Altitudinal variation in life cycle syndromes of California populations of the grasshopper,Melanoplus sanguinipes (F.)

Summary Life cycles of California populations of the grasshopper, Melanoplus sanguinipes, varied along an altitudinal gradient. Temperature records indicate a longer season at low altitude on the coast, based on computation of degree days available for development, even though summer air temperatures are cooler than at high altitude; this is a result of warm soil temperatures. At high and low altitudes there was a high proportion of diapause eggs oviposited, while intermediate proportions of diapause eggs occurred at mid altitudes. The low altitude, and especially sea level, populations diapaused at all stages of embryonic development, while at high altitudes most diapause occurred in the late stages just before hatch. Diapause was more intense at high altitudes. One result of diapause differences was delayed hatching in the sea level population. Nymphal development and development of adults to age at first reproduction were both accelerated at high altitude relative to sea level. At lower temperatures (27° C) there was a tendency for short days to accelerate development of sea level nymphs, but not high altitude nymphs. In both sea level and high altitude grasshoppers, short days accelerated maturation of adults to onset of oviposition at warm temperature (33° C); there was little reproduction at 27° C. Population differences for all traits studied appear to be largely genetic with some maternal effects possible. We interpret diapause variation at low and mid altitudes to be responses to environmental uncertainty and variations in development rates to be adaptations to prevailing season lengths.     

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

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

Diapause induction in the codling moth, Cydia pomonella: effect of prediapause temperatures

The effect of four prediapause temperatures (18, 22, 26 and 30°C) on the photoperiodic response of the codling moth, Cydia pomonella (L.), was studied under controlled conditions. The highest rates of diapause were recorded, for all day-lengths, at temperatures of 22 and 26°C while relatively lower rates of diapause were elicited at 18 and 30°C. The same trend was demonstrated by projecting the values of the critical photoperiod which induces 50% diapause (=CPhP₅₀) over the prediapause temperature. The change in diapause incidence as a function of photoperiod, at all prediapause temperatures, exhibited a response characteristic of long-day insects, i.e. high rates of diapause at short days (12–13.5 h) and a decrease in diapause incidence at long days (14–15 h). The results for temperatures 22, 26 and 30°C support the view that lower prediapause temperatures enhance diapause induction, at a give photoperiod, while higher temperatures tend to avert or diminish the process. On the other hand, the low rates of diapause obtained at 18°C contradict this view. Nevertheless, high correlation was found between the laboratory evidence and field data, indicating the adaptability of the Israeli codling moth to subtropical climate.     

Geographic variation in embryonic development time and stage of diapause in a grasshopper

Embryonic development times and the stage at which embryonic diapause occurs varied dramatically among 23 populations of the Melanoplus sanguinipes/ devastator species complex in California, USA. Grasshoppers were collected from a wide range of latitudes (32°57N to 41°20N) and altitudes (10m to 3031 m), spanning much of the variation in climatic conditions experienced by these insects in California. When reared in a common garden in the laboratory, total embryonic development times were positively correlated to the mean annual temperature of the habitat from which the grasshoppers were collected (varying from about 19 days to 32 days when reared at 27°C). These grasshoppers overwinter as diapausing eggs and the proportion of embryonic development completed prior to diapause was significantly higher in populations collected from cool habitats (>70%) than in populations collected from warm environments (<26%). The length of pre-diapause development time is determined by the stage of embryonic development at which diapause occurs, and varies considerably among populations of these grasshoppers; grasshoppers from warmer environments tend to diapause at very early stages of embryogenesis, while grasshoppers from cooler environments diapause at very late stages. The combined effect of variation in embryonic development times and variation in the stage at which diapause occurs results in a dramatic reduction in the time needed to hatch in the spring; populations from warm environments required up to 20 days (at 27°C) to hatch while populations from cool environments required as few as 5 days to complete embryonic development prior to hatching. Egg size also varied significantly among populations, but tended to be larger in populations with shorter embryonic development times. Significant family effects were observed for development time and stage of diapause, suggesting significant heritabilities for these traits, although maternal effects may also contribute to family level variation. We interpret these findings to support the hypothesis that embryonic development time and the stage of embryonic diapause have evolved as adaptations to prevailing season lengths in the study populations.     

