Role of photoperiod and temperature in the induction of overwintering pupal diapause in the spotted tentiform leafminer,Phyllonorycter blancardella

The role of photoperiod and temperature in the induction of overwintering diapause inPhyllonorycter blancardella (F.) (Lepidoptera: Gracillariidae) was examined in the laboratory and field using leafminers from commercial apple orchards in Ontario, Canada.P. blancardella exhibited a long-day response to photoperiod: long daylengths resulted in uninterrupted development whereas short daylengths induced diapause. The estimated critical photoperiod for diapause induction was L14.25∶D9.75. The larvae of leafminers destined to enter diapause took ca. 3× longer to complete development than the larvae of non-diapausing leafminers. The development prolonging effect of photoperiod decreased with decreasing daylength. Temperature modified the diapause inducing effect of photoperiod. At L14.25∶D9.75, diapause incidence was similar at 15 and 20°C but was lower at 25°C. Photoperiod also altered the normal relationship between development rate and temperature. At L14.25∶D9.75, the duration of larval development of diapausing leafminers was similar at 15, 20 and 25°C. Temperature alone is unlikely to have a role in the induction of diapause because leafminers exposed to natural late summer and fall temperature regimes and L16∶D8 did not enter diapause.     

Feeding,reproductive and locomotor activities in diapausing and non-diapausing adults ofDrosophila

Feeding, reproductive and locomotor activities of fourDrosophila species were studied under short and long daylengths at 15°C. A short daylength induced firm reproductive diapause in experimental strains ofD. subauraria andD. triauraria from northern Japan, but very shallow diapause in those ofD. lutescens andD. rufa from southern Japan. A subtropical strain ofD. triauraria had no diapause. The influence of diapause on feeding activity was detected only in aged (> 12 day old) females; that is, the feeding activity was lower in diapausing females than in non-diapausing ones. Females that do not produce eggs would not require so much energy. On the other hand, young adults of the study species exhibited a high feeding activity and rapidly increased bodyweight irrespective of sex and the diapause state. They would need nutrition to build up their adult body. In males, the feeding activity decreased with age irrespective of the diapause state. Males would not require so much energy for reproductive activity. Diapausing males became heavier than non-diapausing males, perhaps because they accumulated triacylglycerols in fat bodies. However, female bodyweight did not differ by the diapause state, perhaps because diapausing females accumulated triacylglycerols and reproducing females had eggs in their ovaries. InD. triauraria, diapausing individuals exhibited somewhat lower locomotor activity than non-diapausing ones.     

European field collections and Canadian releases ofCeranthia samarensis (Dipt.: Tachinidae), a parasitoid of the gypsy moth

A programme to collect, import and release into Canada the gypsy moth parasitoid,Ceranthia samarensis (Diptera: Tachinidae) is described. The parasitoid's potential for biological control in Canada is also discussed. The parasitoid was collected in Europe by exposing experimental gypsy moth larvae in areas where local gypsy moth populations were at low densities. Following field exposure, the host larvae were returned to the laboratory and parasitoids reared from them. This technique has shown thatC. samarensis is the suffers 7–16% hyperparasitism. From 83–90% of theC. samarensis typically enter diapause as pharate adults within the puparia. Laboratory tests of post-exposure host rearing conditions indicate that constant temperatures disrupt the normal parasitoid diapause and that this effect can not be offset by use of either static long or short photoperiods or natural daylengths. Shipping and cold-storage procedures for puparia are described. Post-storage time to emergence of adultC. samarensis decreased with longer cold storage periods and with higher post-storage incubation temperatures. Emergence requires 112 degree-days above a threshold of 8°C after a period of at least 8 months cold storage. Releases of adultC. samarensis into field cages at four locations in southern Ontario are documented. While dissection of host larvae from the field cages has failed so far to demonstrate evidence of parasitism, we remain hopeful that some establishment of the parasitoid has occurred.      

Male mating activity and genetic aspects in imaginal diapause of Drosophila triauraria

In Drosophila triauraria Bock & Wheeler (Diptera: Drosophilidae) of which females were known to enter reproductive diapause at short daylengths, males also showed reduced mating activity at short daylengths, i.e., males as well as females entered reproductive diapause. The critical daylength for diapause induction did not differ between females and males. Both male and female diapause ended even under short daylengths, but the male diapause was somewhat weaker than the female diapause.The critical daylength and the diapause rate varied geographically in this species. In the cross between diapausing and non-diapausing strains, the critical daylength and the diapause duration inherited in a quantitative manner. On the basis of the present and previous crossing experiments, some models are proposed on the mechanism of diapause induction of this species.

