Thermal response and reversibility of diapause in the eggs of Locusta migratoria

Abstract. Eggs of a local population of Locusta migratoria L. (Orthoptera: Acrididae) collected near Hirosaki (40.5^oN) entered diapause when incubated at temperatures between 20 and 35^oC. For diapause development the optimum temperature was 10^oC. The lower thermal threshold for post-diapause development was 14.7^oC. After chilling at 10^oC for 20 days, the rate of hatching varied with incubation temperature, being 0, 61% and 81% at 20, 25 and 30^oC, respectively. After chilling for 40 days or more, however, almost all eggs hatched at 20–30^oC. Diapause with a reduced intensity seemed to be eliminated easily by a high temperature of 25 or 30^oC.
When eggs chilled at 10^oC for 20 days were kept at 20^oC for 7 days or more before incubation at 25^oC, almost all eggs maintained diapause. Most eggs chilled at 10^oC for three 10 day periods separated by 3 days of warming at 25^oC failed to terminate diapause. Daily alternations of 10^oC (18 h) and 25^oC (6h) decreased the diapause-terminating effect of chilling. These facts suggest that diapause intensity can be restored if eggs chilled insufficiently are exposed to a moderately high temperature. This reversible change in diapause intensity would play an important role in maintaining diapause before winter.     

Diapause induction and clock mechanism in the pine caterpillar Dendrolimus tabulaeformis (Lep., Lasiocampidae)

Abstract:  Dendrolimus tabulaeformis overwinters as third to fourth instar larvae at short days in autumn. Using 24-h light–dark cycles, the photoperiodic response curves were similar at 24 and 28°C. The critical night length was 9 h 20 min at 24°C and 9 h 50 min at 28°C. Under non-24 h light–dark cycles, duration of scotophase proved crucial in the determination of diapause. In night interruption experiments using 24-h light–dark cycle, a 1-h light pulse falling 8 h in the darkness strongly averted diapause in comparison with other light pulses. Nanda–Hamner experiments showed two weak troughs of diapause inhibition, suggesting the possible involvement of the circadian system. However, Bünsow experiments did not support the evidence of the involvement of circadian oscillatory system in photoperiodic time measurement. These results suggest that photoperiodic time measurement in this moth shows a non-oscillatory 'hourglass-like' response model or a rapidly damping oscillator model.     

A mutant strain of Chymomyza costata (Diptera: Drosophilidae) insensitive to diapause-inducing action of photoperiod

ABSTRACT. From a Japanese population of Chymomyza costata which has been known to have a photoperiodic larval diapause, we selected a mutant strain which did not respond to photoperiod. However, about 70% of the individuals of this strain entered diapause at 11^oC irrespective of photoperiod, and about the same percentage of those of the photoperiod-sensitive strain also did so in continuous illumination at 11^oC. This indicates that low temperature induces diapause independently of photoperiod. On the other hand, a temperature drop from 18 or 25^oC to 15^oC and chilling at 4^oC did not induce diapause.     

Effects of photoperiod and low temperature on diapause termination in the yellow-spotted longicorn beetle (Psacothea hilaris)

Abstract. .The effects of photoperiod and low temperature on diapause termination in the yellow-spotted longicorn beetle, Psacothea hilaris (Pascoe) (Coleoptera: Cerambycidae), were examined using a population from Ino, Japan. Diapausing insects obtained by rearing larvae under short daylength (12 or 13 h) at 25^oC were subjected to various treatments. When the photoperiod was changed at the same temperature, diapausing larvae showed a long-day response with a critical daylength between 13.5 and 14h. The diapause was terminated and consequently pupation occurred if the daylength was longer than 13.5 h. Chilling the diapausing larvae at 10^oC for 30 or more days also terminated diapause in most larvae irrespective of the photoperiods during and after chilling treatment. In contrast, the post-chilling photoperiod had a critical effect on development of diapausing larvae chilled for only 15 days.     

