Geographical variation and inheritance of the photoperiodic response controlling larval diapause in two distinct voltine ecotypes of the Asian cornborer Ostrinia furnacalis

The Asian corn borer Ostrinia furnacalis Guenée (Lepidoptera: Crambidae) enters facultative diapause as fully‐grown larvae in response to short day lengths during autumn. As a result of geographical variations in photoperiodic response, the moths from Nanchang (28.8°N, 115.9°E; NC strain) judge both LD 14 : 10 h and LD 15 : 9 h photocycles as long days and develop directly, whereas moths from Haerbin (44.9°N, 127.2°E; HB strain) judge the same photocycles as short days and enter diapause. Crosses between the two strains are used to evaluate the inheritance of diapause. The critical day lengths for diapause induction in the HB strain are significantly longer than those in the NC strain at all temperatures. The critical day length of F₁ progeny is intermediate between the two strains. However, the critical day length in all crosses is significantly longer with HB strain fathers or grandfathers than with NC strain fathers or grandfathers, indicating that the male parent has significantly more influence on the critical day length of subsequent progeny than the female. The results from all crosses under LD 14 : 10 h or LD 15 : 9 h photocycles at 25 °C show that the inheritance of diapause in O. furnacalis does not fit a purely additive hypothesis and that the capacity for diapause is transmitted genetically in the manner of incomplete dominance. The incidence of diapause for F₁ progeny under an LD 14 : 10 h photocycle is significantly higher than that under an LD 15 : 9 h photocycle, suggesting that the induction of diapause can be influenced by interactions between the F₁ genotype and photoperiod.     

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.     

Inheritance of Diapause in Crosses between the Northernmost and the Southernmost Strains of the Asian Corn Borer Ostrinia furnacalis

The northernmost Harbin strain (N strain) of the Asian corn borer, Ostrinia furnacalis enters facultative diapause as fully grown larvae in response to short daylengths; whereas the southernmost Ledong strain (S strain) exhibits almost no diapause under the same light conditions. In the present study, we examined the inheritance of diapause induction and termination by crossing the two strains under a range of environmental conditions. The N strain showed a typical long-day response with a critical daylength of approximately15.88 h at 22°C, 15.72 h at 25°C and 15.14 h at 28°C, whereas the S strain showed a weak photoperiodic response at 22°C. The F1 progeny also showed a long-day response at 22, 25 and 28°C. However, the critical daylengths in S ♀ × N ♂ crosses were significantly longer than those in N ♀ × S ♂ crosses, indicating a sex linkage in the inheritance of diapause induction, with the male parent having more influence on the following F1 progeny. The incidence of diapause in S ♀ × N ♂ crosses was the same as in the N strain under short daylengths of 11-13 h, indicating that diapause trait is completely dominant over the non-diapause trait. The critical daylength in backcross to N was significantly longer than it was in backcross to S, showing a grandfather gene effect. Whether the inheritance of diapause fits an additive hypothesis or not was dependent on the rearing photoperiod, and the capacity for diapause was transmitted genetically in the manner of incomplete dominance. The duration of diapause for the reciprocal crosses under different diapause-terminating conditions showed different patterns of inheritance. The results in this study reveal that genetic and genetic-environmental interactions are involved in diapause induction and termination in O. furnacalis.     

Responses to stepwise photoperiodic changes for the larval diapause of the Indian meal moth Plodia interpunctella

Abstract The Indian meal moth Plodia interpunctella Hübner (Lepidoptera: Pyralidae) diapauses as a last‐instar (fifth) larva. At 30 °C, no larvae enter diapause under any photoperiodic conditions; at 25 °C, the photoperiodic response curve is a long‐day type with a critical length of approximately 13 h light; at 20 °C, diapause is induced moderately even under long days (> 13 h). Cumulative effects of short days or long days on diapause induction are determined by alternate, stepwise and gradually changing regimes of photoperiod at 25 °C. When the larvae are repeatedly exposed to LD 16 : 8 h and LD 12 : 12 h photoperiods every other day, the incidence of diapause is 37%. When the larvae are placed under an LD 16 : 8 h photoperiod for 2 days and then under an LD 12 : 12 h photoperiod for 1 day, it is 38 %. Exposure to an LD 16 : 8 h photoperiod for 1 day and then to an LD 12 : 12 h photoperiod for 2 days induces only 15% diapause. This may indicate that the photoperiodic information is not accumulated in a simple fashion despite the generally accepted hypothesis (i.e. photoperiodic counter). Larvae exposed to an LD 16 : 8 h photoperiod for 5 days after oviposition express a very high incidence of diapause even under short days between an LD 2 : 22 h and LD 12 : 12 h photoperiod. After 10 days exposure to an LD 16 : 8 h photoperiod, however, the short day does not induce diapause strongly. On the other hand, an LD 12 : 12 h photoperiod in the early larval life is highly effective in the induction of diapause. A gradual increase or decrease of photoperiod (2 min day^?1) shows that the direction of photoperiodic change does not affect the diapause determination.     

