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.     

Sexual dimorphism in photoperiodic plasticity of body mass in the green dock leaf beetle <Emphasis Type="Italic">Gastrophysa viridula</Emphasis> (Coleoptera,Chrysomelidae)

Sexual dimorphism in body mass was studied in prepupae, pupae and immature adults of the green dock leaf beetle Gastrophysa viridula under a constant temperature of 24°C and two photoperiods (12L:12D and 22L:2D). Females were significantly heavier compared to males at all the three developmental stages; just after emergence from pupae they weighed, on average, 11.4 mg, while males—10.3 mg. Sex differences in photoperiodic body mass plasticity were revealed: under short-day conditions pupal and adult mass in females was significantly higher compared to long day, but not in males. Such a response to short day (the factor inducing reproductive diapause in this species) is likely to promote accumulation of greater amount of nutrients by females which is necessary for earlier oviposition after overwintering.     

Photoperiodic control of larval development in the semivoltine cockroachPeriplaneta japonica (Blattidae: Dictyoptera)

Periplaneta japonica is semivoltine, entering early diapause in any (except the first) larval instar before the last, and late diapause in the last instar. Early diapause was induced under a short day of 13 h or less at 28°C, and under both short and long daylength (12–16 h) at 20°C. The shorter the daylength and the lower the temperature, the younger the instar was entering early diapause. Early diapause was terminated by a long day (16 h) or a high temperature (28°C), after which larvae grew faster in short days than in long days until the last instar, when they again entered diapause, always in short days and frequently in long days. This late diapause was terminated also by an increase in daylength and was always followed by adult emergence. In this case, 13 and 14 h daylengths after exposure to 12 h daylength were as effective as 16 h daylength. Ourdoor samples collected in late autumn, winter and early spring at Hirosaki (40.5°N) comprised two distinct size groups, corresponding with the early and late diapause instars. Based on these results, the seasonal development strategy and intriguing aspects of the photoperiodic response in this cockroach are discussed.     

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 of diapause induction in the pine caterpillar, Dendrolimus punctatus

The pine caterpillar, Dendrolimus punctatus (Walker) (Lepidoptera: Lasiocampidae), is a multivoltine pest of pine trees in China, overwintering as larvae. Winter diapause was induced by short day length. The critical night length was about 10 h 40 min at 25, 28, and 31 °C in the field, showing a temperature‐compensated diapause induction. Transfer experiments from a short night (L16:D8) to a long night (L12:D12) or vice versa at different times after hatching showed that sensitivity to day length was restricted to the first 14 days; the required day number for a 50% response at 25 °C was about 3.5 days for short nights but 7.5 days for long nights, indicating that short nights are photoperiodically more effective. When four successive short nights (L16:D8) were used to interrupt the long‐night regime (L12:D12) at different development stages and vice versa, the results showed that the highest sensitivity to photoperiod occurred on the 4th?8th day, corresponding to the second larval instar. Experiments of alternating short‐night (L16:D8) and long‐night (L12:D12) cycles during the larval period showed that the information of short nights as well as long nights could be accumulated. By rearing the larvae under conditions other than 24‐h light–dark cycles, we clearly showed that the dark period (scotophase) played a major role in the determination of diapause. The Nanda‐Hamner and Bünsow experiments failed to reveal rhythmic fluctuations with a period of about 24 h in the occurrence of diapause. Therefore, the photoperiodic clock in D. punctatus is an hourglass timer or a damped circadian oscillator.     

Effect of photoperiod on the induction and maintenance of diapause and on development of the predatory bugPodisus maculiventris (Hem.: Pentatomidae)

The predatory bugPodisus maculiventris Say displays a reproductive diapause. In the laboratory, at 23±1°C, when the pre-imaginal stages were reared on larvae ofGalleria melonella L. (Lepidoptera: Pyralidae) under a short photoperiod of 8L: 16D, diapause was induced in all adult females, whereas under a long photoperiod of 16L: 8D none entered diapause. Nymphal development was faster under the diapause-inducing short photoperiod than under the diapause-averting long photo-period. By contrast, embryonic development was faster under the long than under the short photoperiod. Diapause maintenance was also under the control of photoperiod. After a 10-d-period of chilling at 4°C, diapause was quickly terminated in adults kept under long photoperiod whereas it was maintained in most of those kept under short photoperiod. The predation rate of non-diapause or post-diapause adult females was much higher than that of diapause females.     