Influence of thermoperiod and photoperiod on reproductive diapause in Bruchidius atrolineatus (Pic) (Coleoptera, Bruchidae)

Abstract Bruchidius atrolineatus (Pic) is a common pest infesting cowpea seeds ( Vigna unguiculata Walp) in Africa. In the Sahelian zone, the adults are in reproductive diapause during the dry season. Diapause induction depends on the climatic conditions during embryonic and post-embryonic development. A strain of B. atrolineatus originating from the Niamey (13^oN) was reared in different thermoperiodic and photoperiodic conditions. In the thermoperiodic regime 40:25^oC, in continuous darkness, induction of diapause was dependent on the duration of the thermophase. Photoperiod could also influence diapause induction but the response depended on the temperature. When the beetles were reared in conditions of LD 12:12 h, 40:25^oC, for seven generations, the proportion of diapausing adults decreased and their sensitivity to photoperiod increased. In LD 14:10 h, 40:25^CC, the proportion of diapausing adults remained high regardless of the photoperiod. In these conditions, the first emerging adults were sexually active and were used for selection of a strain with a low incidence of diapause. After twelve generations, the proportion of diapausing beetles was lower than 5%. The responses of this selected strain to photo- and thermoperiods were reduced. The sensitivity of B. atrolineatus to photoperiod and thermoperiod could be controlled by genetic systems as observed in other insect species.     

Calling behaviour in M.convecta females under different temperature and photoperiodic conditions

Abstract. . The effects of temperature and photoperiod on calling behaviour in females of the Australian common armyworm, Mythimna corwecta (Walker) (Lepidoptera: Noctuidae), were studied in the laboratory. Age at first calling was greater at 15 and 20^oC compared with 25^oC, but there were no significant differences between 25 and 30^oC. Females kept at 10^oC did not call, but if transferred at 10 days to 20^oC they called after a period similar to the pre-calling period at constant 20^oC. Onset time of calling was earlier at 15 and 20^oC compared with 25 and 30^oC, but there were no significant differences in calling duration. At 20^oC, age at first calling was increased by shorter day length (12h and 14h, compared with 16h), and there was an interaction between daylength and moth age affecting the duration of calling. Onset times advanced with longer daylength, but peak calling was later in relation to the lengtii of the scotophase. These results are discussed in connection with migration in M.convecta. Evidence for a circadian rhythm of calling was found in females entrained for 3 and 8 days after emergence under reverse-cycle conditions and then transferred to constant darkness. However, after 56 h and 80 h respectively of darkness, calling became de-synchronized. Subsequently, it appeared to re-synchronize to a different clock, which approximately correlated with the normal photoperiods the moths had experienced during larval development.     

Circadian Timekeeping for the Photoperiodic Control of Budset in Picea abies (Norway Spruce) Seedlings

Romanian populations of Norway spruce are induced to set terminal buds by four inductive cycles of 8 h light/16 h darkness. To distinguish between circadian and hourglass timekeeping for the photoperiodic control of budset, seedlings were raised in continuous light at 300 µmol m^-2s^-1 at 20°C for 10 weeks. They were then exposed to an extended night regime consisting of three cycles of 8 h light/40 h dark with 4-h or 1-h nightbreaks (120 µmol m^-2s^-1) applied to groups of plants at intervals during the extended night. Following a final cycle of 8 h light/16 h dark to maximize budset, the plants were transferred to continuous light. Budset was delayed when the night-break was applied close to the critical nightlength (CNL) of 6-7 h or about 22-23 h later in the extended night, consistent with circadian rather than hourglass timekeeping. Confidence intervals were calculated for the times to maximum effect of the night-breaks.     

Effect of temperature on photoperiodic response in a selected 'non-diapause' strain of Pyrrhocoris apterus (Heteroptera)

Abstract. The artificially selected 'non-diapause' strain of Pyrrhocoris apterus (L.) (Heteroptera) showed no diapause response to photoperiod at 26°C (Socha & Hodkova, 1994). However, the diapause response to short-day photoperiod (LD 12:12 h) became apparent at lower temperatures of 17°C (70% diapause) or 20°C (41% diapause). Diapause was induced in 60% females by short-day photoperiod combined with thermoperiod of 26/16°C, whereas only 20% diapause was induced by the same thermoperiod under continuous darkness. Thus the time-measuring system was not removed by artificial selection but the diapause response was shifted to lower temperatures. The diapause response to short days seems to be favoured rather by low temperature during scotophase than by low temperature throughout the whole light/dark cycle. If the percentage of diapause at 26°C is compared in F₁ hybrids and in wild and selected parental strains the diapause appears to be dominant at LD 13:11 h but recessive at LD 11:13 h and LD 10:14 h. A hypothesis is proposed that the inheritance of the percentage of diapause in F1 hybrids is determined by interactions of genes controlling the temperature dependence of photoperiodic response.     

