Life history responses of the small brown planthopper Laodelphax striatellus to temperature change.

Temperature is a key environmental factor in ectotherms and influences many life history traits. In the present study, the nymphal development time, sex ratio and wing dimorphism of the small brown planthopper Laodelphax striatellus were examined under the conditions of constant temperatures, naturally varying temperatures (or different generations), and different temperatures combined with different photoperiod. The results showed that from 18 to 28 °C, the developmental time of nymphs was gradually shortened with the increase of temperature, whereas the high temperatures of 30 and 32 °C in the third to fifth instar nymphal stages and high summer temperature of 28.8 and 29.7 °C significantly delayed developmental time and resulted in higher mortality of nymphs. In all treatments, the developmental time was longer in females than males. The nymphs took significantly longer time to develop in the short daylength of 12 h than in longer daylengths of 13, 14, 15 and 16. Differences in developmental time were also found between wing morph, with long-winged individuals being significantly longer than the short-winged individuals at lower temperatures and significantly shorter than the short-winged individuals at higher temperatures. In all treatments, the sex ratio was stable, approaching 1:1, without being affected by temperature, generations and photoperiod. Photoperiod and temperature had significantly influence on the wing dimorphism. Long daylength combined with different temperatures resulted in significantly higher proportions of long-winged morph, whereas the low temperatures combined with the short daylengths in autumn and winter resulted in significantly high proportion of short-winged morph. This study broadens our understanding of the life-history traits of this planthopper and provides basic data for analyzing the effects of climate change on the planthopper reproduction.

Partial reuse of circadian clock genes along parallel clines of diapause in two moth species.

Understanding the molecular basis of repeated evolution improves our ability to predict evolution across the tree of life. Only since the last decade has high-throughput sequencing enabled comparative genome scans to thoroughly examine the repeatability of genetic changes driving repeated phenotypic evolution. The Asian corn borer (ACB), Ostrinia furnacalis (Guenée), and the European corn borer (ECB), Ostrinia nubilalis (Hübner), are two closely related moths displaying repeatable phenological adaptation to a wide range of climates on two separate continents, largely manifesting as changes in the timing of diapause induction and termination across latitude. Candidate genes underlying diapause variation in North American ECB have been previously identified. Here, we sampled seven ACB populations across 23 degrees of latitude in China to elucidate the genetic basis of diapause variation and evolutionary mechanisms driving parallel clinal responses in the two species. Using pooled whole-genome sequencing (Pool-seq) data, population genomic analyses revealed hundreds of single nucleotide polymorphisms (SNP) whose allele frequencies covaried with mean diapause phenotypes along the cline. Genes involved in circadian rhythm were over-represented among candidate genes with strong signatures of spatially varying selection. Only one of two circadian clock genes associated with diapause evolution in ECB showed evidence of reuse in ACB (period [per]), but per alleles were not shared between species nor with their outgroup, implicating independent mutational paths. Nonetheless, evidence of adaptive introgression was discovered at putative diapause loci located elsewhere in the genome, suggesting that de novo mutations and introgression might both underlie the repeated phenological evolution.

A multi-year dormancy strategy in a cabbage beetle population in southeastern China.

The life cycle of the cabbage beetle Colaphellus bowringi in southeastern China is complex due to four options for adult development: summer diapause, winter diapause, prolonged diapsuse, and nondiapause. However, detailed information on the multi-year emergence patterns of diapausing individuals in this beetle has not been documented. In this study, we monitored the adult emergence patterns of diapausing individuals and estimated the influence of the diapause-inducing temperature and photoperiod on the incidence of prolonged diapause under seminatural conditions for several years. The duration of diapause for adults collected from the vegetable fields in different years varied from several months to 5 years. Approximately 25.9%-29.2% of individuals showed prolonged diapause (emergence more than 1 year after entering diapause) over the 5 years of observation. Furthermore, regardless of insect age, the emergence of diapausing adults from the soil always occurred between mid-February and March in spring and between late August and mid-October in autumn, when the host plants were available. The influence of diapause-inducing temperatures (22, 25, and 28°C) combined with different photoperiods (L:D 12:12 h and L:D 14:10 h) on diapause duration was tested under seminatural conditions. Pairwise comparisons of diapause duration performed by the log-rank test revealed that the low temperature of 22°C combined with the long photoperiod of L:D 14:10 h induced the longest diapause duration, whereas the low temperature of 22°C combined with the short photoperiod of L:D 12:12 h induced the highest proportion of prolonged diapause. This study indicates that C. bowringi adopts a multi-year dormancy strategy to survive local environmental conditions and unpredictable risks.

