Diapause termination, post-diapause development and reproduction in the beet webworm, Loxostege sticticalis (Lepidoptera: Pyralidae)

Effects of photoperiod and cold exposure on diapause termination, post-diapause development and reproduction in Loxostege sticticalis were examined. Larvae were reared at diapause inducing condition (22 °C, L:D 12:12) consistently or transferred to long day photoperiod (L:D 16:8) and darkness (L:D 0:24) respectively, after entering into diapause. Diapause was terminated in approximately 40% of the larvae after 36 days, and no significant differences were observed between photoperiods, suggesting larval diapause was terminated spontaneously without being induced by photoperiods. Cold exposure significantly hastened diapause termination. The diapause termination incidence increased significantly with peaks of 98% at both 5 °C and 0 °C exposure for 30 days, as compared to 42% in controls not exposed to cold, while the mortality and number of days required for diapause termination decreased dramatically. The optimal low temperature exposure periods under 5 °C or 0 °C were 20 days and 30 days, showing a higher termination incidence and shorter time for diapause termination. This suggests that the low temperatures in winter play an important role in diapause termination under natural conditions. The threshold temperatures for post-diapause development in prepupae and pupae were 9.13 °C and 10.60 °C respectively, with corresponding accumulations of 125 and 200 degree-days. Adults that experienced larval diapause significantly delayed their first oviposition, oviposition period was prolonged, and the lifetime number of eggs laid was decreased, however both males and females have significantly longer longevity. The field validation of diapause termination, the degree-days model, and the relationship between diapause and migration in L. sticticalis were also discussed.     

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.     

Induction and termination of diapause inOrius predatory bugs

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

光周期和温度对大猿叶虫滞育后成虫生物学特性的影响

大猿叶虫Colaphellus bowringiBaly是我国十字花科蔬菜上的一种重要害虫,以成虫在土壤中越冬和越夏,滞育发生主要受温度和光周期调节。本文就光周期和温度对滞育后成虫生物学特性的影响进行了研究。结果表明,在25℃下,光周期（L14∶D10和L12∶D12）对成虫滞育后的存活率、寿命、日平均产卵量、总产卵量均无显著影响。在长光照（L14∶D10）下,温度18℃、22℃和25℃）对滞育后成虫的生物学特性具有明显的影响;随温度的降低,总产卵量呈下降趋势,平均产卵量显著降低,雌虫滞育后寿命显著延长,18℃下少数个体能被诱导再次滞育。     

Acceptance and suitability of four plant substrates for rearing Orius sauteri (Hemiptera: Anthocoridae)

Orius sauteri (Poppius) is an important hemipterous predator that has been mass-reared for biological control of numerous pests in protected crop-production systems. To find a good oviposition substrate for mass-rearing this predator under insectary conditions [25°C, 65 ± 5% relative humidity, and a photoperiod of 16:8 (L:D) h], we compared kidney bean, soybean, broad bean sprouts and fresh leaves of kidney bean. We found that O. sauteri made more punctures and laid more eggs in kidney bean sprouts than in the other substrates examined. However, there were no significant differences among substrates in the proportion of punctures receiving eggs. Female O. sauteri laid the most eggs (as many as 68 eggs) in kidney bean sprouts and also had the shortest pre-oviposition period on this plant material. In addition, there were no significant differences in total oviposition durations or female longevity among the four plant substrates. The hatch rates of nymphs in the sprouts and leaves of kidney bean (>90%) were higher than those in soybean and broad bean sprouts. Thus, we found that the kidney bean sprout was the most suitable substrate for mass-rearing of O. sauteri.     

光照时间对双尾新小绥螨生长发育及种群参数的影响

【目的】为了进一步明确光照时间对双尾新小绥螨Neoseiulus bicaudus Wainstein生长发育的影响。【方法】在实验室恒温条件下,采用生命表研究方法,研究在不同光照时间下,双尾新小绥螨捕食土耳其斯坦叶螨的生长发育情况。【结果】双尾新小绥螨在不同光照时间段下均能完成世代周期。在光照时数小于12 h时,从卵发育至成螨的各个发育阶段所需时间均呈缩短趋势,当光照时数大于16 h后各个发育阶段所需的时间又呈增长趋势。产卵期在12L︰12D光照时数下最长为20.74 d,且同其他光照条件均存在显著性差异(P<0.05)。产卵后期、寿命、总产卵量均表现在16L︰8D光照时数下最长,分别为11.93 d、31.05 d和42.4粒。双尾新小绥螨种群的净增值率(R_0)在16L︰8D时达到最大值33.88,在8L︰16D时最低为14.97。【结论】光照时间在12~16 h范围内最适合双尾新小绥螨生长发育。     

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.     

