Photoperiodic control of diapause termination, colour change and postdiapause reproduction in the southern green stink bug, Nezara viridula

Abstract.  The effects of day length on adult diapause development, associated with diapause body colour change as well as postdiapause reproduction are studied in Nezara viridula from Japan. Facultative diapause spontaneously terminates under three constant short-day and near-critical photoperiods at 25 °C without low temperature treatment. The period required for body colour change from russet to green and the precopulation and preoviposition periods differ significantly between the photoperiodic treatments, being shortest under LD 13 : 11 h, intermediate under LD 12 : 12 h and longest under LD 10 : 14 h. Photoperiodic conditions do not affect postdiapause reproductive performance: the total egg production, duration of the period of oviposition and other reproductive indices do not differ significantly between the photoperiodic conditions. The total egg production depends on the duration of the period of oviposition but not on how long females remained russet during diapause. It is concluded that diapause in N. viridula does not require low temperature for its successful completion and diapause duration affects winter survival but not postdiapause reproductive performance or longevity. Such independence of the postdiapause reproductive performance from the duration of diapause may have contributed to the continuous worldwide range expansion of this species into temperate zone.     

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.     

Effects of temperature of adults and eggs on the induction of embryonic diapause in the band-legged ground cricket, Dianemobius nigrofasciatus

Abstract.  Eggs laid by adult female Dianemobius nigrofasciatus , reared under long-day (LD 16 : 8 h, 25 °C) or short-day (LD 12 : 12 h, 25 °C) conditions from the nymphal stage, are kept at several constant temperatures. At 22.5–30.0 °C, eggs laid by long-day adults show lower incidences of diapause than those laid by short-day adults. In both eggs laid by adults under long-day conditions and those under short-day conditions, the higher the temperature at which the eggs are kept, the lower the incidence of diapause. When eggs of long-day adults are exposed to a low-temperature pulse (10 °C, 24 h) on the day of deposition (day 0), the incidence of diapause increases. The low-temperature pulse on day 1 does not increase the incidence of diapause. By contrast, when the eggs of short-day adults are exposed to a high-temperature pulse (35 °C, 24 h) on day 0 or day 1, the incidence of diapause decreases. The temperature pulses on day 0 are more effective at diapause prevention. Staining of diapause eggs by the Feulgen–Rossenbeck method shows that the eggs enter diapause at the blastoderm stage, which is on day 1 or day 2 at 25 °C. The exposure of adults to long days and higher temperatures prevents the eggs from entering diapause. In D. nigrofasciatus , embryonic diapause is controlled by maternal effects, adult photoperiod and temperature, and egg temperature before or at diapause.     

PHOTOPERIODIC CONTROL OF ADULT DIAPAUSE IN CHRYSOPERLA SINICA (TJEDER) (NEUROPTERA: CHRYSOPIDAE— I. CRITICAL PHOTOPERIOD AND SENSITIVE STAGES OF ADULT DIAPAUSE INDUCTION

Abstract  Photoperiodic sensitivity for diapause induction of the green lacewing, Chrysoperla sinica (Tjeder) was examined at 22°C. The adult diapause of C. sinica was induced by short-day photoperiods, and the critical photoperiod for its induction was between 12.5L-11.5D and 13L-11D.
Adults developed without diapause under long-day conditions, and entered diapause under short-day conditions. The adult stage was the uppermost sensitive stage for adult diapause induction, adults could go into diapause only when the emerging adults were under diapause-inducing short-day photoperiods. The short-day photoperiodic experience by transferring between 15L: 9D and 9L: 15D at preimaginal stages did not result in adult diapause under 15L: 9D photo regime, although some treatments extended the pre-oviposition period in adult stage. The results showed that the 3rd instar larvae and pre-pupae were more sensitive to the photoperiodic change from 15L: 9D to 9L: 15D photo regime than the other preimaginal stages.     

