Comparison of the circadian eclosion rhythm between non-diapause and diapause pupae in the onion fly,Delia antiqua: the effect of thermoperiod

When non-diapause and diapause pupae of Deliaantiqua were exposed to various thermoperiods where thermophase (T) was 25 °C and the cryophase (C) was 15 or 20 °C (TC₁₅ or TC₂₀) in constant darkness (DD), the majority of both types of flies emerged before the rise in temperature. Eclosion time was delayed at the lower cryophase temperature. Moreover, there was a significant difference in the time of adult eclosion between non-diapause and diapause pupae; diapause pupae eclosed earlier than non-diapause pupae. When the two types of pupae were transferred to a constant low temperature (15 or 20 °C) after having experienced TC₁₅ or TC₂₀ 12:12 h, they showed circadian rhythmicity in eclosion. The free-running period (τ) of the eclosion rhythm changed after transfer to constant low temperatures in both non-diapause and diapause pupae, suggesting that this change represents a transient cycle until the temperature-sensitive oscillator is coupled again to the temperature-insensitive pacemaker. However, diapause pupae tended to show a shorter τ than non-diapause pupae. This observation suggests that the difference in adult eclosion time under thermoperiodic conditions between non-diapause and diapause pupae is related to their different τ s.     

Comparison of the circadian eclosion rhythm between non-diapause and diapause pupae in the onion fly, Delia antiqua: the change of rhythmicity

When pupae of Delia antiqua were transferred to constant darkness (DD) from light-dark (LD) cycles or constant light (LL), the sensitivity to light of the circadian clock controlling eclosion increased with age. The daily rhythm of eclosion appeared in both non-diapause and diapause pupae only when this transfer was made during late pharate adult development. When transferred from LL to DD in the early pupal stage, the adult eclosion was weakly rhythmic in non-diapause pupae but arrhythmic in diapause pupae. However, the sensitivity of the circadian clock to temperature cycles or steps was higher in diapause pupae than in non-diapause pupae; in the transfer to a constant 20 degrees C from a thermoperiod of 25 degrees C (12 h)/20 degrees C (12 h) on day 10 after pupation or from chilling (7.5 degrees C) in DD, the adult eclosion from diapause pupae was rhythmic but that from non-diapause pupae arrhythmic. In a transfer to 20 degrees C from the thermoperiod after the initiation of eclosion, rhythmicity was observed in both types of pupae. The larval stage was insensitive to the effect of LD cycle initiating the eclosion rhythm. In D. antiqua pupae in the soil under natural conditions, therefore, the thermoperiod in the late pupal stage would be the most important 'Zeitgeber' for the determination of eclosion timing.     

Hormonal control of seasonal morph determination in the swallowtail butterfly, Papilio xuthus L. (Lepidoptera: Papilionidae)

Seasonal morphs (spring and sumer forms) of Papilio xuthus L. are determined coincidentally with diapause and non-diapause in pupae by larval exposure to short days and long days respectively. The neuroendocrine principle underlying seasonal-morph determination was studied using surgical operations in P. xuthus.When recipient 0-day old or chilled diapause pupae were joined to donor 0-day old non-diapause pupae, the recipients developed into summer or intermediate morphs. When the same kinds of recipients used above were joined to 0-day old or chilled diapause pupae, there were no significant effects on the adult morph. In contrast, recipient non-diapause pupae all developed into summer morphs, regardless of groups of the type of donors.Furthermore, when the brain of 5th-instar larvae, pharate pupae or pupae, predetermined to be diapause, was transplanted into the abdomen of 0-day old, 30-day old or chilled diapause pupae, the recipients developed into summer or intermediate morphs.The results indicate that the brain of non-diapause pupae secretes a humoral factor producing the summer morph. In non-diapause pupae, the factor may be secreted at about the stage of larval-pupal ecdysis coincidentally with that of prothoracicotropic hormone.     