Thermoperiodic response and effect of photoperiod on thermoperiodic induction of diapause in Colaphellus bowringi

The effects of thermoperiods on diapause induction under continuous darkness (DD), continuous light (LL), and an L12:D12 photoperiod were investigated in the cabbage beetle, Colaphellus bowringi Baly (Coleoptera: Chrysomelidae), a short‐day species. Diapause could be induced by thermoperiod under both LL and DD; however, in the range of 24–30 °C, lower incidences of diapause were observed under LL than under DD. The critical cryophase was found to be dependent on the mean temperature of the thermoperiod applied. Although the thermoperiodic response pattern was similar under LL and DD, the incidence of diapause was higher under LL when the duration of the cryophase did not exceed 12 h. In contrast, when the duration of the cryophase was longer than 12 h, the incidence of diapause under LL was lower or equal to that under DD. When a thermoperiod of 24 °C (cryophase) and 28 °C (thermophase) was applied, the incidence of diapause was higher under LL than under DD, regardless of the duration of the cryophase. Thermoperiodic responses under a photoperiod of L12:D12 and under DD further revealed that induction of diapause was strongly influenced by the photophase temperature. Moreover, the incidence of diapause was lower when the thermophase coincided with the photophase than when the cryophase coincided with the photophase.     

The effect of photoperiod on walking, feeding, and oviposition in the western flower thrips

The western flower thrips, Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae), is a major pest in glasshouses, where light and photoperiod are often manipulated to grow crops. The effect of five photoperiods (L0:D24, L6:D18, L12:D12, L18:D6, and L24:D0) on the behaviour of adult females was investigated in the laboratory at 25 °C. The amount of walking, pollen consumption, and oviposition all increased with increasing photophase. The three behaviours occurred at a low rate in continuous darkness, and are therefore not exclusively diurnal. Three out of 16 thrips (19%) showed weakly rhythmic walking activity in continuous darkness, with an average circadian period of 25.3 ± 1.5 h. It is proposed that the length of the photophase determines the amount of nutrient intake, which in turn determines oviposition rate and thus affects population increase.     

Effect of the photoperiod of the site of oviposition of Aedes mariae (Diptera,Culicidae)]

The overwintering biology of Aedes (Ochlerotatus) mariae (one of the sibling species of the mariae complex) was studied in populations from the Tyrrhenian coast of Central Italy (Sperlonga and Formia). Ae. mariae has in this zone several generations per year and its larvae are commonly found from March to October exclusively in rock pools along the coast. Field and laboratory observations show in these populations an embryonic winter diapause induced by short-day photoperiods (9-12 hours). The photoperiod acts primarily on the parental females during their preimaginal life. The expression of diapause is influenced by temperatures and photoperiods experienced by the embryos. Full embryonic diapause is observed when the eggs are incubated at relatively low temperatures (less than 16 degrees C) and at short photoperiods. Short-day photoperiods also induce a remarkable change in the oviposition behavior of Ae. mariae. Laboratory observations show that the adult females readily oviposit on water surfaces when originating from larvae reared at long-day photoperiod while they are very reclutant to oviposit in the same situation when reared at short day photoperiod. Choice experiments involving four alternative oviposition sites (see Plate 1) demonstrate a preference for outside free water in long-day mosquitoes and for inside moist surfaces in short-day mosquitoes. The behavioral difference persists in successive gonotrophic cycles and it is not apparently affected by the photoperiod acting on the adult females. The above evidence together with recovery of overwintering eggs in holes and crevices of rock pools suggest that the shift in oviposition site shown in the laboratory reflects a similar shift occurring in nature. Such photoperiodically induced change in oviposition behavior seems to have an important adaptive significance in providing more constant microclimates to the diapausing eggs and in protecting them from the mechanical action of winter storms.     

Diapause induction in the thrips predatorAmblyseius cucumeris [Acarina: phytoseiidae] under greenhouse conditions

The predatory miteAmblyseius cucumeris (Oudemans) is an important biological control agent for thrips in commercial greenhouses, but its effectiveness in fall and winter is limited by reproductive diapause induced under short day conditions. Influence of photoperiod and temperature on diapause induction was investigated to provide information for successful management of the predator. Under 8∶16 (L∶D) photoperiods and 22°C photophase temperatures, diapause incidence was inversely related to scotophase temperature, decreasing from 100% diapause at 15°C to no diapause at 21°C. In continuous darkness, diapause was induced by thermoperiods of 20∶10 and 22∶17 and 22∶17 but not 25∶15°C (T∶C) (8h thermophase). Critical daylength for inducing diapause under 22∶17°C (T∶C) was 12.45 h, which was consistent with the trend in diapause incidence in mites collected from an established greenhouse population September to November. MostA. cucumeris diapaused only when exposed to diapause inducing conditions throughout their juvenile development and none stopped laying eggs when transferred from nondiapause to diapause inducing conditions as adults. After 14 generations of genetic selection for a nondiapause strain, diapause incidence was 33.3%.      

