Effects of temperature and photoperiod on termination of pseudopupal diapause in the bean blister beetle, Epicauta gorhami

Larvae of the bean blister beetle, Epicauta gorhami (Coleoptera: Meloidae), feed on grasshopper eggs in soil and undergo hypermetamorphosis. This beetle undergoes larval diapause in the fifth instar as a pseudopupa, a form characteristic of hypermetamorphosis in meloid beetles. This pseudopupal diapause is maintained for a long period if the larvae are kept under continuous diapause-inducing short-day conditions. In the present study, the effects of temperature and photoperiod on termination of pseudopupal diapause in E. gorhami were examined using pseudopupae obtained under 12L-12D at 25°C. Diapause was terminated by exposure to low or high temperature, but not by transfer to long-day conditions. The pseudopupal stage comprises diapause and post-diapause phases despite its unchanging appearance. The optimum low temperature for diapause termination was 10-15°C, and a higher or lower temperature was less effective. There was an individual variation in the low temperature requirement for diapause termination and post-diapause development may not occur until this requirement is satisfied. Although under natural conditions pseudopupae may encounter low temperatures that are effective for diapause termination at different ages, our results show that pseudopupae at various ages respond similarly to low temperature exposure. This response may ensure that resumption of development synchronizes in a population.     

Environmental induction of larval diapause and life-history consequences of post-diapause development in the Large Copper butterfly,Lycaena dispar (Lepidoptera: Lycaenidae)

Many insects in temperate zones withstand the adverse conditions of winter through entering diapause and the two most important environmental stimuli that induce diapause are photoperiod and ambient temperature. The Large Copper butterfly, Lycaena dispar Haworth (Lepidoptera: Lycaenidae), is a Palearctic butterfly that hibernates as larvae. Since this butterfly is a near threatened species in some regions, there has been a growing need for a standardized protocol for mass rearing of this butterfly based on the adequate knowledge of its ecology. In the present study, we first identified that L. dispar larvae were sensitive to the photoperiodic induction of diapause during their first larval instar. We then investigated to what extent the diapause-inducing effects of photoperiod could be modified by ambient temperatures in L. dispar larvae by exposing them to the range of day-lengths (L:D 14:10, 12:12, 10:14 and 8:16) at three different temperatures (15, 20 and 25 °C). All larvae were induced to enter diapause at low ambient temperature (15 °C) regardless of photoperiod, whereas most of them (86 %) exhibited direct development when temperature was high (25 °C). The photoperiodic induction of diapause was evident when day-length was shorter than 14 h at intermediate temperature (20 °C). Pre-diapause development was prolonged at low temperatures. Finally, we found that post-diapause development of L. dispar larvae was determined by both the chilling temperature experienced by diapausing larvae and the duration of larval diapause. Adult emergence was enhanced when larvae were chilled at 8 °C and when they had been under the state of diapause for 20 days before they were treated to terminate diapause.     

Control of summer and winter diapause in the leaf-mining fly Pegomyia bicolor Wiedemann (Dipt., Anthomyiidae)

Effects of photoperiod and temperature on diapause induction and termination were investigated in both aestival and hibernal pupae of Pegomyia bicolor Wiedemann under field and laboratory conditions. In the field, summer diapause had occurred already in part of the first pupal population; the proportion of diapause gradually rose as the day length and temperature increased. This fly is a short-day species with a pupal summer and winter diapause. Summer diapause was induced by both long day-lengths and mild temperatures. The whole larval life is sensitive to photoperiod. Winter diapause was induced mainly by low temperatures, especially in the first 10 days after pupation. High temperatures strongly enhanced summer diapause induction regardless of photoperiod. The diapause-averting influence of short photoperiods was fully expressed only at moderately low temperatures. High temperatures delayed diapause development, resulting in a rather long summer diapause; whereas low temperatures hastened it, leading to a short winter diapause and showing a low thermal threshold for diapause development. In the field, the post-diapause development started in January, the coldest month, suggesting that the thermal requirements for post-diapause development is also low.     