Effects of photoperiod and temperature on diapause induction in Conogethes punctiferalis (Lepidoptera: Pyralidae)

The yellow peach moth, Conogethes punctiferalis (Guenée), a multivoltine species that overwinters as diapausing larvae, is one of the most serious insect pests on maize in China. Effect of photoperiod and temperature on larval diapause was examined under empirical laboratory conditions. Short‐day treatments caused larval diapause at 25°C, and the critical photoperiod was between 12 and 13 h (or 12 h 51 min) light per day. No sensitive instar was identified for diapause induction under alternated short‐ (L : D 11 : 13 h) and long‐day (L : D 14 : 10 h) treatments at different larval stages. However, accumulative treatment of three instars and 10 d under short‐day treatment was required for the induction of 50% larval diapause. All larvae entered diapause at 20°C, whereas less than 3% did so at 30°C, irrespective of the long‐ or short‐day treatment. Furthermore, under the short‐day treatment, more than 90% of larvae went into diapause with temperatures ≤ 25°C, but less than 17% did so at 28°C. In contrast, under the long‐day treatment, less than 19% of larvae went into diapause with temperatures ≥ 23°C. The forward shift (5°C) of critical temperature under the long‐day regime demonstrated the compensatory effect of temperature and photoperiod on diapause induction. In conclusion, C. punctiferalis had a temperature‐dependent type I photoperiodic diapause response; there was no sensitive instar for diapause determination, but the photoperiodic accumulation time countermeasures both of the short‐day cycles and the number of instars exposed, and the photoperiodic diapause response, was a temperature‐compensated phenomenon.     

Photoperiodic response in the subtropical and warm‐temperate zone populations of the southern green stink bug Nezara viridula: why does it not fit the common latitudinal trend?

The photoperiodic response of diapause induction is studied in females of five subtropical and warm‐temperate zone populations of Nezara viridula (L.) (Heteroptera: Pentatomidae) in Japan (26.4–34.7°N; 127.4–135.7°E). Laboratory tests at 25 °C demonstrate that both warm‐temperate and subtropical populations have pronounced photoperiodic responses of adult diapause induction. Under short‐day conditions (LD 10 : 14 h), 73–100% of females enter diapause, whereas, under long‐day conditions (LD 15 : 9 h), 87–100% of females are nondiapause and reproduce. When the critical photoperiod for diapause induction is plotted against the latitude of origin of each population, the data points do not show the expected tendency of increasing critical photoperiod northwards but, instead, vary between 12 h 15 min and 13 h 30 min. It is suggested that adults from different populations of such a highly migratory species move often among subtropical islands (by themselves or being assisted by typhoons), thus constantly destabilizing the photoperiodic responses. Although important for general synchronization of seasonal development in the tropics and subtropics, winter diapause might not be so critical for survival in mild winters in these zones. Such circumstances might make the selective pressure over the photoperiodic response less severe in the tropics and subtropics than in the temperate zone. It is suggested that the current climate warming might contribute to this situation as well by promoting polewards migrations and lessening the selective pressure of overwintering conditions.     

