Effects of temperature and other environmental factors on the post‐diapause development of walnut husk fly Rhagoletis completa (Diptera: Tephritidae)

To more effectively manage walnut husk fly Rhagoletis completa (Diptera: Tephritidae), in California walnut orchards, it is important to understand the factors that affect the timing of adult emergence. In the present study, we examine the effects of incubation temperature, pre‐chill and chill durations, latitude, cultivar and size on the post‐diapause development of R. completa puparia. The lower developmental threshold, upper developmental threshold and optimal temperature for puparial development are estimated to be 4, 34 and 26.6 °C, respectively. The thermal requirement for adult emergence after 120 days of chilling is estimated to be 2024 degree days. Percentage adult emergence declines at both higher and lower incubation temperatures. Chill duration at 5 °C for diapausing puparia has a nonlinear negative effect on the thermal requirement but no effect on percentage emergence. Insufficient chilling leads to poor synchronization of adult emergence. Greater pre‐chill duration at room temperature increases the thermal requirement and slightly decreased percentage emergence. Latitude has a negative effect on the thermal requirement. Puparia from northern California black walnut (Juglans hindsii) have a slightly greater thermal requirement than puparia from cultivated walnut (Julgans regia). There is no significant difference in puparial fresh weight or mean thermal requirement between males and females, although the positive correlation between thermal requirement and puparial fresh weight is stronger for females than males. The effects of temperature and other environmental factors on the post‐diapause development of R. completa are discussed in relation to observations from other Rhagoletis species.     

松毛虫赤眼蜂滞育诱导及解除条件研究

【目的】以柞蚕Antheraea pernyi卵为繁殖寄主,对松毛虫赤眼蜂Trichogramma dendrolim滞育诱导及解除条件进行研究,以解决赤眼蜂工厂化生产和大面积应用中面临的的中、长期储存问题。【方法】通过观测不同发育阶段(寄生柞蚕卵在26℃培养40、96和144 h)、滞育诱导温度(10、13和16℃)和诱导时间对松毛虫赤眼蜂滞育的影响,确定松毛虫赤眼蜂滞育诱导条件;通过观测滞育诱导温度和滞育后的贮藏温度对滞育解除的影响,确定松毛虫赤眼蜂滞育解除条件。【结果】在松毛虫赤眼蜂的不同发育阶段对其进行持续的低温刺激均能使其导入滞育,但以小幼阶段(26℃培养40 h)开始效果最佳,寄生卵在26℃培养40 h后,转入10℃和13℃下连续诱导31 d,滞育率可达100%和99.12%。滞育诱导温度和滞育后的贮藏温度对松毛虫赤眼蜂解除滞育所需时间和解除滞育后的羽化出蜂率有较大影响,10℃诱导滞育后置于1℃冷藏的赤眼蜂解除滞育所需时间最短,解除滞育后的羽化出蜂率和单卵出蜂数更高,更耐储存。此条件下冷藏约30 d开始打破滞育,在正常发育下温度下羽化出蜂,60 d羽化出蜂率达到95.24%,冷藏4个月后羽化出蜂率仍在60%以上,单卵出蜂数高于50头。【结论】松毛虫赤眼蜂最佳滞育诱导条件为26℃培养40 h后,转入10℃连续低温诱导31 d;最佳滞育解除条件为1℃低温储存,但储存期不能超过4个月。     

Diapause induction and termination in a commonly univoltine leaf beetle (Phratora vulgatissima)

Abstract The leaf beetle Phratora vulgatissima (Linnaeus 1758) is commonly univoltine in south‐central Sweden but may sometimes initiate a partial second generation. The current study was set out to investigate under what abiotic conditions the beetles initiate a second generation. Using climate chamber experiments, the beetles were shown to have a facultative reproductive diapause induced by declining day‐length. The critical day‐length (CDL) for diapause induction was estimated to be 18 h and 10 min. In the field, first‐generation beetles developing to adulthood before August in 2009 became reproductively active and produced a second generation, but most individuals emerged later and were in reproductive diapause. P. vulgatissima overwinter as adults and diapause was shown to be maintained until mid‐winter in 2008/2009. The cumulative temperature requirement for oviposition after diapause termination was estimated to be 222 day‐degrees with a 5.5°C temperature threshold. Three different day‐degree models that were developed to predict the phenology of female oviposition in the spring were validated by comparing model results with field data on the timing of oviposition in previous years. The study suggests that P. vulgatissima may initiate a second generation in Sweden if development of the first generation is completed before August. Warmer spring and summer temperatures due to ongoing climate change may cause advanced insect phenology and faster completion of insect life‐cycles at northern latitudes, which will affect the proportion of insects that initiate a second generation.     