Effects of temperature,photoperiod and soil humidity on induction of pseudopupal diapause in the bean blister beetle Epicauta gorhami

Larvae of the bean blister beetle Epicauta gorhami Marseul (Coleoptera: Meloidae) feed on grasshopper eggs in soil and undergo hypermetamorphosis. This beetle undergoes larval diapause in the fifth instar as a pseudopupa, a form characteristic of hypermetamorphosis in meloid beetles. The effects of temperature, photoperiod and soil humidity on larval development of E. gorhami are examined in a population in Miyazaki, Japan, using egg pods of Locusta migratoria L. as food. At lower temperatures (20 and 22.5 °C), all larvae become pseudopupae, regardless of the photoperiod. By contrast, at higher temperatures (27.5 and 30 °C), almost all larvae pupate at the end of the fourth instar, again regardless of the photoperiod. A long‐day photoperiodic response occurs only at an intermediate temperature (25 °C): under an LD 12 : 12 h photocycle, all larvae enter diapause, although the diapause incidence tends to decrease as the day length becomes longer. Pseudopupae are immobile and remain in diapause for ≥120 days when they are kept under the same conditions, except that diapause terminates within a relatively short time at 30 °C. Although lower soil humidity retards post‐feeding development, soil humidity has no effect on the diapause incidence. On the basis of the short developmental period and diapause avoidance under summer conditions, it is suggested that this beetle partially produces two generations a year in southwestern Japan.     

Diapause termination, post-diapause development and reproduction in the beet webworm, Loxostege sticticalis (Lepidoptera: Pyralidae)

Effects of photoperiod and cold exposure on diapause termination, post-diapause development and reproduction in Loxostege sticticalis were examined. Larvae were reared at diapause inducing condition (22 °C, L:D 12:12) consistently or transferred to long day photoperiod (L:D 16:8) and darkness (L:D 0:24) respectively, after entering into diapause. Diapause was terminated in approximately 40% of the larvae after 36 days, and no significant differences were observed between photoperiods, suggesting larval diapause was terminated spontaneously without being induced by photoperiods. Cold exposure significantly hastened diapause termination. The diapause termination incidence increased significantly with peaks of 98% at both 5 °C and 0 °C exposure for 30 days, as compared to 42% in controls not exposed to cold, while the mortality and number of days required for diapause termination decreased dramatically. The optimal low temperature exposure periods under 5 °C or 0 °C were 20 days and 30 days, showing a higher termination incidence and shorter time for diapause termination. This suggests that the low temperatures in winter play an important role in diapause termination under natural conditions. The threshold temperatures for post-diapause development in prepupae and pupae were 9.13 °C and 10.60 °C respectively, with corresponding accumulations of 125 and 200 degree-days. Adults that experienced larval diapause significantly delayed their first oviposition, oviposition period was prolonged, and the lifetime number of eggs laid was decreased, however both males and females have significantly longer longevity. The field validation of diapause termination, the degree-days model, and the relationship between diapause and migration in L. sticticalis were also discussed.     

Adult diapause and cold hardiness in Aulacophora nigripennis (Coleoptera: Chrysomelidae)

We investigated the control of diapause termination and seasonal changes of cold hardiness and polyol content in Aulacophora nigripennis. Adults were ready to start post-diapause development upon transfer to high temperature by late February irrespective of photoperiod. Photoperiod probably functions to maintain diapause before winter because adults resume reproductive development at a long photoperiod in autumn. Adults showed a decreased supercooling point (SCP), increased chill tolerance and high myo-inositol content during winter. Chill tolerance at 0 degrees C appears to be a more suitable indicator of their cold hardiness than SCP because they die at 0 degrees C without freezing and they normally have no chance of being exposed to low subzero temperatures close to their SCP. The temporal pattern for changes in chill tolerance was synchronized with that for fluctuations in myo-inositol content, indicating a possible causal relationship between the two phenomena.     

Diapause development in the chestnut weevilCurculio elephas

Larval diapause development in the chestnut weevilCurculio elephas (Coleoptera, Curculionidae) was studied in the laboratory at different temperatures. The results proved that exposure to low temperatures (3–6°C) in the period December–February is not required to complete diapause. The diapause is terminated in December and from January on, the larvae can initiate post-diapause morphogenesis in the laboratory, if temperatures allow it. In the field developmental rates are negligible during winter cold (4–6°C) and only after March morphogenesis can proceed with no interruption until adult emergence. Diapause and post-diapause quiescence contribute to individual synchronization for initiation of development. The observed spread of adult emergence was 30 days in the laboratory. This variability produced during post-diapause development may be a response to annual variation in the phenology of the chestnuts.     