Effects of photoperiod and temperature on reproductive diapause in Ophraella communa (Coleoptera: Chrysomelidae), a potential biocontrol agent against Ambrosia artemisiifolia

Abstract To investigate the seasonal adaptation strategies of Ophraella communa to new habitats, the effects and regulation mechanisms of photoperiod and temperature on the reproductive diapause in a population collected from Changsha, Hunan were examined. Adults showed obvious reproductive diapause, which was regulated by photoperiod and temperature. At 30°C, there was no adult diapause occurring under either long‐day or short‐day conditions; at 25°C the pre‐oviposition period was short and fecundity was high in adult females under L : D 16 : 8 h, whereas under L : D 12 : 12 h, a few females entered reproductive diapause; at 20°C under short‐day conditions, all female adults entered diapause. The pre‐oviposition period was significantly prolonged when the pupae and adults were transferred from long‐days to short‐days, but the day length influence was not obvious when they were transferred only in the adult stage. However, the fecundity dropped greatly no matter whether the photoperiod shifted to short‐days only in the adult stage or whether the shift occurred in both the pupal and adult stage. The fecundity was extremely low when photoperiod shifted from long‐days to short‐days in both pupal and adult stages. This was an indication that the pupal and adult stages were the photoperiod‐sensitive stage for adult reproductive diapause. This was especially true for the photoperiod in the pupal stage, which has a distinctly significant regulative effect on reproductive diapause. Additionally, this article also addresses the reason for different photoperiodic response patterns in reproductive diapause induction between the Changsha strain and the Tsukuba strain (Japan) of O. communa.     

Photoperiodic and temperature control of nymphal development and induction of reproductive diapause in two predatory Orius bugs: interspecific and geographic differences

Abstract The effects of day‐length and temperature on pre‐adult growth and induction of reproductive diapause are studied in Orius sauteri and Orius minutus (Heteroptera: Anthocoridae) from northern (43.0°N, 141.4°E) and central (36.1°N, 140.1°E) Japan. In the north, at 20 °C, pre‐adult growth is slower under an LD 14 : 10 h photoperiod than under shorter or longer photophases. At 24 and 28 °C, the longer photophases result in shorter pre‐adult periods. Acceleration of nymphal growth by short days in autumn appears to be adaptive. In the central region, this response is less pronounced, suggesting that timing of adult emergence is less critical than in the north. Day length also influences the thermal requirements for pre‐adult development. The slope of the regression line representing temperature dependence of pre‐adult development is significantly smaller and the lower development threshold (LDT) is significantly lower under an LD 12 : 12 h photoperiod than under long‐day conditions. The weaker dependence of nymphal growth on temperature and the lower LDT in autumn might be adaptive. In the north, increased temperature shifts the critical day length of diapause induction and suppresses the photoperiodic response in O. sauteri but not in O. minutus. Further south, the incidence of diapause in both species is low even under short‐day conditions but the same interspecific difference is observed (i.e. increase of temperature affects the response in O. sauteri but not in O. minutus). This suggests seasonally earlier diapause induction with weaker temperature dependence in O. minutus than in O. sauteri.     

Effects of photoperiod and temperature on diapause induction and termination in the swallowtail,Sericinus montelus

Abstract Sericinus montelus overwinters as diapausing pupae. In the present study, the effects of photoperiod and temperature on diapause induction and termination of diapause are investigated. The results obtained demonstrate that high temperature can reverse the effect of short day‐lengths on diapause induction. Under an LD 12 : 12 h photoperiod, all pupae enter diapause at 15, 20 and 25 °C, whereas all pupae develop without diapause at 35 °C. No pupae enter diapause under an LD 14 : 10 h photoperiod when the temperature is above 20 °C. Photoperiodic response curves obtained at 25 and 30 °C indicate that S. montelus is a long‐day species and the critical day‐length is approximately 13 h at 25 °C. At 25 °C, the duration of diapause is shortest when the diapausing pupae are maintained under an LD 16 : 8 h photoperiod and increases under LD 14 : 10 h and LD 12 : 12 h photoperiods. Under an LD 16 : 8 h photoperiod, the duration of diapause is shortest when the diapausing pupae are maintained at 25 °C, followed by 20 and 30 °C, and then at 15 °C. These results suggest that a moderate temperature favours diapause development under a diapause‐averting photoperiod in this species. The duration of diapause induced by an LD 12 : 12 h photoperiod is significantly longer at 25 °C than those at 15, 20 and 30 °C, and is shortest at 15 °C. At 25 °C, the duration of diapause induced by LD 6 : 18, LD 12 : 12 and LD 13 : 11 h photoperiods is similar and longer than 90 days. Thus, the diapause‐inducing conditions may affect diapause intensity and a photoperiod close to the critical day‐length has significant influence on diapause intensity in S. montelus.     