Spatial distribution of resting stages,rate of emergence from diapause and times to adulthood and to the appearance of the first clutch in 3 species of cyclopoid copepods

The spatial distribution of dormant copepodids of 3 species of cyclopoid copepods — Cyclops vicinus, Mesocyclops leuckarti and Thermocyclops crassus — was studied in 4 small lakes in South Germany. The rate of emergence from diapause and times from the resting stage to adulthood and from adulthood to the appearance of the first clutch was studied at 4 constant temperatures (5, 10, 15, 20 °C) in the laboratory. Resting stages of C. vicinus were always concentrated in the deepest parts of the lakes and were found relatively deep in the mud. M. leuckarti- and T. crassus-copepodids preferred shallow areas in deep lakes but were concentrated in the deep areas in shallow lakes. Copepodids of both species were always concentrated in the uppermost mud layers.Rate of emergence from diapause was strongly temperature-dependent. At high temperatures (20 °C) copepodids of all species under study emerged within 2 weeks. At lower temperatures C. vicinus copepodids showed the highest rate of emergence. At 5 to 10 °C only few M. leuckarti- and T. crassus-copepodids had emerged after the investigation period (7 weeks). Both C. vicinus and T. crassus showed the highest rate of emergence at the natural end of diapause but even at that time only few T. crassus-copepodids emerged at 5 °C. Times to adulthood at 5 °C were shortest in C. vicinus. At higher temperatures this species was passed by M. leuckarti. Times from adulthood to the appearance of the first clutch at 5–15 °C were shortest in C. vicinus. T. crassus produced no clutch at 5 and 10 °C.     

Diapause in the egg parasitoid Trichogramma cordubensis: role of temperature

The role of temperature in the induction of diapause in Trichogramma cordubensis (Hymenoptera: Trichogrammatidae), under controlled laboratory conditions, was investigated using Ephestia kuehniella Zeller (Lepidoptera: Pyralidae) eggs as hosts. Results indicate that prestorage temperatures and the duration of exposure of the parasitoids to these temperatures affected the induction of diapause. It was possible to induce diapause in prepupae of T. cordubensis by exposing the preimaginal stages (prior to the prepupal stage) to 10°C for at least 30 days, but adults emerged without diapause when the duration of exposure was of only 10 or 20 days. Parasitoids failed to enter diapause when prestorage temperatures were 7 or 12°C, regardless of the duration of exposure. However, at these two temperatures, preimaginal development of T. cordubensis was delayed, allowing short-term storage (40 days at induction temperatures followed by 30 days at 3°C) by keeping parasitoids in quiescence without reducing the percentages of adult emergence. Good percentages of adult emergence after long-term low-temperature storage (30 or 40 days at 10°C followed by six months at 3°C) occurred only when T. cordubensis was in diapause. The long-term storage of parasitoids in diapause allows an enlargement in the mass rearing potentialities of this species for future biological control releases by allowing producers to stockpile the parasitoids for release in the field season.     

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

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

Cell cycles in embryos of the silkworm,Bombyx mori: G2-arrest at diapause stage

Summary In the silkworm, Bombyx mori, diapause occurs at a specific embryonic stage, i.e. after formation of the germ band with cephalic lobes and telson and sequential mesoderm segmentation. As long as the eggs are incubated at 25° C, cell divisions and morphological development of the embryos cease. To examine changes in percentage of embryonic cells in the G₁, S and G₂ phases during embryogenesis, nuclear fractions were isolated from embryos, stained with propidium iodide and then subjected to flow cytometric analysis. The percentages of embryonic cells in G₁, S and G₂ were 10, 35 and 55%, respectively, at the stage of formation of cephalic lobes, whilst 98% of cells were in G₂ at diapause stage. After termination of diapause by acclimation at 5° C or by a combination of chilling and HCl, cell division resumed in the embryos. During this period, the cells rapidly entered S phase through G₁ from G₂, suggesting that their G₁ phase was short. In eggs in which diapause was averted by HCl-treatment after incubation at 25° C for 20 h after oviposition, embryonic development proceeded continuously for 9.5 days at 25° C until hatching. Along with this development, the G₁ fraction increased to levels of about 90%. These results indicate that embryonic cells are arrested in G₂ at diapause and suggest that, concomitant with further embryonic development, cell cycles become slower in proportion to an increasing length of G₁. Finally, most of the cells may be arrested in G₁, while there is only a small fraction of cells continuously cycling. Offprint requests to: T. Yaginuma     

