Effects of thermoperiod on photoperiodic determination of larval diapause in Ostrinia nubilalis

Some effects of thermoperiods on the photoperiodic determination of larval diapause in the European corn borer, Ostrinia nubilalis are reported. Thermoperiods and photoperiods were shown to interact to a highly significant degree in the induction of diapause. Diapause determination was found to be sensitive to the duration and temperature of the cryoscotophase of the thermophotoperiod; the thermophotophase characteristics exerted only minor influence. Cryoscotophase temperatures of 10°C or lower caused an unexpected increase in the apparent critical nightlength for diapause determination. A 0°C cryoscotophase effectively prevented the photoperiodic determination of diapause. Experimental investigation of the effects of 0°C cryoscotophases produced evidence that such low-temperature cryoscotophases may suppress the biological clock functions regulating the determination of diapause. Photoperiodic regimes in which 0°C pulses were inserted at successive points in 24-h cycle evoked responses suggesting that the early scotophase (saturation phase) was more strongly influenced by the low temperature pulse than was the later scotophase (scotonon); low temperature had little effect on the light-dependent processes (photonon) associated with the photophase.     

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.     

Possible involvement of distinct photoreceptors in the photoperiodic induction of diapause in the flesh fly Sarcophaga similis

Physiological characteristics of the photoreceptors involved in the photoperiodic induction of diapause were investigated in the flesh fly Sarcophaga similis. Both the early and late phases of scotophase were sensitive to light and a light pulse during each of these phases prevented diapause. Certain physiological differences between the phases were, nevertheless, detected. Compared with early scotophase, late scotophase required a light pulse with a long period and a large number of night interruption photoperiodic cycles in order to effectively prevent diapause. The diapause-averting effects of a light pulse during early scotophase were canceled by an additional long dark period, but those during late scotophase were not. Thus, the diapause-averting effects produced during early scotophase are different to those produced during late scotophase. The early scotophase was sensitive to light at wavelengths of 470 nm or shorter, but not to light of 583 nm or longer. In contrast, the late scotophase was sensitive to light of a broad range of wavelengths, ranging from 395 to 660 nm. Furthermore, the early scotophase was considerably more sensitive to monochromatic light with low photon flux density than the late scotophase. These results suggest that different types of photoreceptor are involved in the photoperiodic response.     

Effect of light pulses in early scotophase on resetting of the night-measuring diapause clock of the Indian meal moth, Plodia interpunctella

Abstract.  The Indian meal moth, Plodia interpunctella Hübner (Lepidoptera: Pyralidae) may enter diapause in the last larval instar in response to the photoperiod during the preceding instars. An hourglass-type photoperiodic clock may measure night length for this purpose. The present study describes the resetting of the hourglass by light pulse(s) in the early scotophase and by scanning the subsequent clock phase by another light pulse (P). When the lights-off time of a first light pulse is fixed at 4 h after dusk under LD 4 : 20 h and LD 6 : 18 h photoperiods and its duration is increased from 1 to 3 h, the critical night length (CNL) from dawn is decreased, but that from dusk to P increases. A 3-h first light pulse efficiently resets the time measuring system. If this 3-h light pulse is split into two 1-h light pulses (L₁ and L₂) by 1 h of darkness, the dark-time measuring function appears to be impeded and CNL from P to dawn disappears, but that from L₂ to P is expressed. This indicates that the receptivity to light pulses varies among individual insects.     

The Influence of Light Wavelength on Phase-Dependent Responsiveness of the Photoperiodic Clock in Migratory Blackheaded Bunting

We investigated the phase-dependent effects of light wavelength on photoperiodic clock in the migratory blackheaded bunting. Two experiments were performed, employing a skeleton paradigm (6 hours light : 6 hours darkness : 1 hour light : 11 hours darkness; 6L : 6D : 1L : 11D) at 37 ± 2 lux intensity. In the experiment 1, both 6 and 1 h light pulses were given at the same wavelength, 500 nm (green) or 650 nm (red). A group exposed to both pulses of white light served as control. In the experi-ment 2, the two light pulses were given at two different wavelengths, 6 h at 500 nm (green) and 1 h at 640 nm (red) in one group or vice-versa in the other. There was almost no photoinduction when both light pulses in experiment 1, or 1 h light pulse in experiment 2, were green. On the other hand, birds fattened and testes recrudesced when both the light pulses in experiment 1, or 1 h light pulse in experiment 2, were red. Birds receiving both pulses of white light in experiment 1 showed an intermediate response. Taken together, these results indicate that the photoperiodic clock in buntings is differentially responsive at its various circadian phases to different light wavelengths.     