Influence of photoperiod on low temperature acclimation for cold-hardiness in Drosophila auraria

ABSTRACT. Imagines of Drosophila auraria Peng, a reproductive diapause species, developed cold-hardiness at low temperatures to a greater extent when exposed to a diapause-inducing photoperiod (LD10:14 h) than when exposed to a diapause-preventing photoperiod (LD 16:8h). Imagines kept at 18°C, which was the temperature at which they were reared to eclosion, did not survive a test exposure to -5°C for 8 days regardless of age or photoperiod. When transferred to 10 or 5°C, either from eclosion or from 8 days after eclosion, the survival rate, on testing, rose with time since transfer and rose faster and higher with a photoperiod of LD 10:14h than with LD16:8h. Flies transferred to 15°C only showed improved ability to survive the test if they were kept in LD 10:14h. When cultured at 18°C to the age of 8 days after eclosion, diapause was terminated in about 30% of females even at LD 10:14h. In these post-diapause females the ability to develop cold-hardiness at lower temperatures was somewhat less than in the diapausing females, but apparently greater than in the non-diapause females. These results suggest that the physiological mechanism which promotes cold-hardiness under a diapause-inducing photoperiod is not directly linked to the process causing reproductive diapause.
In Sapporo, flies from a natural population became tolerant to cold in October when they entered diapause and daily mean temperature fell below 15°C and the light/dark cycle fell below LD 12:12h.     

Multiple role of temperature during insect diapause: a review

The review emphasizes that there are multiple pathways to diapause completion. The programmed course of events is modified by environmental cues. Often chilling is not a prerequisite for the completion of hibernation diapause (examples tabulated). Diapause completion progresses well at intermediate or high temperatures, sometimes it is even stimulated by high or increasing temperature. Low temperatures are important, as they (1) conserve metabolic reserves, (2) prevent resumption of post-diapause morphogenesis and thus synchronize the life-cycle, (3) represent contrast to the later increase in temperature. Diapause consists of phases with different prerequisites. There is a principal difference between diapause development and photoperiodic activation as indicated by the subsequent physiological condition of insects.     

Diapause and seasonal life cycle strategy in the house spider, Achaearanea tepidariorum (Araneae, Theridiidae)

Abstract In order to elucidate the mechanism regulating its seasonal life cycle, the photoperiodic response of Achaearanea tepidariorum has been analysed. Nymphal development was faster in long-day and slower in short-day photoperiods. The combined action of low temperature, poor food supply and short daylength induced diapause at an earlier developmental stage than short days alone. Thus, photoperiod is a primary factor inducing nymphal diapause, but the diapausing instar is influenced by both temperature and food supply. Hibernating nymphs became unresponsive to photoperiod in late December. After hibernation, however, sensitivity was restored and the nymphs remained sensitive to photoperiod throughout their life. This spider could also enter an imaginal or reproductive diapause. Photoperiod was again a primary inducing factor and temperature modified the photoperiodic response to some extent. The induction of the reproductive diapause was almost temperature-compensated whereas development was not. So the involvement of a photoperiodic counter system was suggested. Irrespective of whether the nymph had experienced diapause or not, the imaginal diapause was induced in response to a short-day photoperiod after adult moult. Based on these observations, the seasonal life cycle and the adaptive significance of nymphal and imaginal diapause are discussed.     