Differences in the pre‐diapause and pre‐oviposition accumulation of critical nutrients in adult females of the beetle Colaphellus bowringi

In many insect species, the differentiation of development between diapause and reproduction first becomes obvious during the diapause preparation (pre‐diapause) and pre‐oviposition phases. However, the differentiation of nutrient accumulation between these two phases remains unclear. We compared the weights of pre‐diapause and reproductive adult female Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), and measured their triacylglycerol (TAG), protein, and carbohydrate content from emergence until they had fed for 4 days post‐eclosion. We also compared the ovarian development and accumulation of lipid droplets between pre‐diapause and reproductive adult females in order to determine whether we could visually detect differences in nutrient allocation. The weights of both pre‐diapause and reproductive females increased with duration of feeding. The fresh weight and water content of pre‐diapause females was significantly lower than that of reproductive females after feeding for 3 days post‐eclosion. Pre‐diapause females channeled their reserves into TAG in the fat body, whereas reproductive females converted nutrients into proteins and carbohydrates for egg development. These results quantify differences in nutrient accumulation between pre‐diapause and reproductive adult female C. bowringi, and provide clues for understanding the molecular mechanisms underlying differences in the allocation of nutrients between diapause and reproduction in insects.     

Responses in metabolic rate to changes in temperature in diapausing Colorado potato beetle Leptinotarsa decemlineata from three European populations

Many insects survive adverse periods in seasonal environments by entering diapause, a deep resting stage, during which energy consumption is typically low and gas exchange is in the form of a discontinuous gas exchange cycle (DGC). Because insects in high‐latitude environments are severely time constrained during summer, an effective diapause termination with careful regulation of metabolic rate is important. The present study examines whether diapausing Colorado potato beetles Leptinotarsa decemlineata Say originating from three latitudinally different regions in Europe differ in their quantitative or qualitative gas exchange patterns in response to an increasing temperature. Overall production of gaseous CO₂, as well as qualitative patterns relating to the DGC, are measured at a late stage of diapause at four different temperatures in increasing order from 13, 18, 23 to 28 °C. Overall CO₂ production is found to be lower in the two northern populations (61°49′N and 55°75′N) compared with the southernmost population (45°48′N) but increases as a function of temperature in all populations in a similar way. However, in the northern populations, raising the temperature increases the amount of CO₂ discharged during single DGC peaks, whereas the DGC frequency remains relatively unchanged. By contrast, in the southernmost population, the amount of CO₂ discharged during individual DGC peaks remains relatively unchanged, whereas the DGC frequency increases as a function of temperature. The observed differences may relate to water retention benefits or, alternatively, energetic benefits relating to heightened gas exchange efficiency in hypoxic or hypercapnic environments. Overall, the results suggest that, although populations of L. decemlineata may have similar thermal sensitivities, they have different qualitative strategies to regulate metabolic re‐activation at diapause termination.     

Enzymatic capacity for accumulation of polyol cryoprotectants changes during diapause development in the adult red firebug, Pyrrhocoris apterus

Abstract. Diapausing adults of the red firebug, Pyrrhocoris apterus, were maintained in the laboratory at constant conditions of 20 °C and short days (LD 12 : 12 h) for 5 months. Spontaneous termination of diapause is seen in 0.4% of adults. At different times of diapause development, groups of adults were exposed to an 8‐week gradual cold treatment ending at 0 °C. Ribitol and sorbitol contents remain very low at constant 20 °C and their rapid accumulation only occurrs at temperatures below 5 °C. The capacity to accumulate ribitol in response to low temperature stimulus remains relatively stable but the capacity to accumulate sorbitol decreases to zero during the 5 months of diapause development. Glycogen (whole‐body and fat‐body reserves) accumulates relatively rapidly during the early phase (up to 1–2 months) and slowly depletes during the late phase of diapause development under constant conditions. Upon cold treatment, part of the glycogen reserve was depleted. The activities of most enzymes involved in polyol metabolism (namely glycogen phosphorylase, glucose‐6‐P dehydrogenase, phosphofructokinase, aldose reductase, polyol dehydrogenase and ketose reductase) increase relatively rapidly during the early phase of diapause development and, in the late phase, they either become stable or slowly decrease. Cold treatment has either no effect or results in a moderate increase in activity when applied in the early phase of diapause development but results in a more or less obvious decrease of enzymatic activity when applied in the late phase.     

