced significantly in aphids cooled at the adult stage, the number of aphids born per day decreasing as the exposure period of the cold-pulse increased, suggesting that low temperature had affected embryogenesis. All the nymphs born to adults surviving exposure to -5°C for 6 h died within 48 h of birth, indicating that low temperature has a pre-natal effect on mortality.     

Maternal effect of low temperature on handedness determination in C. elegans embryos

C. elegans embryos, larvae, and adults exhibit several left-right asymmetries with an invariant dextral handedness, which first becomes evident in the embryo at the 6-cell stage. Reversed (sinistral) handedness was not observed among > 10,000 N2 adults reared at 16°C or 20°C under standard conditions. However, among the progeny of adults reproducing at 10°C, the frequency of animals with sinistral handedness was increased to ∼0.5%. Cold pulse experiments indicated that the critical period for this increase was in early oogenesis, several hours before the first appearance of left-right asymmetry in the embryo. Hermaphrodites reared at 10°C and mated with males reared at 20°C produced sinistral outcross as well as sinistral self-progeny, indicating that the low temperature effect on oocytes was sufficient to cause reversals. Increased frequency of reversal was also observed among animals developed from embryos lacking the egg shell. Possible mechanisms for the control of embryonic handedness are discussed in the context of these results, including the hypothesis that handedness could be dictated by the chirality of a gametic component. © 1996 Wiley-Liss, Inc.     

Life history and life table parameters of the predatory miceTyphlodromus talbii

The predatory miteTyphlodromus talbii Athias-Henriot occurs in European vineyards and is often associated with economically important species. Neither its role in vineyards nor the factors affecting its population dynamics and relationships with other phytoseiid species are well known. The development and the reproduction ofT. talbii were studied in the laboratory by rearing the predator on different kinds of food (Panonychus ulmi, Eotetranychus carpini, Colomerus vitis, Tydeus caudatus, Mesembryanthemum criniflorum pollen). Overwintered females reared on tydeids survived for long periods and laid eggs, but they died after a few days when spider mites or pollen were provided. Development occurred on all mite species but not on pollen. Developmental times on tydeids were shorter than on the other prey. Oviposition was recorded on tydeids and, to a lesser extent, on eriophyids but not on spider mites or pollen. Experiments on tydeids, which resulted as being the best food, were conducted at two temperatures (20° and 27°C). The highest temperature affected the duration of development and oviposition rates positively, but total fecundity was similar. Predators reared at 27°C consumed more prey than those reared at 20°C. The life table parameters of the species were evaluated onT. caudatus (at 20° and 27°C) and onC. vitis. The highest r_m ofT. talbii was found for individuals reared onT. caudatus at 27°C (0.165). Lower values were obtained on the same prey at 20°C (0.089) or onC. vitis (0.030). The feeding habits ofT. talbii may explain why the species coexists with the generalistAmblyseius aberrans orTyphlodromus pyri.     

Effect of thermoperiod amplitude on the adult eclosion rhythm of Indian meal moth,Plodia interpunctella (Lepidoptera: Pyralidae)

The thermoperiodic time cue is known to be one of several zeitgebers (Zt) that entrain an organism's biological rhythms. This study investigated the adult eclosion rhythm of the Indian meal moth, Plodia interpunctella, (Lepidoptera: Pyralidae) under various thermocycles in constant darkness. The insects were exposed to different thermoperiods (e.g. 25.5°C/24.5°C). Thermophase : cryophase ratios (TC) ranged from 4 h : 20 h to 20 h : 4 h. The peak in adult eclosion rhythm occurred at Zt 18.8 ± 4.4 (mean ± SD) on a given day and at Zt 0.7 ± 3.8 on the subsequent day under TC 4 h : 20 h and TC 20 h : 4 h conditions, respectively (start of temperature increase = Zt 0). Thus, longer thermophase periods delayed the peak in eclosion. The regression line obtained was then compared with that obtained under the temperature cycle of 30°C/20°C, and the two lines were found to lie parallel to each other. The peak in adult eclosion rhythm occurred earlier under the 25.5°C/24.5°C conditions than under the 30°C/20°C conditions. The results strongly suggest that the amplitude of the temperature cycle is an important factor determining the temporal position of the peak in adult eclosion. This is a significant finding among insect species.     