Egg production in response to combined alternative foods by the predator Coccinella transversalis

The peformance of the parasitoid Anagyrus kamali Moursi [Hymenoptera: Encyrtidae], as a function of host density, temperature, and photoperiod was investigated with the objective to optimize a mass-rearing system in the context of a biological control program. The number of hosts parasitized at densities varying from 2–100 hibiscus mealybug (HMB), Maconellicoccus hirsutus Green [Homoptera: Pseudococcidae], corresponded to a type II-III functional response in fixed-time conditions and a type III in variable-time conditions. Twenty-six percent of the oviposited eggs led to progeny emergence with a sex ratio of 0.49±0.102 (M/F), regardless of host density. Fecundity and oviposition period under six abiotic combinations (i.e., two temperatures (26±2 °C and 32±2 °C) and three photoperiods (L0:D24, L12:D12, L24:D0)) were measured. Lifetime fecundity and reproductive life were significantly affected by temperature and photoperiod conditions. Optimum female parasitoid lifetime fecundity was attained at 26±2 °C, L0:D24 with an average of 116.1±17.43 eggs. At 32±2 °C, L24:D0 and L12:D12, an average of 79.4±34.57 and 85.8±35.81 eggs were laid, respectively. Reproductive longevity was maximal at 26±2 °C, L0:D24 with 12±4.85 days of oviposition. Because the parasite A. kamali can be reared optimally without light, this may save tremendous energy costs.     

温度和光周期对绿盲蝽滞育诱导的影响

为了阐明环境因子对绿盲蝽Apolygus lucorum Meyer-Dür卵滞育诱导作用,测定了3个温度和6个光周期组合处理对绿肓蝽的滞育诱导和绿盲蝽光周期感应的敏感虫态,系统调查了绿肓蝽在不同温度和不同光照组合下所产卵的孵化率.结果表明:绿盲蝽的敏感虫态为1龄若虫;在17℃,20℃和23℃3个不同温度下,光照时间小...     

Diapause in the mite Metaseiulus occidentalis: Stages sensitive to photoperiodic induction

Stages of Metaseiulus occidentalis sensitive to photoperiod induction of diapause were determined by transferring various stadia into diapause-inducing conditions, and rearing them until adult females could be scored for reproductive condition. When eggs were transferred to 10 hr light at 19°C from 24 hr light at 25°C and the mites reared to adults, 92 per cent entered diapause. When larvae and all subsequent stages were kept under the inductive conditions, 62 per cent of adult females diapaused. Mites transferred as protonymphs into inductive conditions yielded only 10 per cent in diapause, and mites transferred as deutonymphs or newly emerged females did not enter diapause.However, adult females reared from eggs at 19°C under 12 hr light (which is near the critical photophase of 11·2 hr at 19°C) showed an unexpected sensitivity to photoperiod. Some newly emerged females oviposited upon transfer to an 8 hr photophase at 19°C. Some then stopped ovipositing and apparently entered diapause; these females resumed ovipositing after intervals ranging from 34 to 100 days. This was termed ‘switching’ into diapause. Some females reared under a 16 hr photophase at 19°C ‘switched’ also upon transfer as adults to shorter photophases—either 8 or 12 hr at 19°C. Thus, ‘switching’ may be due to transfer to shorter photophases. Promptness of mating vs delayed mating allowed ‘switching’ to be more easily detected.     

Effects of photoperiod and temperature on reproductive diapause in Ophraella communa (Coleoptera: Chrysomelidae), a potential biocontrol agent against Ambrosia artemisiifolia

Abstract To investigate the seasonal adaptation strategies of Ophraella communa to new habitats, the effects and regulation mechanisms of photoperiod and temperature on the reproductive diapause in a population collected from Changsha, Hunan were examined. Adults showed obvious reproductive diapause, which was regulated by photoperiod and temperature. At 30°C, there was no adult diapause occurring under either long‐day or short‐day conditions; at 25°C the pre‐oviposition period was short and fecundity was high in adult females under L : D 16 : 8 h, whereas under L : D 12 : 12 h, a few females entered reproductive diapause; at 20°C under short‐day conditions, all female adults entered diapause. The pre‐oviposition period was significantly prolonged when the pupae and adults were transferred from long‐days to short‐days, but the day length influence was not obvious when they were transferred only in the adult stage. However, the fecundity dropped greatly no matter whether the photoperiod shifted to short‐days only in the adult stage or whether the shift occurred in both the pupal and adult stage. The fecundity was extremely low when photoperiod shifted from long‐days to short‐days in both pupal and adult stages. This was an indication that the pupal and adult stages were the photoperiod‐sensitive stage for adult reproductive diapause. This was especially true for the photoperiod in the pupal stage, which has a distinctly significant regulative effect on reproductive diapause. Additionally, this article also addresses the reason for different photoperiodic response patterns in reproductive diapause induction between the Changsha strain and the Tsukuba strain (Japan) of O. communa.     

