Inheritance of the photoperiodic response controlling imaginal summer diapause in the cabbage beetle, Colaphellus bowringi

Adults of the cabbage beetle Colaphellus bowringi display a summer diapause in response to the exposure of their larvae to long photoperiods. In the present study, the inheritance of the photoperiodic response controlling summer diapause in C. bowringi by crossing a high diapause strain (D strain) with a laboratory selected nondiapause strain (N strain) was investigated under different photoperiods at 22, 25 and 28 °C. The beetles in both reciprocal crosses and backcrosses showed a clear short-day response for the induction of diapause at all temperatures, similar to that of the D strain, suggesting that photoperiodic response of this beetle is heritable. The diapause incidences in the progeny from all the crosses under LD 15:9 or LD 12:12 at 25 °C suggest that genetic and genetic-environmental interactions are involved in diapause induction. The incidence of diapauses in F₁ progeny was significantly lower than that in the D × D strain but significantly higher than that in the N × N strain, indicating that the diapause capability is inherited in an incomplete dominant manner. The incidence of diapause was greater among the offspring of hybrid females when those females had a D strain mother or grandmother rather than a N strain mother or grandmother, indicating that maternal effects on diapause induction are stronger than paternal effects. The laboratory selected nondiapause strain also showed a short-day photoperiodic response at a low temperature of 22 °C, indicating that the photoperiodic photoreceptor and photoperiodic clock still function in the nondiapause strain.     

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.     

Diapause induction, maintenance and termination in the rice stem borer Chilo suppressalis (Walker)

The rice stem borer, Chilo suppressalis, enters facultative diapause as fully grown larvae in response to short-day conditions during the autumn. Our results showed that the critical night length for diapause induction in C. suppressalis was between 10 h 22 min and 10 h 45 min at 22, 25 and 28 °C, 11 h 18 min at 31 °C, and between 10 h 5 min and 10 h 20 min under field conditions (average temperature ranged from 27.2 to 30.7 °C). The diapause incidence declined in ultra-long nights (18-22 h scotophases) and DD, and increased in ultra-short nights (2-6 h scotophases) and LL. Moreover, we found that the third instar was the stage most sensitive to the photoperiod, and night length played an essential role in the initiation of diapause. Night-interruption experiments with a 1-h light pulse at LD 12:12 (light 12:dark 12) exhibited two troughs of diapause inhibition, with one occurring in early scotophase and the other in late scotophase. Field observations for six years showed that most larvae entered winter diapause in August in response to declining day lengths, despite the high temperatures prevailing during August. By periodically transferring the field-collected overwintering larvae to different photoperiods and temperatures, the results showed that photoperiod had a significant influence on diapause development during the early phase of diapause, while high temperature significantly accelerated the termination of larval diapause.     

Seasonal patterns of the thermal response in relation to sugar and polyol accumulation in overwintering adults of elm leaf beetle, Xanthogaleruca luteola (Coleoptera: Chrysomelidae)

Elm leaf beetle, Xanthogaleruca luteola (Muller), is one of the key pests of elm trees that survive winter in reproductive diapause in sheltered locations. Overwintering adults of the elm leaf beetle showed a complex sugar/polyol cryoprotectant system. The major components of the multiple systems were glucose, myo-inositol and trehalose. In this study, we investigated the seasonal profile of low molecular weight compounds and glycogen in natural population and also in response to thermal constant regimes (5 and 15 °C). Among these components, a remarkable seasonal pattern of accumulation/depletion was observed in myo-inositol over the course of hibernation with the development of diapause progress. Incubating at 5 °C only elicited a strong response in myo-inositol synthesis during diapause. It suggests that the elm leaf beetle accumulates myo-inositol not only in relation to entering diapause but also in response to low temperatures and their interactions. The laboratory acclimation experiments showed that adults exposed to 15 °C had no chance for accumulation of low molecular weight carbohydrate even during diapause. The results of this study illustrated that overwintering adults of elm leaf beetle produce myo-inositol as the primary substance which plays a specific role in some biochemical adjustments in overwintering adults of X. luteola.     

