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.     

The predatory mite Hypoaspis miles: temperature dependent life table characteristics on a diet of sciarid larvae, Bradysia paupera and B. tritici

Life table characteristics of Hypoaspis miles Berlese (Acarina: Hypoaspidae) fed on a mixture of Bradysia paupera Tuomikoski (Diptera: Sciaridae) and B. tritici Coquillet larvae were investigated in laboratory experiments at 4 temperatures (15, 20, 25, 30 °C) for development time, juvenile mortality, sex ratio, preoviposition period, oviposition period, postoviposition period, age-specific fecundity, and adult longevity. Juvenile development time decreased with increasing temperature from 46 days at 15 °C to 10 days at 30 °C. The lower temperature threshold was 9.9 °C and development required 205 °D. Juvenile mortality decreased from 52% at 15 °C to 3% at 25 °C and then increased to 24% at 30 °C. Preoviposition period varied with temperature from 12 days at 15 °C to 3 days at 25 °C and then increased to about 4 days at 30 °C. Oviposition period decreased with increasing temperature from 58 days at 15 °C to 25 days at 30 °C. The mean number of eggs per female per day increased from 0.4 at 15 °C to 2.3 at 25 °C and decreased to 1.3 at 30 °C. Age-specific fecundity was described by a temperature dependent model from which the maximum daily fecundity rate could be estimated to be attained at 25.6 °C. Female longevity was significantly shorter than for males, and decreased from 90 days at 15 °C to 34 days at 30 °C. Sex ratio was female-biased at all 4 temperatures and increased with temperature up to 25 °C, decreasing at 30 °C. Estimates of net reproductive rate, intrinsic rate of increase, finite rate of increase, mean generation time and doubling time were obtained. The r _m -value increased with temperature from 0.031 day^-1 at 15 °C to 0.133 day^-1 at 25 °C, after which it decreased to 0.112 day^-1 at 30 °C. The study showed that H. miles can develop and reproduce at temperatures between 15 and 30 °C. H. miles and sciarids have approximately the same optimum temperature and thresholds for development and reproduction and H. miles can be used for biological control of sciarids within the temperature range where the pest occurs.     

Distinct effects of different low temperatures on the induction of NAD-sorbitol dehydrogenase activity in diapause eggs of the silkworm,Bombyx mori

Summary Diapause eggs of the silkworm, Bombyx mori, exposed to 5°C and 0.5°C from 2 or 30 days after oviposition, were examined for changes in contents of glycogen, sorbitol and glycerol. Cold acclimation did not alter the profile of accumulation of sorbitol from that in eggs kept continuously at 25°C. However, acclimation at 5°C resulted in conversion of sorbitol to glycogen, while acclimation at 0.5°C was not accompanied by the utilization of sorbitol. NAD-sorbitol dehydrogenase (NAD-SDH; EC 1.1.1.14) activity was examined in the cold-acclimated eggs. The activity was induced by acclimation at 5°C but not at 0.5°C. Incubation at 0.5°C suppressed any further increase in the activity that had been induced. Temperature-directed changes in NAD-SDH activity paralleled those in sorbitol content. Hatching of the diapause eggs was monitored after cold acclimation for various periods of time and subsequent transfer to 25°C. Incubation at 0.5°C was less effective than 5°C at breaking diapause. The time required for the eggs to hatch in synchrony after acclimation at 5°C coincided with that required for the induction of NAD-SDH activity. These results show that different effects result from acclimation at 5°C and near 0°C with respect to the control of NAD-SDH activity, that utilization of sorbitol is controlled by NAD-SDH activity, and that induction of this activity is temperature-dependent. Furthermore, induction of NAD-SDH activity is involved in the termination of diapause in B. mori.Abbreviations DH diapause hormone - NAD nicotinamide-adenine-dinucleotide - NAD-SDH NAD-sorbitol-dehydrogenase     

Entering diapause is a prerequisite for successful cold-acclimation in adult Graphosoma lineatum (Heteroptera: Pentatomidae)

In diapause adults of Graphosoma lineatum overwintering in a field-cage, high chill-tolerance (CT) developed gradually, within 5 months from August to December. In laboratory-acclimation experiments, the diapause state appeared to be an essential pre-condition for successful cold-acclimation and overwintering. First, diapause prevented elevation of the median supercooling point (SCP) by about 5.5°C that accompanies the onset of reproductive activity in non-diapause specimens. Second, diapause allowed subsequent physiological changes resulting in cold-acclimation during a gradual (18-day) decrease of temperature from 25 to 0°C. No, or very modest, cold-acclimation was observed in non-diapause specimens. Decrease of temperature led to a rapid loss of ca. 1/3 of the body water in both non-diapause and diapause specimens. Approximately 0.1 M of trehalose accumulated in tissues of diapause specimens only, and haemolymph osmolality rose from 347 mOsm (at 25°C) to 444 mOsm after an 18-day cold-acclimation and to 764 mOsm during further storage at 0°C for 100 days. Upon transfer of cold-acclimated diapause specimens back to 25°C for one week (de-acclimation), the high CT was lost, the SCP elevated by about 2.5-3°C, and the levels of trehalose, water content and haemolymph osmolality returned to pre-acclimation or non-diapause levels.     