Effect of hydration,photoperiod and temperature on diapause termination in eggs ofPetrobia (Tetranychina) harti (Acari: Tetranychidae)

Petrobia harti (Ewing) displays a facultative summer diapause in the egg stage. An adult female will lay only either diapause or non-diapause eggs throughout her life. In the laboratory, diapause eggs are laid by females which develop on detachedOxalis articulata leaves under long-day photoperiods and a relatively low temperature of 19±1°C.Diapause occurs in a stage of advanced embryonic development, in which the embryo appears U-shaped when observed from the egg's ventral side. Embryonic development ceased at this stage, and no further growth occurred when the eggs were kept under a relative humidity of about 70% in various photoperiod and temperature conditions. However, when the eggs were hydrated by placing them on wet cotton wool, development in some embryos (apparently in those which had completed their diapause development) proceeded beyond the U-stage at a rate similar to that in non-diapause embryos and the eggs hatched.Under LD 168 and 19±1°C or 26±1°C, the later from oviposition the period of egg hydration started, the higher the percentage of diapause termination. Under LD 168 and 26±1°C, diapause termination occurred mostly during the first week of hydration, while at 19±1°C mostly during the second and third week.At 26±1°C, in eggs hydrated 15 days but not 30 days from oviposition, the percentage of diapause termination was higher under a long-day than under a short-day photoperiod.Under LD 168, when the eggs were hydrated continuously from oviposition or starting 15, 30 and 45 days from it, the percentage of diapause termination was higher at 26±1°C than at 19±1°C.The percentage of diapause-laying adult females and the intensity of egg diapause were higher when the pre-imaginal mites grew at LD 1212 and 19±1°C, than when they grew at LD 168 and 26±1°C. This maternal effect on egg diapause intensity was expressed when the eggs were maintained at LD 1212 and 19±1°C but not at LD 168 and 26±1°C.     

An analysis of diapause and resistance in the egg stage of Floronia bucculenta (Araneida: Linyphiidae)

Summary Floronia bucculenta hibernates in the egg stage; the egg sacs are deposited on the leaves of grass tussocks without any shelter. The morphogenesis of the eggs was divided into 10 arbitrarily chosen stages, in order to test the dependence of embryonic development on temperature in the laboratory. The eggs developed slowly at 23°C, 16°, 12.5°; embryogenesis stopped after 70–45 days, when prosomal appendage rudiments began to form. At 10°, 7.5°, 5°, 0° complete embryogenesis was possible until the emergence of the first complete stage. The eggs developed most rapidly at 5° (mean developmental time 203 days). The egg development was normal at 5° and 0°, when compared with the timetable of the embryogenesis of the linyphiid Bathyphantes gracilis, a species which has no egg diapause. At 7.5° and 10° the embryogenesis was strongly delayed during the median phases of development (elongation of the germ band, formation of prosomal appendages); after reversion the development was accelerated (postdiapause phase). After long exposure to low temperatures (-10° to +10°) the diapause was terminated. A temperature of 0° was optimal (minimal time of exposure 8–9 weeks). The time required for embryonic development of postdiapause eggs decreased hyperbolically with increasing temperature. In the field the median phases of embryogenesis were retarded by low ambient temperatures; diapause was terminated from late December to mid-January. The spread of hatching in spring was 7–15 days.During the diapause phase the O₂-consumption of the eggs at 25° was depressed. It rose from 1.55 (in late diapause) to 4.21 ml/100 eggs·h at the onset of postdiapause, whereas O₂-utilization did not change significantly at 5° (from 0.54 to 0.61 ml/100 eggs·h just after the termination of diapause).The diapause phase was not characterized by higher resistance to cold, drought, or flooding. As compared with single eggs removed from the cocoon, the silken wall of the intact egg sac did not affect the survival of postdiapause eggs exposed to-15° (LD₅₀=28 days); it raised, however, the survival time of eggs exposed to a R.H. of 32% (at 5°) or flooding by distilled water (at 5°): from LD₅₀=37 to 68 days at drought, from LD₅₀=30 to 92 days at flooding.Diapause is important for synchronizing the life-cycle of F. bucculenta with the seasonal fluctuations of environment. The egg stage is highly tolerant to the extreme factors of the winter. Some implications of the relation of the studied spider to its habitat are discussed.     