Temperature and photoperiodic control of diapase induction in the ant <Emphasis Type="Italic">Lepisiota semenovi</Emphasis> (Hymenoptera,formicidae) from Turkmenistan

The ants L. semenovi has been found to belong to species with endogenous-heterodynamic seasonal life cycles with the obligate diapause induced predominantly by factors internal for a colony, whereas external ecological factors (photoperiods and temperature) produce merely modifying effects by accelerating or delaying the diapause onset. The photoperiodic and temperature regulation of diapause induction in larvae and queens is shown. Under effect of short days and low temperature the periods of larval pupation and queen oviposition in a colony are shortened markedly, i.e., the diapause of larvae and queens occurs earlier. The daily rhythms of temperature 15/25°C and particularly 20/30°C as compared with constant temperatures 20 and 25°C that correspond to the mean circadian temperatures of the thermorhythm, inhibit manifestations of the short day effects by stimulating the non-diapause development and increasing duration of the seasonal development cycle of ant colonies. The L. semenovi photoperiodic reaction is quantitative, as development and pupation of larvae and egg-laying of queens cease sooner or later under both the short and the long days, but in the latter case significantly later. Thus L. semenovi is one more example among very rare ant species that are revealed to have the photoperiodic regulation of the colony development seasonal cycle.     

Larval diapause of Monochamus urussovi and photoperiodic effects on larval development in tree bolts

To determine the larval diapause and the effect of photoperiod on development in Monochamus urussovi (Coleoptera: Cerambycidae), larvae were reared on Abies sachalinensis and Picea jezoensis logs and bolts. Larvae stopped developing in the final instar at 25°C and 16L : 8D (16 h light and 8 h dark) whereas an exposure to 5°C in the dark (134 days) following acclimation at 12°C under natural daylength led to adult emergence. When larvae were reared under 8L : 16D or 16L : 8D at 25°C with an intervening period of chilling at 5°C in the dark (112 days), a photoperiod of 8L : 16D induced a shorter time required for adult emergence after being returned to 25°C, and smaller adult body size than 16L : 8D.     

Photoperiodic Regulation of the Daily Rhythm of <Emphasis Type="Italic">Trichogramma</Emphasis> Adult Emergence (Hymenoptera,Trichogrammatidae)

Laboratory experiments demonstrate that the adult emergence rhythms of two Trichogramma species (T. telengai Sor. and T. principium Sug. et Sor.) significantly depend on photoperiod. The synchronicity of adult emergence is maximal at day lengths of 12–16 h. Under longer days (18–20 h) adult emergence is extended, whereas under very short days (4–8 h) a substantial proportion of adults emerge before light-on. The pattern of T. telengai emergence under day length of 4 h suggests the possibility of the existence of two fractions (the first that emerges in the dark and the second that waits for light-on) which results in two peaks of emergence. As a whole, these data agree well with the hypothesis of the periods of “potential readiness” and “obligate activity” forwarded by V.B. Chernyshev. The results of the study can be used for optimization of the methods for laboratory and mass rearing of Trichogramma species.     

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.     

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.     

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

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

Univoltine seasonal cycle and obligate diapause in the noctuid moth <Emphasis Type="Italic">Charanyca trigrammica</Emphasis> Hufn. (Lepidoptera,Noctuidae)

Charanyca trigrammica is a univoltine species over its whole distribution range. To study the diapause-inducing factors, caterpillars were reared at 24°C and day lengths of 12, 13, 14, 15, 16, 17, and 18 h. In all the studied regimes, pupation was not recorded, suggesting an obligate diapause. However, the development was faster under short days. The threshold of this short-day photoperiodic response was ca 17 h. This type of environmental regulation of development may be considered a special feature of diapause occurring in the last larval instars. Hence, the obligate diapause in the last larval instars determines the univoltine cycle of Ch. trigrammica in Belgorod Province of Russia, ensuring successful hibernation of all individuals.     