Control of diapause in codling moth larvae

Diapause in a New Zealand strain of codling moth (Cydia pomonella Linnaeus [Lepidoptera: Olethreutidae]) was induced in larvae by photoperiods of 15 h or less. Once diapause had been initiated, it could not be terminated by any combination of conditions tested for at least 20 days after cocooning. In diapausing larvae a low rate of pupation occurred at 25 °C under a long day (18 h) photoperiod. A high rate of pupation was achieved under a long day regime when larvae were decocooned, and provided with apple as nourishment. Diapause could be terminated predictably in 94–100% of larvae by 1) conditioning at 15 °C and constant darkness for periods of 40–100 days, then 2) chilling at 2±2 °C and constant darkness for 20–50 days followed by 3) any post-chill condition periods at 25 °C, 18 h photoperiod. Complete diapause termination was achieved when 100 days conditioning was followed by 30 days or 50 days post-chill period. Under these conditions, 76% termination occurred in the post-chill period after 10 days, and 93% after 25 days.To terminate diapause in codling moth larvae, we recommend that a 100 days conditioning followed by 30 days chilling and 50 days post chilling periods be used.     

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.     

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

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

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.     

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

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

Diapause development and acclimation regulating enzymes associated with glycerol synthesis in the Shonai ecotype of the rice stem borer larva,Chilo suppressalis walker

Overwintering larvae of the Shonai ecotype of the rice stem borer, Chilo suppressalis, enter diapause in early September and terminate diapause at the end of October. Cold acclimation at 0°C did not influence glycerol, trehalose or glycogen content in larvae collected on 22 September. Acclimation at 0°C increased the glycerol content and reduced the glycogen content significantly in larvae collected on 2 October and 22 November compared with acclimation at 15°C. These results indicate that overwintering larvae at different phases of diapause development respond differently to the low temperature stimulus for glycerol synthesis. Thus, we evaluated the metabolic rearrangements associated with glycerol synthesis during diapause development and after temperature acclimation. Larvae collected on 2 October were acclimated at 15°C for 15 and 60 days. Some of those acclimated at 15°C were then moved to 0°C for 15 days. The larvae acclimated at 15°C for 15 days were in deep diapause and accumulated little glycerol, while larvae acclimated at 15°C for 60 days were nearly ready to emerge from diapause and accumulated glycerol at 155.5 μmol/g. When larvae acclimated to 15°C for 15 days were transferred to 0°C, glycerol accumulation was stimulated to the same extent (ca 140 μmol/g) as it was in larvae that were acclimated to 15°C for 60 days and then transferred to 0°C. These results indicate that low temperature has a cumulative effect on glycerol production in larvae at different phases of diapause development. Glycerol accumulation was accomplished by activation of glycogen phosphorylase and inhibition of fructose-1,6-bisphosphatase, and activation of enzymes associated with glycerol synthesis, mainly glyceraldehyde-3-phosphatase and polyol dehydrogenase with glyceraldehyde activity.     

Biology of the shore fly Scatella stagnalis in rockwool under greenhouse conditions

The development times and survival of immature stages in rockwool and the fecundity and longevity of adult Scatella stagnalis were determined and stage-specific life-tables constructed for the species at constant 20 and 25 °C and at a fluctuating temperature (23–34 °C, mean 28.5 °C). Development time from egg to adult decreased with temperature, being 15.9±0.1 days at 20 °C, 11.4±0.1 days at 25 °C and 10.1±0.2 days at fluctuating temperature with mean of 28.5 °C. The lower threshold for egg-to-adult development was 6.4±2.7 °C and the total quantity of thermal energy required to complete development was 212.8±.0 °C. The proportion of females in two populations studied was 0.521. High temperature increased the mortality of pupae from 7% (20 °C) and 10% (25 °C) to 29% at 28.5 °C. At 25 °C, female longevity was 15.5±0.7 days and fecundity 315±19 eggs/female (20.4 eggs/female/day). Males lived for 22.0±1.1 days. At constant 25 °C, the net reproductive rate was 126.1 female eggs/female, generation time was 18.4 days, the doubling time of the population 5.3 days, and the intrinsic rate of increase (r _m) 0.263 day^–1.     

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.     

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.     