Photoperiod- and temperature-dependent induction of larval diapause in a multivoltine bruchid, Bruchidius dorsalis

The wild bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and overwinters in several developmental stages in the middle part of Japan. We investigated the incidence of diapause under different conditions of photoperiod (from L8:D16 to L16:D8) and temperature (at 20 °C and 24 °C). Our experiments revealed the following results: (1) B. dorsalis entered diapause at the final (late fourth) instar larva under short photoperiods, (2) the larval diapause incidence was dependent on temperature (critical photoperiods were 12.5 h at 20 °C and 12 h at 24 °C), (3) some individuals did not enter diapause under short-photoperiod conditions at 24 °C, and (4) the sensitive stages to the photoperiod were from the late egg stage to the early first instar larva. Based on these results, we discuss not only the evolution of a complex overwintering strategy inB. dorsalis but also the domestication process of stored-bean pests.     

Diapause and overwintering of two spruce bark beetle species

Diapause, a strategy to endure unfavourable conditions (e.g. cold winters) is commonly found in ectothermic organisms and is characterized by an arrest of development and reproduction, a reduction of metabolic rate, and an increased resistance to adversity. Diapause, in addition to adaptations for surviving low winter temperatures, significantly influences phenology, voltinism and ultimately population growth. We review the literature on diapause and overwintering behaviour of two bark beetle species that affect spruce‐dominated forests in the northern hemisphere, and describe and compare how these strategies can influence population dynamics. The European spruce bark beetle Ips typographus (L.) (Coleoptera, Curculionidae) is the most important forest pest of Norway spruce in Europe. It enters an adult reproductive diapause that might be either facultative or obligate. Obligate diapausing beetles are considered strictly univoltine, entering this dormancy type regardless of environmental cues. Facultative diapausing individuals enter diapause induced by photoperiod, modified by temperature, thus being potentially multivoltine. The spruce beetle Dendroctonus rufipennis (Kirby) (Coleoptera: Curculionidae) infests all spruce species in its natural range in North America. A facultative prepupal diapause is averted by relatively warm temperatures, resulting in a univoltine life cycle, whereas cool temperatures induce prepupal diapause leading to a semivoltine cycle. An adult obligate diapause in D. rufipennis could limit bi‐ or multivoltinism. We discuss and compare the influence of diapause and overwinter survival on voltinism and population dynamics of these two species in a changing climate and provide an outlook on future research.     

Thermal response and reversibility of prolonged larval diapause in the chestnut weevil, Curculio sikkimensis

Curculio sikkimensis undergoes prolonged larval diapause that is terminated by chilling and warming cycles. To examine the effects of warming temperatures and their duration on diapause termination, we exposed diapause larvae that had not been reactivated after chilling at 5 °C to 20 or 25 °C and chilled them again before incubation at 20 °C. With increasing warming duration at 20 °C, diapause termination after chilling increased and shorter chilling durations became effective. In contrast, few or no larvae warmed at 25 °C terminated diapause after chilling, irrespective of the warming duration. To investigate the effect of warming temperature on diapause intensity, larvae with diapause weakened by initial incubation at 20 °C after the first chilling were subsequently incubated at 15, 20, or 25 °C, then chilled at 5 °C before incubation at 20 °C. Diapause termination increased significantly after the larvae were treated at 15 or 20 °C but decreased significantly after they were treated at 25 °C. The intensification of prolonged diapause at 25 °C was reversed when the larvae were transferred to 20 °C. Diapause intensity in C. sikkimensis therefore decreases at 20 °C, increases at 25 °C, and can be reversed by alternately exposing diapause larvae to 20 and 25 °C. In C. sikkimensis, prolonged diapause does not always proceed in one direction, and its intensity fluctuates in response to ambient temperature conditions.     