Factors affecting the natural duration of diapause and post-diapause development in the Mediterranean corn borer Sesamia nonagrioides (Lepidoptera: Noctuidae)

Sesamia nonagrioides responds to photoperiod for diapause termination at high temperature with a Type II response curve, and only photoperiods longer than 12:12 terminate diapause. However, these photoperiods never occur in the field when diapausing larvae are competent to terminate diapause. Under a temperature similar to the natural field temperatures diapause terminates spontaneously in approximately 4 months, which ensures that the larvae reach the middle of winter without pupation. S. nonagrioides larvae pupate after going through a specific number of light-dark cycles or days: the required day number (RDN) for diapause completion. This RDN could be modulated by temperature but more research is necessary to clarify this point. In post-diapause development, when a suitable temperature threshold is considered, the absolute accumulation of heat is more important than whether the temperature received is fluctuating or constant. The temperature threshold for diapause and post-diapause development was lower than the temperature threshold of larvae in continuous development. This is important for adjusting phenological models in S. nonagrioides and in other species, and may explain why in many cases adults appear in the field when the supposed temperature threshold for development has not been attained.     

Induction and termination of reproductive diapause in a neotropical beetle, Chelymorpha alternans (Coleoptera: Chrysomelidae)

The presence of a reproductive diapause in the life cycle of the neotropical cassidine beetle, Chelymorpha alternans , was investigated by exposing groups of beetles to conditions differing in photoperiod and humidity. Diapause was characterized by the absence of egg-laying in females up to 70 days after emergence and was induced in response to a short photoperiod of 12:12 h L: D but averted under a longer photoperiod of 13:11 h L:D. High (>90% RH) and low (55–75% RH) humidity conditions did not influence diapause induction. Males appeared to court and attempt to mate with females under all conditions between 16 and 30 days after emergence, but declined after this time in short photoperiods and it is not known if matings in these groups were successful. Adults induced to diapause by exposure to a 12:12 h L:D photoperiod and subsequently exposed to a 13:11 h L: D photoperiod 70 or 72 days after emergence did not show a rapid response by commencing egg-laying. However, all diapausing groups in long and short photoperiod and high and low humidity, in both experiments performed, had commenced egg-laying after 128 days, suggesting an endogenous rhythm for diapause termination. Long photoperiod and high humidity combined may hasten diapause termination since egg-laying began after only 95 days in this group in experiment two. This strategy of induction and termination is discussed with reference to the seasonality of the natural environment of C. alternans in Panama.     

白蛾周氏啮小蜂滞育诱导及滞育后发育

本研究针对人工繁育白蛾周氏啮小蜂Chouioia cunea Yang过程中出现的小蜂滞育现象, 对其滞育诱导的光周期反应及敏感光照虫态进行了调查。结果表明: 沈阳地区的白蛾周氏啮小蜂属长日照型昆虫, 以老熟幼虫进入滞育状态, 但在不同的温度条件下诱导滞育的临界光周期不同, 在18℃时诱导滞育的临界光周期处于13L∶11D和14L∶10D之间; 在21℃和24℃时诱导滞育的临界光周期变短, 处于12L∶12D和13L∶11D之间。白蛾周氏啮小蜂滞育诱导的敏感光照虫态为幼虫期, 且以幼虫的后期最为敏感, 但整个幼虫期接受短光照对滞育的形成更为有利。通过观察白蛾周氏啮小蜂滞育后在18℃, 21℃, 24℃和30℃的恒温条件下的发育历期, 由最小二乘法计算出白蛾周氏啮小蜂老熟幼虫滞育后发育起点温度和有效积温分别为14.60±0.31℃和209.38±8.72日·度。这些结果可为进一步研究白蛾周氏啮小蜂的种蜂长期保存技术和指导商品蜂生产, 正确把握放蜂时机提供理论依据。     

低温和光周期对绿盲蝽越冬卵滞育解除和发育历期的影响

为了探讨温度和光周期对绿盲蝽Apolygus lucorum Meyer-Dür越冬卵滞育解除和发育历期的影响, 系统调查了绿盲蝽越冬卵在不同温度和不同光照组合下的孵化率和孵化时间, 结果显示：绿盲蝽的越冬卵均为滞育卵,低温和光周期对绿盲蝽越冬卵的滞育解除均有影响。2℃的低温处理能够显著促进其滞育解除,在0~65 d范围内,随着低温处理时间增长,其滞育解除时间缩短,未经低温处理的越冬卵T₅₀为68.5 d,低温处理65 d的T₅₀为12.25 d,绿盲蝽越冬卵在2℃低温处理65 d后完全解除滞育;在0~40 d范围内,低温处理时间越长,绿盲蝽越冬卵的孵化率越高,在25℃、全光照的条件下不经低温处理的孵化率为68.65%,低温处理40 d后在25℃的条件下的孵化率达到99.46%。在20~26℃范围内,绿盲蝽越冬卵的滞育后发育历期随着温度的上升而缩短, 随着光周期的延长而缩短。结果说明低温处理能够提高绿盲蝽越冬卵滞育解除率,但不是其滞育解除的必要条件,低温处理与自然变温对绿盲蝽滞育解除的作用相似;高温和长光照能够促进绿盲蝽的滞育解除,缩短发育历期。     