The Influence of Light Wavelength on Phase-Dependent Responsiveness of the Photoperiodic Clock in Migratory Blackheaded Bunting

We investigated the phase-dependent effects of light wavelength on photoperiodic clock in the migratory blackheaded bunting. Two experiments were performed, employing a skeleton paradigm (6 hours light : 6 hours darkness : 1 hour light : 11 hours darkness; 6L : 6D : 1L : 11D) at 37 ± 2 lux intensity. In the experiment 1, both 6 and 1 h light pulses were given at the same wavelength, 500 nm (green) or 650 nm (red). A group exposed to both pulses of white light served as control. In the experi-ment 2, the two light pulses were given at two different wavelengths, 6 h at 500 nm (green) and 1 h at 640 nm (red) in one group or vice-versa in the other. There was almost no photoinduction when both light pulses in experiment 1, or 1 h light pulse in experiment 2, were green. On the other hand, birds fattened and testes recrudesced when both the light pulses in experiment 1, or 1 h light pulse in experiment 2, were red. Birds receiving both pulses of white light in experiment 1 showed an intermediate response. Taken together, these results indicate that the photoperiodic clock in buntings is differentially responsive at its various circadian phases to different light wavelengths.     

The effect of photoperiod on the calling behaviour of virgin females of the true armyworm,Pseudaletia unipuncta (Haw.) (Lepidoptera: Noctuidae)

The effect of photoperiod on the calling behaviour of Pseudaletia unipuncta virgin females was examined under five different photoperiodic regimes at 25°C, 65% r.h. The age at which females called for the first time following emergence varied with photoperiod; generally calling was later under long-scotophase conditions. However under a 6 h scotophase there was also a delay in calling and >63% of the females tested never called. There was a considerable variability in the daily calling patterns between the different photoperiods, and the mean onset time of calling was not constant with either “lights on” or “light off”. There was however a constancy of the mean onset time of calling relative to the mid-point of both the photo- and scotophase at all photoperiods tested, indicating that females could measure the absolute duration of either the photo- or scotophase. Transferring females from one photoperiodic condition to another once calling had been initiated, determined that it was the “lights off” signal that P. unipuncta females used to phase set the clock governing circadian calling behaviour.Females subjected to a decrease of 4 or 6 h in the length of the scotophase following the initiation of calling required several days to adjust to the new photoperiodic regime and a high proportion of females did not call during the night following the transfer. A 4 or 6 h increase in the scotophase did not inhibit calling on the night following the transfer but females still required several days to adjust completely. However, females experiencing a 2 h increase or decrease in the duration of the scotophase were able to maintain normal calling behaviour. The results of these experiments are discussed in relation to the seasonal biology of the true armyworm and the hypothesis that this is a migrant species.     

Capacity of cytochrome and alternative path in coupled and uncoupled root respiration of Pisum and Plantago

A critical duration of darkness must be exceeded for the photoperiodic induction of flowering in short-day plants. This requires detection of the light/dark transition at dusk and the coupling of this information to a time-measuring system.
Lowering the P_fr/P_tot, ratio photochemically at the end of the day did not accelerate the onset of dark timing in Pharbitis nil Choisy cv. Violet. Time-measurement was initiated when, with no change in spectral quality, the irradiance fell below a threshold value. Thus, if the light/dark transition at dusk is sensed by a reduction in P_fr, this reduction can be achieved as rapidly through thermal reactions as through photochemical ones. When given at hourly intervals during a 6-h extension of a 24-h main light period in white light, pulses of red light were as effective as continuous red light in delaying the onset of timing; pulses every 2 or 3 h were less effective. The effectiveness of intermittent red light indicates that phytochrome is the photoreceptor and the requirement for frequent exposures suggests that P_fr is lost rapidly in the dark. However, the red light pulses could not be reversed by far-red light, which argues against this hypothesis. An alternative explanation is that the perception of light as being continuous occurs only when "new" P_fr is regenerated sufficiently frequently.
The nature of the coupling of the dusk signal to the time-measuring system is discussed and it is suggested that the effect of each red light pulse is to delay the phase of the photoperiodic rhythm by 1–3 h.     