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Résumé Les femelles de D. triauraria Bock & Wheeler (Dipt. Drosophilidae) sont connues pour présenter une diapause reproductrice aux photophases courtes. Les mâles eux aussi ont révélé une activité sexuelle réduite aux photophases courtes, c'est-à-dire qu'ils sont entrés comme les femelles en diapause reproductive. Les photophases critiques pour l'induction de la diapause des mâles et des femelles n'ont pas présenté de différences. Les diapause des mâles et des femelles s'achèvent même sous courtes photophases, mais la diapause mâle était quelque peu plus faible que la diapause femelle.La photophase critique et le taux de diapause ont varié en fonction de l'origine géographique dans l'espèce actuelle. Lors de croisements entre lignées diapausantes et non-diapausantes, la photophase critique et la durée de la diapause ont été héritées quantitativement. A partir de ces expériences et d'expériences précédentes de croisements (Kimura, 1983), quelques modèles de méchnisme d'induction de la diapause de cette espèce sont proposés.

     

Life history of Galerucella nymphaeae and implications of reproductive diapause for rearing univoltine chrysomelids

Abstract. Galerucella nymphaeae L. , a chrysomelid that feeds on Lythrum spp., water lily and water chestnut, is closely related to two European species that were recently introduced into North America for biological control of L.salicaria , purple loosestrife. To develop a paradigm for continuously rearing these and other univoltine chrysomelids, we conducted field and laboratory studies on G. nymphaeae's development , reproduction and diapause.
A high incidence of reproduction without diapause occurred when fourth instars, pupae, and adults (held as pairs) experienced very long daylengths (LD18:6 h), i.e. longer than those G.nymphaeae encounters in nature. Under similar photoperiodic conditions, adults maintained in groups showed a significantly higher rate of reproductive diapause than those held as pairs. Females laid three to seven egg masses/week, and the size of egg masses varied between nine and nineteen eggs. During their reproductive lifetimes, individual females showed a highly significant propensity to lay a consistent number of eggs/egg mass.
After diapausing under short daylengths and low temperature (LD 10:14h, 5C), adults transferred to long days (LD 18:6h at 21C) had high rates of diapause termination and postdiapause oviposition. In contrast, those transferred to short daylengths (LD 10:14h at 21C) had low rates of reproduction.
Laboratory-derived heat-degree models accurately predicted egg and pupal, but not larval, development in the field. In nature, most females in the summer generation entered reproductive diapause without ovipositing; a small proportion of females that emerged relatively early (by mid June) oviposited before entering diapause. The overwintering population consists of adults from the first-generation and a small number from the second generation.     

In vivo and in vitro effects of 20-hydroxyecdysone and methoprene on diapause maintenance and reproductive development in Musca autumnalis

Abstract. Face flies overwinter as adults in reproductive diapause. Administration of 20-hyroxyecdysone and/or methoprene induced reproductive development in diapausing flies which were maintained in a diapause-inducing environment. Hormone effects were additive and female flies were more sensitive than males. Release of vitellogenin from cultured fat body was stimulated by 20-hydroxyecdysone or methoprene. Transfer of flies from diapause to diapause-breaking environments induced some to break diapause, but this decreased with the time flies had been in a diapause-inducing environment. In contrast, topical application of methoprene to diapausing flies induced reproductive development irrespective of their ages even when they were kept in the diapause-inducing environment for 80 day degrees above a 12°C base temperature (14.5 days). Therefore diapause induction must depend on hormone levels less than some threshold level. The putative threshold varied according to diapause propensities of different genetic lines. Lines showing high frequencies of diapause required greater amounts of methoprene for reproductive development in diapause conditions than did lines showing low frequencies of diapause.     