Effects of photoperiod and temperature on the termination of diapause in the univoltine seed wasp Eurytoma plotnikovi

Abstract Mummified pistachios containing fully grown diapause larvae of Eurytoma plotnikovi Nikol'skaya (Hym., Eurytomidae) were collected in early August and late September in coastal northern Greece and subjected to various photoperiod and temperature treatments, then maintained at 19 or 26°C and a long-day (LD 16:8 h), a changing, or a short-day (LD 10:14 h) photoperiod until pupation. In larvae of early August (beginning of diapause) subjected for 20 weeks to 19°C under a long, a changing, or a short photophase, followed by 19°C and a long photophase, 50% of the larvae pupated after 24, 18 and 13 weeks respectively. After exposure for 20 or even 12 weeks to a short photophase and low temperatures (10 or 4°C), pupation occurred after only 7–8 weeks and was more synchronous. The ranges of temperature for diapause development and post-diapause morphogenesis overlap. After exposure for 12 weeks to short days and low temperature, larvae of late September pupated much sooner under long days than under short days and sooner at 26° than at 19°C. E.plotnikovi depends on both temperature and photoperiod for diapause development, low temperature having a strong favourable effect on the earlier part and long day on the later part of diapause. In a few larvae of another pistachio seed wasp, Megastigmus pistaciae Walker, after a long enough period of low temperatures, diapause was terminated normally at 26°C and long days, or at 19°C and long or short days.     

Completion of diapause in field populations of the cabbage root fly (Delia radicum)

The various diapause and post-diapause stages entered by cabbage root fly pupae during the overwintering period are shown schematically. Although diapause induction started in mid-Aug., the early-pupating insects did not develop further but were maintained in diapause by the warm autumn temperatures. Therefore, diapause development was simultaneous in all Wellesbourne pupae, whether of second or third generation origin. Diapause development started only in mid-Oct., when mean soil temperatures fell below 10°. In the field, 90% of the overwintering population of cabbage root fly pupae had completed pleted diapause by 5 March 1980, 17 Feb. 1981 and 18 Feb. 1982. This was equivalent to a duration of 19 weeks from mid-Oct. onwards, during the winters of 1979–80, 1980–81 and 1981–82 respectively. A further break between the completion of diapause and the warm conditions required to start post-diapause development also helps to condense the emergence of flies in the spring. Hence, an accurate forecast of the time of spring attack by populations of flies similar to those at Wellesbourne should be possible.This study was financed partly by the Commission of the European Communities as CEC Contract No. 0771.     

Predicting spring emergence of blackcurrant leaf midge (Dasineura tetensi) from air temperatures