Geographic variation in photoperiodic diapause induction and diapause intensity in Sericinus montelus (Lepidoptera: Papilionidae)

Abstract Due to the risk of extinction and ornamental value of the swallowtail butterfly, Sericinus montelus Gray (Lepidoptera: Papilionidae) in China, knowledge about local adaptations is important for the conservation and economical utilization of the species. In the present study, photoperiodic diapause induction and diapause intensity of S. montelus populations from Jiamusi (46°37′N), Beijing (40°15′N), Zibo (36°48′N), Fangxian (32°36′N), Wuhan (30°33′N) and Huaihua (27°33′N) were characterized at 25°C. Logistic regression analysis revealed a significant population × hours of light interaction, confirming that photoperiodic responses varied among populations. The critical photoperiod was positively correlated with latitude and increased toward the north at a rate of about 1 h for each 6.67 degrees of latitude. Survival analyses indicated that survival time of diapausing pupae before adult eclosion differed significantly among populations at 25°C and 16 : 8 L : D h. The mean duration of pupal diapause was also positively correlated with latitude. Our study reveals geographic variation in the critical photoperiod for diapause induction and in diapause intensity of S. montelus. These results provide useful information for our general understanding about seasonal adaptation in insects and may also be used to predict how geographic populations respond to climate warming.     

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 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.     

Maternal influence on diapause induction in Trichogramma (Hymenoptera: Trichogrammatidae): the dynamics of photosensitivity

It is known that the prepupal diapause in Trichogramma is dependent on temperature conditions of embryonic and larval development and on the photoperiodic conditions of preimaginal development of maternal females. However, the relative photosensitivity of different preimaginal stages has been never investigated. We studied the position of the photosensitive period over the preimaginal development of maternal females in T. embryophagum Htg. and T. principium Sug. et Sor. by transferring preimaginal stages between diapause‐averting ‘long’ day (L : D = 20 : 4) and diapause‐inducing ‘short’ day (L : D = 12 : 12). Results showed that the influence of maternal photoperiod on progeny diapause was determined during the late pupal stage (last days of preimaginal development at 20°C). During this time, the critical duration of the photoperiodic induction was extremely small: one short or long day caused almost the same effect as the permanent development under these conditions. As a whole, photoperiodic response in Trichogramma was found to be unusually rapid, labile and easily reversible which is probably explained by extremely small size and fast development of these egg parasitoids. The results of this study could be used for elaboration of optimal methods for Trichogramma mass rearing and storage and for prediction of its seasonal cycles under natural conditions.     

Biological characteristics of a non‐photoperiodic‐diapause strain of the cabbage beetle Colaphellus bowringi (Coleoptera: Chrysomelidae)

Comparison of biological characteristics between diapausing and non‐diapausing strains of insects provides some insights into the mechanisms regulating diapause. In this study, biological characteristics, especially diapause characteristics and life‐history traits, of a non‐photoperiodic‐diapause (NPD) strain of the cabbage beetle Colaphellus bowringi were compared with those of a normal, high‐diapause (HD) strain that enters diapause in response to either long day length or low temperature. The NPD strain did not enter diapause at any photoperiod at 22°C or higher, but still had a capacity to enter diapause at low temperatures. Although diapause could be induced in both strains by exposure to 20°C, the proportion of individuals having shorter diapause duration was greater in the NPD strain compared to the HD strain. The NPD strain had significantly lower hatching and larval survival rates, longer developmental duration of immature stages, smaller body size and lower longevity and female fecundity compared to the HD strain. The NPD strain of this species is a promising model for investigating diapause regulation in insects in general.     

Photoperiodic Control of Pre-Adult Development and Adult Diapause Induction in Zoophytophagous Bug Dicyphus errans (Wolff) (Heteroptera,Miridae)