温度在柑橘凤蝶蛹滞育解除中的作用

调查温度在柑橘凤蝶Papillio xuthus L.滞育蛹解除中的作用。结果显示,羽化温度在15,20,25和30℃时,滞育蛹最早羽化分别在第45,16,11和第5天,发育历期分别为60.3,26.0,15.2和11.6d,羽化持续时间分别为35,19,14和15d。同时低温处理(10℃)显示,25℃时,低温处理0,20和40d后,滞育蛹分别在处理后第38,第14和第13天开始羽化,羽化时间持续分别为40,71和16d,发育历期分别为60.9,28.5和19.0d。结果表明,随着羽化温度升高和低温处理时间延长,滞育蛹羽化时间提前,发育历期缩短。但当羽化温度超过25℃后,滞育蛹发育历期差异已不显著。     

Influences of daylength and temperature on the period of diapause and its ending process in dormant larvae of burnet moths (Lepidoptera,Zygaenidae)

The onset of larval diapause in the burnet moth Zygaena trifolii is clearly characterized by the larva molting into a specialized dormant morph. In a potentially bivoltine Mediterranean population (Marseille) two types of diapause can occur within 1 year: firstly, a facultative summer diapause of 3–10 weeks, and secondly, an obligate winter diapause, which can be lengthened by a period of thermal quiescence to several months in temperatures of 5°C. For the first time, three successive physiological periods have been experimentally distinguished within an insect dormancy (between onset of diapause and molting to the next non-diapause stage), using chilling periods of 30–180 days at 5°C, and varying conditions of photoperiod and temperature. These stages are: (1) a continuous Diapause-ending process (DEP); (2) thermal quiescence (Q); and finally, (3) a period of postdiapause development (PDD) before molting to the next larval instar. The result of transferring dormant larvae from chilling at 5°C to 20°C depended on the length of the chilling period. After chilling for 120–180 days, molting to the next instar occurred after 6–10 days, independent of daylength. This period corresponds with the duration of PDD. After shorter chilling periods (90, 60, 30 days and the control, 0 days) the period to eclosion increased exponentially, and included both the latter part of the previous diapause process and the 6–10 day period of PDD. However, photoperiod also influences the time to eclosion after chilling. Short daylength (8 h light / 16 h dark: LD 8/16) lengthened the diapause in comparison to long daylength (16 h light / 8 h dark: LD 16/8). Short daylength had a similar effect during chilling at 5°C, as measured by the longer time to eclosion after transfer. The shorter time to eclosion resulting from longer chilling periods (30–90 days) demonstrates that the state of diapause is continuously shortened at 5°C, and corresponds to the neuroendocrine controlled DEP. Presumably the DEP has already started after the onset of diapause. When chilling was continued after the end of the DEP, which ranged between 90 and 120 days, thermal quiescence (Q) followed (observed maximum 395 days). Different photoperiodic conditions during the pre-diapause inductive period modified diapause intensity (measured as the duration of diapause), in that a photoperiodic signal just below the critical photoperiod for diapause induction (LD 15/9) intensified diapause. Experiments simulating the summer diapause showed that PDD occurred in the range of 10–25°C. Higher temperatures (15 and 20°C) shortened the DEP at LD 16/8, so that at 20°C many individuals had already terminated diapause after 10–40 days and had molted after the 6–10 days of PDD. A temperature of 25°C unexpectedly lengthened the DEP to 110 days in several individuals. The ecological consequences and the adaptive significance of variation in the duration of the diapause are discussed in relation to the persistence of local populations predictably variable and rare climatic extremes throughout the year.     