Diapause pupal color diphenism induced by temperature and humidity conditions in Byasa alcinous (Lepidoptera: Papilionidae)

We investigated whether diapause pupae of Byasa alcinous exhibit pupal color diphenism (or polyphenism) similar to the diapause pupal color polyphenism shown by Papilio xuthus. All diapause pupae of B. alcinous observed in the field during winter showed pupal coloration of a dark-brown type. When larvae were reared and allowed to reach pupation under short-day conditions at 18 °C under a 60 ± 5% relative humidity, diapause pupae exhibited pupal color types of brown (33%), light-brown (25%), yellowish-brown (21%), diapause light-yellow (14%) and diapause yellow (7%). When mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C and 25 °C under a 60 ± 5% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 91.2, 8.8 and 0.0% at 10 °C, and 12.2, 48.8 and 39.0% at 25 °C, respectively. On the other hand, when mature larvae reared at 18 °C were transferred and allowed to reach pupation at 10 °C, 18 °C and 25 °C under an over 90% relative humidity after a gut purge, the developmental ratio of brown and light-brown, yellowish-brown, and diapause light-yellow and diapause yellow types was 79.8, 16.9 and 3.3% at 10 °C, 14.5, 26.9 and 58.6% at 18 °C, and 8.3, 21.2 and 70.5% at 25 °C, respectively. These results indicate that diapause pupae of brown types are induced by lower temperature and humidity conditions, whereas yellow types are induced by higher temperature and humidity conditions. The findings of this study show that diapause pupae of B. alcinous exhibit pupal color diphenism comprising brown and diapause yellow types, and suggest that temperature and humidity experienced after a gut purge are the main factors that affect the diapause pupal coloration of B. alcinous as environmental cues.     

Pupal diapause of Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae) mediated by larval host plants: pupal weight is important

Facultative diapause, a strategy that allows insects to initiate additional generations when conditions are favorable or to enter diapause when they are not, has a profound effect on the ecology and evolution of species. Most previous studies have concentrated on the role of photoperiod and temperature in inducing facultative diapause in insects. In contrast, here we studied pupal diapause mediated by larval host plants in the cotton bollworm Helicoverpa armigera, and confirmed that pupal weight is a critical factor. Two groups of third instar H. armigera larvae, kept at 25 °C with L:D = 8:16 and 20 °C with photoperiod of L:D = 8:16, respectively, were fed on six host plants and on artificial diet (as a control) to determine how larval host plants affect diapause incidence and related traits (such as pupal weight and developmental duration). The data showed larval host plants affected diapause incidence significantly and the effects could be masked by low temperature. Further analysis showed that pupal size, not the length of the sensitive stage, affected the decision to enter diapause. In a further experiment, third-instar to final-stage larvae deprived of artificial diet for 2 days demonstrated a direct relationship between pupal weight and diapause incidence. These results suggest that larval host plants, by affecting pupal size, may influence diapause occurrence in H. armigera. This has important adaptive significance for both over-wintering survival and the possibility for completing an additional generation.     

The role of photoperiod and temperature in determination of summer and winter diapause in the cabbage beetle,Colaphellus bowringi (Coleoptera: Chrysomelidae)

The cabbage beetle, Colaphellus bowringi, is a short-day species undergoing an imaginal summer and winter diapause. Its photoperiodic response highly depends on temperature. All adults entered diapause at ≤ 20 °C regardless of photoperiods. High temperatures strongly weakened the diapause-inducing effects of long daylengths. The diapause-averting influence of short daylengths was expressed only at high temperatures (above 20 °C). This indicates that the beetle has a cryptic ability to reproduce in summer. In fact, summer and winter diapause were induced principally by relatively low temperatures in the field, whereas photoperiod had less influence on diapause induction. The critical daylength for the autumnal population was between 12 h and 13 h. By transferring from a long day to a short day or vice versa at different times after hatching, it was shown that the sensitive stage with regard to photoperiod was the larva, whereas a long day was photoperiodically more potent than a short day. The sensitive stage to temperature encompassed the larval, pupal and adult stages. This different response pattern serves to ensure that the beetle enters summer and winter diapause in time. The selections for non-diapause trait under laboratory (at 25 °C) and natural conditions (at >24 °C) showed that the beetle could lose its sensitivity to photoperiod very rapidly.     