Effects of temperature on life-history traits of the newly invasive fall armyworm, Spodoptera frugiperda in Southeast China.

In mid-May, 2019, the fall armyworm (FAW) Spodoptera frugiperda invaded Jiangxi Province, China, and caused extensive damage to corn crops. However, little attention has been given to the life-history traits of the FAW. In the present study, we systematically investigated the life-history traits of the newly invasive FAW on corn leaves at 19, 22, 25, 28, and 31°C under a photoperiod of LD 15:9 hr. The FAW thrived on the corn leaves with short developmental periods, high survival rates of larvae and pupae, very high mating success rates, and high fecundity. The pupal developmental stage was significantly longer in males than females at all temperatures, thus resulting in a protogyny phenomenon. The pupal weight was heaviest after a relatively shorter larval development stage at a higher temperature (25°C); thus, the FAW did not follow the temperature-size rule. Females were smaller than males, indicating sexual size dimorphism. A small proportion of females delayed their pre-oviposition period and began to lay eggs on the 7th to 9th day after adult emergence. There were positive relationships between pupal weight and larval developmental time and between adult weight and fecundity. There was a negative relationship between fecundity and longevity. These findings can help us to predict the population dynamics of the FAW on corn and to develop a suitable and practical management strategy.

Inheritance of key life-history traits in crosses between northern and southern populations of the cabbage beetle Colaphellus bowringi (Coleoptera: Chrysomelidae).

A southern population (S) from Xiushui County (29°1'N, 114°4'E) and a northern population (N) from Shenyang city (41°48'N, 123°23'E) of the cabbage beetle, Colaphellus bowringi vary greatly in their life-history traits, and may serve as an excellent model with which to study the inheritance of life-history traits. In the present study, we performed intraspecific hybridization using the two populations, comparing the key life-history traits (fecundity, development time, body weight, growth rate, and sexual size dimorphism (SDD)) between the two populations (S♀ × S♂ and N♀ × N♂) and their two hybrid populations (S♀ × N♂ and N♀ × S♂ populations) at 19, 22, 25, and 28°C. Our results showed that there were significant differences in life-history traits between the two parental populations, with the S population having a significantly higher fecundity, shorter larval development time, larger body weight, higher growth rate, and greater weight loss during metamorphosis than the N population at almost all temperatures. However, these life-history traits in the two hybrid populations were intermediate between those of their parents. The life-history traits in the S × N and N × S populations more closely resembled those of the maternal S population and N population, respectively, showing maternal effects. Weight loss for both sexes was highest in the S population, followed by the S × N, N × S, and N populations at all temperatures, suggesting that larger pupae lost more weight during metamorphosis. The changes in SSD with temperature were similar between the S and the S × N populations and between the N and the N × S populations, also suggesting a maternal effect. Overall, our results showed no drastic effect of hybridization on C. bowringi, being neither negative (hybrid inferiority) nor positive (heterosis). Rather, the phenotypes of hybrids were intermediate between the phenotypes of their parents.

Expression of alternative developmental pathways in the cabbage butterfly, Pieris melete and their differences in life history traits.

The seasonal life cycle of the cabbage butterfly, Pieris melete is complicated because there are three options for pupal development: summer diapause, winter diapause, and nondiapause. In the present study, we tested the influence of temperature, day length, and seasonality on the expression of alternative developmental pathways and compared the differences in life history traits between diapausing and directly developing individuals under laboratory and field conditions. The expression of developmental pathway strongly depended on temperature, day length, and seasonality. Low temperatures induced almost all individuals to enter diapause regardless of day length; relatively high temperatures combined with intermediate and longer day lengths resulted in most individuals developing without diapause in the laboratory. The field data revealed that the degree of phenotypic plasticity in relation to developmental pathway was much higher in autumn than in spring. Directly developing individuals showed shorter development times and higher growth rates than did diapausing individuals. The pupal and adult weights for both diapausing and directly developing individuals gradually decreased as rearing temperature increased, with the diapausing individuals being slightly heavier than the directly developing individuals at each temperature. Female body weight was slightly lower than male body weight. The proportional weight losses from pupa to adult were almost the same in diapausing individuals and in directly developing individuals, suggesting that diapause did not affect weight loss at metamorphosis. Our results highlight the importance of the expression of alternative developmental pathways, which not only synchronizes this butterfly's development and reproduction with the growth seasons of the host plants but also exhibits the bet-hedging tactic against unpredictable risks due to a dynamic environment.