A comparison of photoperiodic control of diapause between aestivation and hibernation in the cabbage butterfly Pieris melete

In the cabbage butterfly, Pieris melete, summer and winter diapause are induced principally by long and short daylengths, respectively; the intermediate daylengths (12-13 h) permit pupae to develop without diapause. In this study, photoperiodic control of summer and winter diapause was systematically investigated in this butterfly by examining the photoperiodic response, the number of days required to induce 50% summer and winter diapause and the duration of diapausing pupae induced under different photoperiods. Photoperiodic response curves at 18 and 20 degrees C showed that all pupae entered winter diapause at short daylengths (8-11 h), the incidence of diapause dropped to 82.3-85.5% at 22 degrees C without showing a significant difference between short daylengths, whereas the incidence of summer diapause induced by different long daylengths (14-18 h) was varied and was obviously affected by temperature. By transferring from various short daylengths (LD 8:16, LD 9:15, LD 10:14 and LD 11:13) to an intermediate daylength (LD 12.5:11.5) at different times after hatching, the number of cycles required to induce 50% winter diapause (7.28 at LD 8:16, 7.16 at LD 9:15, 7.60 at LD 10:14 and 6.94 at LD 11:13) showed no significant difference, whereas by transferring from various long daylengths (LD 14:10, LD 15:9, LD 16:8 and LD 17:7) to an intermediate daylength (LD 12.5:11.5) at different times, the number of cycles required to induce 50% summer diapause (5.95 at LD 14:10, 8.02 at LD 15:9, 6.80 at LD 16:8, 7.64 at LD 17:7) were significantly different. The intensity of winter diapause induced under different short daylengths (LD 8:16, LD 9:15, LD 10:14 and LD 11:13) was not significantly different with an average diapause duration of 87 days at a constant temperature of 20 degrees C and 92 days at a mean daily temperature of 19.0 degrees C, whereas the intensity of summer diapause induced under different long daylengths (LD 14:10, LD 15:9, LD 16:8 and LD 17:7) was significantly different (the diapause duration ranged from 75 to 86 days at a constant temperature of 20 degrees C and from 76 to 88 days at a mean daily temperature of 19.0 degrees C). All results suggested that photoperiodic control of diapause induction and termination is significantly different between aestivation and hibernation.     

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

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

Photoperiodic sensitivity and diapause induction during ovarian,embryonic and larval development of the flesh fly,Sarcophaga argyrostoma

Sensitivity to the daily photoperiod, particularly with respect to pupal diapause induction, was studied during ovarian, embryonic, and larval development of the flesh flySarcophaga argyrostoma. Large flies were shown to have a greater number of primary follicles in their ovaries and to be capable of limited ovarian maturation in the absence of exogenous protein (autogeny). Such ovarian development occurred independently of photoperiod. However, long days experienced during embryogenesis caused more rapid development, and earlier larviposition, than short days. Short days during embryonic and subsequent larval development also induced pupal diapause, whereas long days led to continuous or non-diapause development of the pupae. Pupal diapause could not be induced by photoperiods during the vitellogenic phase of ovarian development. InSarcophaga argyrostoma, a maternal effect preventing pupal diapause among the progeny of files with a diapause history was not observed.     

Life-History Variation inChrysoperla carnea:Implications for Rearing and Storing a Mexican Population

A population ofChrysoperla carneafrom San Pedro, Coahuila, Mexico, showed phenological responses that differ from those of previously studied North American biotypes. Short daylengths that induce diapause in other biotypes resulted in low incidences of diapause in the San Pedro population. Diapause induction required a decrease in daylength (from long to short days). Constant long days produced variable reproductive responses, whereas an increase in daylength (from short to long day) resulted in high incidences of reproduction. Providing prey to adults that had been reared under short days consistently increased the incidences of reproduction. Storage of either diapausing or nondiapausing adults at low temperature (5°C) for 13 weeks yielded better survivorship and reproductive performance than in unstored (24°C) controls. For efficient production, the San Pedro population ofC. carneamay be reared either with an increase in daylength or with the regular intervention of a storage period. To induce diapause in individuals destined for storage, larvae may be reared under long-day photoperiod and the prepupae transferred to short days.     

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.     

The influence of physiological age of Argas reflexus larvae (Acari: Argasidae) and of temperature and photoperiod on induction and duration of diapause