Development times and fecundity of three important arthropod pests of strawberry in the United Kingdom

The development rates and fecundity of three important pests of strawberry in the UK were determined over a range of temperatures. Development time of the strawberry tarsonemid mite, Phytonemus pallidus, from egg lay to adult, ranged from a mean of 28.4 days at 12.5°C to 8.8 days at 25°C. No nymphs developed to adult at 10°C. Females lived for up to 45 days and laid a mean of 24.3 and 28.5 eggs at 20°C and 25°C respectively. Total development time from egg lay to adult for the strawberry blossom weevil, Anthonomus rubi, ranged from a mean of 95.7 days at 10°C to 18.2 days at 25°C. Mean fecundity at 20°C was 157.6 eggs, and the oviposition period averaged 71.6 days. When nymphs were reared on strawberry, development of the European tarnished plant bug, Lygus rugulipennis, from egg lay to adult, ranged from 83.8 days at 15°C to 28.8 days at 25°C. Development times on groundsel were shorter and ranged from 65.6 to 22.2 days at 15°C and 25°C. Only two nymphs developed to adults at 10°C; no eggs hatched at that temperature. Mean fecundity at 20°C was 75.4 eggs, but ranged from 23 to 179. Under a fluctuating temperature regime of 10°C for 12 h:20°C for 12 h, nymphs of L. rugulipennis took 40.3 days to become adult on strawberry, and 33.4 days on groundsel. Simple linear models fitted the developmental rate ‐ constant temperature relationship well for all species, accounting for 95–98% of the total variation in observed developmental rates. Development under fluctuating temperatures illustrated the potential problem of extrapolating linear models beyond the conditions of the experiment.     

Influence of larval rearing temperature on the quality of cold‐stored Oomyzus sokolowskii Kurdjumov (Hymenoptera: Eulophidae)

Oomyzus sokolowskii, an important parasitoid of Plutella xylostella, has great potential for use in biological control. Storage at suboptimal temperature is valuable for increasing the shelf‐life of insect parasitoids. In this study, O. sokolowskii larvae were reared at 30/25, 25/25 and 25/20°C light/dark (65 ± 5% RH, 16 : 8 h L : D) until pupation. The pupae were then cold‐stored at 4 ± 1°C (60 ± 5% RH, full darkness). The pupae were removed out from the storage at 10, 20, 30 and 40 days after storage (DAS) and maintained at 25 ± 2°C until adults emerged or pupae died. Quality of the emerging adults and their F₁ offspring were assessed. Incidence of parasitism by O. sokolowskii was higher at 30/25°C than at 25/20°C. Cold storage of O. sokolowskii pupae greatly affected the fitness of the parasitoid: adult emergence rates were lower in the 40 DAS treatment than in other treatments; when O. sokolowskii larvae developed at 25/25°C, female proportions of the emerged adults were lower in the 40 DAS treatment than in the 0 and 10 DAS treatments. Larval rearing temperature mildly affected the adult emergence rate, post‐storage developmental time and female proportion with a few exceptions. Number of parasitoids emerged per host pupa, and incidence of parasitism by the females were neither affected by larval rearing temperature nor cold storage duration. Trans‐generational effects on F₁ offspring were evident in adult emergence rate, egg‐adult developmental time and female proportion which were negatively affected by long duration of storage (40 days), but not by larval rearing temperature with a few exceptions. In conclusion, O. sokolowskii pupae could be stored at 4°C for up to 30 days without significant fitness loss.     

Development and Survival of Chironomus tepperi Skuse (Diptera: Chironomidae) at a Range of Constant Temperatures

Chironomus tepperi Skuse were reared individually at a range of constant temperatures from 12.5 to 37.5°C (2.5°C intervals), with development and survival monitored at regular intervals. C. tepperi is protandrous, with males developing significantly faster than females at the majority of temperatures examined, due primarily to a shorter final instar. Some individuals completed development at all temperatures, however emergent adults failed to successfully inflate their appendages at 37.5°C. Developmental rate increased with increasing temperature up to 32.5°C, but fell at 35°C. Low adult emergence at 37.5°C precluded a reliable estimate of total development time at that temperature. Survival to adult emergence varied from 10 to 60%, with highest mortality in the pupal stage at all temperatures. Degree-days (DD) and developmental zero (DZ) estimates for egg to adult development are 150.5 DD and 10.5°C for males and 167.1 DD and 10.3°C for females. DD and DZ estimates are presented for each developmental stage. The significance of wide thermal tolerances in a colonist midge species is discussed.     