Effect of rearing conditions on the correlation between larval development time and pupal weight of the rice stem borer, Chilo suppressalis.

A strong positive correlation between development time and body size is commonly assumed. However, the evidence is increasing that the correlation between the two traits can be positive, zero or negative, depending on whether the two traits are under antagonistic or synergistic selection. In the present study, we examined the relation between larval development time and pupal weight of the rice stem borer Chilo suppressalis under laboratory and field conditions. For individuals reared at constant temperatures (22, 25, 28 and 31°C), a longer larval period tended to result in larger pupae, showing a positive correlation between larval development time and pupal weight; whereas for those reared under field conditions, a longer larval period tended to result in smaller pupae at 23.5 and 29.8°C, showing a negative correlation between the two traits. There was no correlation between the two traits at the mean daily temperature of 31°C. At constant temperatures, larval development time shortened significantly as rearing temperature increased, whereas pupae tended to become larger at higher temperatures, although no significant difference was detected among temperatures for pupal weight. Under field conditions, larval development time decreased significantly as the mean daily temperature increased, whereas pupal weight of females increased significantly with the increase in the mean daily temperature, which is an example of the reverse temperature-size rule. Feeding method significantly affected larval development time and pupal weight. For individuals fed on live rice plants, larval development time shortened significantly and pupal weight increased significantly compared with those reared on fresh rice stems.

Life-history responses of the rice stem borer Chilo suppressalis to temperature change: Breaking the temperature-size rule.

Temperature is a key environmental factor for ectotherms and affects a large number of life history traits. In the present study, development time from hatching to pupation and adult eclosion, pupal and adult weights of the rice stem borer, Chilo suppressalis were examined at 22, 25, 28 and 31°C under L18:D 6. Larval and pupal times were significantly decreased with increasing rearing temperature and growth rate was positively correlated with temperature. Larval and pupal developmental times were not significantly different between females and males. The relationship between body weight and rearing temperature in C. suppressalis did not follow the temperature-size rule (TSR), both males and females gained the highest body weight at 31°C. Females were significantly larger than males at all temperatures, showing a female biased sex size dimorphism (SSD). Contrary to Rensch's rule, SSD and body weight in C. suppressalis tended to increase with rising temperature. Male pupae lost significantly more weight at metamorphosis compared to females. We discuss the adaptive significance of the reverse-TSR in the moth's life history.

Variation of life-history traits of the Asian corn borer, Ostrinia furnacalis in relation to temperature and geographical latitude.

Life-history traits from four geographical populations (tropical Ledong population [LD], subtropical Guangzhou [GZ] and Yongxiu populations, and temperate Langfang population [LF]) of the Asian corn borer, Ostrinia furnacalis were investigated at a wide range of temperatures (20-32°C). The larval and pupal times were significantly decreased with increasing rearing temperature, and growth rate was positively correlated with temperature. The relationship between body weight and rearing temperature in O. furnacalis did not follow the temperature-size rule (TSR); all populations exhibited the highest pupal and adult weights at high temperatures or intermediate temperatures. However, development time, growth rate, and body weight did not show a constant latitudinal gradient. Across all populations at each temperature, female were significantly bigger than males, showing a female-biased sexual size dimorphism (SSD). Contrary to Rensch's rule, the SSD tended to increase with rising temperature. The subtropical GZ population exhibited the largest degree of dimorphism while the temperate LF exhibited the smallest. Male pupae lose significantly more weight at metamorphosis compared to females. The proportionate weight losses of different populations were significantly different. Adult longevity was significantly decreased with increasing temperature. Between sexes, all populations exhibit a rather female-biased adult longevity. Finally, we discuss the adaptive significance of higher temperature-inducing high body weight in the moth's life history and why the moth exhibits the reverse TSR.

Photoperiod and temperature effects on the adult eclosion and mating rhythms in Pseudopidorus fasciata (Lepidoptera: Zygaenidae).