The occurrence of diapause and quiescence was investigated in Argas reflexus engorged larvae, nymphs I and nymphs II. For diapause experiments, larvae were maintained at five different locations: at constant 20°C long day (LD; 17 h light:7 h dark) or short day (SD; 10 h light:14 h dark), at two locations with natural photoperiod and temperature and at one location with natural photoperiod but constant 15°C. At 20°C, diapause incidence was low in physiologically young larvae, increased with larval age, and then decreased to zero in specimens of increased physiological age. This pattern, observed both at constant LD and SD, suggests that the propensity to diapause changes with the physiological age of the unfed larva. The duration of diapause decreased with increasing larval physiological age at all locations, resulting in a seasonally synchronized moulting pattern. The results suggest that A. reflexus larvae are photoperiodically sensitive both before and after feeding and that decreasing daylengths may be particularly strong inductive stimuli. The developmental zero and thermal constant of the larvae were determined as 13.24°C and 220 degree-days, respectively. Degree-day measurements revealed that larval A. reflexus may enter a diapause of different length when fed between August and December and kept at natural daylength. Development of engorged nymphs I and nymphs II, but not of larvae, was ultimatively restricted at a temperature of 37.5°C, but immediately resumed at 25°C, demonstrating the occurrence of quiescence at high temperatures. Similarly, at a low temperature of 15°C, many nymphs I and II did not develop within 58 months, but did so successfully after transfer to 25°C, without additional food intake. Received: 20 May 1997 / Accepted: 4 August 1997     

A study of the diapausing behaviour of Rhipicephalus appendiculatus and R. zambeziensis under quasi-natural conditions in Zambia

Abstract. The diapausing behaviour of Rhipicephalus appendiculatus was studied under quasi-natural conditions in the Eastern Province of Zambia. Newly-moulted adults of ticks indigenous to the study area entered a behavioural diapause when exposed to daylengths below a critical photoperiod, provisionally a daylength between 11 h 20min and 11 h 45min. In the Eastern Province of Zambia the diapause was apparently not terminated by a long-day signal, but by a weakening of the photoperiodic maintenance of the diapause because of increasing age of the ticks. Adults of a reference stock indigenous to Kenya also entered a diapause when exposed to daylengths below the same threshold and maintained this diapause for the same length of time. Adults of a reference stock of Rhipicephalus zambeziensis indigenous to Zimbabwe did not enter a diapause when exposed to the same daylengths. The relevance of the findings is discussed in relation to the distributions of the two species.     

Diapause affects cuticular hydrocarbon composition and mating behavior of both sexes in Drosophila montana

Envipnmental cues,mainly photoperiod and temperature,are known to control female adult reproductive diapause in several insect species.Diapause enhances female survival during adverse conditions and postpones progeny production to the favorable season.Male diapause(a reversible inability to inseminate receptive females)has been studied much less than female diapause.However,if the males maximized their chances to fertilize females while minimizing their energy expenditure,they would be expected to be in diapause at the same time as females.We investigated Drosophila montana male mating bchavior under short-day conditions that induce diapause in females and found the males to be reproductively inactive.We also found that males reared under long-day conditions(reproducing individuals)court reproducing postdiapause fermales,but not diapausing ones.The diapausing fies of both sexes had more long-chain and less short-chain hydrocarbons on their cuticle than the reproducing ones,which presumably increase their survival under stressful conditions,but at the same time decrease their attractiveness.Our study shows that the mating behavior of females and males is well coordinated during and afier overwintering and it also gives support to the dual role of insect cuticular hydrocarbons in adaptation and mate choice.     

First evidence of pupal summer diapause in Pieris brassicae L.: the evolution of local adaptedness

Pieris brassicae L. is adapted to a wide variety of local climates throughout the Palaeartic by means of geographic variation in daylengths and temperatures triggering winter diapause. We report for the first time a population from Sotogrande in southern Spain (strain S) that additionally exhibits a pronounced summer diapause in the pupal stage as a response to long daylengths. As well as having diapause triggered by long daylengths, this population develops directly at short, ecologically relevant daylengths. This unusual short-day type response is expressed at temperatures of 21 degrees C and above. The absence of cryptic summer diapause in a population from southern France indicates that this summer diapause evolved locally in Spain or Northern Africa in a population of P. brassicae which lacked this trait. The PPRs of strains S, H (Denmark) and B (France) and crosses between them suggest that summer diapause is inherited as a recessive allele which has a clinal decrease of frequency in populations northwards. The population from southern France shows that this simple pattern of inheritance is further modified by an unknown genetic mechanism controlling the expression of summer diapause near the border between aestivating and non-aestivating populations. On the basis of the photoperiodic response and climatic data we predict P. brassicae to have five or more generations each year in southern Spain.     