Diapause development in early and late emerging phenotypes of Delia floralis

The turnip fly, Delia floralis Fall6n （Diptera： Anthomyiidae） is an important insect pest of brassica vegetable crops in the holarctic region. Different populations have strongly varying temperature requirements for fly emergence, a challenge for accurate prediction of activity. This study focused on diapause development in one early and one late emerging phenotype. The physiological state after various treatments was deduced from emergence data. Our results showed a slow diapause progression at chilling conditions for both populations and diapause ended about 7 months after pupae were formed for the early population. For the late population held at 4℃ diapause did not end, no matter how long the duration of chilling. These pupae required a period with elevated temperatures above 6~C to continue development. At constant non-chilling conditions （18℃） from the time pupae were formed both populations completed diapause most rapidly. These results indicate that chilling delayed, rather than accelerated development and was not a prerequisite for diapause development. For post-diapause, results indicated a linear relationship between rate of development and temperature within the range of 6-18℃and a theoretical base temperature for development of about 2℃ for both populations. In conclusion, D. floralis pupae are in diapause throughout a long winter period, and delayed emergence of the late population appears to be caused by prolonged diapause regulated by a developmental temperature threshold. The study has added information on the biology of turnip fly populations, a prerequisite for improved pest control.     

Transfer of life‐history phenology from mothers to progeny in a solitary univoltine parasitoid

Among univoltine insects that experience diapause, differences in emergence timing between adult males and females are expected to be dictated by sex‐specific developmental factors. In multivoltine insects without a diapause, there is often an additional relationship between the date of oviposition and the date of adult emergence. Differences between male and female emergence timing in the latter case can therefore be influenced by female sex‐allocation decisions. In the present study, it is shown that eggs of a univoltine parasitoid wasp Diachasma alloeum Muesebeck (Hymenoptera: Braconidae) that are laid earlier also eclose earlier during the subsequent year, independent of (although complementary to) sex‐related differences in development time. The implications of this pattern for sex allocation decisions by female univoltine parasitoids are discussed.     

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.     

Seasonal regulation of dormancy in Chrysopa carnea (Neuroptera)

In Ithaca, New York, diapause incidence in Chrysopa carnea exceeds 50 per cent among adults emerging between 31 August and 3 September. Apparently, decreasing late summer day lengths, which exceed the stationary critical photoperiod in the laboratory, act on the sensitive stages and induce diapause. Full diapause intensity occurs about 2 weeks after adult emergence; complete winter coloration appears approximately 2 weeks later. Photoperiodic stimili in combination with the geographic strain largely determine diapause depth.Decreasing autumn day lengths decelerate diapause development and maintain diapause until the winter solstice. Subsequently, short day lengths and cold temperatures slow diapause development. In nature, neither long days, increasing day lengths, nor chilling hasten diapause termination. Diapause duration is largely determined by the interaction between inducing photoperiod, maintaining photoperiodic and thermal stimuli, and geographic strain. In Ithaca, diapause ends (all response to photoperiod ceases) between 22 January and 12 March.After diapause ends, the animals retain their diapause characteristics until temperatures exceed 4°C. Post-diapause reproductive development in females requires approximately 100 heat units above 4°C; however, below 8°C mating and oviposition are absent. Although the winter coloration remains, mating occurs within 1 day after transfer from 4 to 24°C. Temperature determines the rate at which the green summer colour returns.     