Effects of temperature on the development,growth, and survival of larvae and pupae of a north-temperate chrysomelid beetle

Summary Developoment, growth, and survival of larvae and pupae of the red turnip beetle, Entomoscelis americana Brown, were studied in 10 constant and four alternating temperature regimes (10 to 32.5° C), in field-cages, and in natural populations in Manitoba. This beetle has a northtemperate distribution in North America. Larval and pupal development occurs in spring and normally is completed before the end of June. Growth and development occurred at all constant temperatures tested, but survival was low at the extreme temperatures. Therefore, the threshold and upper limit were near 10 and 32.5° C. The developmental times of the sexes did not differ and decreased with temperature, except possibly at 32.5° C. The average weight of adult females increased with temperature up to 32.5° C and those of males up to 25° C. Considering developmental rate, survival, adult weight, and incidence of malformed adults, the optimum temperature was estimated to be near 27.5° C.Development was accelerated significantly (6 to 9%) in alternating regimes with temperatures differing by 10° C, but not in regimes differing by 5 and 15° C. All alternating regimes increased adult weight, 5 to 17% for females and 2 to 10% for males. Field cage studies confirmed the increase in adult weight, but not the acceleration in development.A three-parameter normal function described accurately the relationship between developmental rate and constant temperature. A computer simulation model based on this equation estimated developmental times in field cages to within one to five days. For natural populations the model overestimated the developmental times by five to 16 days. The discrepancies between model estimates and observed developmental times in natural populations apparently were due to the elevation of larval and pupal body temperatures above air temperatures by behavioral thermoregulation. The elevation of body temperature was estimated to be equivalent to the addition of 5 to 6° C to the maximum daily air temperature. The adaptations and responses of this beetle to the cool spring temperatures of the north-temperate region are discussed.Contribution No. 1164, Agriculture Canada, Research Station, Winnipeg, Manitoba, Canada     

Geographic variation in diapause induction and termination of the cotton bollworm, Helicoverpa armigera Hübner (Lepidoptera: Noctuidae)

Overwintering diapause in Helicoverpa armigera, a multivoltine species, is controlled by response to photoperiod and temperature. Photoperiodic responses from 5 different geographical populations showed that the variation in critical photoperiod for diapause induction was positively related to the latitudinal origin of the populations at 20, 22 and 25 °C. Diapause response to photoperiod and temperature was quite different between northern and southern populations, being highly sensitive to photoperiod in northern populations and temperature dependence in southern populations. Diapause pupae from southern population showed a significantly shorter diapause duration than from northern-most populations when they were cultured at 20, 22, 25, 28 and 31 °C; by contrast, overwintering pupae from southern populations emerged significantly later than from northern populations when they were maintained in natural conditions, showing a clinal latitudinal variation in diapause termination. Diapause-inducing temperature had a significant effect on diapause duration, but with a significant difference between southern and northern populations. The higher rearing temperature of 22 °C evoked a more intense diapause than did 20 °C in northern populations; but a less intense diapause in southern population. Cold exposure (chilling) is not necessary to break the pupal diapause. The higher the temperature, the quicker the diapause terminated. Response of diapause termination to chilling showed that northern populations were more sensitive to chilling than southern population.     