Daily distributions of eclosion and mating activities of Pseudopidorus fasciata Walker (Lepidoptera: Zygaenidae) were recorded under natural and various laboratory conditions. Eclosion of this insect exhibited circadian gating in constant darkness (DD) but not in constant light (LL) at 28°C. Under natural conditions, the majority of adults emerged in midmorning with an eclosion peak around 1000 hours. The eclosion distribution was significantly affected by ambient temperature but not by photoperiod under laboratory conditions. Eclosion was more spread out at 22°C than at higher temperatures, and peak eclosion times were advanced at higher temperatures up to 30°C. Under natural and laboratory diurnal cycles, adults of P. fasciata preferred to mate at dusk, within a few hours before the start of the scotophase. Photoperiod and ambient temperature interacted in regulating the mating distribution in P. fasciata. Mating rhythmicity disappeared under DD and LL, under which the insect either mated arrhythmically (DD) or barely mated (LL). Overall, eclosion rhythm in this insect was predominantly regulated by temperature rather than photoperiod, whereas photoperiod appeared to be more influential than temperature in rhythmic gate of mating patterns.

A comparison of the life-history traits between diapause and direct development individuals in the cotton bollworm, Helicoverpa armigera.

In order to understand the differences of life-history traits between diapause and direct development individuals in the cotton bollworm, Helicoverpa armigera (Hübner) (Lepidoptera: Noctuidae), the development time, body size, growth rate, and adult longevity were investigated between the two populations, which were induced under 12:12 L:D and 16:8 L:D photoperiods, respectively, at 20, 22, and 25°C. The results indicated that the larval development time, pupal weight, adult weight, and growth rate were significantly different between diapause and direct developing individuals. The diapause developing individuals had a significantly higher pupal and adult weight and a longer larval time compared with direct developing individuals. However, the growth rate in diapause developing individuals was lower than that in the direct developing individuals. Analysis by GLM showed that larval time, pupal and adult weight, and growth rate were significantly influenced by both temperature and developmental pathway. The pupal and adult weights were greater in males than females in both developmental pathways, exhibiting sexual size dimorphism. The dimorphism in adult weight was more pronounced than in pupal weight because female pupae lost more weight at metamorphosis compared to male pupae. Protogyny was observed in both developmental pathways. However, the protogyny phenomenon was more pronounced at lower temperatures in direct developing individuals, whereas it was more pronounced in diapause developing individuals when they experienced higher temperatures in their larval stage and partial pupal period. The adult longevity of diapause developing individuals was significantly longer than that of direct developing individuals. The results reveal that the life-history strategy was different between diapause and direct developing individuals.

Comparison of sexual compatibility in crosses between the southern and northern populations of the cabbage beetle Colaphellus bowringi.

It is widely accepted that the genetic divergence and reproductive incompatibility between closely related species and/or populations is often viewed as an important step toward speciation. In this study, sexual compatibility in crosses between the southern XS population and the northern TA population of the polyandrous cabbage beetle Colaphellus bowringi was investigated by testing their mating preferences, mating latency, copulation duration, and reproductive performances of post-mating. In choice mating experiments, the percentages of matings were significantly higher in intra-population crosses than in inter-population crosses. Both isolation index (I) and index of pair sexual isolation (IPSI ) indicated partial mating incompatibility or assortative mating in crosses between the two different geographical populations. In single pair mating experiments, XS females in inter-population crosses mated significantly later and copulated significantly shorter than those in intra-population crosses. However, TA females in inter-population crosses mated significantly earlier and copulated longer than those in intra-population crosses, suggesting that larger XS males may enhance heterotypic mating. The lifetime fecundity was highest in XS homotypic matings, lowest in TA homotypic matings, and intermediate in heterotypic matings between their parents. The inter-population crosses resulted in significantly lower egg hatching rate and shorter female longevity than intra-population crosses. These results demonstrated that there exist some incompatibilities in premating, postmating-prezygotic, and postzygotic stages between the southern XS population and northern TA population of the cabbage beetle Colaphellus bowringi.

The influence of female age on male mating preference and reproductive success in cabbage beetle, Colaphellus bowringi.