Summer diapause and nymphal growth in a subtropical cockroach: response to changing photoperiod

Abstract.  The effects of photoperiod on nymphal growth and adult reproduction were investigated in a small cockroach, Margattea satsumana, living on the subtropical, Hachijo island (33°N), Japan. Nymphal development is slow under constant photoperiods at 25 °C. The shortest mean duration of nymphal development (176 days) is observed at LD 14 : 10 h, followed by LD 12 : 12 h (221 days) and LD 16 : 8 h (309 days). Nymphal development is further prolonged when the nymphs are transferred from LD 12 : 12 to LD 16 : 8 h at 90 days after hatching. However, rapid and synchronized development is observed when nymphs were transferred in the opposite direction. A decreased change in photoperiod from LD 14 : 10 to LD 12 : 12 h also reduces the duration of nymphal development, and this cannot be explained by the results obtained at constant photoperiods. Similarly, nymphs reared at LD 16 : 8 h during the first 60 days mature more rapidly when transferred to LD 12 : 12 h than when transferred to LD 14 : 10 h. The developmental suppression induced by long days may represent a form of summer diapause that is terminated rapidly by short days. Based on these observations and field-census data, it is suggested that this cockroach has a univoltine life cycle overwintering as nondiapause adults, and that this life cycle is stabilized by the response to changing photoperiod.     

Bimodal life cycle of the burying beetle Nicrophorus quadripunctatus in relation to its summer reproductive diapause

Abstract 1. Under natural conditions in Kyoto, Japan, the reproductive activities of Nicrophorus quadripunctatus Kraatz (Coleoptera: Silphidae) decreased in summer and the species showed a bimodal life cycle.
2. In the laboratory, most adult pairs raised at 20 °C under a LD 12:12 h regime reproduced when provided with a piece of chicken. In adults raised at 20 °C under a LD 16:8 h regime, however, both reproductive behaviour and ovarian development were reduced. It is concluded that these adults entered a reproductive summer diapause.
3. High temperature (25 °C) also suppressed the reproductive behaviour even under a favourable LD 12:12 h regime. In the field, therefore, adults reduce their reproductive activity in summer because of diapause induced by long-day photoperiods and direct inhibition of reproduction by high temperatures.
4. When the temperature was changed from 20 °C to 25 °C immediately after hatching of larvae, they reached the wandering stage in 95% of adult pairs. When the temperature was changed from 20 °C to 25 °C immediately after oviposition, however, no larvae hatched in 85% of pairs. Egg mortality was significantly higher at 25 °C than at 20 and 22.5 °C; no eggs hatched at 27.5 °C. The physiological mechanisms for reducing reproduction probably prevent the beetles from inefficient oviposition in summer.     

Effects of photoperiod and temperature on the development of Bonagota cranaodes

Abstract.  The Brazilian apple leafroller, Bonagota cranaodes (Meyrick) (Lepidoptera: Tortricidae) is reared in the laboratory under a long-day (LD 14 : 10 h) and a short-day (LD 7 : 17 h) photoperiod at 22 °C, and under two different temperatures (10–13 °C and 21–22 °C). The development time from larval to adult eclosion do not differ between the two photoperiods, but did between the two temperature regimes. However, the larvae do not enter diapause, even under short day conditions and low temperatures. The number of adults obtained does not differ with temperature and light conditions. Field captures with pheromone traps show that Brazilian apple leafroller occurs in apple orchards throughout the year and the population densities are lower in winter. Accordingly, control measures should be taken during the off-season.     