Endocrine changes during diapause development in the cabbage army moth Mamestra brassicae

Many insects undergo diapause to survive adverse seasons. Although the mechanism of diapause induction is the subject of extensive study, that of diapause termination remains poorly understood. In the present study, we show the endocrine processes leading to the termination of pupal diapause in Mamestra brassicae. Diapause of this insect is terminated if the pupae are exposed to a low temperature for several weeks. During this period, the prothoracic glands (PGs) of pupae acquire the potential to secrete sufficient ecdysteroids necessary for inducing adult development. The main endocrine changes observed under the low temperature conditions are: (i) the increase in activity of the PGs in two steps; (ii) the increase in responsiveness of the glands to prothoracicotropic hormone (PTTH); and (iii) two‐step increase in PTTH gene expression in the brain. The timing of the first and second increases in PG activity roughly coincides with that of the two steps of increase in PTTH gene expression, and the timing of the increase in the responsiveness of the PGs to PTTH coincides with the second, larger increase in PTTH gene expression. The ablation of the PGs prior to cooling pupae does not affect the increase in PTTH gene expression, whereas brain removal results in a failure to increase PG activity, strongly suggesting that PTTH is the master regulator of diapause development and termination.     

Control of diapause and supercooling by the retrocerebral complex inPyrrhocoris apterus

Effects of external (photoperiod, temperature) and internal (retrocerebral complex, diapause condition) factors on supercooling were studied inPyrrhocoris apterus (L.) (Heteroptera: Pyrrhocoridae) adults. An increase in supercooling capacity is associated with the induction of diapause by short-day photoperiod or cardiacallatectomy of long-day non-diapause insects in spite of a high temperature of 26°C. The induction of diapause is a prerequisite for a further increase in supercooling capacity by cold acclimation. Post-diapause adults show low values of the supercooling point also in late January or early February, although their developmental potential is fully restored. Evidently the ability to supercool associated with diapause induction is maintained by a low ambient temperature in spite of diapause termination. The supercooling point increases rapidly when these adults are transferred to a high temperature of 26°C and cold re-acclimation is then no longer possible. The inhibition of morphogenesis (maturation of gonads) by cardiacallatectomy does not prevent the supercooling point from increasing in post-diapause adults at the high temperature.     

The effect of cold temperature exposure and long-day photoperiod on the termination of the reproductive diapause of newly emerged female Pissodes strobi (Coleoptera: Curculionidae)

Abstract

• 1 There is confusion in the literature concerning a possible reproductive diapause in the adult white pine weevil Pissodes strobi.
• 2 We evaluated the effects of temperature, photoperiod, feeding substrate and mating status on the sexual maturation and oviposition of female white pine weevils.
• 3 Less than 30% of female P. strobi became sexually mature and laid eggs without experiencing dormancy under a temperature regime of 2 °C for 4 weeks.
• 4 Among the females that experienced a cold temperature treatment after emergence, 80% laid eggs after dormancy when exposed to a long‐day (LD 16 : 8 h) photoperiod and 17.6% laid eggs when exposed to a short‐day (LD 8 : 16 h) photoperiod.
• 5 Significantly more eggs were laid by all the females (with and without a cold treatment) when subjected to a long‐day photoperiod compared with a short‐day photoperiod.
• 6 A period of cold temperature followed by exposure to a long‐day photoperiod with warmer temperatures is required to break reproductive diapause and to obtain a good oviposition response in female P. strobi.
• 7 This study reveals the existence of much intraspecific variation in the response of the white pine weevil to temperature and photoperiod with respect to the induction and termination of reproductive diapause.

     

Effect of temperature on the termination of diapause in the univoltine almond seed wasp Eurytoma amygdali

Diapausing larvae of Eurytoma amygdali Enderlein (Hymenoptera, Eurytomidae) were collected in early August and late September. They were subjected to various photoperiod and temperature regimens for up to 20 weeks, then kept at L16:D8 and 19 °C for another 14 to 26 weeks for diapause to be terminated and pupation to take place. Photoperiod did not affect diapause completion. It was confirmed that the two morphologically distinct diapause stages have different temperature requirements for their completion. The first diapause stage was completed synchronously at temperatures between 16 and 19 °C. A higher temperature of 26 °C delayed diapause development. The second stage required lower temperatures between 4 and 10 °C. Spontaneous termination of diapause was observed at constant 19 °C. When applied to the first diapause stage for 20 weeks, low temperatures made the larvae refractory to subsequent intermediate temperatures. The first stage was thus maintained until a higher temperature of 26 °C made the larvae regain their ability to respond to the intermediate temperatures and complete this stage. Larvae grown in Retsou almonds had a higher diapause intensity than larvae grown in Truoito almonds. The results suggest that, in nature, the high temperatures of late summer and early autumn are likely to maintain the first diapause stage. Subsequently, the less warm temperatures of autumn allow the completion of the first stage by late autumn, and the low temperatures of late autumn and of winter allow the completion of the second diapause stage by mid winter.     