Thermoperiod-photoperiod interactions in the determination of diapause in Ostrinia nubilalis

Photoperiodic determination of larval diapause in Ostrinia nubilalis is shown to be strongly influenced by superimposed thermoperiods, here termed “thermophotoperiods.” Thermoperiodic cryophase temperatures (0–25°C) exert effects that are markedly different from responses evoked by thermophotoperiodic cryoscotophases of the same temperatures. The differences are attributed to the presence or absence of light (thermophotophase or thermophase). Perturbations of 12 h dark: 12 h light and 10 h dark: 14 h light photoperiods by low-temperature pulses of 4 h duration demonstrated that the biological clock reactions associated with both the scotophase and photophase are sensitive to temperature. Statistically significant thermoperiod-photoperiod interactions have been demonstrated.     

Phase-shifting effects of a light pulse interrupting scotophase on locomotor activity rhythm and photoperiodic time measurement for diapause in the onion fly, Delia antiqua

When a light pulse of 1 h duration was given 3 h after lights off in a photoperiod of 11 h light : 13 h dark (LD 11 : 13) at 20°C, the phase of the major peak of locomotor activity rhythm in Delia antiqua was delayed for approximately 0.6 h. In contrast, it was advanced by approximately 0.6 h by a light pulse given 9 h after lights off. It is suggested that in the circadian clock, a pulse falling in the early scotophase is taken as a new dusk and a pulse falling in the late scotophase is taken as a new dawn. Although a sharply defined critical photoperiod did not exist in the diapause response to photoperiod in D. antiqua, the percentage of pupal diapause decreased by these pulses in LD 11 : 13 at 20°C. The effect of a 15 min light pulse on both locomotor activity rhythm and pupal diapause induction was stronger at 3 h than at 9 h after lights off, while a 1 min light pulse was ineffective at both times. The parallel effects of light pulse on locomotor activity rhythm and diapause response might be based on the same chronobiological functions.     

Night-interruption experiments and action spectra for dawn and dusk in relation to the photoperiodic clock of the cabbage whitefly,Aleyrodes proletella (Homoptera: Aleyrodidae)

The dark period (scotophase) is the most photoperiodically important part of a light-dark cycle in Aleyrodes proletella. Night-interruption studies have revealed three distinct dark stages: the photosensitive stage 1 lasts for about 3 h after dusk and 1-h light breaks both stop and re-set the photoperiodic clock; stage 2 also lasts about 3 h, but is photorefractory to some degree; stage 3 is photosensitive, but short light breaks do not re-set the clock although a 4-h light break (equivalent to a main photophase) does restore the capacity to respond to a normal critical night length in the post-interruption scotophase.Action spectra revealed peak photoperiodic sensitivity to blue light (410–430 nm) with 50% responses., at 1.5 μWcm⁻² and 2.5 μWcm⁻² for the dusk and dawn peaks respectively. These data are consistent with the view that the photopigment is a carotenoprotein.The results are interpreted in terms of the photoperiodic clock in A. proletella operating on the hour glass principle.     

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.     

Evidence for ‘dawn’ and ‘dusk’ hourglasses in Pimpla photoperiodic clock

Abstract

Resonance experiments (Nanda‐Hamner protocol) conducted at two temperatures for diapause termination in Pimpla instigator (Hymenoptera: Ichneumonidae) do not support the view that the photoperiodic clock has an oscillatory component, but suggest the presence of a non‐rhythmic timer or hourglass mechanism. These results are best explained by a two hourglasses model, one of which starts at light‐on and measures the photophase and the other is initiated by light‐off and measures the scotophase. The most likely hypothesis is that the ratio of photophase to scotophase lengths is the determining element. Good agreement is obtained between results predicted by two hourglasses model and results observed in Pimpla. The diurnal hourglass continues to run for long time (several months) in constant condition (LL) and does not require to be ‘turned over’ by D/L transition, in contrary to the classical model of hourglass which executes a single act of time measurement in extented phase and then stops. The most simple explanation is that some essential factor of diapause termination is synthesized during photophase and degraded during scotophase. Therefore an independent photoperiodic counter (for sommation of daily informations) is not necessary. The two hourglasses system serves as photoperiodic clock and accumulation of product as counter.     