Influences of daylength and temperature on the period of diapause and its ending process in dormant larvae of burnet moths (Lepidoptera,Zygaenidae)

The onset of larval diapause in the burnet moth Zygaena trifolii is clearly characterized by the larva molting into a specialized dormant morph. In a potentially bivoltine Mediterranean population (Marseille) two types of diapause can occur within 1 year: firstly, a facultative summer diapause of 3–10 weeks, and secondly, an obligate winter diapause, which can be lengthened by a period of thermal quiescence to several months in temperatures of 5°C. For the first time, three successive physiological periods have been experimentally distinguished within an insect dormancy (between onset of diapause and molting to the next non-diapause stage), using chilling periods of 30–180 days at 5°C, and varying conditions of photoperiod and temperature. These stages are: (1) a continuous Diapause-ending process (DEP); (2) thermal quiescence (Q); and finally, (3) a period of postdiapause development (PDD) before molting to the next larval instar. The result of transferring dormant larvae from chilling at 5°C to 20°C depended on the length of the chilling period. After chilling for 120–180 days, molting to the next instar occurred after 6–10 days, independent of daylength. This period corresponds with the duration of PDD. After shorter chilling periods (90, 60, 30 days and the control, 0 days) the period to eclosion increased exponentially, and included both the latter part of the previous diapause process and the 6–10 day period of PDD. However, photoperiod also influences the time to eclosion after chilling. Short daylength (8 h light / 16 h dark: LD 8/16) lengthened the diapause in comparison to long daylength (16 h light / 8 h dark: LD 16/8). Short daylength had a similar effect during chilling at 5°C, as measured by the longer time to eclosion after transfer. The shorter time to eclosion resulting from longer chilling periods (30–90 days) demonstrates that the state of diapause is continuously shortened at 5°C, and corresponds to the neuroendocrine controlled DEP. Presumably the DEP has already started after the onset of diapause. When chilling was continued after the end of the DEP, which ranged between 90 and 120 days, thermal quiescence (Q) followed (observed maximum 395 days). Different photoperiodic conditions during the pre-diapause inductive period modified diapause intensity (measured as the duration of diapause), in that a photoperiodic signal just below the critical photoperiod for diapause induction (LD 15/9) intensified diapause. Experiments simulating the summer diapause showed that PDD occurred in the range of 10–25°C. Higher temperatures (15 and 20°C) shortened the DEP at LD 16/8, so that at 20°C many individuals had already terminated diapause after 10–40 days and had molted after the 6–10 days of PDD. A temperature of 25°C unexpectedly lengthened the DEP to 110 days in several individuals. The ecological consequences and the adaptive significance of variation in the duration of the diapause are discussed in relation to the persistence of local populations predictably variable and rare climatic extremes throughout the year.     

Life history study of Thrips setosus

Development, reproduction and population growth of Thrips setosus Moulton (Thysanoptera, Thripidae), reared on a leaf of kidney bean, was studied under six different constant temperatures, and the effect on reproduction of short photoperiod during immature stages was examined. Survival rates from hatch to adult were more than 67.5% at temperatures between 17.5 and 27.5 °C, but less than 55% at 30 °C. Developmental rates increased linearly as rearing temperature increased. A total of 181.1 degree-days, above a developmental zero of 12.5 °C, were required to complete development from egg to adult oviposition. These data were related to records of field temperatures in Kurashiki in western Japan, and an estimate produced that, under outdoor conditions, a maximum of between seven and 12 generations could have developed annually between 1990 and 1999. There were no significant differences in mean adult longevity and mean fecundity among three temperatures (20, 22.5 and 25 °C). The intrinsic rate of natural increase (r _m) was 0.1997 at 25 °C. Reproductive diapause was induced by a photoperiod less than 12 h at 20 °C.     