Responses to stepwise photoperiodic changes for the larval diapause of the Indian meal moth Plodia interpunctella

Abstract The Indian meal moth Plodia interpunctella Hübner (Lepidoptera: Pyralidae) diapauses as a last‐instar (fifth) larva. At 30 °C, no larvae enter diapause under any photoperiodic conditions; at 25 °C, the photoperiodic response curve is a long‐day type with a critical length of approximately 13 h light; at 20 °C, diapause is induced moderately even under long days (> 13 h). Cumulative effects of short days or long days on diapause induction are determined by alternate, stepwise and gradually changing regimes of photoperiod at 25 °C. When the larvae are repeatedly exposed to LD 16 : 8 h and LD 12 : 12 h photoperiods every other day, the incidence of diapause is 37%. When the larvae are placed under an LD 16 : 8 h photoperiod for 2 days and then under an LD 12 : 12 h photoperiod for 1 day, it is 38 %. Exposure to an LD 16 : 8 h photoperiod for 1 day and then to an LD 12 : 12 h photoperiod for 2 days induces only 15% diapause. This may indicate that the photoperiodic information is not accumulated in a simple fashion despite the generally accepted hypothesis (i.e. photoperiodic counter). Larvae exposed to an LD 16 : 8 h photoperiod for 5 days after oviposition express a very high incidence of diapause even under short days between an LD 2 : 22 h and LD 12 : 12 h photoperiod. After 10 days exposure to an LD 16 : 8 h photoperiod, however, the short day does not induce diapause strongly. On the other hand, an LD 12 : 12 h photoperiod in the early larval life is highly effective in the induction of diapause. A gradual increase or decrease of photoperiod (2 min day^?1) shows that the direction of photoperiodic change does not affect the diapause determination.     

Environmental and genetic control of the larval diapause of the Southwestern corn borer, Diatraea grandiosella

ABSTRACT. The interaction of photoperiod and temperature in the regulation of the induction and termination of the larval diapause of the Southwestern corn borer, Diatraea grandiosella Dyar (Lepidoptera), was examined. A population originating from south-eastern Missouri had critical daylengths for diapause induction of about 15h 5min (ecological threshold) and llh (physiological threshold). The ecological threshold was more stable than was the physiological threshold at temperatures lower than 25°C. Above 25°C the diapause response was suppressed. The insect appears to measure photoperiods in a stationary manner since a stepwise increase or decrease in photoperiod did not affect the incidence of diapause. In the critical region of the photoperiodic response curve, a higher incidence of diapause was found among females than among males. Females entered diapause later than did males, but resumed active development earlier than males. The rate of diapause development was more temperature dependent than was the rate of diapause induction, yet it was also clearly under photoperiodic control. The temperature coefficient (Q₁₀) for this process was about 4. Several other factors including sex-linkage, age, and geographic adaptations are involved in controlling the rate of diapause development, even more so than they are in controlling diapause induction. In the laboratory, the intensity of diapause declined gradually without larvae being exposed to non-diapause inducing conditions. Incubation of field-collected larvae revealed that their sensitivity to diapause maintaining photoperiods had ended by January. Three generations of selection of a Mississippi population of D. grandiosella at 30°C and LD 12:12 led to the production of an essentially diapause-free strain and a diapause strain.     

Photoperiodic control of diapause in Pseudopidorus fasciata (Lepidoptera: Zygaenidae) based on a qualitative time measurement

In the zygaenid moth, Pseudopidorus fasciata, both larval diapause induction and termination are under photoperiodic control. In this study, we investigated whether photoperiodic time measurement (with a 24-h light-dark cycle) in this moth is qualitative or quantitative. Photoperiodic response curves, at 22, 25, and 28 degrees C indicated that the incidence of diapause depended on whether the scotophases exceeded the critical night length (CNL) or not. All scotophases longer than the CNL-induced diapause; all scotophases shorter than the CNL-inhibited diapause. The CNL was 10.5h at 25 and 28 degrees C, and 10h at 22 degrees C. By transferring from various short photoperiods (LD 8:16, LD 9:15, LD 10:14, LD 11:13, LD 12:12, and LD 13:11) to a long photoperiod (LD 16:8) at different times, the number of light-dark cycles required for 50% diapause induction at 25 degrees C was 7.14 at LD 8:16, 7.2 at LD 9:15, 7.19 at LD 10:14, 7.16 at LD 11:13, and 7.13 at LD 12:12, without showing a significant difference between the treatments. Only at LD 13:11 (near the CNL), the number of light-dark cycles was significantly increased to 7.64. The intensity of diapause induced under different short photoperiods (LD 8:16, LD 9:15, LD 10:14, LD 11:13, and LD 12:12) at 25 degrees C was not significantly different with an average diapause duration of 36 days. The duration of diapause induced under LD 13:11 was significantly reduced to 32 days. All results indicate that the night-lengths are measured as either "long" or "short" compared with some critical value and suggest that photoperiodic time measurement for diapause induction in this moth is based on a qualitative principle.     

