Interaction between photoperiod, temperature, and chilling in dormant larvae of the tree-hole mosquito, Toxorhynchites rutilus Coq.

1. Unchilled, diapausing larvae of Toxorhynchites rutilus rely on photoperiod for the maintenance of diapause. The photoperiodic clock is temperature-compensated between 16.5 degrees and 25 degrees C, maintaining both a similar set-joint and inherent accuracy over this range. The rates of development among larvae terminating diapause are dependent upon both temperature and photoperiod. 2. Chilling of dormant Toxorhynchites rutilus can promote response to progressively shorter daylengths, thus decreasing the critical photoperiod. Chilling can also accelerate response to long days, thereby decreasing the depth of diapause and, after prolonged exposure, can eventually terminate diapause directly, leaving subsequent morphogenesis independent of photoperiod. 3. The optimal temperature for these effects of chilling is above 4 degrees C, below 16.5 degrees C, and may lie around 7 degrees C. 4. Temperatures between 5 degrees and 15 degrees C are vernal and autumnal rather than hibernal. The interaction between chilling and photoperiod may then represent an adaptive compromise between selection due to long-term climatic trends and the vagaries of spring weather.     

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.     

Effect of photoperiod on the development and diapause of the green lacewing Chrysopa pallens (Neuroptera: Chrysopidae)

To investigate the physiology of Chrysopa pallens, the effect of photoperiod on diapause and development was examined in a Japanese population (33.4°N). The response stage for diapause of C. pallens was considered to be the prepupal stage. The critical photoperiod for diapause induction at 20.0°C was between 13 h light : 11 h dark (LD 13:11) and LD 14:10. The larval developmental period was affected by photoperiod: larvae in diapause took longer to complete their development. This difference of larval developmental period in relation to photoperiod was considered to be an adjustment of larval diapause timing.     

Geographic variation in critical photoperiod for diapause induction and its temperature dependence in Hyphantria cunea Drury (Lepidoptera: Arctiidae)

The fall webworm, Hyphantria cunea Drury (Lepidoptera: Arctiidae), was introduced from North America to Japan half a century ago. The critical photoperiod for diapause induction and its temperature dependence, as defined by the difference in the critical photoperiod between 20 and 25°C, were investigated in order to understand the mechanisms behind a shift from bi- to trivoltine life cycles. The critical photoperiod for diapause induction was shorter in the southern trivoltine populations than in the northern bivoltine populations, and this was more marked at 25°C than at 20°C. Although the critical photoperiod showed a positive correlation with the original latitude, the correlation was relatively low at both temperatures. Conversely, temperature dependence of the critical photoperiod for diapause induction correlated negatively with the original latitude. The trivoltine populations showed greater temperature sensitivity than the bivoltine populations. These results suggest that an increase in temperature sensitivity of the diapause response to photoperiods was involved in the shift to a trivoltine life cycle. The crossing experiments suggested that the photoperiodic control of diapause induction and its temperature dependence are under polygenic control without sex-linkage. Received: 29 October 1996 / Accepted: 26 February 1997     

光周期和温度对亚洲玉米螟滞育诱导的影响

亚洲玉米螟Ostrinia furnacalis (Guenée)属兼性滞育昆虫, 其滞育特性与种群利用适生境的最大化和季节性活动密切相关。解析季节性因素如光周期和温度对其滞育的诱导作用可为预测种群发生动态提供科学依据。本实验在HPS 500型和HPG-320H型自动控制环境气候箱中, 测定了RH 80%下温度20℃, 27℃和30℃与11个光周期组合对吉林农安（NA）、 河北衡水（HS）、 广东惠州（HZ）和海南海口（HK）等4个地理种群的滞育诱导作用。结果表明： 光周期、 温度及其交互作用均对亚洲玉米螟滞育诱导具有重要影响, 其中光周期在滞育诱导中起主导作用。在20℃条件下, 其光周期反应曲线属典型的长日照反应型, 临界光周期随种群分布的地理纬度降低而缩短, 4个地理种群NA, HS, HZ和HK的临界日昼长依次为14 h 3 min, 13 h 59 min, 13 h 32 min和13 h 7 min, 最有效的滞育诱导光周期是日昼长12 h。另一方面, 随种群分布纬度升高, 其对光周期诱导滞育的敏感性降低。温度为27℃时, 其光周期反应曲线为短日照 长日照反应型, 各种群有两个临界光周期, 临界日昼长依次为12 h 50 min和13 h 32 min, 11 h 35 min和13 h 8 min, 12 h 58 min和13 h 1 min, 以及11 h 50 min和12 h 26 min, 且最有效的滞育诱导光周期范围的滞育率明显低于20℃。一定温度范围内（20℃～27℃）, 随温度升高, 临界光周期缩短; 温度达到30℃时则滞育显著被抑制, 滞育率仅为4.3%或更低。这些结果说明温度对亚洲玉米螟滞育的光周期诱导具有很强的补偿作用。因此, 在自然界亚洲玉米螟的滞育属具有温度补偿作用的长日照反应型昆虫, 临界光周期随地理纬度北移而增长; 种群滞育的短日照 长日照反应型特性可能是产生局部世代种群的重要内因。     

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.     