Temperature effect on entrainment,phase shifting,and amplitude of circadian clocks and its molecular bases

Effects of temperature and temperature changes on circadian clocks in cyanobacteria, unicellular algae, and plants, as well as fungi, arthropods, and vertebrates are reviewed. Periodic temperature with periods around 24 h even in the low range of 1-2 degrees C (strong Zeitgeber effect) can entrain all ectothermic (poikilothermic) organisms. This is also reflected by the phase shifts-recorded by phase response curves (PRCs)-that are elicited by step- or pulsewise changes in the temperature. The amount of phase shift (weak or strong type of PRC) depends on the amplitude of the temperature change and on its duration when applied as a pulse. Form and position of the PRC to temperature pulses are similar to those of the PRC to light pulses. A combined high/low temperature and light/dark cycle leads to a stabile phase and maximal amplitude of the circadian rhythm-when applied in phase (i.e., warm/light and cold/dark). When the two Zeitgeber cycles are phase-shifted against each other the phase of the circadian rhythm is determined by either Zeitgeber or by both, depending on the relative strength (amplitude) of both Zeitgeber signals and the sensitivity of the species/individual toward them. A phase jump of the circadian rhythm has been observed in several organisms at a certain phase relationship of the two Zeitgeber cycles. Ectothermic organisms show inter- and intraspecies plus seasonal variations in the temperature limits for the expression of the clock, either of the basic molecular mechanism, and/or the dependent variables. A step-down from higher temperatures or a step-up from lower temperatures to moderate temperatures often results in initiation of oscillations from phase positions that are about 180 degrees different. This may be explained by holding the clock at different phase positions (maximum or minimum of a clock component) or by significantly different levels of clock components at the higher or lower temperatures. Different permissive temperatures result in different circadian amplitudes, that usually show a species-specific optimum. In endothermic (homeothermic) organisms periodic temperature changes of about 24 h often cause entrainment, although with considerable individual differences, only if they are of rather high amplitudes (weak Zeitgeber effects). The same applies to the phase-shifting effects of temperature pulses. Isolated bird pineals and rat suprachiasmatic nuclei tissues on the other hand, respond to medium high temperature pulses and reveal PRCs similar to that of light signals. Therefore, one may speculate that the self-selected circadian rhythm of body temperature in reptiles or the endogenously controlled body temperature in homeotherms (some of which show temperature differences of more than 2 degrees C) may, in itself, serve as an internal entraining system. The so-called heterothermic mammals (undergoing low body temperature states in a daily or seasonal pattern) may be more sensitive to temperature changes. Effects of temperature elevation on the molecular clock mechanisms have been shown in Neurospora (induction of the frequency (FRQ) protein) and in Drosophila (degradation of the period (PER) and timeless (TIM) protein) and can explain observed phase shifts of rhythms in conidiation and locomotor activity, respectively. Temperature changes probably act directly on all processes of the clock mechanism some being more sensitive than the others. Temperature changes affect membrane properties, ion homeostasis, calcium influx, and other signal cascades (cAMP, cGMP, and the protein kinases A and C) (indirect effects) and may thus influence, in particular, protein phosphorylation processes of the clock mechanism. The temperature effects resemble to some degree those induced by light or by light-transducing neurons and their transmitters. In ectothermic vertebrates temperature changes significantly affect the melatonin rhythm, which in turn exerts entraining (phase shifting) functions.     

An “hourglass-like” feature in the photoperiodic time measurement in Diatraea grandiosella (Pyralidae)

Photoperiodic time measurement of Diatraea grandiosella, a Pyralid moth, was investigated for its role in the determination of diapause by using various night-interruption protocols. The photoperiodic-response curve showed a temperature dependence under short days, whereas at long days it was stable in the range between 20 and 30°C. A light pulse averted diapause most effectively when it was placed 6 h after lights-off. Earlier pulses were less effective but gradually increased in effectiveness as the time of the pulse approached the critical time, i.e. 6 h after lights-off. A strong conversion in response appeared as the pulse passed the critical time. The same response pattern was observed both in diapause induction and termination. The species required a long light pulse to avert diapause completely, even if the light pulse was placed during the critical time; 45 min was required to reverse diapause in 50% of the individuals. The most crucial event for photoperiodic time measurement in this species was whether a night phase 6–7.5 h after lights-off was illuminated or not. This hourglass-like feature was also shown in more complex night-interruption protocols with 2–3 light pulses. A possible relation of this phenomenon to the cricadian pacemaker was sought. The Dual System Theory failed to account for most of the features for the photoperiodic time measurements of D. grandiosella found in this study.     

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.     

The behaviour and coupling of the photoreceptor and hour-glass photoperiod timer at low temperature in the aphid Megoura viciae

In Megoura viciae photoperiodic time is measured during the scotophase by an hour-glass mechanism which is temperature compensated within the range 6–20°C. At lower temperatures the clock “loses time”, running at about half the normal rate at 0°C. The present experiments, based on the analysis of critical night lengths, showed that for short periods of scotophase chilling (4h or less) the same diminished rate is maintained irrespective of the position of the low temperature period within the scotophase. This occurs despite the known changes in spectral sensitivity of the photoreceptor. With longer periods of chilling (>6h) a normal critical response was not observed unless several hours of darkness intervened between the end of the chilling period and the temperature-modified critical night length. This is interpreted as a “recovery” process involving the photoreceptor but not the hour-glass. Additional data indicated that the sequence of changes in the photoreceptor, including the early resetting phase, is closely coupled to the progression of the timer even when photosensitivity is temporarily impaired by low temperature.     