Timing of diapause induction outside the natural distribution range of a species: an outdoor experiment with the bean bug Riptortus clavatus

The phytophagous bug Riptortus clavatus (Thunberg) (Heteroptera: Alydidae) produces two or three generations per year in Central Japan and overwinters in the adult stage. In bugs from the Kyoto population (35°00 N, 135°45 E), we studied (1) the effects of day-length on the nymphal and preoviposition periods under constant photoperiod at 20.5 °C, and (2) photoperiodic induction of adult diapause at 20.5 °C and under a combination of constant photoperiod and natural daily rhythm of temperature in the forest-steppe zone of Russia (50°38 N, 35°58 E). Then, we examined (3) the timing of diapause induction under quasi-natural conditions in the same region, far outside the species' natural geographical range. At 20.5 °C, the nymphal period in both males and females was significantly longer under regimes with shorter photophases than under those with longer photophases. The preoviposition period in females was significantly longer under the near-critical long-day regime L14:D10 than under typical long-day regimes (L15:D9, L16:D8, and L17:D7). The critical day-length for diapause induction was shorter under conditions of natural daily rhythm of temperature than those reported at constant 20, 25, and 30 °C. Under quasi-natural conditions in the forest-steppe zone, R. clavatus entered diapause in September, much later than the local populations of true bugs studied to date. This experiment showed that R. clavatus was maladapted to new environmental conditions: diapause was induced too late with the result that all or most nymphs hatched in late August or early September will die.     

Photoperiodic induction of diapause in the spider mite Tetranychus urticae: qualitative or quantitative time measurement?

Abstract. Insects and mites may measure photoperiods eitfier by classifying them as long or short relative to a critical value (qualitative time measurement) or by using the absolute value (quantitative time measurement). The spider mite Tetranychus urticae is thought to use a qualitative mechanism of time measurement. In this paper we present the results of experiments with an inbred line of the spider mite (to keep genetic variation in photoperiodic responses small), to test whether quantitative aspects also play a role. Differences in diapause incidence in different long-night photoperiods at different temperatures may be an indication of quantitative responses to photoperiod. The effect of temperature on the photoperiodic response curve was studied at 16^oC, 19^oC and 22^oC. The response curves appeared to be similar at 16^oC and 19^oC, with a critical nightlength between 10 and 11 h. At 22^oC, diapause induction was less than 100% in all long-night regimens and die critical nightlength had shifted to 12 h. Maximum diapause induction (93%) occurred in a light-dark cycle with a 16 h dark phase (LD 8:16 h). Diapause induction was lowest in long-night photoperiods with dark phases of 20 h and longer. The number of light-dark cycles needed for 50% diapause induction at 19^oC varied. between 12.1 and 14.7 for LD 6:18 h, between 10.9 and 12.5 for LD 8:16 h, between 10.6 and 11.6 for LD 10:14 h, and between 10.1 and 10.7 for LD 12:12 h. Independent of die light-dark regimen, diapause induction took place in some individuals after receiving 8 cycles and virtually all individuals entered diapause after 16 cycles. No effect was found of the photoperiodic treatment during prediapause development (LD 6:18 h, LD 8:16 h, LD 10:14 h, LD 12:12 h) on diapause duration. The average diapause duration at LD 10:14 h and 19^oC was 61 days over all four treatments. We explained the results by hypothesising that nightlengths are assessed qualitatively and mat the photoperiodic clock operates more accurately near the critical nightlength.     