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.     

Diapause decision in the small tortoiseshell butterfly,Aglais urticae

Insects in temperate areas spend the inhospitable winter conditions in a resting stage known as diapause. In species that diapause in the larval or pupal stage, the decision whether to diapause or develop directly is customarily taken during the late instars, with long days (i.e., long light phases) and high temperatures promoting direct development. Among butterflies that overwinter as adults, data are rare and variable, but imply that the larval daylength conditions can affect the pathway decision. We studied the small tortoiseshell, Aglais urticae L. (Lepidoptera: Nymphalidae, Nymphalini), which is partially bivoltine from Central Scandinavia and southwards, and tested whether the pathway decision is taken in the larval or adult stage. We reared larvae under long‐day (L22:D2) or short‐day (L12:D12) photoperiods, and recorded the pathway taken by the eclosing adults by scoring their propensity to mate and produce eggs. We also tested whether the larval photoperiod influenced adult ability to diapause by assessing adult survival. The results clearly indicate that (1) there is no detectable effect of larval photoperiod treatment on the pathway decision taken by adults whether to enter diapause or to develop directly, (2) some individuals are obligately univoltine and insensitive to photoperiod during adulthood, whereas (3) other individuals can facultatively enter diapause or direct development, depending on the photoperiod experienced after adult eclosion.     

Effects of photoperiod and temperature on pupal diapause induction of grape berry moth Lobesia botrana

Abstract. The grape berry moth, Lobesia botrana Denn. & Sciff. (Lepidoptera: Tortricidae), one of the most injurious pest of grape berries in Greece, is a multivoltine species that overwinters as diapausing pupae. The effects of several diel and non-diel photoperiods and of temperature, experienced by eggs and larvae, on pupal diapause induction were investigated. The diapause response curve was of Type I (long day type) and the determining factor was the duration of scotophase (> 11 h), regardless of the duration of photophase. However, at very short (< 4 h) photoperiods, the incidence of diapause was also high. Diapause was positively and significantly correlated with the egg-larval developmental time, pupal mortality and the duration of the pupal stage. Eggs and larvae reared under LD 12 : 12 h photoperiod and various temperatures (from 12 to 30 °C) produced diapausing pupae (almost 100%), but the duration of the pupal stage (intensity of diapause) increased with increasing temperature. Under continuous darkness, however, the percentage diapause decreased with increasing temperature. Single and double 1-h light pulses were applied systematically at various times during the scotophase of six diapause-inducing diel photoperiods. Two photosensitive points in time (called A and B) were revealed, during which illumination resulted in a significant decrease of diapause induction. The decrease was much greater during the first sensitive period (early in scotophase) rather than in the second (late in the scotophase).     

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

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

Summer dormancy ensures univoltinism in the predatory bug Picromerus bidens

The seasonal cycle of Picromerus bidens L. (Heteroptera: Pentatomidae) is usually considered to be univoltine with an obligatory winter egg diapause. Seasonal adaptations of the species were studied in the laboratory and in field experiments. When reared under short-day photoperiodic conditions (L12:D12 and L14:D10), all females began to lay eggs synchronously soon after their emergence. However, in the females reared under long-day conditions (L18:D6 and L20:D4) and outdoors in June–July, oviposition was significantly delayed. This delay in reproduction induced by photoperiodic conditions and then spontaneously terminated was considered to be aestivation. Egg batches laid by females in the laboratory and in the field were kept at 25 °C for two months. From 30.8 to 93.8% of batches contained eggs which hatched without cold treatment between day 14 and 60 after oviposition. The proportion of eggs hatched was 17.7 to 20.9% in the short-day regimes, while it was significantly less (5.7 to 6.0%) under long-day conditions. It is concluded that in some eggs diapause is of low intensity and that if under natural conditions the first batches had been laid at the end of June, nymphs would have hatched at least from some eggs during the same season even without cold treatment. Such untimely hatching would have resulted in the death of nymphs and adults unprepared for overwintering. A photoperiodic response which induces aestivation in the early emerging adults in June–August may prevent early oviposition and occurrence of a second generation and thus maintains univoltinism in P. bidens.