Phenological adaptations of a colonizing insect: The southwestern corn borer,Diatraea grandiosella

Summary The developmental rate, critical photoperiod, and diapause intensity were determined for three populations of the southwestern corn borer, Diatraea grandiosella, from Missouri, Mississippi and Kansas. Mississippi larvae grew at the highest rate and Missouri larvae grew at the lowest rate. The zero developmental temperatures (°C) for the Missouri population were estimated from regression lines as follows: 10.5° (eggs), 10.8° (diapausing larvae), 13.3° (non-diapausing larvae) and 11.4° (pupae). The required heat units were: 85° (eggs), 588° (diapausing larvae), 333° (non-diapausing larvae) and 149° days (pupae). However, the observed low temperature limit for larval growth under constant temperature regimes was approximately 17°C.The critical day lengths for diapause induction observed at 25°C were: 15 h 11 min (Missouri); 15 h 20 min (Mississippi); and 15 h 22 min (Kansas). The photoperiodic response of the Mississippi larvae was more or less retained at 30°C, whereas the response of the Missouri larvae was completely suppressed at this temperature. Diapause was most easily terminated in the Kansas larvae. The most intense diapause was observed in the Mississippi larvae.Model seasonal life cycles of the three geographic populations were constructed using photothermograms. Although the models showed good agreement with the field situation for the Missouri and the Kansas populations, some unknown factor(s) remains to account for an extremely long critical photoperiod in the Mississippi population.Contribution from the Missouri Agricultural Experiment Station, as journal series no. 9001     

Combined Effects of Algal (<Emphasis Type="Italic">Chlorella vulgaris</Emphasis>) Food Level and Temperature on the Demography of <Emphasis Type="Italic">Brachionus havanaensis</Emphasis> (Rotifera): a Life Table Study

We evaluated the combined effects of food (0.5 × 10⁶, 1.0 × 10⁶ and 2.0 × 10⁶ cells ml⁻¹ of Chlorella vulgaris) and temperature (15, 20 and 25 °C) on life history variables of B. havanaensis. Regardless of Chlorella density there was a steep fall in the survivorship of B. havanaensis at 25 °C. Both food level and temperature affected the fecundity of B. havanaensis. At any given food level, rotifers cultured at 15 °C showed extended but low offspring production. At 25 °C, offspring production was elevated, the duration of egg laying reduced and the fecundity was higher during the latter part of the reproductive period. The effect of food level was generally additive, at any given temperature, and higher densities of Chlorella resulted in higher offspring production. Average lifespan, life expectancy at birth and generation time were 2–3 times longer at 15 °C than at 25 °C. At 20 °C, these remained at intermediate levels. The shortest generation time (about 4 days) was observed at 25 °C. Gross and net reproductive rates and the rate of population increase (r) increased with increasing temperature and generally, at any given temperature, higher algal food levels contributed to higher values in these variables. The r varied from 0.11 to 0.66. The survival patterns and lower rates of reproduction at 15 °C suggest that the winter temperatures (10–15 °C) prevailing in many waterbodies in Mexico City allow this species to sustain throughout the year under natural conditions.     

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.     

A laboratory study on the life cycle of Sericostoma personatum (Kirby & Spence), and light dark-dependent food consumption

The life cycle of Sericostoma personatum (Spence) was studied at 6 °C, 10 °C and 14 °C and at each temperature at 8 and 14 hrs daylength. Embryogenesis was not temperature dependent in the 12°–18°C range. Only 7 of 38 (app. 18%) had a direct development, the rest remained in diapause with partly developed larvae. Hatching success of single egg masses was over 95%. At 6 °C at both LDs, about 452 days are required for larval development. At 10 °C 370 days (LD 8/16), or 320 days (LD 14/10) and at 14° C 319 days (LD 8/16) and 295 days (LD 14/10) were required. Duration of instars III and IV was longer at 6 °C (both LDs), compared with all other groups. Vth instar larvae of the 14 °C (LD 14/10) group grew fastest. Instar VI larvae of the 10 °C short day group developed faster than all others. Instar VII larvae of both 14 °C groups and of the 10 °C long day group develop faster than the rest. Duration of pupal instar is only temperature dependent, regardless of light regime. The field life cycle of S. personatum may require 2–5 years. Larvae are night active. They feed on Coarse Particular Organic Material (CPOM) on the sediment surface at night. They release faeces (Fine Particular Organic Material, FPOM) into the sediment where they rest by day at a few cm depth. Their burrowing behavior thus contributes to the retention of FPOM in the stream channel. Daily food consumption at constant 10 °C is significantly dependent on night length (r ² = 0.979, p < 0.05). Two factors thus may limit food consumption: in winter, low temperatures, and in summer short nights. The species thus avoids competition by day-active shredders and predation by day-active predators.     

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

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

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.     

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