Zoophytophagous plant bugs (Heteroptera, Miridae) increasingly attract interest as agents of biological plant protection. In the laboratory experiment, the effects of the day length and temperature on the duration of the pre-adult period and on induction of facultative winter adult diapause were studied in Dicyphus errans (Wolff, 1804) collected in Italy. The experiment demonstrated that at 20°C the duration of the pre-adult period of D. errans significantly depended on the day length. On average, females developed 1.3 days longer than males and, at the same time, the day length equally influenced the duration of the pre-adult period in both sexes. The pre-adult period was the shortest under short-day conditions (10 to 12 h of light per day), reached its maximum at day length of 14 h, but then decreased at 15 h, and at day length of 16 h it was as short as under short-day conditions. Also, a pronounced long-day type photoperiodic response of adult diapause induction was recorded in females of D. errans at 20°C: under short-day conditions (10 to 14 h of light per day) almost all females entered diapause, whereas under long-day conditions (15 and 16 h of light per day) about 90% of females were mature. The threshold of this photoperiodic response was close to 14 h 30 min. The mean (± S.D.) egg load of mature females was 6.3 ± 4.0 eggs per female and did not depend on the day length at which the female was reared before and after the final molt. When photoperiodic response of adult diapause induction was observed at two constant temperatures (20 and 25°C), the proportion of mature females depended significantly on the day length but not on the temperature: the shapes of the photoperiodic response curves of diapause induction were almost the same within the near-threshold zone at 20 and 25°C, i.e., the photoperiodic response was thermostable. The set of two photoperiodic responses manifested at different stages of the species’ life cycle has an obvious adaptive significance. In Central Europe, D. errans has 2 or 3 generations per year and hibernates at the adult stage. Due to the thermostable photoperiodic response, females enter diapause always at the same time at the end of summer, regardless of the weather conditions of a particular year. When oviposition and pre-adult development are extended over a prolonged period in summer, nymphs from the later eggs might not be able to molt to adults in due time and then fully prepare for stable winter diapause. Under such circumstances, the photoperiodic response controlling the rates of pre-adult development acquires apparent adaptive meaning: with an autumnal shortening of the day length to 10–12 h, even under conditions of seasonal decrease in temperature, the rates of nymphal development increase and, thus, the chances of nymphs from the later eggs to molt to adults and properly prepare for overwintering also increase. The new data should be taken into account when analyzing the seasonal cycle of D. errans and developing the programs of mass rearing of this zoophytophagous mirid as an agent of biological plant protection.

     

Geographic variation in embryonic diapause,cold‐hardiness and life cycles in the migratory locust Locusta migratoria (Orthoptera: Acrididae) in China

To investigate geographic adaptation of the migratory locust Locusta migratoria in China, locusts were collected from six localities, ranging from 47.4°N to 19.2°N. Using offspring from the various populations, we compared embryonic diapause, reproductive traits, cold‐hardiness and adult body size. The incidence of embryonic diapause was influenced by the genetic makeup, parental photoperiod, and incubation temperature of the eggs. The northern strain (47.4°N) produced diapause eggs under all photoperiodic conditions, whereas the other strains produced a higher proportion of diapause eggs when exposed to a short photoperiod. The incubation temperature greatly influenced diapause induction. At a low temperature, all eggs entered diapause, even some of those from a tropical strain (19.2°N) in which no diapause was induced at high temperatures. Photoperiodic changes during the parental generation affected the incidence of embryonic diapause. Diapause intensity decreased with decreasing original latitude. Cold hardiness was compared by exposing eggs in diapause to either ?10 or ?20°C for various periods; the northern strain was more cold‐hardy than the southern strain, although some eggs in the tropical strain were probably not in a state of diapause. Adult body size and head width showed a complicated pattern of variation along the latitudinal gradient, whereas egg pod size (egg pod width and egg number) and hatchling weight tended to decrease with decreasing latitude. These results reveal that L. migratoria has adapted to local environments and that the latitudinal gradient appears to play an important role in shaping L. migratoria life cycle and development.     

Effect of photoperiod on winter and summer diapause of the soybean pod borer Leguminivora glycinivorella (Lepidoptera: Tortricidae)

To understand the geographical differences between diapause systems and synchronization of adult occurrence in the soybean pod borer Leguminivora glycinivorella (Lepidoptera: Tortricidae), we examined the timing of winter diapause termination and intensity of summer diapause using univoltine and potentially bivoltine individuals in Iwate, Japan. In laboratory rearing experiments of mature larvae maintained at constant temperature (20 °C), winter diapause intensity weakened by January without photoperiodic responses. Meanwhile, summer diapause was maintained by the long day length and presumably terminated with the photoperiodic transition from long to short day length. The intensity of summer diapause was stronger for cocoons that transitioned from a 16 h light to 8 h dark (LD 16:8) to a LD 15:9 photoperiod than for those that transitioned from LD 15:9 to LD 14:10. These results suggest that populations distributed in relatively low-latitude areas, with partly or potentially bivoltine individuals, would have a weaker summer diapause or none at all. Moreover, sexual differences in the number of days to emergence were not detected when individuals experienced a photoperiodic transition from long to short day length, suggesting that the summer diapause system may function to synchronize the emergence of males and females in the population examined.     