Life table and heat tolerance of Acyrthosiphon pisum (Hemiptera: Aphididae) in subtropical Taiwan

The effect of temperature on the life table of Acyrthosiphon pisum reared on Pisum sativum was evaluated under laboratory conditions using temperatures of 10, 15, 20, 25, 30, and 35°C. The development time of juvenile A. pisum decreased with increasing temperature (from 21.3 days at 10°C to 4.7 days at 35°C). Adult longevity also decreased with increasing temperature (from 53.2 days at 10°C to 2.3 days at 35°C). Interestingly, 70% and 25% of A. pisum nymphs reared at 30°C and 35°C, respectively, successfully developed into adults. These temperatures have previously been considered unsuitable for A. pisum development. However, adult aphids reared at 30°C and 35°C failed to reproduce. Linear regression analysis revealed that the lower development threshold of A. pisum was 153.1 degree‐days above 1.9°C. Maximal average reproductive capability was observed at 10°C for A. pisum adults, with each adult producing more than 120 nymphs. The intrinsic rate of increase (r_m) of A. pisum increased from 0.124/day at 10°C to 0.337/day at 25°C, whereas opposite trends were observed for the net reproductive rate (R₀) and the mean generation time (GT). At 20°C and 25°C, the intrinsic rate of increase of A. pisum was significantly higher than at 10°C and 15°C (P < 0.0001), indicating that 20°C and 25°C are within the optimal range for the growth of A. pisum, and that 30°C is beyond the upper threshold limit for reproduction, which involves a temperature range that is narrower than that of the survival range (upper limit is unknown, but above 35°C).     

Alternative life cycles controlled by temperature and photoperiod in the oligophagous bug, Dybowskyia reticulata

Abstract. Effects of temperature and photoperiod on the induction and re-induction of adult diapause were examined in Dybowskyia reticulata (Dallas) (Heteroptera: Pentatomidae). Adults collected from the field after overwintering in early summer continued oviposition under long-day conditions of LD 16:8 h at 20 or 25°C, while they re-entered diapause under short-day conditions of LD 12:12 h at 25, 27.5 or 30°C. By contrast, adults reared in the laboratory from eggs at 20 or 25°C entered diapause under both long-day and short-day conditions, whereas those reared at 27.5 and 30°C entered diapause only under short-day conditions. Under quasi-natural conditions in 1993, when summer temperature was low, most adults of the first generation entered diapause in late July. However, in the warmer summer of 1996, oviposition was recorded in many females that ecdysed into adults from July to early August. Even though the seeds of the host plants occur in a restricted period from early summer to early autumn, in warmer years D. reticulata may produce a second generation. The response to temperature with a threshold between 25 and 27.5°C in D. reticulata brings about a switch between the univoltine and bivoltine life cycles.     

Comparison of cold hardiness and sugar content between diapausing and nondiapausing pupae of the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae)

Abstract. To understand overwintering of the cotton boll worm Helicoverpa armigera, cold hardiness and sugar content are compared between diapausing and nondiapausing pupae. Diapausing and nondiapausing pupae reared at 20 °C under short and long photoperiods are acclimatized with a reduction of 5 °C per 5 days to 0 °C. When the acclimation temperature reaches 0 °C, the survival of diapausing pupae is assessed. The survival gradually decreases as the period of treatment progresses and approximately half survive for 112 days. However, nondiapausing pupae survive only 14 days after exposure to 0 °C. The surpercooling points of nondiapausing, diapausing and acclimatized pupae are approximately −17 °C. The major sugars contained in pupae are trehalose and glucose. Even though trehalose contents in diapausing pupae (initial level: 0.6 mg 100 mg⁻¹ fresh weight) increase significantly during cold acclimation and continue increasing until 58 days after exposure to 0 °C (maximum level: 1.8 mg 100 mg⁻¹), glucose is maintained at low levels (0.02 mg 100 mg⁻¹) for 56 days at 0 °C. However, glucose contents increase (maximum level: 0.8 mg 100 mg⁻¹) with decreasing contents of trehalose 84 days after exposure to 0 °C. Glycogen content gradually decreases during cold acclimation. When nondiapausing pupae are acclimatized with a reduction of 5 °C per 5 days to 5 °C from the beginning of pupation until the eyespots move, trehalose content increases (maximum level: 1.0 mg 100 mg⁻¹). Glucose contents in nondiapausing pupae increase before eclosion (0.09 mg 100 mg⁻¹). From these results, diapausing pupae of H. armigera can overwinter in regions where average winter temperatures are higher than 0 °C, but nondiapausing pupae cannot.