Interacting effects of photophase and scotophase on the diapause response of the Indian meal moth, Plodia interpunctella

The diapause-programming response to photoperiod in Plodia interpunctella was analyzed by exposing larvae to various 24-h and non-24-h regimes of light and darkness. The response to 24-h regimes indicated three photoperiodic parameters—a critical scotophase, a minimal photophase, and a minimal scotophase for a full expression of the response. The critical response was based on dark-time measurement, because disruption of the scotophase abolished the response and the diapause incidence varied as a function of scotophase in non-24-h regimes. The critical scotophase varied with the duration of the preceding photophase. Prevention of diapause by single or double-night interruptions of long scotophases could be explained by resetting of the dark-time measurement. The effect of a light pulse was modified by the quantitative interaction of light and dark reactions. The sensitivity to resetting by a light pulse seemed to be decreased in the early scotophase with an increasing duration of the preceding light period. Therefore, the significance of light in the photoperiodic response was something more than delimiting scotophase for the time measurement.     

Djungarian hamsters: a species with a labile circadian pacemaker? Arrhythmicity under a light-dark cycle induced by short light pulses

In most cases, phase-shifting effects of light pulses are studied in animals kept in constant darkness (DD) or in animals released into DD following the stimulus. In this study, the authors exposed Djungarian hamsters (Phodopus sungorus) to short light pulses during the dark phase of a 16:8 light-dark (LD) cycle and thus obtained a type VI phase response curve. Light pulses early in the night caused phase delays of the activity onset as well as phase advances of the activity offset, whereas light pulses later in the night resulted in phase advances of the activity offset only. A combination of two 15-min light pulses-the first one given late in the scotophase and the second given early in the dark phase of the following night-led to a strong compression of the activity phase alpha. In 75% of all animals, daily rhythms were no longer visible after complete alpha compression, and long-term arrhythmicity (up to 145 days) persisted despite continued exposure to an LD cycle. Because three independent output rhythms of the clock (i.e., activity, body temperature, and melatonin rhythms) were equally affected, the authors conclude that overt arrhythmicity was due not merely to disrupted output pathways but to an altered state of the central pacemaker. The authors suggest a qualitative two-oscillator model to explain this phenomenon. Their hypothesis assumes that, due to loose coupling, the pacemaker of Djungarian hamsters can be driven to a state of zero phase difference between the two oscillators, with zero amplitude of their outputs.     

暗期干扰对棉铃虫两个不同地理种群滞育抑制作用的比较

至今,所测试昆虫的光周期反应均表明,光周期反应对暗期干扰高度敏感,短暂的光脉冲都可在不同程度上逆转长夜效应,抑制滞育的发生。在研究了棉铃虫Helicoverpa armigera泰安种群(36.15°N,116.59°E)和喀佐种群(41.34°N,120.27°E)光周期反应的基础上,在滞育诱导的短光周期下(L12:D12和L9:D15),分别测试了暗期不同时段1h光脉冲对这两个不同地理种群滞育抑制的影响。25和22℃下的光周期反应显示了泰安种群在长暗期11—14 h的滞育率均显著低于喀佐种群;泰安种群的临界暗长分别为11.7 h和11.5 h,喀佐种群分别为10.5 h和10.3 h,泰安种群均比喀佐种群长1.2 h。在所测试的暗期干扰实验中,除了极少数光脉冲干扰点外,泰安种群蛹滞育率显著低于喀佐种群,但两者的滞育反应曲线基本相似。在短光周期L9:D15下,泰安种群和喀佐种群均显示了光脉冲落入暗期的第9—11小时最有效地抑制了滞育的发生。在短光周期L12∶D12下,泰安种群和喀佐种群在25℃时均显示了光脉冲落入暗期的第3—4小时和第10小时导致了最低的滞育发生;但在22℃时,喀佐种群只在暗期的第3—4小时显示了最高的滞育抑制。这些结果揭示了偏南的泰安种群对暗期干扰的敏感性强于偏北的喀佐种群,但这两个地理种群的最高光敏感位点基本相同。