Effect of daylength and temperature on the larval diapause of the sunflower moth, Homoeosoma electellum

The effect of daylength and temperature on the regulation of the larval diapause of a central Missouri population of the sunflower moth, Homoeosoma electellum, was examined. Fully grown fourth-instar larvae exhibit a facultative diapause. Measurements of the effect of photoperiod on diapause induction revealed critical photoperiods of about 13 h 30 min light/day at 20°C, and between 11 h 45 min and 12 h light/day at 23°C. Third and fourth-instar larvae were shown to be the main sensitive stages for diapause determination. Daylength was also shown to be an important regulator of the rate of diapause development. A short day of LD 10:14 h permitted only a low rate of diapause development, whereas long days of LD 14:10 h and LD 16:8 h accelerated diapause development at 25 and 30°C. When long days were alternated with short days at 30°C the accelerating effect of long days on diapause development was not found. Systematic transfers of chilled diapausing larvae revealed an accelerated diapause development in groups transferred from 10 to 30°C LD 10:14 h, but diapause development was not accelerated in groups transferred from 10 to 30°C LD 16:8 h.     

Effects of thermoperiods on the reproductive activity of females and on the maternal induction of larval diapause in the blowfly, <Emphasis Type="Italic">Calliphora vicina</Emphasis> R.-D. (Diptera,Calliphoridae)

The interaction of thermoperiod and photoperiod in their influence on the reproductive maturation of females and on the induction of the maternal effect determining larval diapause of the progeny of the blowfly, Calliphora vicina, was first investigated under laboratory conditions. Under the combination of a day length of 12 h with a thermoperiod (the alternation of 12 h long periods with temperatures of 10 and 20°C) the reproductive maturation of females was faster than at the corresponding mean constant temperature of 15°C. Under the “natural” thermoperiod, when the period with a temperature of 10°C coincided with “night-time” (the dark phase of the diurnal light-dark cycle) the maturation of females was slower than that under the “inverted” thermoperiod, when the period with a temperature of 10°C coincided with “day-time” (the light phase of the diurnal light-dark cycle). The proportion of diapausing individuals was maximal in the progeny of females kept at 20°C and decreased with the increase in temperature. Under thermoperiods (the alternations of 12 h long periods with temperatures of 20 and 26°C) the proportion of diapausing progeny was lower than that under the corresponding mean constant temperature of 23°C, but under the inverted thermoperiod with a high night temperature this effect was much stronger. In combination with the results of our previous studies, these data support the hypothesis that the effects of “night” and “day” temperatures are substantially different only when the thermal response interacts with a strong photoperiodic response.     

Photoperiodic responses of pupa and adult determining maternal influence on progeny diapause in two species of the genus Trichogramma (Hymenoptera, Trichogrammatidae)

The adult photoperiodic responses of two Trichogramma species were investigated by exposing adults during 3 days to one of the seven light: dark regimes: L: D = 4: 20, 8: 16, 12: 12, 14: 10, 16: 8, 18: 6, and 20: 4. The preimaginal stages of these individuals developed under short (L: D = 12: 12), long (L: D = 20: 4) or intermediate photoperiods (L: D = 14: 10 and L: D = 16: 8 for Trichogramma principium and T. embryophagum, respectively). The progeny of these females developed under short day L: D = 12: 12 and at two moderately diapause-inducing temperatures (13 and 14°C for T. principium, 14 and 15°C for T. embryophagum). In both Trichogramma species which developed at both temperatures, the percentage of diapausing prepupae was significantly dependent both on the photoperiodic conditions of the preimaginal development of the maternal generation and on the photoperiod which influenced the adult females. The adults showed a typical long-day photoperiodic response with a threshold day length of ca 13 h in T. principium and ca 16 h in T. embryophagum, which practically coincided with the thresholds of the pupal photoperiodic responses of these species revealed in our previous studies. However, the ultra-short photoperiods (L: D = 4: 20 and 8:16) caused a relatively stronger diapause-inducing effect on the progeny when applied to the adult females than when it was applied to the pupae. Thus, in both the Trichogramma species studied, the patterns of photoperiodic responses of pupa and adult were somewhat different although they almost coincided in the “ecologically significant” part of the photoperiodic scale.     