Photoperiodic and temperature effects on the intensity of larval diapause in Sesamia nonagrioides

Abstract. The intensity of larval diapause in Sesamia nonagrioides Lef (Lepidoptera: Noctuidae) was investigated under laboratory conditions. Newly hatched larvae were exposed to different stationary photoperiods (from LD 7 : 17 h to LD 14 : 10 h), at a constant temperature of 25 °C. Diapause incidence was higher when larvae were exposed to daylengths shorter than the critical value (LD 12 : 12 h), whereas the within‐treatment variation in the larval period appeared to be significantly correlated with the photoperiod applied. The incidences of diapause and the duration of larval development were also measured after exposing larvae to short photoperiods (LD 8 : 16 h, LD 10 : 14 h or LD 12 : 12 h) in combination with various temperatures (20, 22.5 or 25 °C). Although an increase in the incidence of diapause appeared with the lowering of the temperature, no statistical differences were observed in the time needed for pupation within the photoperiodic treatments at the temperatures of 20 and 22.5 °C. Furthermore, when diapausing larvae were transferred to the long photoperiod of LD 16 : 8 h, they immediately proceeded to pupation, regardless of the photoperiod or the temperature to which they had been previously exposed, indicating that there were no differences in the intensity of diapause. Photoperiodic changes from LD 10 : 14 h to LD 12 : 12 h or to LD 14 : 10 h at different larval ages reduced the intensity of diapause with (a) early age of transfer and (b) increase of daylength. By contrast, when larvae were transferred from the long photoperiod of LD 14 : 10 h to shorter, such as LD 10 : 14 h or LD 12 : 12 h, a small increase in the intensity of diapause with the shortening of the daylength was apparent. These results support the hypothesis that insects may compare the duration of the photoperiod and could classify them as either longer or shorter in relation to the critical value.     

Reproductive diapause and quiescence in a tropical rice buh,Leptocorisa oratorius

Experiments were carried out to analyze the effects of food supply and photoperiod on the induction of dormancy inLeptocorisa oratorius Fabricius (Heteroptera: Alydidae) adults in Alor Setar (latitude 6.1^o N), West Malaysia. When the adults were deprived of food, they ceased oviposition, became tolerant to starvation, and their oxygen consumption rate decreased, but they did not accumulate lipid. Their oxygen consumption rate again increased when they were allowed to resume feeding. Therefore, food limitation induced quiescence in the adult. On the other hand, diapause was induced by short daylength (critical photoperiod was 12L:12D), leading to immature ovaries, hypertrophied fat bodies, lipid accumulation and low oxygen consumption rate even when food was available. However, in the study area most of the bugs probably did not enter diapause at any time of the year, because the shortest outdoor daylength including civil twilight was longer than the critical daylength for diapause induction.     

Photoperiodic control of adult diapause in the predacious mite, Amblyseius potentillae: repeated diapause induction and termination

ABSTRACT. Photoperiodic control of facultative reproductive diapause was examined in females of the phytoseiid mite, Amblyseius potentillae (Garman). Full diapause was induced by rearing the mites from egg to adult under short-day photoperiods. Egg-laying females, however, which had experienced a long-day photoperiodic regime during pre-imaginal development, could still be induced to enter diapause when they were transferred to a short-day regime. Diapause development proceeded slowly under a short-day photoperiod, but was accelerated under long days. During diapause development under short days the photoperiodic sensitivity of the females diminished gradually and ultimately disappeared completely. However, after completion of diapause, sensitivity to photo-period reappeared. A second diapause could be induced in post-diapause females under short-day photoperiods and completed again under long-day photoperiods. These results show that A. potentillae remains sensitive to diapause-inducing and diapause-averting daylengths during the adult stage and that a second diapause may be induced after completion of the first one.     

Mode of inheritance of face fly diapause and its correlation with other developmental traits

Abstract.

• 1 Adult face flies, Musca autumnalis De Geer, respond to short photoperiods by entering diapause. Diapause responses to different photoperiods at a mean temperature of 17°C varied among seven laboratory face fly strains. Old strains lost much of their diapause responses.
• 2 To analyse the nature of diapause inheritance, hybrids and their back-crosses were made between two strains that showed the greatest difference in diapause introduction at 10:14 h(L:D), and 17°C.
• 3 Responses of hybrids to the short photoperiod were intermediate to those of their parents. Joint scaling tests showed that diapause did not fit an additive model and the lack of fit was attributed to incomplete dominance. No cytoplasmic effect was detected, but diapause was partially sex linked.
• 4 Diapause heritabilities were 47% and 30% in the broad and narrow senses, respectively.
• 5 Estimates of the minimum number of diapause genes suggested a small number.
• 6 Developmental rates of egg to adult, ovarian follicle growth, and pterin accumulation were inversely correlated with diapause propensities, suggesting pleiotropy or linkage.