Genetic analysis of diapause capability and association between larval and pupal photoperiodic responses in the flesh fly Sarcophaga similis

Abstract A variant of the flesh fly Sarcophaga similis, which does not enter pupal diapause even under diapause‐inducing conditions, is established by artificial selection. Interline crosses of the wild‐type and the variant revealed that the diapause capability is inherited in an incomplete dominant manner. Neither sex linkage nor a maternal factor is involved in the mode of inheritance. Genetic and genetic–environmental interactions are involved in the induction of diapause. In the wild‐type, the post‐feeding larval period is prolonged in response to short days. However, in the variant, the duration of the larval period under short‐day conditions is identical to that under long‐day conditions. Long‐day responses under short‐day conditions at both the larval and pupal stages in the variant indicate that the responses are established by common factors in both stages. The factors causing the long‐day responses under short‐day conditions at different developmental stages in the variant remain to be identified. However, it is plausible to hypothesize that common factors regulate ecdysteroid release at both the larval and pupal stages, and that malfunction of the system regulating these factors triggers such ecdysteroid release, irrespective of the photoperiod. Hence, the photoperiodic responses disappear in the variant in both stages. The adult stage of the variant has a functional circadian clock, suggesting that the nondiapause phenotype is not necessarily involved in malfunction of the circadian clock.     

Maternal thermal and photoperiodic effects on the progeny diapause in Trichogramma telengai Sorokina (Hymenoptera: Trichogrammatidae)

Although maternal photoperiodic and maternal thermal effects on the progeny diapause have been demonstrated in a number of insect species, their interaction has been rarely studied. We investigated this interaction in Trichogramma telengai. In a series of experiments, maternal females were reared at day lengths of 12–18 h and at temperatures of 17, 20, 25 and 30°C. Their progeny developed under day length of 12 h and temperatures of 13, 14 and 15°C. The experiments showed that both short day and low temperature experienced by the maternal generation significantly increased the proportion of diapausing progeny. In particular, the threshold of the maternal photoperiodic response decreased with temperature. Under combinations of photoperiod with daily thermoperiod, the role of the “night” temperature in the induction of diapause in the progeny was much more important than that of the “day” temperature. We conclude that the interaction pattern between the photoperiodic and thermal maternal effects in T. telengai is generally the same as that between the photoperiodic and thermal responses directly influencing diapause induction in other long‐day insects. The threshold temperature of the maternal thermal response of T. telengai was about 25–27°C, while diapause can be induced if larvae develop at temperatures not higher than 15–16°C. This suggests that, at least in the studied Trichogramma species, the maternal thermal effect has no ecological value. In the practice of biocontrol, however, rearing of Trichogramma wasps at high temperature can drastically reduce the proportion of diapausing progeny.     

Photoperiodic induction of prepupal diapause and its role in synchronization with host phenology in the hibiscus caterpillar,Xanthodes transversa

The hibiscus caterpillar, Xanthodes transversa (Lepidoptera: Noctuidae), is a multivoltine insect that is an important pest of Malvaceae plants such as the okra, Abelmoschus esculentus, and the common rose mallow, Hibiscus mutabilis, in Japan. In the present study, the effects of photoperiod and temperature on the induction of prepupal diapause and the adaptive significance of this diapause were examined in a local population of X. transversa in Miyazaki, Kyushu, southwestern Japan. Larvae showed a long‐day photoperiodic response for controlling the induction of prepupal diapause with a critical day length between 13 and 14 h at 20 to 25°C. Under long‐day conditions larvae rapidly pupated from the sixth instar, but under short‐day conditions they entered diapause in the prepupal stage. Diapause occurrence in the field increased in late September, which was consistent with the laboratory results in terms of the photoperiodic response. Non‐diapause development after this time is maladaptive because most larvae of the next generation could not reach the critical stage (prepupae) before winter and died during early winter in outdoor experiments. Larvae suffered from a high rate of mortality when fed leaves collected late in autumn. The photoperiodic response for controlling the induction of diapause in this insect may play an important role in synchronizing the life cycle with the seasonal changes in food and temperature conditions.