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.     

Effects of low temperature on diapause termination and body colour change in adults of a stink bug, Plautia stali

Abstract. Diapause adults of Plautia stali Scott maintained at 20°C under short day conditions (LD 12:12 h) were exposed to four temperatures of 5–20°C to examine the effect on diapause development which was assessed in terms of oviposition. Diapause adults kept at 20°C under short day conditions changed their body colour gradually from brown to green and started egg laying after a prolonged preoviposition period. Those transferred to either 10 or 15°C also showed colour change but did not lay eggs. Bugs exposed to 5°C underwent neither body colour change nor oviposition and died more rapidly than those kept at higher temperatures. When 30-day-old diapause adults were chilled at 5, 10 or 15°C for 30 or 60 days and returned to 20°C and long day conditions (LD 16:8 h), the preoviposition period varied primarily depending on the chilling, but not on the temperature. On the other hand, when 60day-old diapause adults chilled for 30 days were observed at 20°C and long day conditions, their preoviposition period tended to be longer as the chilling temperature was lower In this case, a temperature of 10°C appeared to intensify diapause. Therefore, the effect of chilling on diapause development varied depending on the age at which insects were chilled. When chilled bugs were transferred to short day conditions at 20°C, most females failed to lay any eggs and some turned green, then after a while, some green bugs changed to brown again. These results indicate that bugs remained sensitive to short day conditions even after a 60-day chilling at 10 or 15°C.     

Inheritance of photoperiodic induction of larval diapause in the Asian corn borer Ostrinia furnacalis

The Asian corn borer Ostrinia furnacalis (Guenée) enters facultative diapause as fully‐developed larvae in response to short‐day conditions. As a consequence of geographical variation in photoperiodic response, moths from Nanchang (28°46′N, 115°50′E) enter diapause in response to short day‐lengths (D strain), even at the high temperatures whereas moths from Ledong (18°47′N, 108°89′E) exhibit almost no diapause under the same conditions (N strain). In the present study, crosses between the two strains are used to evaluate the inheritance of diapause under different photoperiods at temperatures of 22, 25 and 28 °C. The moths, both reciprocal crosses and backcrosses, show a clear long‐day response, similar to that of the D strain, suggesting that the photoperiodic response controlling diapause in this moth is heritable. However, the critical day‐length for induction of diapause is shorter in hybrids than in the D strain. The N strain also shows a short‐day photoperiodic response at the lower temperature of 22 °C, indicating that the N strain still has the capacity to enter a photoperiodically‐induced diapause, depending on the rearing temperature. The incidence of diapause in all crosses is highest with D strain fathers or grandfathers and lowest with N strain fathers or grandfathers, indicating that the male parent has significantly more influence on the incidence of diapause of subsequent progeny than the female. The results obtained from all crosses under LD 12 : 12 h or LD 13 : 11 h photocycles at 25 °C show that inheritance of diapause in O. furnacalis does not fit an additive hypothesis and that the capacity for diapause is transmitted genetically in the manner of incomplete dominance.     

The diapause syndrome in the seed-eating caterpillar of Coleophora alticolella

Adult emergence in Coleophora alticolella held at 15 °C was accelerated by exposure to L18:D6 in autumn and midwinter. The effect decreased during winter and exposure of individuals, held at low temperature over winter, to L18:D6 or L6:D18 at 15 °C at the end of March resulted in the same mean emergence date. Long daylength experienced at 5 °C did not promote emergence nor did exposure to low temperature during winter. The number of adults emerging increased with the length of time cultures were held on short day but was always below 50% of the larvae. When larvae were exposed to L18:D6 and L6:D18 at 15 °C at the end of March, on long day 61% adults emerged and 39% remained in diapause, whereas on short day, 25% became adult and 75% remained diapausing larvae. The possibility of cohort splitting, with some individuals undergoing prolonged diapause, is discussed.