The influence of female age on male mating preference and reproductive success has been studied using a promiscuous cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae). In a simultaneous choice test, middle-aged females had significantly greater mating success than young and old females. In single pair trials, when paired with middle-aged virgin males, middle-aged females mated faster, copulated longer, and had greater fecundity and fertility than young or old females, while the longevity of males was not significantly affected by female age. This study on C. bowringi suggests that middle-aged females are more receptive to mating, which can result in the highest male reproductive success.

Effect of photoperiod and temperature on the intensity of pupal diapause in the cotton bollworm, Helicoverpa armigera (Lepidoptera: Noctuidae).

The intensity of pupal diapause in the cotton bollworm, Helicoverpa armigera (Hübner) was investigated under both laboratory and natural conditions. By transferring diapausing pupae induced under LD 11:13, LD 12:12 and LD 13:11 at 20, 22 and 25 °C to 25 °C combined with LD 15:9 to terminate diapause the rearing day length of 11 h evoked greater intensity of diapause than did 12 and 13 h at 25 °C; whereas the rearing temperature of 25 °C evoked more intense diapause than did 20 and 22 °C under LD 11:13. By transferring diapausing pupae induced under LD 12:12 at 20 and 22 °C to six temperatures of 18, 20, 22, 25, 28 and 31 °C combined with LD 15:9 to terminate diapause, the duration of diapause was significantly shortened from 146 days at 18 °C to 24 days at 31 °C, showing that high temperatures significantly accelerate diapause development. Furthermore, the duration of diapause was significantly longer at the rearing temperature of 22 °C than that at 20 °C when the diapause-terminating temperatures were 20 and 22 °C. Chilling at 5 °C did not shorten the duration of diapause but lengthened it when chilling period was included. However, chilling plays an important role in synchronizing adult emergence. Rearing temperature of 22 °C also evoked more intense diapause than did 20 °C in most chilling treatments. When the overwintering pupae were transferred at different times from natural temperatures to 25 °C, it was found that the earlier the transfer took place, the earlier the adults emerged when the time spent under natural conditions was included. However, cool temperatures before March showed an enhanced effect on diapause development at 20 °C, suggesting that the high diapause-terminating temperature can offset the effect of chilling on diapause development. The result of diapause termination under natural conditions suggests that the developmental threshold for post-diapause development in H. armigera should be around 17.5 °C.

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. All results demonstrate that H. armigera is not genetically homogeneous throughout its range, but rather is composed of distinct populations genetically adapted to local environmental conditions despite the potential for gene flow via seasonal migration of adults.

Inheritance of photoperiodic control of pupal diapause in the cotton bollworm, Helicoverpa armigera (Hübner).

Pupae of the cotton bollworm, Helicoverpa armigera display a diapause in response to the exposure of their larvae to short photoperiods and relatively low temperatures. Due to geographic variation in photoperiodic response, moths from a northern population, Langfang (39°32'N, 116°41'E), enter diapause in response to short daylengths (D strain) while moths from a southern population, Ledong (18°28'N, 108°53'E), exhibit no diapause under the same conditions (N strain). In the present study, crosses between the two strains are utilized to evaluate the inheritance of diapause under different photoperiods at temperatures 20, 22 and 25°C. The moths in both reciprocal crosses and backcrosses to D strain showed a clear long-day response, similar to that of the D strain, suggesting that the photoperiodic response controlling diapause in this moth is heritable. The incidences of diapause for all F(1) hybrids were intermediate between those of their parents. However, the incidences of diapause at 20°C in F(1) (N×D) strain were significantly higher than those in F(1) (D×N) strain, indicating that the male parent plays a more important role in the determination of diapause. The N strain also showed a short-day photoperiodic response at the lower temperature of 20°C, indicating that the N strain still has the capability to enter a photoperiodically induced diapause, depending on the rearing temperature. Results from all crosses under photoperiods LD 12:12 or LD 13:11 at 22°C showed that inheritance of diapause in H. armigera did not fit an additive hypothesis and that the capacity for diapause was transmitted genetically in the manner of incomplete dominance with non-diapause characteristic partially dominant over the diapausing. Diapause duration in hybrid pupae was also influenced by their inheritance from both parents. Diapause duration in hybrid pupae was intermediate between those of their parents. These results reveal that both diapause induction and duration are under the control of polygene.

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 F(1) 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.

Diapause induction, maintenance and termination in the rice stem borer Chilo suppressalis (Walker).