Diapause and prolonged development in the embryo and their ecological significance in two cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata

The seasonal timing mechanism of egg hatching was examined in two cicadas, Cryptotympana facialis and Graptopsaltria nigrofuscata, with different but overlapping geographical distributions. These species lay eggs in summer, and nymphs hatch in the summer of the following year after egg durations of 10-12 months. When eggs were maintained at 25 °C from oviposition, both the species entered embryonic diapause within 60 days irrespective of photoperiod, but at different developmental stages between the two species. The optimal temperature for diapause development was approximately 15 °C in both the species. The development rate for postdiapause morphogenesis increased linearly with temperature in the range of 20-27.5 °C in C. facialis, and of 15-25 °C in G. nigrofuscata. The lower development threshold and the sum of effective temperatures were computed as 14.3 °C and 715.3 day-degrees in C. facialis and 12.1 °C and 566.6 day-degrees in G. nigrofuscata, respectively. The hatching dates predicted by these large thermal constants accorded with the hatching dates observed in the field, i.e., late June and mid-July in G. nigrofuscata and C. facialis, respectively. Therefore, the high thermal requirements for postdiapause development compel the cicadas to hatch in summer.     

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.     

Bionomics and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum, at constant temperatures

Development, survival, reproduction and population growth statistics of apterous virginoparae of woolly apple aphid, Eriosoma lanigerum (Hausmann) (Hemiptera: Aphididae) at constant temperatures of 10, 13, 15, 20, 25, 30 and 32°C are reported. The developmental times of all life stages were inversely related to temperature ranging from 10 to 30°C. Span of total development (time from birth to adulthood) decreased from 57.8 days at 10°C to 11.7 days at 30°C and increased to 16.8 days at 32°C. A good linear model fit (R²>0.96) between developmental rate and temperature in the range 10–25°C was observed for all life stages. The lower developmental threshold was estimated at 5.8°C for instar I, 4.8°C for instar II, 4.9° for instar III and 4.4°C for instar IV. The lower temperature threshold for total development was estimated at 5.2°C. The upper developmental limit was found to be 32°C. Mean degree-day accumulations required for completion of instars I, II, III, IV and total development were: 125.6, 51.0, 47.7, 50.7 and 267.6, respectively. Fecundity, larviposition period and adult longevity were reduced with increasing temperature. Net reproductive rate was greatest at 15°C whereas intrinsic rate of increase peaked at 25°C. Optimal performance, as measured by fecundity, survival and intrinsic rate of increase, ocurred in the range 13–25°C.     

Quantitative short-day photoperiodic response in larval development and its adaptive significance in an adult-overwintering cerambycid beetle, Phytoecia rufiventris

The chrysanthemum longicorn beetle, Phytoecia rufiventris, overwinters in the adult stage and reproduces in spring. Larvae of this beetle develop during summer inside a host stem or root. In the present study, photoperiodic control of larval development and its adaptive significance were examined in this beetle using an artificial diet. Larvae showed a short-day photoperiodic response at 25 °C with a critical day length of around 14 h; larvae reared under short-day conditions pupated, whereas those reared under long-day conditions entered summer diapause with some supernumerary molts and did not pupate. A similar response was found at 30 °C, but with a shorter critical day length. Below the critical day length, a shorter day length corresponded to a shorter larval period. Larvae transferred from long-day conditions to various photoperiods showed a similar quantitative response. Field rearing of larvae starting at various times of year showed that pupation occurs within a relatively short period in early autumn. Field rearing of pupae and adults at various times indicated that only pupation in early autumn results in a high survival rate until winter. Earlier or later pupation led to a low survival rate due to death before overwintering in the adult and pupal stages, respectively. Thus, in P. rufiventris, timing of pupation regulated by the quantitative short-day photoperiodic response is vital for survival. Relatively lower developmental threshold in the pupal stage supports this hypothesis.     