Circadian Timekeeping for the Photoperiodic Control of Budset in Picea abies (Norway Spruce) Seedlings

Romanian populations of Norway spruce are induced to set terminal buds by four inductive cycles of 8 h light/16 h darkness. To distinguish between circadian and hourglass timekeeping for the photoperiodic control of budset, seedlings were raised in continuous light at 300 µmol m^-2s^-1 at 20°C for 10 weeks. They were then exposed to an extended night regime consisting of three cycles of 8 h light/40 h dark with 4-h or 1-h nightbreaks (120 µmol m^-2s^-1) applied to groups of plants at intervals during the extended night. Following a final cycle of 8 h light/16 h dark to maximize budset, the plants were transferred to continuous light. Budset was delayed when the night-break was applied close to the critical nightlength (CNL) of 6–7 h or about 22–23 h later in the extended night, consistent with circadian rather than hourglass timekeeping. Confidence intervals were calculated for the times to maximum effect of the night-breaks.     

Experimental evidence for a non-clock role of the circadian system in spider mite photoperiodism

In the spider mite Tetranychus urticae photoperiodic time measurement proceeds accurately in orange-red light of 580 nm and above in light/dark cycles with a period length of 20 h but not in 'natural' cycles with a period length of 24 h. To explain these results it is hypothesized that the photoperiodic clock in the spider mite is sensitive to orange-red light, but the Nanda-Hamner rhythm (a circadian rhythm with a free-running period tau of 20 h involved in the photoperiodic response) is not and consequently free runs in orange-red light. To test this hypothesis a zeitgeber was sought that could entrain the Nanda-Hamner rhythm to a 24-h cycle without inducing diapause itself, in order to manipulate the rhythm independently from the orange-red sensitive photoperiodic clock. A suitable zeitgeber was found to be a thermoperiod with a 12-h warm phase and a 12-h cold phase. Combining the thermoperiod with the long-night orange-red light/dark regime, both with a cycle length of 24 h, resulted in a high diapause incidence, although neither regime was capable of inducing diapause on its own. The conclusion is that the Nanda-Hamner rhythm is necessary for the realization of the photoperiodic response, but is not part of the photoperiodic clock, because photoperiodic time measurement takes place in orange-red light whereas the rhythm is not able to 'see' the orange-red light. It is speculated that the Nanda-Hamner rhythm is involved in the timely synthesis of a substrate for the photoperiodic clock in the spider mite.     

Effect of temperature and photoperiod on the calling behaviour of a migratory insect, the oriental armyworm Mythimna separata

Abstract The pre-calling period (PCP), between emergence and first release of pheromone, determines the number of nights over which female Mythimna separata (Walker) (Lepidoptera: Noctuidae) can express their migratory potential. In its seasonal migrations in eastern Asia, this armyworm penetrates to latitudes at which it is unable to over-winter but at which reliable photoperiodic and temperature cues anticipate the deterioration of conditions. The effect of these environmental factors on PCP was examined in moths descended from samples collected in Nanjing (32^oN) during the summer migration. Under LD 14:10h, a reduction in temperature, from 23^oC during larval development to 18^oC from 24 h after pupation, resulted in a twofold increase in PCP compared with insects held at 23^oC. At 23^oC, short days (LD 12:12 h) extended PCP but the magnitude of the response was increased under a regime of decreasing photoperiod, from LD 15:9h, to LD 12:12 h through larval, pupal and adult development. Further experiments demonstrated that the response depended on a decrease in photoperiod from pre-pupal stage to adult emergence (the pupal period is spent below ground). A reduction of 50–60min over the 13–15 day pupal period at 23^oC was effective. The significance of decreasing photoperiods, irrespective of absolute levels, as a cue for species whose migrations traverse substantial ranges of latitude is considered and the results are discussed in relation to the genetic determination of PCP in M.separata and to its seasonal migrations in eastern China.     

Photoperiodic control of diapause induction and termination in Ostrinia nubilalis: two different clocks?

Both diapause induction and diapause termination are under photoperiodic control in the lepidopteran, Ostrinia nubilalis. In the present study, induction of diapause was maximal in light-dark (LD) cycles that contained 12 hr of light alternating with 12 hr of darkness (LD 12:12). Termination of diapause was maximal in LD 16:8. Diapause termination also occurred rapidly in non-24-hr LD cycles that possessed an 8-hr dark phase. In each of these cases, the period of the LD cycle was not important. Diapause termination did not, however, occur rapidly in non-24-hr LD cycles that lacked an 8-hr dark phase. Thus, the clock mechanism underlying the termination response resembles an hourglass in its behavior. This is in contrast with what is known about induction of diapause. Here it has been demonstrated that the circadian system is somehow involved. It is thus possible that two different physiological clocks underlie these responses.     