The rice stem borer, Chilo suppressalis, enters facultative diapause as fully grown larvae in response to short-day conditions during the autumn. Our results showed that the critical night length for diapause induction in C. suppressalis was between 10 h 22 min and 10h 45 min at 22, 25 and 28 degrees C, 11 h 18 min at 31 degrees C, and between 10 h 5 min and 10 h 20 min under field conditions (average temperature ranged from 27.2 to 30.7 degrees C). The diapause incidence declined in ultra-long nights (18-22 h scotophases) and DD, and increased in ultra-short nights (2-6 h scotophases) and LL. Moreover, we found that the third instar was the stage most sensitive to the photoperiod, and night length played an essential role in the initiation of diapause. Night-interruption experiments with a 1-h light pulse at LD 12:12 (light 12:dark 12) exhibited two troughs of diapause inhibition, with one occurring in early scotophase and the other in late scotophase. Field observations for six years showed that most larvae entered winter diapause in August in response to declining day lengths, despite the high temperatures prevailing during August. By periodically transferring the field-collected overwintering larvae to different photoperiods and temperatures, the results showed that photoperiod had a significant influence on diapause development during the early phase of diapause, while high temperature significantly accelerated the termination of larval diapause.

Photoperiodism of diapause induction in Thyrassia penangae (Lepidoptera: Zygaenidae).

Thyrassia penangae enters winter diapause as a prepupa in a cocoon. Photoperiodism of diapause induction was systematically investigated in this moth. The photoperiodic response curves under 24-h light-dark cycles showed that this insect is a typical long-day species. The critical daylength was 13h 30min at 25 degrees C, 13h at 30 degrees C and 12h 20min at 28 degrees C. Transferring experiments from a short day (LD 12:12) to a long day (LD 15:9) or vice versa indicated that photoperiodic sensitivity mainly occurs during the larval period. In experiments using non-24-h light-dark cycles, when the length of photophase exceeded the critical daylength (13.5h), was diapause inhibited effectively, even when the length of scotophase exceeded the critical nightlength (10.5h). Only when a long scotophase was combined with a short photophase, diapause was induced effectively. This result suggests that daylength measurement is more important than nightlength measurement in T. penangae. Night interruption experiments under 24-h light-dark cycles exhibited two points of apparent light sensitivity, but the photosensitive position was highly influenced by temperature and the length of scotophase. Nanda-Hamner experiments failed to reveal the involvement of a circadian system in this photoperiodic time measurement. All light-dark cycles from LD 12:12 to LD 12:72 resulted in a short day response, and all cycles from LD 14:4 to LD 14:72 resulted in a long day response, suggesting that photoperiodic time measurement in this moth is performed by a day-interval timer or an hourglass-like clock.

Diapause induction and clock mechanism in the cabbage butterfly Pieris melete Ménétriés.

Photoperiodic control of diapause induction was systematically investigated in the cabbage butterfly, Pieris melete, which enters summer and winter diapause as a pupa. Summer and winter diapause are induced principally by short and long scotophases, respectively; the intermediate scotophases (11-12 h) permit pupae to develop without diapause. Photoperiodic responses under 24-h light-dark cycles at 16.9, 18, 20 and 22 degrees C showed that the hibernation response was temperature compensated, whereas aestivation response was strongly temperature-dependent. The incidence of diapause for both aestivation and hibernation showed a decline at the ultra-short and ultra-long scotophases. Experiments using non-24-h light-dark cycles showed that the length of the scotophase played an essential role in the determination of diapause. The highest photosensitivity differed under hibernation and aestivation conditions. With a 3 x LD 12:12 interruption, a maximal inhibition of aestivation occurred in the L3/2 stage, and of hibernation it occurred in the L4/0 stage. A long-night of LD 10:14 induced hibernation diapause but inhibited aestivation diapause and, conversely, a short-night of LD 14:10 inhibited hibernation diapause but induced aestivation diapause. With a 1-h light pulse at LD 11:13, a maximal inhibition of hibernation occurred 3 h before lights-on (late scotophase), whereas, with a 1-h light pulse at LD 12.5:11.5, a maximal induction of aestivation occurred 2-3 h after the onset of darkness (early scotophase). Nanda-Hamner and Bünsow experiments failed to reveal the involvement of a circadian system, suggesting that the photoperiodic time measurement for diapause induction in this butterfly resembles an hourglass-like timer or a damped circadian oscillator.