Influence of photoperiod on low temperature acclimation for cold-hardiness in Drosophila auraria

ABSTRACT. Imagines of Drosophila auraria Peng, a reproductive diapause species, developed cold-hardiness at low temperatures to a greater extent when exposed to a diapause-inducing photoperiod (LD10:14 h) than when exposed to a diapause-preventing photoperiod (LD 16:8h). Imagines kept at 18°C, which was the temperature at which they were reared to eclosion, did not survive a test exposure to -5°C for 8 days regardless of age or photoperiod. When transferred to 10 or 5°C, either from eclosion or from 8 days after eclosion, the survival rate, on testing, rose with time since transfer and rose faster and higher with a photoperiod of LD 10:14h than with LD16:8h. Flies transferred to 15°C only showed improved ability to survive the test if they were kept in LD 10:14h. When cultured at 18°C to the age of 8 days after eclosion, diapause was terminated in about 30% of females even at LD 10:14h. In these post-diapause females the ability to develop cold-hardiness at lower temperatures was somewhat less than in the diapausing females, but apparently greater than in the non-diapause females. These results suggest that the physiological mechanism which promotes cold-hardiness under a diapause-inducing photoperiod is not directly linked to the process causing reproductive diapause.
In Sapporo, flies from a natural population became tolerant to cold in October when they entered diapause and daily mean temperature fell below 15°C and the light/dark cycle fell below LD 12:12h.     

Effects of photoperiod and temperature on diapause induction and termination in the swallowtail,Sericinus montelus

Abstract Sericinus montelus overwinters as diapausing pupae. In the present study, the effects of photoperiod and temperature on diapause induction and termination of diapause are investigated. The results obtained demonstrate that high temperature can reverse the effect of short day‐lengths on diapause induction. Under an LD 12 : 12 h photoperiod, all pupae enter diapause at 15, 20 and 25 °C, whereas all pupae develop without diapause at 35 °C. No pupae enter diapause under an LD 14 : 10 h photoperiod when the temperature is above 20 °C. Photoperiodic response curves obtained at 25 and 30 °C indicate that S. montelus is a long‐day species and the critical day‐length is approximately 13 h at 25 °C. At 25 °C, the duration of diapause is shortest when the diapausing pupae are maintained under an LD 16 : 8 h photoperiod and increases under LD 14 : 10 h and LD 12 : 12 h photoperiods. Under an LD 16 : 8 h photoperiod, the duration of diapause is shortest when the diapausing pupae are maintained at 25 °C, followed by 20 and 30 °C, and then at 15 °C. These results suggest that a moderate temperature favours diapause development under a diapause‐averting photoperiod in this species. The duration of diapause induced by an LD 12 : 12 h photoperiod is significantly longer at 25 °C than those at 15, 20 and 30 °C, and is shortest at 15 °C. At 25 °C, the duration of diapause induced by LD 6 : 18, LD 12 : 12 and LD 13 : 11 h photoperiods is similar and longer than 90 days. Thus, the diapause‐inducing conditions may affect diapause intensity and a photoperiod close to the critical day‐length has significant influence on diapause intensity in S. montelus.     

Diapause and development in the tobacco budworm,Heliothis virescens: A comparison of haemolymph ecdysteroid titres

The signal to induce diapause in H. virescens comes early in development (prior to the third instar in most insects), but the signal to break diapause can come shortly after entrance into diapause at pupation. Haemolymph ecdysteroid titres in both diapause-bound and non-diapause-bound Heliothis virescens larvae were similar in the first two thirds of the last-larval instar, when similar changes in morphology and behaviour occurred. However, the number of stepwise increases in titre and the timing of the steps was different in the two groups of larvae. Haemolymph ecdysteroid titres in the last third of the instar were approx, five times higher in non-diapause than in diapause-bound larvae. In diapausing pupae, haemolymph ecdysteroid titres dropped to levels found in larvae which had completed two thirds of the last instar. When diapausing pupae were warmed to break diapause, haemolymph ecdysteroid titres rose again. However, 2 of the 4 high ecdysteroid levels detected in pupae developing after diapause break were considerably lower than those detected for non-diapause pupae.     