Parental and direct effects of photoperiod and temperature on the induction of larval diapause in the blow fly Lucilia sericata

Abstract.  Lucilia sericata (Meigen) (Diptera: Calliphoridae) shows a facultative diapause in the third, and final larval, instar after the cessation of feeding. The effects of photoperiod and temperature on the induction and duration of diapause were examined in parental (G0) and current (G1) generations. Insects of the G0 generation were reared under four combinations of conditions, involving two photoperiods, LD 16 : 8 and LD 12 : 12 h, and two temperatures, 25 and 20 °C. The G1 generation, present in the eggs laid by these insects, were transferred to 10 combinations of conditions, involving the above two photoperiods and five temperatures, 25, 20, 17.5, 15 and 12.5 °C. In the G1 generation, the time from hatching to cessation of feeding was significantly affected by temperature only, whereas the induction of diapause was influenced by both photoperiod and temperature experienced by the G0 as well as the G1 generation. Short-day and low-temperature conditions in the G0 and in the G1 generation had diapause-inducing effects. In this species, it is likely that, for purposes of acquiring reliable seasonal information, induction of diapause is sensitive to environmental factors both in the G0 and G1 generations. The function of high-intensity diapause, induced by short-day conditions and high temperature in the parental generation, appeared to be the prevention of accidental pupariation in warm autumn weather.     

Induction and termination of diapause inOrius predatory bugs

Photoperiodic induction of reproductive diapause at 18°C was investigated in fourOrius [Heteroptera: Anthocoridae] species.Orius insidiosus (Say) displayed a long-day response with a critical photoperiod between L11:D13 and L12:D12. Diapause in this species was terminated rapidly when the temperature and/or the daylength were increased.Orius majusculus (Reuter) also displayed a long-day response. The critical photoperiod fell between L14:D10 and L16:D8. Diapause in this species was not terminated within 14 days when both temperature and daylength were increased. InOrius albidipennis (Reuter) no diapause could be induced at photoperiods varying from L8:D16 to L16:D8. InOrius tristicolor (White) a high proportion of diapause was found at all photoperiods tested. The effect of temperature on photoperiodic induction of diapause was studied inO. insidiosus at L10:D14. Diapause occurred at 18°C, 21°C and 25°C, but not at 30°C. Again, diapause was terminated rapidly after transfer to 25°C/L16:D8. Exposing only the nymphal instars 1–5 to short daylength was not enough to induce diapause in the whole population ofO. majusculus. Orius predatory bugs are used as biocontrol agents against western flower thrips,Frankliniella occidentalis (Pergande) [Thysanoptera: Thripidael, in greenhouses. The consequences of photoperiodic induction of diapause for the success of early season releases ofOrius are discussed.     

Induction and termination of prepupal summer diapause in Pseudopidorus fasciata (Lepidoptera: Zygaenidae)