Photoperiodic clock of diapause induction in Pseudopidorus fasciata (Lepidoptera: Zygaenidae)

Pseudopidorus fasciata enters diapause as fourth instar larvae at short day lengths. Using 24-h light-dark cycles, the photoperiodic response curves in this species appeared to be similar with a critical night length of 10.5h at temperatures below 30 degrees C. At an average temperature of 30.5 degrees C, the critical night length had shifted to between 15 and 17h. In experiments using non-24-h light-dark cycles, it was clearly demonstrated that the dark period (scotophase) was the decisive phase for a diapause determination. In night interruption experiments using 24-h light-dark cycles, a 1-h light pulse at LD12:12 completely reversed the long night effect and averted diapause in all treatments. At LD 9:15 light pulses of 1-h, 30- or 15-min also averted diapause effectively when both the pre-interruption (D(1)) or the post-interruption scotophases (D(2)) did not exceed the critical night length. If D(1) or D(2) exceeded the critical night length diapause was induced. The most crucial event for the photoperiodic time measurement in this species is the length of the scotophase. A 10-min light pulse placed in the most photosensitive phase reversed diapause in over 50% of the individuals. Night interruption experiments under non-24-h light-dark cycles indicated that the photoperiodic clock measured only D(1) regardless of the length of D(2), suggesting that the most inductive cycles are often those in which L+D are close to 24h. In resonance experiments, this species showed a circadian periodicity at temperatures of 24.5 or 26 degrees C, but not at 30.5 and 23.3 degrees C. On the other hand, Bünsow and skeleton photoperiod experiments failed to reveal the involvement of a circadian system in this photoperiodic clock. These results suggest the photoperiodic clock in this species is a long-night measuring hourglass and the circadian effect found in the final expression of the photoperiodic response in the resonance experiments may be caused by a disturbing effect of the circadian system in unnatural regimes.     

The effect of temperature and light pulses on the induction of diapause in the Toyama strain of the Indian meal moth, Plodia interpunctella

Abstract.The photoperiodic response in Plodia interpunctella collected at Toyama (36.7°N) was of long-day type and highly sensitive to temperature. The critical photoperiod giving 50% diapause was between 14 and 16 h at 20°C, between 12 and 14 h at 25°C and between 6 and 8 h at 30°C. Effects of night interruption by a 2-h light pulse on the diapause response were examined at 25°C on different background photoperiods ranging from LD 12:12 h to LD 2:22 h. Percentage diapause was very low when the middle portion of dark period was interrupted, so that U- or V-shaped response curves were obtained with background scotophases longer than 12 h. In these curves, the descending slopes were less steep than the ascending slopes. The critical dark period measured from dusk to an interrupting light pulse was about 1.5 h longer than the critical dark period ( c . 10 h) in the normal photoperiodic response. The critical dark period from the interrupting light pulse to dawn, on the other hand, was not parallel to dawn but shorter than the normal critical period in LD 12:12 h and LD 10:14 h and longer than that in LD 7:17 h to LD 4:20 h, indicating that the priming effects of the light pulse might be under the influence of the photophase.     

Light and temperature entrainment of a locomotor rhythm in honeybees

Abstract. The circadian locomotor (walking) rhythms of forager honeybees (Apis mellifera ligustica L.) were entrained to eight different 24 h light-dark cycles. The phases of activity onset, peak activity, and offset were correlated with the lights-off transition, suggesting lights-off as the primary zeitgeber for the rhythm. Further support for this hypothesis was provided by LD 1:23 experiments, in which entrainment occurred when the light pulse was situated at the end, but not at the beginning, of the subjective photophase. Steady-state entrainment of the locomotor rhythm was achieved with square-wave temperature cycles of 10^oC amplitude under constant dark: most of the activity occurred within the early thermophase. Smaller amplitude temperature cycles yielded relative coordination of the rhythm. Interactions of temperature and light-dark cycles resulted in entrainment patterns different from those elicited in response to either cycle alone or those formed by a simple combination of the two separate responses. Furthermore, temperature cycles having amplitudes insufficient for entrainment of the rhythm nevertheless modified the pattern of entrainment to light - dark cycles, suggesting a synergism of light and temperature effects on the underlying circadian clock system.     