Effects of constant and changing temperature conditions on diapause induction in Helicoverpa armigera (Lepidoptera: Noctuidae)

The effects of photoperiod and temperature on the induction and termination of facultative pupal diapause in Helicoverpa armigera (Lepidoptera: Noctuidae) were investigated under laboratory conditions. Exposing H. armigera larvae to both constant and fluctuating temperature regimes with a mean of 25°C and 20°C resulted in a type-III photoperiodic response curve of a short-long day insect. The long-day critical daylengths for diapause induction were ten hours and 12 hours at the constant temperatures of 25°C and 20°C, respectively. Higher incidences of diapause and higher values both for the longer and the shorter critical photoperiods for diapause induction were observed at fluctuating regimes compared with the corresponding constant ones. At alternating temperatures, the incidence of diapause ranged from 4.2% to 33.3% and was determined by the temperature amplitude of the thermoperiod and by the interaction of cryophase or thermophase with the photoperiod. Helicoverpa armigera larvae seem to respond to photoperiodic stimuli at temperatures >15°C and <30°C; all insects entered diapause at a constant temperature of 15°C, whereas none did so at a constant temperature of 30°C under all the photoperiodic regimes examined. Although chilling was not a prerequisite for diapause termination, exposure of diapausing pupae to chilling conditions significantly accelerated diapause development and the time of adult emergence. Therefore, temperature may be the primary factor controlling the termination of diapause in H. armigera.     

Effects of nutrition and methoprene treatment upon reproductive diapause in Caloptilia fraxinella (Lepidoptera: Gracillariidae)

Abstract Female Caloptilia fraxinella exhibit a prolonged reproductive diapause immediately post adult emergence in mid‐summer until the next spring when mating, egg development and oviposition on fresh Fraxinus spp. leaflets occur. Factors that effect the termination of reproductive diapause are investigated in this species. Caloptilia fraxinella diapausing adults held in overwintering conditions (2 °C, LD 0 : 24 h) for 24 weeks terminate diapause after placement for 2 weeks in simulated summer conditions (24 °C, LD 16 : 8 h) only if they are provided with 10% sugar water. Exogenous application of the Juvenile Hormone (JH) analogue methoprene to moths in both early‐ (summer) and mid‐ (autumn) reproductive diapause demonstrates that JH affects diapause termination but a carbohydrate nutrition source also mediates mating and vitellogenesis. Mating between moth pairs early in diapause occurs only after treatment with methoprene and provision with sugar water. However, there is no impact of mating on the propensity of females to produce vitellogenic oöcytes. Moths collected in the autumn in mid‐diapause respond in a dose‐dependent fashion to methoprene treatment and the response is greater than that of moths early in diapause collected in the summer. Treatment with methoprene and access to sugar water results in vitellogenic oöcytes in 18.75% of females from mid‐diapause moth pairs treated with 0.01 μg methoprene per insect and in all females from pairs treated at the two highest doses of methoprene (0.1 and 1 μg per insect). Mating occurs only between moths in mid‐diapause treated with the two highest doses of methoprene and these doses induce 91% and 100% mating, respectively. Both control and methoprene‐treated males in mid‐diapause held under summer conditions mate successfully and pass a spermatophore to their methoprene‐treated female partner. These data demonstrate that female C. fraxinella undergo a prolonged reproductive diapause in which termination is dependent on JH and further mediated by a carbohydrate nutrition source. The production of vitellogenic oöcytes is independent of mating. These data also provide evidence that response of moths in diapause to exogenous applications of methoprene differs throughout the diapause period and between male and female C. fraxinella.     

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.