The seasonal life cycle of the zygaenid moth, Pseudopidorus fasciata is complicated by two different developmental arrests: a winter diapause as a fourth larval instar and a summer diapause as a prepupa in a cocoon. Both larval diapause induction and termination are under photoperiodic control. Short days induce larval diapause with a critical daylength of 13.5h and long days terminate diapause with a critical daylength of 14h. In the present study photoperiodic control of summer diapause was investigated in Pseudopidorus fasciata. Under long photoperiods ranging from LD 14:10 to LD 18:6, only part of the population entered summer diapause, the rest continued to develop. The lowest number of prepupae entered diapause at LD 14:10, followed by LD 16:8 and LD 17:7. The highest incidence of diapause occurred with photoperiods of LD 15:9 and LD 18:6. By transferring the diapausing prepupae induced by various long photoperiods (LD 14:10, LD 15:9, LD 16:8, LD 17:7, LD 18:6) to LD 13:11, 25 degrees C, the duration of diapause induced by LD 14:10 was significantly shorter than those induced by longer photoperiods. By keeping aestivating prepupae induced by LD 15:9, 28 degrees C or by natural conditions at short photoperiods (LD 11:13 and LD 13:11) and at a long photoperiod (LD 15:9), the duration of diapause at LD 15:9 was more than twice as long as than those at LD 11:13 and LD 13:11. Moreover, adult emergence was highly dispersed with a high mortality at LD 15:9 but was synchronized with low mortality at LD 11:13 and LD 13:11. When the naturally induced aestivating prepupae were kept under natural conditions, the early aestivating prepupae formed in May exhibited a long duration of diapause (mean 126 days), whereas the later-aestivating prepupae formed in July exhibited a short duration of diapause (mean 69 days). These results indicate that aestivating prepupae require short or shortening photoperiod to terminate their diapause successfully. By transferring naturally induced aestivating prepupae to 25, 28 and 30 degrees C, the duration of diapause at the high temperature of 30 degrees C was significantly longer than those at 25 and 28 degrees C, suggesting that high temperature during summer also plays an important role in the maintenance of summer diapause in Pseudopidorus fasciata. All results reveal that summer diapause can serve as a "bet hedging" against unpredictable risks due to fluctuating environments or as a feedback mechanism to synchronize the period of autumn emergence.     

Diapause induction and coloration in the Senegalese grasshopper, Oedaleus senegalensis

Abstract. Embryonic diapause induction in the Senegalese grasshopper, Oedaleus senegalensis Krauss (Orthoptera: Acrididae), is influenced both by the photoperiod and the temperature experienced by females. High temperatures (40°C) and long photoperiods (LD 14:10h), which characterize the beginning of the rainy season in the Sahel, cause non-diapausing eggs to be laid. Lower temperatures (25°C) and shorter photoperiods (LD 12:12h), which occur at the end of the rains, result in the production of diapausing eggs. At 30°C and constant photoperiods, O. senegalensis exhibited a long-day-short-day response with critical photoperiods of c. 13 h and c. 20 h, only the former value being of ecological significance. The photoperiodically sensitive stages to diapause induction in females occurred from the fifth stadium onwards. Temperature also affected the coloration of both nymphs and adults. Dark-black and pale-white individuals were produced by low (25°C) and high (40°C) temperatures respectively, whereas an intermediate temperature (30°C) produced individuals which were greyish brown. These results are discussed in relation to the ecology of O. senegalensis.     

Diapause‐inducing signals prolong nymphal development in the two‐spotted spider mite Tetranychus urticae

Female two‐spotted spider mite Tetranychus urticae are grown under different photoperiods and the photoperiodic regulation of diapause is examined. The photoperiodic response curve for diapause induction was of the long day–short day type, with critical day lengths (CDLs) of 2 and 12.5 h; diapause was induced between these CDLs. The preimaginal period is significantly longer in diapausing females than in non‐diapausing females; moreover, a significant positive correlation is detected between diapause incidence and deutonymphal period. Diapause incidence is high when long‐night photoperiods are applied against a background of continuous darkness in the stages including the deutonymph; this stage appears to be the most sensitive to photoperiod. These observations suggest that diapause‐inducing conditions inhibit nymphal development, particularly in the deutonymphal stage when photoperiodic time measurement for determination of reproduction or diapause is carried out.     

Photoperiodic response in the subtropical and warm‐temperate zone populations of the southern green stink bug Nezara viridula: why does it not fit the common latitudinal trend?