Diapause and polyphenism of life-history of Lagria hirta

This paper is focused on diapause and polyphenism of development of Lagria hirta L. and tries to unravel its mechanism of life-history adaptation. Lagria hirta, distributed widely in Europe, has a strictly univoltine life cycle. The results showed that larval diapause and moulting polymorphism were the deciding factors that made L. hirta maintain its univoltinism and keep a flexible relation between seasonal changes and life-history phases. In the laboratory, larvae of this species were not able to pupate if kept at constant temperatures of 5 °, 10 °, 15 °, 20 °, 25 ° or 30 °C combined with a photoperiod of either LD (L16:D8) or SD (L8:L16). Pupation only occurred if larvae were reared at 15 °–25 °C when intervened by a three-month chilling at 5 °C in stages L₃, L₄, L₅ or L₆. A chilling treatment was shown to be obligatory for the termination of its larval diapause and had an accelerating and synchronizing effect on larval development. Larval diapause of L. hirta was characterized by no pupation and more moulting in advanced instars, longer duration of each single stage, and moulting desynchronization. Larval development was found to be variable with respect to the total number of instars: most larvae underwent a total of seven or eight moults; some larvae might even moult once or twice more, but they seldom pupated. It seemed that the choice for the 7-instar or the 8-instar development did not directly relate to any of the external conditions, such as temperature, photoperiod, and stages with chilling treatment. This polyphenism was observed in the same group under identical conditions and even in a single egg clutch. In L. hirta, overwintering in different stages of L₃–L₆, and choosing the 7- or 8-instar pathway of development are two features that increase the plasticity and flexibility in coordinating its life cycle with seasonal change, that varies unpredictably from year to year.     

The response of development rate to temperature in the univoltine cranefly,Tipula subnodicornis Zetterstedt

Summary The cranefly, Tipula subnodicornis, emerges as an adult in the spring and has an annual life-cycle in the British Isles. This is maintained partly through the presence of a winter diapause but the response of development rate to temperature also acts to preserve the timing of the cycle. During development under constant temperature conditions in the laboratory the optimum temperature (taken as the temperature which promoted the most rapid development) dropped from 25°C, or above, in the egg stage to below 20°C in the late larval stages. It is suggested that at the warmer, southern limits of the geographical range rapid early development may be compensated by a retardation in late larval growth. In addition, the response of growth rate to change in temperature was small in the fourth, final, instar and resulted in low Q ₁₀ values; 2.4 between 7° and 10°C, 1.5 between 10° and 15°C and 0.9 between 15° and 20°C. As the fourth instar comprises the greater part of the growth period, this has the effect of minimising the effect of temperature differences which are the result of differences of latitude or altitude. Even at optimum temperatures the growth period was prolonged and larvae in the field do not reach maximum weight, and the photosensitive stage, until late autum when short daylength promotes diapause. Subsequent development in the spring, before pupation and during the pupal period, showed a reversion to the higher Q ₁₀ figures of the early stages in development.The development of final instar Tipula subnodicornis larvae is contrasted with that of Tipula melanoceros. Tipula melanoceros emerges as an adult in September and it is likely that it has an egg diapause. Consequently larval development is confined to a short period between April and late July and growth must be rapid during this period. Under constant temperature conditions in the laboratory the growth of final instar larvae showed a marked contrast to that of Tipula subnodicornis in that the response to temperature was large and remained positive over a wider temperature range.     

Diapause termination, post-diapause development and reproduction in the beet webworm, Loxostege sticticalis (Lepidoptera: Pyralidae)

Effects of photoperiod and cold exposure on diapause termination, post-diapause development and reproduction in Loxostege sticticalis were examined. Larvae were reared at diapause inducing condition (22 °C, L:D 12:12) consistently or transferred to long day photoperiod (L:D 16:8) and darkness (L:D 0:24) respectively, after entering into diapause. Diapause was terminated in approximately 40% of the larvae after 36 days, and no significant differences were observed between photoperiods, suggesting larval diapause was terminated spontaneously without being induced by photoperiods. Cold exposure significantly hastened diapause termination. The diapause termination incidence increased significantly with peaks of 98% at both 5 °C and 0 °C exposure for 30 days, as compared to 42% in controls not exposed to cold, while the mortality and number of days required for diapause termination decreased dramatically. The optimal low temperature exposure periods under 5 °C or 0 °C were 20 days and 30 days, showing a higher termination incidence and shorter time for diapause termination. This suggests that the low temperatures in winter play an important role in diapause termination under natural conditions. The threshold temperatures for post-diapause development in prepupae and pupae were 9.13 °C and 10.60 °C respectively, with corresponding accumulations of 125 and 200 degree-days. Adults that experienced larval diapause significantly delayed their first oviposition, oviposition period was prolonged, and the lifetime number of eggs laid was decreased, however both males and females have significantly longer longevity. The field validation of diapause termination, the degree-days model, and the relationship between diapause and migration in L. sticticalis were also discussed.     