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.     

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.     

Role of photoperiod and temperature in the induction of overwintering pupal diapause in the spotted tentiform leafminer,Phyllonorycter blancardella

The role of photoperiod and temperature in the induction of overwintering diapause inPhyllonorycter blancardella (F.) (Lepidoptera: Gracillariidae) was examined in the laboratory and field using leafminers from commercial apple orchards in Ontario, Canada.P. blancardella exhibited a long-day response to photoperiod: long daylengths resulted in uninterrupted development whereas short daylengths induced diapause. The estimated critical photoperiod for diapause induction was L14.25∶D9.75. The larvae of leafminers destined to enter diapause took ca. 3× longer to complete development than the larvae of non-diapausing leafminers. The development prolonging effect of photoperiod decreased with decreasing daylength. Temperature modified the diapause inducing effect of photoperiod. At L14.25∶D9.75, diapause incidence was similar at 15 and 20°C but was lower at 25°C. Photoperiod also altered the normal relationship between development rate and temperature. At L14.25∶D9.75, the duration of larval development of diapausing leafminers was similar at 15, 20 and 25°C. Temperature alone is unlikely to have a role in the induction of diapause because leafminers exposed to natural late summer and fall temperature regimes and L16∶D8 did not enter diapause.     

Some aspects of induction and termination of diapause in a Greek strain of the miteTetranychus cinnabarinus (Boisduval) Boudreaux, 1956 (Acari: Tetranychidae)

A Greek strain of the miteTetranychus cinnabarinus, collected from ivy (Hedera spec.) in Thessaloniki (41 °N), exhibits a facultative, imaginal diapause. Diapause is induced by photoperiod and the photoperiodic response is of the long-day type. The critical daylength is 12.5 h at 19 °C. A period of chilling is not necessary for the termination of diapause under long-day conditions. Diapausing females are sensitive to photoperiod at least during the first 11/2 month of diapause.     

Photoperiodic responses during larval development and diapause of two geographic ecotypes of the rice stem maggot, Chlorops oryzae

Chlorops oryzae is bivoltine in northern Japan but trivoltine in the southern part of the country. In the bivoltine strain, both the egg and larval stages were found to be sensitive to photoperiod. When the egg stage was exposed to a long-day photoperiod (16L:8D), larval development showed a short-day type response, and mature third-instar larvae entered a summer diapause under a long-day photoperiod (15L:9D). When eggs experienced short days, the first-instar larvae entered a winter diapause under short-day conditions, and the critical photoperiod in the larval stage ranged from about 14L:10D to about 12L:12D as the photoperiod experienced by the eggs increased from 12L:12D to 14L:10D. However, the development of the larvae after overwintering was not influenced by the photoperiod. In the trivoltine strain, larval development was retarded under a 14L:10D photoperiod but not under either shorter or longer photoperiods, when larvae had spent the egg stage under a 16L:8D photoperiod. The critical photoperiod of the larval stage for the induction of a winter diapause in the first instar was about 12L:12D, though it varied to some extent with the photoperiod during the egg stage. Thus, Chlorops oryzae was able to adapt itself to the local climatic conditions by the development of variable and complicated photoperiodic responses.     

Diapause induction in the codling moth, Cydia pomonella: effect of prediapause temperatures

The effect of four prediapause temperatures (18, 22, 26 and 30°C) on the photoperiodic response of the codling moth, Cydia pomonella (L.), was studied under controlled conditions. The highest rates of diapause were recorded, for all day-lengths, at temperatures of 22 and 26°C while relatively lower rates of diapause were elicited at 18 and 30°C. The same trend was demonstrated by projecting the values of the critical photoperiod which induces 50% diapause (=CPhP₅₀) over the prediapause temperature. The change in diapause incidence as a function of photoperiod, at all prediapause temperatures, exhibited a response characteristic of long-day insects, i.e. high rates of diapause at short days (12–13.5 h) and a decrease in diapause incidence at long days (14–15 h). The results for temperatures 22, 26 and 30°C support the view that lower prediapause temperatures enhance diapause induction, at a give photoperiod, while higher temperatures tend to avert or diminish the process. On the other hand, the low rates of diapause obtained at 18°C contradict this view. Nevertheless, high correlation was found between the laboratory evidence and field data, indicating the adaptability of the Israeli codling moth to subtropical climate.     

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.     