Diapause induction as an interplay between seasonal token stimuli,and modifying and directly limiting factors: hibernation in Chymomyza costata

Larvae of wild type (WT) strain of Chymomyza costata Zetterstedt (Diptera: Drosophilidae) enter diapause (stop developing) in response to short‐day signal at a constant 18 °C, whereas larvae of a non‐photoperiodic‐diapause (NPD) strain do not respond to photoperiodic signalling and continue in larval development irrespective of daylength. The present study shows that WT larvae also respond reliably to thermoperiodic signalling (daily cycles of temperature) under constant darkness, whereas the NPD larvae do not, suggesting that the pathways transducing the environmental token stimuli (photoperiod and thermoperiod) onto the diapause developmental programme might merge functionally in the central biological clock system known to be mutated in NPD strain. Temperature and larval population density modify the output of token stimuli signalling. High temperatures (>24 °C) tend to avert, whereas low temperatures (<18 °C), especially in combination with constant darkness, stimulate diapause induction in WT strain. Overcrowding (>200 larvae per 5 g of larval diet) lengthens the duration of larval development and induces a ‘diapause‐like’ developmental arrest of relatively weak intensity in up to 60% of larvae of both strains. At high temperatures (>30 °C), all WT larvae continue direct development but subsequently die during the pupal stage. Low temperature exposure (<12 °C) causes quiescence in the majority of the larvae of both strains. Starvation blocks development and causes mortality when applied in larvae younger than day 3 of the third instar. Older larvae survive starvation and their photoperiodically‐induced developmental pre‐programming is not affected. Collectively, the results show that diapause induction in C. costata is a result of various interacting effects of multiple environmental factors.     

Effect of temperature regime on diapause intensity in an adult-wintering Hymenopteran with obligate diapause

Osmia lignaria is a solitary bee that over-winters as a fully eclosed, cocooned, unfed adult. Our objective is to understand the effect of wintering temperature on diapause maintenance and termination in this species. We measure respiration rates and weight loss in individuals exposed to various wintering temperatures (0, 4, 7, 22 °C, outdoors) and durations (28, 84, 140, 196, 252 days). We use time to emerge and respiration response (respiration rate measured at 22 °C) as indicators of diapause intensity. Adults spontaneously lower their respiration rates to ∼0.1 ml/g h within 1 month after adult eclosion, indicating obligatory diapause. Non-wintered individuals maintain low respiration rates, but lose weight rapidly and die by mid-winter. In wintered adults, two phases can be distinguished. First, respiration response undergoes a rapid increase and then reaches a plateau. This phase is similar in bees wintered at 0, 4 and 7 °C. In the second phase, respiration response undergoes an exponential increase, which is more pronounced at the warmer temperatures. Composite exponential functions provide a good fit to the observed respiration patterns. Adults whose respiration response has reached 0.45 ml/g h emerge promptly when exposed to 20 °C, indicating diapause completion. Individuals wintered for short periods do not reach such respiration levels. When exposed to 20 °C these individuals lower their metabolic rate, and their emergence time is extended. The relationship between respiration rates and emergence time follows a negative exponential function. We propose two alternative models of diapause termination to interpret these results.     

Expression of heat shock protein 70a mRNA in Bombyx mori diapause eggs

In an effort to understand whether heat shock protein 70 (Hsp70) participates in the environmental 5 °C signal reception/transduction toward breaking embryonic diapause of the silkworm Bombyx mori, we isolated a cDNA for Hsp70a and examined the expression of Hsp70a mRNA in B. mori diapause and nondiapause eggs by quantitative real-time PCR. Hsp70a mRNA gradually increased in diapause eggs continuously kept at 25 °C after oviposition to maintain diapause. When diapause eggs were exposed to the diapause-terminating condition of 5 °C beginning at 2 days post-oviposition, Hsp70a mRNA increased beginning at 5 days post-cold treatment. Even in nondiapause eggs, Hsp70a mRNA increased slightly with exposure to 5 °C. These results suggest that Hsp70a is involved in reception/transduction of the diapause-terminating (5 °C) signal via gene activation. The expression patterns of Hsp70a mRNA are discussed in relation to those of the cold-response gene Samui.