The photoperiodic response of diapause induction is studied in females of five subtropical and warm‐temperate zone populations of Nezara viridula (L.) (Heteroptera: Pentatomidae) in Japan (26.4–34.7°N; 127.4–135.7°E). Laboratory tests at 25 °C demonstrate that both warm‐temperate and subtropical populations have pronounced photoperiodic responses of adult diapause induction. Under short‐day conditions (LD 10 : 14 h), 73–100% of females enter diapause, whereas, under long‐day conditions (LD 15 : 9 h), 87–100% of females are nondiapause and reproduce. When the critical photoperiod for diapause induction is plotted against the latitude of origin of each population, the data points do not show the expected tendency of increasing critical photoperiod northwards but, instead, vary between 12 h 15 min and 13 h 30 min. It is suggested that adults from different populations of such a highly migratory species move often among subtropical islands (by themselves or being assisted by typhoons), thus constantly destabilizing the photoperiodic responses. Although important for general synchronization of seasonal development in the tropics and subtropics, winter diapause might not be so critical for survival in mild winters in these zones. Such circumstances might make the selective pressure over the photoperiodic response less severe in the tropics and subtropics than in the temperate zone. It is suggested that the current climate warming might contribute to this situation as well by promoting polewards migrations and lessening the selective pressure of overwintering conditions.     

Diapause and seasonal life cycle strategy in the house spider, Achaearanea tepidariorum (Araneae, Theridiidae)

Abstract In order to elucidate the mechanism regulating its seasonal life cycle, the photoperiodic response of Achaearanea tepidariorum has been analysed. Nymphal development was faster in long-day and slower in short-day photoperiods. The combined action of low temperature, poor food supply and short daylength induced diapause at an earlier developmental stage than short days alone. Thus, photoperiod is a primary factor inducing nymphal diapause, but the diapausing instar is influenced by both temperature and food supply. Hibernating nymphs became unresponsive to photoperiod in late December. After hibernation, however, sensitivity was restored and the nymphs remained sensitive to photoperiod throughout their life. This spider could also enter an imaginal or reproductive diapause. Photoperiod was again a primary inducing factor and temperature modified the photoperiodic response to some extent. The induction of the reproductive diapause was almost temperature-compensated whereas development was not. So the involvement of a photoperiodic counter system was suggested. Irrespective of whether the nymph had experienced diapause or not, the imaginal diapause was induced in response to a short-day photoperiod after adult moult. Based on these observations, the seasonal life cycle and the adaptive significance of nymphal and imaginal diapause are discussed.     

Inheritance of the photoperiodic response controlling imaginal summer diapause in the cabbage beetle, Colaphellus bowringi

Adults of the cabbage beetle Colaphellus bowringi display a summer diapause in response to the exposure of their larvae to long photoperiods. In the present study, the inheritance of the photoperiodic response controlling summer diapause in C. bowringi by crossing a high diapause strain (D strain) with a laboratory selected nondiapause strain (N strain) was investigated under different photoperiods at 22, 25 and 28 °C. The beetles in both reciprocal crosses and backcrosses showed a clear short-day response for the induction of diapause at all temperatures, similar to that of the D strain, suggesting that photoperiodic response of this beetle is heritable. The diapause incidences in the progeny from all the crosses under LD 15:9 or LD 12:12 at 25 °C suggest that genetic and genetic-environmental interactions are involved in diapause induction. The incidence of diapauses in F₁ progeny was significantly lower than that in the D × D strain but significantly higher than that in the N × N strain, indicating that the diapause capability is inherited in an incomplete dominant manner. The incidence of diapause was greater among the offspring of hybrid females when those females had a D strain mother or grandmother rather than a N strain mother or grandmother, indicating that maternal effects on diapause induction are stronger than paternal effects. The laboratory selected nondiapause strain also showed a short-day photoperiodic response at a low temperature of 22 °C, indicating that the photoperiodic photoreceptor and photoperiodic clock still function in the nondiapause strain.