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

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

Physiological aspects of diapause and cold tolerance during overwintering in Pieris brassicae

Abstract The relationship between diapause-associated metabolic suppression and carbohydrate metabolism linked with cold tolerance was investigated in pupae of Pieris brassicae L. Cold tolerance was assessed by measuring the crystallization temperature (T_c) and by estimates of pre-freeze mortality. Metabolic suppression was measured using ³¹P nmr and carbohydrates by GLC. Sorbitol (a possible cryoprotectant) accumulated from the onset of diapause in October until December reaching a concentration of c. 40 mMolal in both years of the study, but then declined from January until adult eclosion in May. The T_c remained between -23 and -25^oC throughout the winter except for a slight rise before eclosion in May. The absence of a significant T_c suppression is as predicted from the low concentration of sorbitol accumulated. The pre-freeze mortality experiments indicate that pupae are most cold tolerant in the period December-February when sorbitol concentration is high, suggesting an alternative cryoprotective role for sorbitol. Glycogen declined at the beginning of diapause until February after which there was some recovery suggesting that it may be the source of carbon for sorbitol synthesis. Diapause-associated metabolic suppression is evident in the low ³¹P nmr resonances of ATP during November-February compared with non-diapause pupae and diapause pupae soon after pupation. The suppression of metabolism at this time may have a direct role in cryoprotection and by itself (rather than sorbitol) account for the increased pre-freeze cold tolerance. ATP appears to increase slowly from February until a sharp increase occurs shortly before eclosion. Arginine phosphate remains high during diapause until late February-March when it begins a decline which continues until eclosion. A period of change in energy and carbohydrate metabolism is apparent at the same time which may indicate diapause termination and related changes in cold tolerance mechanisms. It is argued that in P.brassicae sorbitol accumulates as a result of metabolic suppression and may have no cryoprotective role. However, for species living in, or colonizing, low temperature environments it is a short evolutionary step to exploit this pathway and accumulate high concentrations of polyols as a specialized cold tolerance strategy.     

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

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

LIGHT AT NIGHT ALTERS THE PARAMETERS OF THE ECLOSION RHYTHM IN A TROPICAL FRUIT FLY,DROSOPHILA JAMBULINA

We investigated the effects of natural light at night (LAN) in the field and artificial LAN in the laboratory on the circadian rhythm of pupal eclosion in a tropical wild type strain of Drosophila jambulina captured at Galle, Sri Lanka (6.1^oN, 80.2^oE). The influence of natural LAN, varying in intensity from 0.004 lux (starlight intensity) to 0.45 lux (moonlight intensity), on the entrainment pattern of the circadian rhythm of eclosion at 25^o?±?0.5^oC was examined by subjecting the mixed-aged pupae to natural cycles of light and darkness at the breeding site of this strain in the field. The eclosion peak was ～2?h prior to sunrise, and the 24?h rhythmicity was the most robust. Effects of artificial LAN at 25^o?±?0.5^oC were determined in the laboratory by subjecting pupae to LD 12:12 cycles in which the light intensity of the photophase was 500 lux in all LD cycles, while that of the scotophase was either 0 lux (complete darkness, DD), 0.5, 5, or 50 lux. In the 0 lux LAN condition (i.e., the control experiment), the eclosion peak was ～2?h after lights-on, and the 24?h eclosion rhythm was not as strong as in the 0.5 lux LAN condition. The entrainment pattern in 0.5 lux LAN was strikingly similar to that in the field, as the 0.5 lux LAN condition is comparable to the full moonlight intensity in the tropics. LAN at 0.5 lux dramatically altered both parameters of entrainment, as the eclosion peak was advanced by ～4?h and the 24?h eclosion rhythm was better than that of the control experiment. LAN at 5 lux, however, resulted in a weak eclosion rhythm that peaked in the subjective forenoon. Interestingly, the 50 lux LAN condition rendered the eclosion events unambiguously arrhythmic. After-effects of LAN on the period (τ) of the free-running rhythm and the nature of eclosion rhythm were also determined in DD by a single LD 12:12 to DD transfer. After-effects of the LAN intensity were observed on both the τ and nature of the eclosion rhythm in all four experiments. Pupae raised in 0.5 lux LAN exhibited the shortest τ (20.6?±?0.2?h, N?=?11 for this and subsequent values) and the most robust rhythm, while pupae raised in 50 lux LAN had the longest τ (29.5?±?0.2?h) and weakest rhythm in DD. Thus, these results demonstrate the intensity of LAN, varying from 0 to 50 lux, profoundly influences the parameters of entrainment as well as free-running rhythmicity of D. jambulina. Moreover, the observed arrhythmicity in LD 12:12 cycles caused by the 50 lux LAN condition appeared to be the masking effect of relatively bright light at night, as the LD 12:12 to DD transfer restored the rhythmicity, although it was rather weak. (Author correspondence: drdsjoshi08@gmail.com)     