The mean duration of post-diapause development of overwintered Dasineura tetensi larvae (in cocoons) was 72.8 (SD=11.4), 45.9 (SD=8.6), 28.7 (SD=6.0), 15.9 (SD=4.3), 10.4 (SD=1.9) and 10.2 (SD=1.8) days at constant temperatures of respectively 10, 12.5, 15, 17.5, 20 and 25 °C in the laboratory. No perceptible development occurred at 5 or 7.5 °C and complete mortality occurred when larvae were held at 30 °C for prolonged periods. The relationship between development rate (r days^–1) and temperature (T °C) was sigmoidal between 10 and 25 °C, the logistic equation r=0.0158+0.085/(1+exp(–0.696(T–17.0))) accounting for 98% of the variation. Larvae entered the winter in diapause. Populations of cocoons were greatest in the surface soil in the centre of bushes adjacent to the crown, 69, 15, 9 and 6% of cocoons occurring in the top 0–1, 1–2, 2–3 and 3–4 cm of the soil, respectively. The time of termination of diapause in the field varied greatly between individuals and from season to season but a significant proportion (>40%) had broken diapause by the end of January in each of the three seasons studied. Diapause was not terminated in the laboratory by chilling over-wintered larvae in cocoons at –2.5, 2.5 or 10 °C for up to 28 days nor when held in a L16:D8 photoperiod. A computer-based phenological forecasting model was constructed using the development rate values (using the INSIM software developed at The Agricultural University, Wageningen, The Netherlands). The model accumulated daily development amounts calculated from daily maximum and minimum air temperatures from 1 February, the end of the coldest period of the year on average and before significant post-diapause development occurred. The model uses boxcar trains to simulate dispersion. The model predicted the time of first emergence of D. tetensi adults in spring at HRI-East Malling generally to within 6 days of the observed time of emergence, and to within 11 days at worst. There was poorer agreement between observed and predicted times of emergence when daily maximum and minimum soil temperatures (depth ca. 3 cm) were used. The use of the model to time insecticidal sprays in relation to the flowering time of blackcurrant is discussed.     

Induction and development of winter larval diapause in a drosophilid fly,Chymomyza costata

Photoperiodic response during induction of larval hibernal diapause of Chymomyza costata was characterized and the course of diapause development was analyzed in the laboratory. C. costata becomes sensitive to photoperiodic stimuli during an unspecified stage of its early development (embryo, 1st larval instar); the sensitivity gradually increases during the 2nd and early 3rd larval instars and reaches its maximum just before the moment when it abruptly ceases at the age of 15-19 days after oviposition. Diapause intensifies during a period of 2-3 weeks after induction and, later, is maintained without apparent development until death (between 150 and 250 days) under 18 degrees C and a short-day photoperiod (L10:D14, SD). Diapause may be terminated in a horotelic process by exposure to a low temperature (2 degrees C) during which larvae subsequently (1) synchronize their post-diapause development (requires up to 14 days of chilling), (2) lose photoperiodic sensitivity (2 months), and finally (3) terminate diapause (5 months). Alternatively, diapause may be terminated in a tachytelic process by exposure to a high temperature (18 degrees C) and long-day photoperiod (L16:D8, LD) during which no synchronization occurs and pupariation takes place after a mean of 25.2 days (with a broad range from 8 to more than 50 days). Larvae that are transferred from LD to SD during their sensitive period switch their developmental programming from pupariation to diapause. Proliferation of adult primordial structures (imaginal discs, neuroblasts) slows down within 1 day after transfer. In contrast, whole body growth continues for at least 3 days before its rate slows down and matches the rate characteristic for SD conditions.     

昆虫滞育后的生物学特性

从昆虫滞育后性比、寿命、生殖力和重复滞育等方面对昆虫滞育后的生物学特性进行了概括,并分析了影响昆虫滞育后生物学特性数量表达的因子。这些因子包括:(1)滞育前昆虫的发育速率、取食行为和个体大小;(2)滞育期间的环境条件及昆虫取食行为;(3)滞育持续期;(4)滞育后的取食需求;以及(5)滞育后的温度和光周期。     

Larval diapause duration and fat metabolism in three geographical strains of the blow fly,Calliphora vicina

Diapausing larvae of the blow fly, Calliphora vicina, from three geographical strains exposed, as adults, to short days, were maintained under identical conditions (darkness, 11-12 degrees C) and examined for changes in wet weight, dry weight, water and fat content during diapause development to the emergence of post-diapause adults. Larvae produced by flies originating from northern Finland (Nallikari, 65 degrees N) showed a longer, more intense, diapause than those from localities further south (Edinburgh, Scotland, 55 degrees N and Barga, Italy, 44 degrees N), but all three strains showed similar rates of loss of the parameters measured. This was also the case for post-diapause adults, flies of the Barga strain with its relatively short diapause emerging with greater residual fat reserves than flies from the Edinburgh or Nallikari strains with their more protracted diapause. It was concluded that the rates of water and fat loss were functions of the conditions used for diapause larval maintenance (probably temperature) rather than the maternally programmed degree of diapause incidence, or of its 'depth' or 'intensity'.