Diapause induction and photoperiodic clock in Chilo suppressalis (Walker) (Lepidoptera: Crambidae)

The mature larvae of the rice stem borer, Chilo suppressalis Walker (Lepidoptera: Crambidae) enters facultative diapause in response to short‐day conditions in the autumn (August–September). Diapause induction and photoperiodic clock mechanism were investigated in C. suppressalis larvae reared on an artificial diet in the present study. The critical night length for diapause induction was about 9 h 53 min to 10 h 39 min at 22 to 28°C. The third‐instar larvae were found to be relatively sensitive to diapause induction. Photoperiodic response under non‐24‐h light–dark cycles showed that scotophase length played an essential role in the induction of larval diapause in C. suppressalis, and consecutive exposure to long‐night cycles was necessary for a high diapause incidence. In the Nanda–Hamner experiment, diapause incidence peaked at scotophase of 12 h and dropped rapidly at scotophases > 24 h. In the Bünsow experiment, diapause incidence was clearly suppressed, especially at the light pulse located 8 h in the scotophase. Both the Nanda–Hamner and Bünsow experiments showed no rhythmic fluctuations with a period of about 24 h; thus the photoperiodic clock in C. suppressalis is a non‐oscillatory hourglass timer or a rapidly damping circadian oscillator.     

Induction of winged sexuparae in root-inhabiting colonies of the aphid Pemphigus betae

Abstract. Growth chamber experiments were conducted to determine what environmental factors induce the development of sexuparae in root-feeding colonies of the aphid Pemphigus betae Doane (Aphididae: Pemphiginae). Experimental colonies were derived from clones that had been reproducing for at least 15 months under constant laboratory conditions. At constant 20–23^oC, sexuparae were almost never produced regardless of the level of crowding or photoperiod. In contrast, every culture of several clones produced sexuparae when subjected to temperatures alternating between 20 and 15^oC during the light and dark phases respectively. This fluctuating temperature regime resulted in sexupara production regardless of photoperiod. All cultures also produced some sexuparae when cultures were kept in constant darkness at temperatures that were either alternated between 20 and 15^oC or kept constant at 15^oC. Constant 17^oC resulted in sexupara production in most cultures. At 17^oC, experimental manipulation of densities within cultures showed that crowding promotes sexupara production. Thus, within the range of conditions occurring at the time of sexupara induction in the field, photoperiod has no effect on sexupara production, temperature below a threshold between 17 and 20–23^oC triggers sexupara production, and crowding promotes sexupara production but only if temperature is low enough.     

Reproductive and developmental consequences of a diapause maternal effect in the flesh fly, Sarcophaga bullata

Abstract Progeny of Sarcophaga bullata produced from mothers with a history of short day will not enter pupal diapause even if they are reared in a strong diapause-inducing environment (LD 12:12 h at 20^oC). Short-day exposure and diapause commitment are normally inseparable, but this maternal effect provides a tool for examining separately the effect of photoperiod and diapause commitment. Duration of the wandering period of the third instar is longer in diapause-destined larvae than in non-diapause-destined larvae, and fecundity of flies that have experienced pupal diapause is lower than in long-day flies that have not been through diapause. The puparia of diapausing pupae contain more hydrocarbons than puparia of nondiapausing pupae, and this contributes to higher rates of net transpiration for the nondiapausing flies. Flies showing the maternal effect (short-day experience but no diapause) show an intermediate response: length of wandering, fecundity rate and quantities of puparial hydrocarbon are between the extremes observed in the other two groups of flies. Thus, the maternal effect switches the developmental programme to nondiapause, but the progeny retain some characteristics of diapause. Evidence from reciprocal crosses indicates that the photoperiodic history of the female, rather than the male, is responsible for the influence on fecundity.     

Assessment of phosphine fumigation as a plant quarantine measure against Spodoptera littoralis

In laboratory tests first instar larvae of Spodoptera littoralis were killed by phosphine fumigations of 24 and 48 h at a concentration x time product (CTP) of 36 mg h/1. Eggs aged 1–2 days were, however, tolerant to phosphine at a CTP of 144 mg h/1 and this CTP caused severe damage to chrysanthemum cuttings (cv. Freedom). When eggs aged < 1 day and 1–2 days old were preconditioned at 20 ^oC for 72 h before phosphine fumigation, complete mortality occurred at CTPs at and above 36 mg h/1. Preconditioning for 48 h at 20 ^oC and for 48 and 72 h at 15 ^oC did not produce 100% mortality at a CTP of 288 mg h/1. It was suggested that eggs of S. littoralis pass through a phase of tolerance to phosphine which at 20 ^oC lasts about 4 days after oviposition.
There was no indication that phosphine was toxic to chrysanthemum cuttings (cv. Pollyanne) at concentrations of 0–75 mg/1 (CTPs of 18 and 36 mg h/1). Storage at 15 ^oC was not damaging to cuttings but at 20 ^oC some damage became apparent.     

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

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