Ontogenetic patterns of cold-hardiness and glycerol production in Sarcophaga crassipalpis

Developmental patterns of low-temperature tolerance and glycerol production were determined for larval, pupal and adult stages of the flesh fly Sarcophaga crassipalpis Macquart (Diptera: Sarcophagidae). Both diapause and non-diapause-destined flies were reared at relatively high temperatures, 20° or 25°C, prior to testing. Cold tolerance was greatest for diapause pupae aged 12–35 days after pupariation. Among non-diapause-destined flies, pupae exhibited a greater level of low temperature tolerance than larvae or adults. Although diapause pupae were more tolerant than non-diapause pupae maximal cold tolerance was not attained in either group until 10 days after pupariation. Non-diapause-destined feeding and wandering larvae had higher glycerol levels than larvae destined for diapause. During the first 6 weeks after pupariation glycerol titres increased steadily in diapause pupae. Rapid loss of glycerol is associated with the termination of pupal diapause.     

Hormonal control of seasonal morphs by the timing of ecdysteroid release in Araschnia levana L. (Nymphalidae: Lepidoptera)

Araschnia levana L. occurs in two seasonal morphs. Larvae reared under short-day conditions become diapause pupae and emerge as red spring-morph butterflies. Long-day larvae become non-diapause pupae, which emerge as black and white summer morphs. Pupae reared under these different conditions were joined in parabiosis. Both underwent adult development without diapause and the long-day animals developed into the summer morph as normal. The morph of short-day animals depended on the time of parabiosis. When they were joined to fresh long-day pupae 1 day after their own pupation, summer morphs resulted. When parabiosis began 4 days after pupation or later, spring morphs resulted. Extirpation of the brain-corpora cardiaca-allata complex from long-day pupae affected neither non-diapause development nor the summer morph. Adult development could be prevented by removal of head and prothorax. When adult development was initiated in the remaining bodies by 20-hydroxyecdysone 14 days after pupation, the spring morph resulted.—Injection of 20-dydroxyecdysone into 3-day-old short-day pupae initiated adult development and led to the summer morph. Injections into 10-day-old short-day pupae led to the spring morph. The same was true in diapause pupae deprived of their brain-corpora cardiaca-allata complex. These results indicate that seasonal diphenism in A. levana is controlled only by the timing of ecdysteroid release, which initiates adult development. There is no direct influence of the brain on wing coloration.     

棉铃虫蛹期在极端湿度下的失水动态

研究了极端相对湿度 (0%、9%、22.5%、80%、90%和100%) 对棉铃虫Helicoverpa armigera蛹期发育、存活和水分动态的影响。发育蛹在25℃下,相对湿度≤9%时, 不能羽化;湿度为22.5%时,羽化率不足20%; 高湿不影响它们的存活。在同样温度下,湿度≥9%时,滞育蛹在一个月内都极少死亡;在此期间,滞育终止率随湿度降低而升高。各湿度处理组发育蛹和滞育蛹从1日龄起的累计失水率都与其日龄呈线性相关。三个低湿处理组发育蛹中死亡个体在死亡前的平均累计失水率都在32%以上。滞育蛹经0%湿度处理一个月,平均仅失水22.4%;在湿度≥90%时的同期失水率不超过3.6%。在30℃下,发育蛹在4 h内测定的表皮渗透力最大,分别是9.0(♀)和10.7(♂) μg/(cm²·h·mm Hg); 滞育蛹的相应值出现在2 h内, 分别为 4.7(♀)和5.4(♂) μg/(cm²·h·mm Hg)。     