Temperature dependent sorbitol utilization in diapause eggs of the silkworm,Bombyx mori

Summary A series of experiments was conducted to determine whether the interconversion of glycogen and sorbitol at the initiation and termination of diapause in eggs of the silkworm,Bombyx mori was influenced by low temperatures (5°C and 1°C). The conversion of glycogen to sorbitol and glycerol at the initiation of diapause was not affected by exposure to 5°C and 1°C. Chilling diapause eggs at 5°C for over 70 days resulted in a progressive conversion of sorbitol to glycogen. Transfer to 1°C after chilling to 5°C led to a decrease in sorbitol and glycerol conversion. While continuous chilling at 1°C completely inhibited sorbitol utilization for at least 400 days, sorbitol began to decrease immediately following transfer to 5°C. In contrast, glycerol utilization was not prevented by acclimation to 1°C, but a delayed decrease occurred. Continuous exposure to 1°C delayed hatching for a period of 440 days, whereas acclimation to 5°C resulted in 100 per cent hatching by about 100 days. A low, or high acclimation temperature 1°C or 5°C produce different effects on sorbitol utilization and termination of diapause in silkworm eggs.     

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

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

Diapause termination and thermal requirements for postdiapause development inAphelinus mali at constant and fluctuating temperatures

Diapause termination under natural and simulated overwintering conditions, the effect of subzero temperature on postdiapause development and the relationship between postdiapause development rate and constant and fluctuating temperatures was studied in a Dutch population ofAphelinus mali Hald. (Hymenoptera: Aphelinidae).The rate of diapause termination was similar in larvae overwintering under natural and simulated conditions. Most larvae had terminated diapause by the last week of February. Some female larvae may have remained in diapause until the end of March. The exposure of postdiapause larvae to –10°C for two weeks did not affect their survival or postdiapause development rate.PostdiapauseA. mali larvae could complete development and the adults emerge from their mummified aphid hosts at constant temperatures from 12 to 24°C. Although some larvae completed postdiapause development at 10°C, few emerged. The theoretical threshold temperature (t_o) for postdiapause development was 9.4°C and the thermal constant (K) 136.4 degree-days. K was 121.4 and 134.8 for first and 50% emergence, respectively.The number of heat units accumulating above 9.4°C to 1st and 50% emergence was similar under constant and fluctuating temperatures.

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Fin de la diapause et exigences thermiques pour le développement après la diapause d'Aphelinus mali soumis à des températures constantes ou à des thermopériodes

Résumé L'achèvement de la diapause en conditions naturelles ou simulant l'hiver, les effets des températures inférieures à zéro sur le développement après la diapause et les relations entre la vitesse de développement après la diapause et les températures constantes ou en thermopériodes ont été examinés sur des populations néerlandaises d'A. mali (Hymenop.; Aphélinidae).Les taux d'achèvement de la diapause de larves hivernantes étaient semblables en conditions naturelles ou simulées. La plupart des larves ont terminé leur diapause la dernière semaine de février. Quelques larves femelles sont restées en diapause jusqu'à fin mars. L'exposition pendant 2 semaines des larves sorties de diapause à –10 °C ne compromet pas leur survie ou leur taux de développement après la diapause.Les larves ayant diapause peuvent terminer leur développement et les adultes émerger des pucerons momifiés aux températures constantes comprises entre 12 et 24 °C. Bien que quelques larves achèvent leur développement à 10 °C, peu émergent. La température seuil théorique de développement après la diapause (t_o) a été de 9,4 °C et la constante thermique (K), 136,5 degrés-jours. Pour la première émergence et pour 50% d'émergences, les valeurs de K étaient respectivement: 121,4 et 134,8.Le nombre d'unités thermiques pour la première émergence et pour 50% d'émergences était le même à température constante ou avec une thermopériode.

     

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

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

Diapause in a tropical species,Cothonaspis boulardi (Parasitic hymenoptera)

Summary The strain of Cothonaspis boulardi originating from a field collection at Guadeloupe (16° lat. N-West Indies) exhibits a facultative diapause that occurs at the final larval instar. The diapause was induced by low temperature (17.5° C) and was inhibited at 25° C. Diapause was independent of photoperiod. The termination of diapause was hastened in about 10 days by exposing larvae to a temperature of 25° C. Although a succession of sufficient cold days for diapause occurs only rarely in the collection area (Petit-Bourg), at higher elevations within 20 km of Petit-Bourg conditions that could induce diapause occur annually.This study is supported by research grant of C.N.R.S. (ATP Ecophysiologie, n^o 51-3571)