The role of photoperiod and temperature in determination of summer and winter diapause in the cabbage beetle,Colaphellus bowringi (Coleoptera: Chrysomelidae)

The cabbage beetle, Colaphellus bowringi, is a short-day species undergoing an imaginal summer and winter diapause. Its photoperiodic response highly depends on temperature. All adults entered diapause at ≤ 20 °C regardless of photoperiods. High temperatures strongly weakened the diapause-inducing effects of long daylengths. The diapause-averting influence of short daylengths was expressed only at high temperatures (above 20 °C). This indicates that the beetle has a cryptic ability to reproduce in summer. In fact, summer and winter diapause were induced principally by relatively low temperatures in the field, whereas photoperiod had less influence on diapause induction. The critical daylength for the autumnal population was between 12 h and 13 h. By transferring from a long day to a short day or vice versa at different times after hatching, it was shown that the sensitive stage with regard to photoperiod was the larva, whereas a long day was photoperiodically more potent than a short day. The sensitive stage to temperature encompassed the larval, pupal and adult stages. This different response pattern serves to ensure that the beetle enters summer and winter diapause in time. The selections for non-diapause trait under laboratory (at 25 °C) and natural conditions (at >24 °C) showed that the beetle could lose its sensitivity to photoperiod very rapidly.     

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

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

山楂叶螨滞育的初步研究

本文采用海绵水盘法研究了山楂叶螨Tetranychus viennensis Zacher的滞育与温度、光照和寄主营养等环境因子的关系。结果表明21℃下,临界光周期为12 h 32 min,光敏感螨态为幼螨至第二若螨之间的连续两个螨态,单独一个发育阶段对短光照不敏感或仅有微弱的感应。在相同寄主（梨树）叶片上,病叶和老叶诱导的滞育率比嫩叶高,分别为69.7%、55.3%和20%。相同叶片,短光照(12.5 h/d)条件下随螨密度的增大滞育率升高。相同短光照下,低温促进滞育,在15℃、18℃和21℃下滞育率分别为100%、93.1%和66.7%;高温（24℃）抑制滞育,滞育率仅为13.3%;温度为27℃时不滞育。     

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

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

Environmental and physiological factors for diapause induction and termination in the aphid parasitoid,Aphidius ervi (Hymenoptera: Aphidiidae)

In the aphid parasitoid Aphidius ervi Haliday, the incidence of diapause around the critical photoperiod was dependent on temperature. At a photoperiod of 12 h light:12 h dark, a significantly higher proportion of diapausing individuals was found at 12 degrees C than at 15 degrees C. The second larval stage of the parasitoid was most sensitive to photoperiod and localised-illumination experiments showed that the parasitoid responded independently of the aphid host. The sex ratio of the population which emerged after rearing in short-days and/or after diapause was male biased. Topical application of the juvenile hormone-mimic, pyriproxyfen, before or during diapause delayed emergence of Aphidius ervi but did not induce diapause or influence its termination. The ecdysteroid mimic RH 5849 induced diapause in a few treated individuals but did not affect diapause termination.     

Geographical variation in photoperiodic induction of larval diapause in the bruchid beetle, Bruchidius dorsalis: polymorphism in overwintering stages

The bruchid beetle, Bruchidius dorsalis Fahraeus (Coleoptera: Bruchidae), has a multivoltine life cycle and shows geographical variation of overwintering stages in Japan. Our previous study found that B. dorsalis enters larval diapause in the final instar under short photoperiods. In cooler areas, we observed that most individuals overwinter in the final larval stage in diapause, whereas beetles at different developmental stages (non‐diapausing young instars, diapausing instars, and adults) were overwintering in warmer areas. In this study, we investigated geographical variation in the photoperiodic response for induction of larval diapause at 20 °C (three populations) and 24 °C (two populations) to clarify the overwintering strategy of B. dorsalis. We observed that (1) diapause incidence at 20 °C changed sharply from ca. 100% to 0% with a change in photoperiod in all the populations, (2) critical photoperiod was longer at 20 °C in populations from cooler areas, and (3) critical photoperiod at 24 °C was shorter than at 20 °C and a fraction of the larvae did not enter diapause, even under short photoperiods. Overwintering stages estimated from these results were consistent with those actually observed in the field. This study indicates that the geographical variation of overwintering stages is likely to reflect adaptive diapause induction in each local environment.     

Induction and termination of reproductive diapause in a neotropical beetle, Chelymorpha alternans (Coleoptera: Chrysomelidae)

The presence of a reproductive diapause in the life cycle of the neotropical cassidine beetle, Chelymorpha alternans , was investigated by exposing groups of beetles to conditions differing in photoperiod and humidity. Diapause was characterized by the absence of egg-laying in females up to 70 days after emergence and was induced in response to a short photoperiod of 12:12 h L: D but averted under a longer photoperiod of 13:11 h L:D. High (>90% RH) and low (55–75% RH) humidity conditions did not influence diapause induction. Males appeared to court and attempt to mate with females under all conditions between 16 and 30 days after emergence, but declined after this time in short photoperiods and it is not known if matings in these groups were successful. Adults induced to diapause by exposure to a 12:12 h L:D photoperiod and subsequently exposed to a 13:11 h L: D photoperiod 70 or 72 days after emergence did not show a rapid response by commencing egg-laying. However, all diapausing groups in long and short photoperiod and high and low humidity, in both experiments performed, had commenced egg-laying after 128 days, suggesting an endogenous rhythm for diapause termination. Long photoperiod and high humidity combined may hasten diapause termination since egg-laying began after only 95 days in this group in experiment two. This strategy of induction and termination is discussed with reference to the seasonality of the natural environment of C. alternans in Panama.