Interacting effect of thermoperiod and photoperiod on the eclosion rhythm in the onion fly, Delia antiqua supports the two-oscillator model

Daily light and temperature cycles entrain adult eclosion rhythms in many insect species, but little is known about their interaction. We studied this problem in the onion fly, Delia antiqua. Pupae were subjected to various combinations of a photoperiod of 12L:12D and thermoperiods. The thermoperiods consisted of 12 h warm phase (W) and 12 h cool phase (C), giving a mean temperature of 25 °C with different temperature steps of 8, 4 and 1 °C. As the phase relation of the two Zeitgebers was varied, the phase of eclosion rhythm was shifted, depending on the phase angle with the light cycle and the amplitude of the temperature cycle. When the temperature step in the thermoperiod was 8 °C (WC 29:21 °C), the eclosion rhythm was entrained mainly to thermoperiod rather than photoperiod. In the regime with a 4 °C temperature step (WC 27:23 °C), both thermoperiod and photoperiod affected eclosion rhythm, and a phase jump of the eclosion rhythm occurred when the warm phase of thermoperiod was delayed 15-18 h from light-on. In regimes with a 1 °C temperature step (WC 25.5:24.5 °C), the eclosion rhythm was completely entrained to photoperiod. The observed interacting effect of light and temperature cycle on the eclosion rhythm in D. antiqua can be explained by the two-oscillator model proposed by Pittendrigh and Bruce (1959).     

The adult eclosion rhythm in Hyphantria cunea (Lepidoptera: Arctiidae): Endogenous and exogenous light effects

Abstract

Endogenous and exogenous effects of light on adult eclosion in Hyphantria cunea were tested by exposure to various light regimes. Regression analysis showed that the position of the eclosion peak after lights on was proportional to the length of the photophase, and that the peak was influenced by the timing of both lights‐on and lights‐off. Under photoperiods of 2–12 hours LD cycle, the eclosion peak was situated after lights‐off, but moved into the light phase as the photophase increased to 22 hours. Pupae were exposed to 3 “skeleton”; photoperiods of LDLD2:2:6:14, 4:2:4:14 and 6:2:2:14. Under the first of these, most adults emerged at the start or just before the longest dark period. Under the second and third skeleton regimes, 20% and 70% respectively of pupae emerged during the shorter dark period. When the compound eyes of the pharate adults were covered, adults smerged 1–4 hours before lights‐off under LD10:14, compared to a control group which emerged just after lights‐off. When pupae were transferred from LD to LL or DD conditions, the eclosion peak occurred approximately every 24 hours after the last LD peak. Results suggest that light received by the compound eyes influences the eclosion rhythm, either through an exogenous masking effect, or by altering the phase of the pacemaker controlling eclosion.     

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.     

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.     

Effects of transgenic overexpression of diapause hormone and diapause hormone receptor genes on non-diapause silkworm

Diapause is a state of developmental arrest that is most often observed in arthropods, especially insects. The domesticated silkworm, Bombyx mori, is a typical insect that enters diapause at an early embryonic stage. Previous studies have revealed that the diapause hormone (DH) signaling molecules, especially the core members DH and DH receptor 1 (DHR1), are crucial for the determination of embryonic diapause in diapause silkworm strains. However, whether they function in non-diapause silkworm strains remains largely unknown. Here, we generated two transgenic lines overexpressing DH or DHR1 genes in a non-diapause silkworm strain, Nistari. Our results showed that developmental expression patterns of DH and DHR1 are quite similar in transgenic silkworms: both genes are highly expressed in the mid to late stages of pupae and are most highly expressed in day-6 pupae but are expressed at very low levels in other developmental stages. Moreover, the overexpression of DH or DHR1 can affect the expression of diapause-related genes but is not sufficient to induce embryonic diapause in their offspring. This study provides new insights into the function of DH and DHR1 in a non-diapause silkworm strain.